Great Basin Naturalist

Volume 55 Number 2 Article 17

4-21-1995

Full Issue, Vol. 55 No. 2

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

Recommended Citation (1995) "Full Issue, Vol. 55 No. 2," Great Basin Naturalist: Vol. 55 : No. 2 , Article 17. Available at: https://scholarsarchive.byu.edu/gbn/vol55/iss2/17

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

GREAT BASINB A S I1 N naturalistnaturalist

A

VOLUME 55 n2na 2 APRIL 1995

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

associate editors MICHAEL A BOWERS PAUL C MARSH blandy experimental farm university of center for environmental studies arizona virginia box 175 boyce VA 22620 state university tempe AZ 85287 J R CALLAHAN STANLEY D SMITH museum of southwestern biology university of department of biology new mexico albuquerque NM university of nevada las vegas mailing address box 3140 hemet CA 92546 las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university department of environmental resource sciences university heights OH 44118 university of nevada reno 1000 valley road reno NV 89512 BORIS C kondratieff department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia university Morganmorgantowntown WV 26506612526506 6125

editorial board berranjerran T flinders chairman botany and range science duke S rogers zoology william hess botany and range science 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication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 1995 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 all back issues are in print and available for sale all matters pertaining to subscriptions back issues or other busi- ness should be directed to the editor great basin naturalist 290 MLBM PO box 20200 brigham young university provo UT 84602020084602 0200 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian 6385 HBLL PO box 26889 brigham young university provo UT 84602688984602 6889

editorial production staff joanne abel technical editor jan spencer assistant to the editor

copyright C 1995 by brigham young university ISSN 001736140017 3614 official publication date 21 april 1995 4-49595 750 13821 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 55 30 APRIL 1995 no 2

great basin naturalist 552 0 1995 appp 95 104

DIETS OF YOUNG COLORADO SQUAWFISH AND OTHER SMALL FISH IN backwaters OF THE GREEN RIVER COLORADO AND UTAH

robert T muthlmuth1muchl and darrel E snyderlsnyderssnyder1

ABSTRACT we compared diet of young of year colorado squawfish ptychocheilus lucius an endangered cyprinid with diets of other fish 75 minmm total length TL collected from backwaters of the green river between river kilome- ters 555 and 35 during summer and autumn 1987 species included native rhinichthysrhimchthys osculus catostomus discobolus and C latipinnislatipinmslatiiatipinnis and nonnative cyprinellaCyp rinella lutlutrensisrensis notropis stramineusstrastramineousmineus pimephales kromelaspropromelasmelas ictalurus punctatuspunctatus and Lepolepomisinis cyancyanellusellus for each species diet varied with size and between upper and lower river reaches but not between seasons for fish of similar size larval chironomids and ceratopogonids were principal foods of most fishes Copecopepodscopepodapods and cladoceranscladoceianscladocerans were important in diets ofofeF lucius 21 mm TL and L cyancyanellusellus 31 minmm TL catostoniuscatostomus discobolus was the only species that ate moderate amounts of algae fish all larvae were in digestive tracts of only 10 P lucius 21 73 mm TL about 1 ofofeP lucius analyzed high diet overlap occurred between some size reach groups ofofeP lucius and C lutlutrensisrensis R osculus C latipinnislatipmnislatiiatipinnis LI1 punctpunctatusatus and L cyancyanellusellus potential for food competition between young of year P lucius and other fishes in backwaters appeared greatest with the very abundant C lutlutrensisrensis

key words ptychocheilus lucius Cypcyprinellarinella lutienlutiutlutiensislutrensisrensissis nonnative fishes young of year diets diet overlap backwaters green river wild populations of federally endangered colorado squawfish nesler et al 1988 haines colorado squawfish ptychocheilus lucius per- and tyus 1990 tyus and haines 1991 ichthyo- sist only in the upper colorado river basin fauna of these backwaters is dominated by they are most abundant in the green and nonnative fishes especially red shiner capricypri yampa rivers of eastern utah and northwest- nella lutlutrensisrensis tyus et al 1982 haines and ern colorado tyus 1991a decline of this and tyus 1990 this observation has led to a hy- other native fishes in the colorado river basin popothesisthesis that nonnativenormative fishes adversely affect has been attributed to habitat alterations survival of young colorado squawfish through caused by water development and introduc- competition or predation stanford 1993 sug- tion and proliferation of nonnative fishes gested that strong food web interactions be- carlson and muth 1989 minckley 1991 tween native and nonnativenormative fishes probably backwaters of the green river below its occur but dietary relationships have not been confluence with the yampa river are impor- adequately documented haines and tyus tant nursery areas for young of year YOY 1990 ruppert et al 1993 our objectives

larvalI1 fish laboratory department of fishery and wildlife biology colorado state university foitroitroltfort collins CO 80523

95 96 GREAT BASIN naturalist volume 55 were to 1 describe diets of YOY colorado contributed by each taxon to total volume of squawfish and other small fish in backwaters food in each digestive tract larimore 1957 of the green river and 2 examine diet overlap mathur 1977 for diet analyses food item and potential for competition with colorado taxa total of 124 were grouped into 20 family squawfish order or broader based categories sometimes divided according to habitat eg aquatic or METHODS terrestrial data for each fish species were stratified samples of small fish were provided by the according to length lomm10 mm TL or larger inter- US fish and wildlife service field station at vals by season summer or autumn within river vernal UT these were collected from back- reach upper or lower only subsets with at waters of the green river during summer 30 least six fish containing food were included in june 27 august and autumn 22 september analyses diet measures calculated for each 10 december 1987 the study area extends subset were 1 mean percentage each food from confluence of the green and yampa category contributed to total volume of food in rivers in echo park dinosaur national monu- each digestive tract mean of volume percent- ment CO to turks head in canyonlandsCanyon lands ages and 2 percentage of all digestive tracts national park UT river kilometer RK 555 in which each food category occurred per- to 35 above confluence with the colorado centage of occurrence wallace 1981 evalu- river upper and lower reaches are divided at ated several diet measures and concluded that sand wash RK 346 UT a convenient access mean of volume percentages is the best mea- point just above desolation canyon each sure for calculating overlap however per- river reach began with a rocky high gradient centage of occurrence is useful for describing 13131.3 212.1 mkmmam segment and continued with a general variations in diet wallace 1981 sand and silt laden low gradient 02og020.2 040.4 bowen 1983 mkmmam segment known for relatively high similarities in diet by subset between colo- catches of YOY colorado squawfish haines rado squawfish and other fishes were evaluat- and tyus 1990 tyus and haines 1991 the ed by schoenerschooner s 1970 resource overlap river was further divided into 8 km sections index starting from a random location within each n reach to help assure an even distribution of oc 1 05pxi05yosy I1 axipxi pyl collection sites backwaters were defined as shallow typltypi- il where n is the number of food categories axipxi cally 0.5 m 05os05 in maximum depth ephemeral is the proportion of food category i expressed embaymentsembay with ments negligible water velocity as mean of volume percentages in the diet of consisted of silt and sand silt substrates or and species x colorado squawfish and pyl is the mud sometimes overlaying or interspersed proportion of food category i in the diet of with gravel or cobble backwaters had little or species y other fishes values range from 000.0oo no rooted aquatic vegetation but some had no overlap to 101.0io complete overlap when dense mats of algae two backwaters were data on resource availability are absent weekly each 8 sampled in km section during schoenerschooner s index is one of the best indices daylight 1000 1800 h using im2ima1 m2ma seines 08os080.8 available for calculating resource overlap mma mm2 mesh in summer and 1 m X 3 m seinesselnes hurlbert 1978 linton et al 1981 wallace 32 mm X 48 mm mesh in autumn fish 1981 diet overlap is useful in helping to elu- were killed and fixed in 10 formalin immedi- cidate food relationships among species and ately after collection has been considered biologically important up to five specimens 20 mm total length when values exceed 0600.60ogo zaret and rand TL and five 20 minmm TL of each fish species 1971 matthews and hill 1980 galat and representing graded size series were selected vucinich 1983 from each sample each digestive tract from esophagus to vent was removed opened and RESULTS visually assessed for percent fullness food items were identified to lowest practical taxon digestive tracts from 2554 fish represent- and a visual estimate was made of percentage ing 15 species were examined for food items 199519951 DIETS OF FISHES IN backwaters 97

3 were empty mostly from fish 13 mm head sucker 21 30 or 31 40 mm TL and in TL after subsets with 6 specimens contain- flannelmouth sucker channel catfish and ing food were eliminated from the data set green sunfish 40 mm TL mean percent full- 2297 specimens representing nine species ness of digestive tracts was highest in fish remained for diet analyses native fish includ- 21 30 or 3131101040 mm TL for all species ed 972 colorado squawfish 75757.5 73073.0 mm TL aquatic insects were a principal part of mean 19119.1igligi 35 speckled dace rhinichthys diets for all fishes except fathead minnow and osculus 23123.1 39839.8 mm TL mean 28128.1 42 blueheadbluebeadblue head sucker of identifiable insects blueheadbluebeadbluehead sucker catostomus discobolus immature dipterdipteransans especially larval chiranochirono 23023.0 58958.9 mm TL mean 35935.9 and 21 flan- mids were predominant in digestive tracts nelnelmouthmouth sucker C latilatipinnispinnis 32032.0 64364.3 mm larval chironomids were represented by at least TL mean 47947.9 nonnative fish included 21 genera the most common being chironomus 729 red shiner 11311311.3 74574.5 mm TL mean followed by rheotanytarsus eukieffetiellaeukiejferiella poly 29129.1 92 sand shiner notropis stramineusstrastramineousmineus pedpedilumilum tanytarsusTanytarsus cricotopusCricotopus and micronmicrop 22222.2 53253.2 mm TL mean 31031.0 330 fathead lectrasectra representative families of other imma- minnow pimephales kromelaspropromelasmelas 110llolio11.0 65965.9gsg mm ture dipterdipteransans were in order of importance TL mean 32532.5 58 channel catfish ceratopogonidae simuliidae dolichopodidae ictalurus punctpunctatusatus 22522.5 70070.0too mm TL mean empididae muscidae and tipulidae propor- 42942.9 and 18 green sunfish lepomis tional contribution of immature dipterdipteransans to cyancyanellusellus 20720.7907 56856.8 mm TL mean 39639.6 diets of red shiner sand shiner speckled dace and flannelmouth sucker was higher in the of characterization diets lower than upper reach relative importance no major or consistent seasonal differences of immature dipterdipteransans in diets of red shiner in diet measures were observed within species sand shiner and speckled dace decreased and for fish of similar size accordingly summer utilization of other insects increased as fish and autumn data were combined for species length increased conversely relative impor- and lengths by river reach trends in values of tance of immature dipteransdipter ans in diets of proportional importance of each food category colorado squawfish and channel catfish were similar between the two diet measures increased or remained high with increasing for all fishes therefore only means of volume fish length corixidsCorixids larval and adult aquatic percentages are reported coleoptercoleopteransans predominantly dytiscidae elmi diets consisted mostly of insects zooplank- dae haliplidae and hydrophilidae trichopter- ton algae seeds and organic and inorganic an larvae mainly hydropsychidaeHydropsychidae and hydrophydroa debris but relative importance of these food tilidaetilidae and ephemeropteran nymphs pre- categories varied among fishes or subsets dominantly baetidaeBaetidae and heptageniidae were within species table 1 based on total num- minor components of diets for all fishes 10 ber of food categories included in the diet of of food volume except larger red shiner each fish species diets of colorado squawfish speckled dace and green sunfish and red shiner were the most varied 18 and red shiner and sand shiner ate more semi- 17 food categories respectively followed by aquatic or terrestrial insects than other fishes speckled dace 15 fathead minnow channel semiaquatic insects consumed were primarily catfish and green sunfish 12 each sand shin- larval and adult coleoptercoleopteransans predominantly er 11 flannelmouth sucker 9 and blueheadbluebeadbluehead Heterocercheterocercidaeidae and staphylinidae and adult sucker 6 variety of food consumed was hymenopterhymenopteransans scelionidae terrestrial greater in the lower than upper reach for red insects consumed were primarily hemipterhemipteransans shiner colorado squawfish flannelmouth and formicideformicidsformicids sucker channel catfish and green sunfish all fishes ate zooplankton but it was partic- whereas diets of sand shiner fathead minnow ularly important in diets of colorado squaw- and speckled dace were more varied in the fish 31 mm TL especially 21 mm TQTL upper reach diet of blueheadbluebeadbluehead sucker was ana- green sunfish 31 mm TL and to a lesser lyzed for fish from the upper reach only diet extent red shiner and channel catfish 31 mm variety relative to fish length was greatest in TL and flannelmouth sucker cladoceransCladocerans many red shiner sand shiner fathead minnow identified as daphnia Euryeurycercuscercus and macro colorado squawfish speckled dace and blue thrixthrax and especially cyclopoid copepodscopecopepodapods 98 GREAT BASIN naturalist volume 55

TABLE 1 diets by totaltotetotoll lengthh interintervalsvalsvais mimmn of nineiline fish species calleccollectedcollec ted duringduning summersunlnierinner anandd autumnautriautuinn 1981987 7 from suiesulesure is mean percentage contalcontributedcontnl utedbuted byby each fbodfood caitegorycategory tcto total volumealumevolume of food in each digestiveve tract mean of vol colcoicoloradoorado redreds linerhinerbinershiner sandsaisalid shineshiner r fatheadd minnowV squawfishsquirabish food category 11 20 21 30 31 40 40 21 30 31 40 40 11 20 21 30 31 40 40 11 11 20

upperTT n reachaalaqiaqla insects unidentifiable partspaipal ts 11 9 25 37 3 9 22 2 1 1 semiaquatic or teitel restealrestnalterrestrial 1 4 6 12 5 1 diptera maturesimimmaturesimmatured 30 27 29 13 25 19 3 7 4 4 13 70 chnahnchironomidaeonomidae adults 1 3 3 10 9 4 9 anisoptera nymphs aquatic coleoptera 3 10 corixidae 1 1 10 trichoptera larvae 1 1 1 1 ephemeroptera nymphs 1 2 zooplankton cladocera and copepoda 7 6 3 3 1 1 26 16 rotlrotiferaRotifeiabeiabela 6 1 1 1 1 16 1 ostracoda 1 1 Gammangammaridaedae 1 hydiacarmahydracarinaHydraHydiacarinaearlnacarmaearma 2 1 inveiinvertebrateinmei tebratetebrake eggs I1 I1 1 1 1 1 3 1 nematoda 1 1 6 1 fish plant seeds 7 9 1 1 1 1 4 1 algae 2 4 1 1 2 2 2 2 2 organic inorganic debusdebris 35 32 22 26 50 60 64 87 93 90 31 10

number offishot fishhishbish 45 140 63 17 40 19 7 0 108 75 26 40 305 mean fullness of digestive tracts 52 64 44 38 81 77 73 83 84 75 40 59

lower reacha insects unidentifiable parts 3 13 16 23 12 14 6 2 2 1 semiaquatic or terrestrial 9 8 10 11 2 1 diptera immatures 55 36 32 28 37 31 27 9 9 4 1 19 47 chironomidaechii onomidae adults 1 3 5 2 aquatic coleoptera 2 1 3 6 corixidae 1 trichopteraa larvae 1 1 1 6 1 ephemeroptera nymphs 1 I1 anisoptera nymphs zooplankton cladocera and copepoda 6 5 3 1 I1 I1 1 1 29 35 rotlrotiferaRoti feiafelaoela 1 1 1 7 3 ostracoda 1 1 1 2 1 gammaridae hydracarinahydracdrmaHydracarinacarlna invertebrate eggs 1 I1 1 3 5 2 nematoda I1 1 9 18 1 1 1 fish plant seeds 9 7 4 5 1 2 algae 1 1 1 1 2 1 1 1 2 3 1 oiorganicgame inorganic debusdebris 18 23 26 28 40 32 59 88 87 93 95 33 10

nuinulnumberabernber of fish 62 208 138 56 10 10 6 9 31 49 32 27 301 mean fullness of digestive tracts 58 60 49 36 66 52 37 72 85 80 78 45 70

uppciauplcl lachinchlaehreach confluence ofofgicengreen and yampa liversrivers at RK 555 river kilometers above confluence of the green and Coloiacoloradodo riverss in echo park dinosaur national monument CO to sand wash urrk346UT RK 346 lower i eachreach sand wash to turks head canyonlandsCanyonlands national park UTutrk35RK 35 199511995 DIETS OF FISHES IN backwaters 99 backwaters in two reaches of the green river below its confluence with the yampa river colorado and utah diet mea- ume percentages colorado flannelmouth green squawfish speckled dace bluebeadBluehead suckersueker sucker channel catfish sunfish 21 30 3131101040 40 21 30 31 40AO 21 30 31 40 40 31031 40 40 21 30 31 40 40 21 30 40 upper reacha

10 34 4 2 1 72 52 54 31 5 1 28 22 70 66 8

8 13 2 3 12 4 2

1 8 13 1 1 1 1 14

2 6 5 1 1 6 10 25 29 8 2 I1 1 11 11 10 1 13 10 10 3 82 88 90 64 52 14 6

46 12 11 18 0 14 19 9 8 6 0 0 8 0 6

61 50 57 73 88 78 81 77 63 78 71

lower reacha

1 1 9 8 8 2 13 10 1 1 6 1 3 5 61 66 78 65 49 52 55 83 68 48 19 1 4 5 1 1 11 1 12 2 2 7 15 1

3 34

18 13 1 3 7 13 2 44 2 1 1 3 14 5 1 4 1 2 6 1 3 1 1 1 2 5 1 8 3 1 1 4 1 10 10 10 7 8 34 13 13 11 8 4

174 37 19 11 6 0 0 0 0 7 8 16 26 6 6

80 54 59 79 71 75 88 73 71 77 59 100 GREAT BASIN naturalist volume 55

many identified as cyclops represented most diet overlap zooplankton found in digestive tracts identified degree of diet overlap between YOY colo- genera fers of rotirotifersrotifera included brachionusBrachionus rado squawfish and other fishes was influenced Cephalodella Ke Mono cephalodella ratellaseratellakeratella lecanebecane monostylastyla mainly by zooplankton and especially imma- polyarthra and trichocercaTricho proportion of cerca ture dipterdipteransans table 2 within each reach zooplankton in diets of all fishes tended to diet overlap for all length intervals of decrease with fish length increasing colorado squawfish generally decreased as blueheadbluebeadBluehead sucker was the only species that lengths of other species increased degree of ate moderate amounts of algae 10 30 of diet overlap among fish of similar size was food volume other fishes consumed minor generally greater in the lower than upper amounts algae consisted mostly of six diatom reach overlap values were 060ogo0600.60 range genera cymbella fragilaria cyrosigmagyrosigmaCyroGyro sigma 0100.10 0590.59 for most comparisons generally navicula surirellaSurirella and synedrdsynedra one desmid values were lowest for comparisons with fat- genus closterium and to a lesser extent head minnow and blueheadbluebeadbluehead sucker range pediastrum a colonial green alga most diges- 0100.10olo 0440.44 biologically important overlap tive tracts contained debris that accounted for values 060ogo0600.60 occurred only between moderate or large proportions of gut contents colorado squawfish 10 mm TL and some 30 in all fishes except speckled dace and size reach groups of native speckled dace and green sunfish it was over 80 of gut content flannelmouth sucker and nonnative red shiner in fathead minnow and blueheadbluebeadbluehead sucker debris green sunfish and especially channel catfish consisted of fibrous particles of vascular plant these higher overlap values were primarily tissue usually mixed with large amounts of attributed to high proportions of larval chi clay particles and sand grains suggesting bot- ronoronomidsmids in diets and secondarily especially tom feeding seeds many identified as tama- for diet overlap with green sunfish 40 mm risk tamarix fallicagalgallicalica were eaten by all fishes TL upper reach and 21 30 mm TL lower especially red shiner 31 mm TL reach to proportions of zooplankton degree two observations were unique to colorado of diet overlap was greatest with channel cat- squawfish fish larvae were found in digestive fish and green sunfish tracts of 10 colorado squawfish about 1 of total examined I1 was 21 mm TL 8 were discussion 36 48 mm TL and I1 was 73 mm TL probably among food a yearling no fish were detected in digestive comparisons habits investiga- difficult of differences in tracts of other species the 18 fish larvae tions are because of study design location and season found most were too digested for species however our observations on diets of native and nonna- identification or accurate length measurement tive fishes in backwaters of the green river but all were cypriniforms mostly cyprincyprinidsids generally agree with results of prior studies in and probably 10 fish larvae 6 9 mm TL six the upper colorado river basin eg vanicek identified as red shiner and mm TL were one and kramer 1969 jacobi and jacobi 1982 about 8 mm TL as fathead minnow interest- mcada and tyus 1984 and reported food ingly the smallest colorado squawfish had habits of the nonnative species within their four prey fish all red shiner whereas only native ranges eg carlander 1969 1977 one or two fish were found in digestive tracts pflieger 1975 harlan et al 1987 larger YOY of the others gut contents of six colorado or yearling red shiner sand shiner speckled squawfish 36 48 mm TL and the 73 mm TL dace flannelmouth sucker channel catfish specimen were exclusively fish those for the and green sunfish eat mainly immature aquatic remaining specimens were 70 80 fish insects diets of larger YOY or yearling fathead digestive tracts of six colorado squawfish minnow and blueheadbluebeadbluehead sucker consist mostly of contained 2 6 cestode parasites probably algae and organic debris diet of YOY colo- proteocephalus ptychocheilus flagg 1982 rado squawfish consists primarily of zooplank- cescestodescustodestodes were not found in guts of other fish- ton and immature insects especially chirono- es colorado squawfish infested with cescestodescustodestodes mid larvae and occasionally includes fish were larger than 27 minmm TL and were collect- reported size at which wild colorado ed from both river reaches in autumn squawfish shift to a more piscivorous diet 199511995 DIETS OF FISHES IN backwaters 101 varies but generally fish become an important erallybrally considered biologically important table food item after colorado squawfish attain a 2 although not conclusive these compar- length of 40 mm osmundson and kaeding isons suggest either general resource parti- 1989 suggested that slower growth and poor- tioning or differences in diet preferences diet er condition of YOY and especially yearling overlap values were considered biologically colorado squawfish in grow out ponds with important only for comparisons with certain lower densities of appropriate size forage fish size interval river reach groups of five fishes might have been caused by higher reliance on because interspecific demand for resources insect forage identifiable fish reported in might not exceed supply bowen 1983 noted digestive tracts of YOY colorado squawfish that even extensive diet overlap is not conclu- here and by mcada and tyus 1984 and sive evidence for competition accordingly grabowski and hiebert 1989 were either red mcada and tyus 1984 who also used shiner or fathead minnow larvae these non- schoenerschooner s index to examine diet overlap native species are short lived fractional spawn between YOY colorado squawfish and nonna- ers gale and buynak 1982 gale 1986 and tive fishes in the green river suggested that are typically present in high numbers and at high diet overlap they observed between appropriate forage sizes in backwaters of the colorado squawfish 22 40 mm TL and chan- green river throughout summer and autumn nel catfish 19 55 mm TL overlap value tyus et al 1982 karp and tyus 1990 karp 0600.60ogo and especially red shiner 15 69 mm TL and tyus 1990 suggested that although the overlap values 0700.70 080oso0.80 might reflect shared abundance of small nonnativenormative prey fishes in use of abundant resources primarily imma- the green river might benefit growth of ture dipteransdipterans rather than competition the young colorado squawfish the benefit might same may be true for higher diet overlaps we be countered by the aggressive nature of some observed ward et al 1986 reported that chi nonnativenormative fishes which could have negative ronoronomidsmids the principal food category result- effects on growth and survival of young ing in high diet overlap were among the more colorado squawfish in their laboratory exper- common benthic invertebrates in the colo- iments on behavioral interactions karp and rado river basin tyus observed that red shiner fathead min- we observed that overlap values were gen- now and green sunfish shared activity sched- erally higher and for most fishes diet variety ules and space with colorado squawfish and was greater in the lower than upper reach exhibited antagonistic behaviors toward small- perhaps because food resources were more er colorado squawfish abundant and diverse in backwaters of the we could not effectively evaluate competi- lower reach based on observations during tion for food between YOY colorado squaw- summer and autumn 1979 1988 haines and fish and other fishes because study design did tyus 1990 found that backwaters in the not provide for estimation of resource abun- upper and lower reaches were similar in mean dance and availability intraspecific diet selec- surface area but that those in the lower reach tivity and effects of interspecific use of impor- were shallower and warmer conditions that tant resources direct evidence for interspecific may favor higher productivity also within the competition should be determined through upper reach grabowski and hiebert 1989 experiments demonstrating that shared use of noted that during summer and autumn a limited resource negatively affects one or 1987 88 concentrations of backwater nutri- more of the species schoener 1983 under- ents particulate organic matter phytoplank- wood 1986 wiens 1992 additionally we ton zooplankton and benthic macroinverte assume gut contents represented food con- brates particularly chironomid larvae in- sumed in the backwaters of capture but this creased progressively downstream they sug- might not always have been the case tyus gested this trend was due to attenuation of 1991b observed that although young colo- flow releases from flaming gorge reservoir rado squawfish in the green river were found located near the wyoming utah border at mostly in backwaters some moved to or from downstream sites that reduced the degree of other habitats during 24 h periods we found water exchange between the main channel and that diet overlap for most comparisons with backwaters and allowed for greater backwater colorado squawfish was below the level gen warming and stability 102 GREAT BASIN naturalist volume 55

tahrtahetaurtableTABLL 2 diet overlap by total length TQTL intervals mm between young of year colorado squawfish and eight other confluence with the yampa rivelriver colorado and utah overlap values were calculated using schoenerschooner s 1970 index asterisk fathead red shiner sand shiner minnow TL ofot upper reachreacha lower reachreacha upper lower upper Coloiacoloradodo squawfish 11 20 21 30 314031 40 40 11 20 21 30 314031 40 40 21 30 314031 40 40 21 30 31 40 40 214021 30 31031 40 40 11 059 054 040 043 042 049 050 047 046 047 035 053 055 052 040 037 038

11 20 049 045 043 031 063 053 047 038 037 031 014 049 045 038 018 015 015

21 30 055 051 053 041 074 057 051 042 042 043 027 049 047 038 023 019 023

31 40 040 039 040 027 073 057 052 042 035 029 012 050 045 038 017 013 015

40 039 037 039 035 065 047 044 037 034 028 012 053 047 037 017 013 014 autperreachuppci ic ch confluencec fluencfluence ofofgiecnG n andnd yampa rivers at RK 555 aveinverrivernvei kilometers above confluence of green and Coloiacoloradodo rivers inin echo parkpaik dinosaur national monument CO losuidwishto sand was ulur RKrk346346 1loccilowcicreeachcrreachbicichicich sandS md wishwash to turks head CanyonC inyonldndscanyonlandslands national park UTutrk35RK 35

alternatively greater diet overlap and vari- minckley 1973 greger and deacon 1988 ety in the lower reach might have been a rinne 1991 reflection of a difference in backwater avail- competition might also be a factor between ability between the upper and lower reaches smaller specimens of both colorado squawfish tyus and haines 1991 reported about 150 and other fishes few specimens 21 mm TL more backwaters per kilometer in the upper other than red shiner and fathead minnow than lower reach fishes in the lower reach 11 20 mm TL were available for comparisons might have been more crowded in available with colorado squawfish however as for backwaters resulting in greater shared use smaller colorado squawfish zooplankton and broader intraspecific use of available food would likely be an important component of mcada and tyus 1984 attributed reduc- their diets joseph et al 1977 and corre- tions in diet overlap between colorado squaw- spspondingonding overlap values would be high espe- fish 40 mm TL and red shiner or channel cially for specimens 11 mm TL although catfish to decreased consumption of immature dense populations may develop in backwaters dipterdipteransans and increased consumption of fish zooplankton may be limited under certain by colorado squawfish however ruppert et conditions because plankton communities in al 1993 reported fish larvae in digestive rivers are subject to dramatic spatial or tempo- tracts of 15 of adult red shiner 36 79 mm ral fluctuations in abundance and diversity TQTL from ephemeral shoreline embaymentsembay ments hynes 1970 welcommeWelcomme 1985 ward 1989 in near confluence of the green and yampa support of this generalization grabowski and rivers unlike our study they sampled on a hiebert 1989 reported that zooplankton den- diel basis and killed fish with an overdose of sities were higher in backwaters than in main anesthetic before preservation to minimize channel habitats within the upper reach and possible regurgitation their results suggest documented both spatial and temporal fluctu- that high diet overlap between young ations in zooplankton abundance they also colorado squawfish 40 mm TL and red observed higher concentrations of zooplank- shiner might reoccur or continue with larger ton in more confined backwaters than those piscivorous red shiner although we docu- with a broad connection to the river and sug- mented high diet overlap between young gested that densities were influenced by colorado squawfish 10 mm TL and other extent of water exchange between backwaters fishes in backwaters of the green river espe- and the main river cially channel catfish table 2 only red shiner in conclusion we found high diet overlap because of its extreme abundance haines and between YOY colorado squawfish and several tyus 1990 is likely to be a serious competitor small size groups of other fish species in green for food with young colorado squawfish red river backwaters because of the extreme shiner has often been implicated in decline of abundance of red shiner we speculate that native fishes of the american southwest eg diet overlap could result in food competition 199519951 DIETS OF FISHES IN backwaters 103 fish species collected during summer and autumn 1987 from backwaters in two reaches of the green river below its with mean of volume percentages as the diet measure values 0 60 biologically important overlapoveilovellap are marked with an

fathead flannelmouth green minnow speckled dace blueheadbluebeadBluehead sucker sucker channel catfish sunfish lower upper lower upper upper lower upper lower upper lower

11 2021 30 31 40 40 21 30 21 30 31 40 21 30 31 40 40 314031 40 40 40 40 21 30 31 40 40 40 21 30 40 044 043 037 033 021 036 034 039 035 034 052 057 059 034 046 0340 34 0300 30 0210 21 0560 56 0250 25 022 021 015 011 037 059 057 016 012 011 046 045 065 083 072 061ogi061057057 073 091 026

021 021 015 012 052 076 061 020 016 019 042 036 069 089 081 075075075 075 073 075 027

021 021 014 010 042 078 061 014 011 011 038 031 069 068 081 079082079 082 061 069 027

021 020 014 010 035 074 058 014 010 010 038 031 063 064 069 089077089 077 068 057 024

and might have a negative impact on colorado 1 1977 handbook of freshwater fishery biology press ames squawfish growth condition or survival volume 2 iowa state university 431 PP studies are needed to better assess the type CARLSON C A AND R T MUTH 1989 the colorado and strength of interactions between native river lifeline of the americanamerleanameiamer ican southwest pages and nonnative fishes in backwater food webs 220 239 in D P dodge editor proceedings of the flow and to international large river symposium canadian under present regulated regimes special publication of fisheries and aquatic define factors affecting these interactions sciences 106 FLAGG R 1982 disease survey of the colorado river acknowledgments fishes pages 177 184 in colorado river fishery proj- ect final report part 3 contracted studies USU S fish and wildlife service and bureau of reclamation S initiated H tyus C karp and lanigan salt lake city UT this study and provided samples and field galazCALATGALAT D L AND N VUCINICH 1983 food partitioning data H copeland J piccolo and PE sikoski between young of the year of two sympatric tui chub assisted with analysis of gut contents H tyus morphsmorphe transactions of the american fisheries 486 K society 112 497 and C karp reviewed data analyses GALE W F 1986 indeterminate fecundity and spawning bestgen D beyers J deacon G haines J behavior of captive red shiner fractional crevice hawkins C karp H tyus and R valdez spawner transactions of the american fisheries reviewed drafts of the manuscript this proj- society 115 429 437 recovery GALE W FE AND G L BUYNAK 1982 fecundity and ect was funded by the implemen- spawning frequency of the fathead minnow frac- tation program for endangered fish species tional spawner transactions of the american in the upper colorado river basin the pro- fisheries society 111 354035 40 gram is a effort of the US fish and GRABOWSKI S J AND S D HIEBERT 1989 some aspects joint selected backwaters and of reclamation of trophic interactions in wildlife service US bureau the mainmaln channel of the green river utah final western area power administration states of report of USU S bureau of reclamation research and colorado utah and wyoming upper basin laboratory services division applied sciences water users and environmental organizations branch environmental sciences section denver CO for USU S bureau of reclamation upper colorado this paper is contribution no 75 of the colo- regional office salt lake city UT 131 appp rado state university larval fish laboratory GREGER P D AND J E DEACON 1988 food partitioning among fishes of the virgin river copela 1988 314 323 CITED literature HAINES G B AND H M TYUS 1990 fish associations and environmental variables in ageageo 0 colorado squaw- BOWEN S H 1983 quantitative description of diet fish habitats green river utah journal of freshwater pages 325 336 in L A nielsen and D L johnson ecology 5 427 435 editors fisheries techniques american fisheries HARLAN J R E B SPEAKER AND J MAYHEW 1987 iowa society bethesda MD fish and fishing iowa department of natural CARLANDERCARLANDEB K D 1969 handbook of freshwater fishery resources des moinesmolnes 323 appp biology volume 1 iowa state university press HURLBERT S H 1978 the measurement of niche overlap ames 752 appp and some relatives ecology 59 67 77 104 GREAT BASIN naturalist volume 55

HYNES H B N 1970 the ecology of running water 1983 field experiments on interspecific competi- university of toronto press ontario canada 555 tion american naturalist 122 240 285 PP STANFORD J A 1993 instreamInstream flows to assist the recov- JACOBI G Z AND M D JACOBI 1982 fish stomach con- ery of endangered fishes of the upper colorado tent analysis pages 285 324 in colorado river fish- river basin review and synthesis of ecological infor- ery project final report part 3 contracted studies mation issues methods and rationale final report USU S fish and wildlife service and bureau of of flathead lake biological station university of reclamation salt lake city UT montana poisonpolson for USU S fish and wildlife service JOSEPH T W J A SINNING R J BEHNKE AND P B region 6 denver CO 89 appp appendices HOLDEN 1977 an evaluation of the status life histo- TYUS H M 1991a ecology and management of colorado ry and habitat requirements of endangered and squawfish pages 379 402 in W L minckley and threatened fishes of the upper colorado river J E deacon editors battle against extinction system USU S fish and wildlife service FWSOBS university ofarizonaof arizona press tucson 7762 169 appp 1991991bib movements and habitat use of young colo- KARP C A AND H M TYUS 1990 behavioral interac- rado squawfish in the green river utah journal of tions between young colorado squawfish and six fish freshwater ecology 6 43 51 species copeial990copela 1990 25 34 TYUS H M AND G B HAINES 1991 distribution habi- LARIMORE W R 1957 ecological life history of the war- tat use and growth of ageageo 0 colorado squawfish in mouth centrarchidae illinois natural history the green river basin colorado and utah survey bulletin 27 1 83 transactions of the american fisheries society 120 LINIONLINTON L R R W DAVIES AND FE J WRONA 1981 79 89 resource utilization indices an assessment journal TYUS H M B D BURDICK R A VALDEZ C M of animal ecology 50 283 292 HAYNES T A LYTLE AND C R BERRY 1982 fishes MATHUR D 1977 food habits and competitive relation- of the upper colorado river basin distribution ships of the bandfinbandzin shiner in halawakeeHalawakee creek abundance and status pages 12 70 in W H miller alabama american midland naturalist 97 89 100 H M tyus and C A carlson editors fishes of the MATTHEWS W J AND L G HILL 1980 habitat partition- upper colorado river system present and future ing in the fish community of a southwestern river western division of the american fishery society southwestern naturalist 25 51 66 bethesda MD mcada C W AND H M TYUS 1984 resource overlap UNDERWOOD T 1986 the analysis of competition by field of ageageo 0 colorado squawfish with other fish species experiments pages 240 268 in J kikkawa and D J in the green river fall 1980 proceedings of the anderson editors community ecology pattern and bonneville chapter american fisheries society process blackwell scientific publications oxford 1984441984 44 54 england MINCKLEY W L 1973 fishes of arizona arizona game VANICEK C D AND R H KRAMER 1969 life history of and fish department phoenix 293 appp the colorado squawfish ptychocheilus lucius and 1991 native fishes of the grand canyon region the colorado chub gila robusta in the green river an obituary pages 124 178 in colorado river ecolo- in dinosaur national monument 1964 1966 gy and dam management national academy press transactions of the american fisheries society 98 washington DC 193 208 WALLACE R K 1981 of overlap NESLER T P R T MUTH AND A F WASOWICZ 1988 JR an assessment diet nhsllr indexes evidence for baseline flow spikes as spawning cues transactions of the american fisheries for colorado squawfish in the yampa society 110 72 76 in river pages colorado american fisheries society symposium 5 WARD J V 1989 riverine wetland interactions 385 400 R R sharitz and gibbons 68 79 in shantz J W editors freshwater wetlands indandand wildlife USU S department OSMUNDSON D B AND L R KAEDING 1989 colorado of energy symposium series 61 USU S of squawfish and razorazorbackiback sucker grow pond department out energy office of scientific and technical studies as part of measures for the conservation information oak ridge TN green mountain and ruedi reservoir water sales WARD J V H J ZIMMERMAN AND L D CLINE 1986 S grand final report of USU fish and wildlife service lotic zoobenthos of the colorado system pages junction CO 57 appp 403 422 in B R davies and K FE walker editors piPFLIEGERLIEGER W L 1975 the fishes of missouri missouri the ecology of river systems dr W junk dordrechtDordrecht department of conservation jefferson city 343 appp the netherlands RINNE N 1991 habitat use by spikedacespikedace medafulgidaMeda fulgida J WIENS J A 1992 ecology of bird communities pices the apices cyprinidae in southwestern streams with volume 2 cambridge university press new york reference to probable habitat competition by red NY 316 appp shiner notropis lutlutrensisrensis apicespices cyprinidae south- WELCOMME R L 1985 river fisheries FADFAO fisheries western naturalist 36 7 13 technical paper 262 330 appp RUPPERT J B R T MUTH AND T P NESLER 1993 ZARET T M AND A S RAND 1971 competition in tropi- predation on fish larvae by adult red shiner yampa cal stream fishes support for the competitive exclu- green and rivers colorado southwestern naturalist sionslon principle ecology 52 336 342 3839738 397 399 SCHOENER T W 1970 synchronousnonsynchronousnon spatial overlap received 21 april 1994 of lizards in patchy habitats ecology 51 408 418 accepted 15 september 1994 great basin naturalist 552 0 1995 appp 105 116 invertebrate FAUNA OF wastewater PONDS IN southeastern IDAHO karen L Cieminskicieminski12ciemiyiskil212 and lester D flakel3flake13

ABSTRACT water column invertebrates were sampled with 383883813 881L activity traps in 15 sewage industrial and radioactive wastewater ponds at the idaho national engineering laboratory in southeastern idaho one collection was made per pond per month during all months the ponds were ice free from june 1990 through july 1991 in addition nutrient and selected heavy metal concentrations in pond water were determined inm july 1991 arsenic barium boron lead selenium and mercury were detected in ponds sewage ponds generally had higher nitrogen and phosphorus lev- els than industrial and radioactive ponds of the 30 aquatic invertebrate taxa collected the most ubiquitous were rotifera daphnidaeDaphnidae eucopepoda ostracoda acariacarlaearlacan baetidaeBaetidae corixidae notonectidae dytiscidae and chironomidae activity trap samples from sewage ponds contained more rotifera daphnidaeDaphnidae and notonectidae whereas industrial ponds yielded more chydondaechydoridae acarlacariaearlacan and Baetbaetidaeidae numbers of oligochaeta eucopepoda ostracoda corixidae dytiscidae and chironomidae collected were not significantly different between sewage and industrial ponds compared with natural systems these ponds had fewer taxa but a greater number of individuals of most taxa the high number of invertebrates collected is attributed to the lack of fish in wastewater ponds and the high levels of nitrogen and phosphorus particularly in sewage ponds

key words aquatic invertebrates sanitary wastewater industrial wastewater idaho national engineering laboratory

constructed ponds have been a common birds blackbirds and swallows use the ponds tool in wastewater treatment for decades heavily feeding partially or exclusively on gloyna et al 1976 wastewater ponds are aquatic invertebrates and on invertebrates constructed in a variety of manners and used that have emerged from the ponds millard et in various treatment procedures from settling al 1990 cleminsCieminscieminskild 1993 ponds to ponds with various aquatic macro most studies of macromacroinvertebratesinvertebrates espe- phytespaytes that enhance removal of nutrients and cially insects in conjunction with waste treat- break down organic materials brix 1993 ment have been limited to studies of benthic recently constructed wetlands have also been invertebrate assemblages in streams receiving incorporated into many wastewater treatment raw sewage or effluent from sewage treatment systems associated with municipalities and plants eg klotz 1977 kownacki 1977 duda industry task force on natural systems 1990 et al 1982 kondratieff and simmons 1982 moshiri 1993 wastewater ponds and wet- kondratieff et al 1984 chadwick et al 1986 lands are also associated with federal research lewis 1986 crawford et al 1992 literature sites such as the idaho national engineering on plankton and nekton in constructed ponds laboratory INEL in southeastern idaho and focuses mainly on pathogens and microscopic the hanford site in south central washington flora and fauna important in waste decomposi- wastewater ponds at INEL receive sani- tion such as bacteria protozoa and algae tary industrial and radioactive waste pro- goulden 1976 task force on natural duced at the facility other than wildlife systems 1990 watering cisterns and ephemeral rain pools because the invertebrate fauna of waste- waste disposal ponds are usually the only sur- water ponds attracts wildlife it is important to face water at INEL and as such attract understand invertebrate communities of the wildlife halford and millard 1978 howe and ponds as well as if and how they differ from flake 1989 millard et al 1990 cieminski natural communities our objectives were to 1993 migrating and resident waterfowl shore 1 provide baseline data on invertebrate

department of wildlife and fisheriesFishenes sciences south dakota state university box 2140b brookings SD 57007 resent address national park service 13025 riley s lock road Poopoolesvillelesville MD 20837 address reprint requests to this author

105 106 GREAT BASIN naturalist volume 55 resources available to migrating birds in con- METHODS strucstructedted waste ponds and 2 determine if nutrients and selected heavy metals in ponds water samples were collected at ponds in influence invertebrate populations july 1991 and analyzed for nutrients nitrogen and phosphorus and selected heavy metals STUDY SITE arsenic barium beryllium boron lead sele- nium and mercury that could influence pres- the 231600 ha INEL lies in butte ence of invertebrates water ph was taken bonneville bingham clark and jefferson once at each pond at the same time water counties ID on the western edge of the samples were collected further heavy metal snake river plain near the foothills of the lost and nutrient sampling was prohibitively river leahilemhi and bitterroot mountain ranges expensive and time consuming water samples fig 1 topography at INEL is flat to rolling were analyzed at the US geological survey s with elevation ranging from 1463 in to 1829 in national water quality laboratory at arvada big lost river little lost river and birch CO collection and analysis methods were as creek drainages terminate in playasolayas on or per brown et al 1970 and fishman and near INEL flow is intermittent and largely friedman 1989 data on heavy metals for diverted for agriculture during this study no pond aalianliAN li acronyms and names of pools are surface water flowed onto INEL plant com- included in tables 1 and 5 were taken from munitiesmunities are dominated by big sagebrush analyses conducted in 1988 artemisia tridentata low sagebrush A arbus- benthic samples were not taken because cula and three tipped sagebrush A triparti most ponds had lined bottoms or because ta mcbride et al 1978 sediment sampling was not permitted for other we column INEL lies in a semiarid cold desert annual reasons collected water inverte- each month to obtain gross temperatures range from 42 C to 39 C brates once esti- 42c 39c mates of invertebrate populations additional average annual precipitation is 19119.1 cm 40 collections and identification were time and of which falls from april through june clawson cost prohibitive given our concurrent collection et al 1989 precipitation levels are lowest in 1 of bird and mammal count data at these ponds july snowfall averages 71371.3 cineinelncm per year andana for a related project nevertheless we felt that from december through snow cover can persist invertebrates influenced bird use of ponds march thus the need for estimates of invertebrate ponds contained san- wastewater on INEL abundance itary waste eight ponds industrial waste water column invertebrates were collected three four ponds or radioactive waste at all radioactivenonnonradioactive ponds in months the ponds fig 1 because two radioactive ponds ponds were ice free from june 1990 through analy- also contained industrial waste in most may 1991 because of restricted access to ses radioactive ponds were grouped with in- radioactive waste ponds they were sampled dustrialdustrial ponds as industrial ponds for com- only once during july 1991 invertebrates parison with sewage ponds were collected in 38381388 L activity traps ross ponds were grouped around INEL facili- and murkin 1989 suspended horizontally 535.3 ties which were 4 36 kinkm apart generally cm under the water surface for approximately each facility had between one and four sewage 24 h modifications on the technique of ross ponds and an industrial waste pond sewage and murkin 1989 were necessary since most ponds ranged from 0040.04 to 2202.20 ha and were ponds had artificial liners therefore jars could 060.6og 2 in deep industrial waste ponds ranged not be suspended from a pipe driven in the from 0200.20 to 2242.24 ha and were 030.3 454.5 in deep pond bottom instead jars were suspended seven of the sewage ponds and one industrial from floats and attached to a 50 to 300 cm pond were lined to prevent infiltration into long piece of PVC pipe anchored on the surrounding soil four ponds all industrial pond s shore the first sample was taken at the andor radioactive supported emergent plant southeast corner of each pond subsequent growth A more thorough description of the monthly sample locations were chosen ran- ponds can be found in cieminski 1993 domly based on a single digit number of paces 199519951 invertebrates IN wastewater PONDS 107

bitterootbitterrootBitteroot idaho I1 rangerango containment test facility disposal pond cefictfi birch creekcrook technical support facility

INEL

leahilemhi ngengo CTF TSF disposal pond esfirtsfir uffie lost navalnavai reactors facility riverriper

lost river industrial waste range ditch 4 8 12 arfinrfi NRF kilometers

ANL sewage pond NRFs TRA 1cpp big lost r

argonne national laboratory west secondary sewage I1I1 pondvunaygna AMT e test reactor area anubanu2 prunariprunary sewage pond anasianlsi

orth sewage pond jras2 idaho chemical processing plantpiant south sewage industrial waste pond annANU pond cirallcirasl sewage ponds 141 4 warm waste cpps1 cfps4 pond alarmlar north coldcoid waste pond 0ma12 south coldcoid waste Z pond trallluauitra11lujui east percolation pond cppir2 west percolation pond cppirlCPPirl

fig 1 map of the idaho national engineering laboratory indicating location of facilities and wastewater ponds where invertebrate fauna was sampled waste type is indicated by lowercase letter in the pond code s sewage i industrial r radioactive counterclockwise from the previous sample brates were removed first samples from shal- site where dense emergent vegetation cov- low ponds with unlined bottoms often con- ered the near shore zone the activity trap was tained sediment to these rose bengal stain placed in the nearest open water was added to aid in sorting micromieromicroinmicroinvertebratesmicropinvertebratesinvertebrates activity trap contents were strained through mason and yevich 1967 samples in which a tsim75 amim no 200 sieve and preserved in 80 zooplankton was estimated to exceed 300 indi- propanol in the laboratory macroinverte viduals were subsamplessubsampledsub sampled to subsample 108 GREAT BASIN naturalist volume 55 samples were diluted to 500 or 1000 ml and tebratetebrake numbers between ponds with heavy stirred while 1 of the volume was drawn out metal concentrations greater than EPA criteria with I1 and 2 ml henson stemple pipettes and those with heavy metal concentrations invertebrate fauna were counted and iden- within EPA chronic exposure standards data tified to family with the exception of the were again pooled over all ponds years and orders oligochaeta acari araneae eucope- months radioactive waste ponds were elimi- poda ostracoda and lepidoptera and the nated from median tests because only one phyla nematoda and rotifera invertebrates sample was taken from them were identified using keys in pennak 1989 for non insects merritt and cummins 1984 RESULTS for aquatic insects and borror and delong 1971 for terrestrial insects B mcdaniel water chemistry plant south dakota science department heavy metal concentrations in most ponds state university brookings identified terres- were below criteria established by the EPA trial invertebrate families and verified other environmental protection agency 1987 identifications US 1 mercury the only metal found in data were not normally distrib- table was because that might affect aquatic life uted nonparametricnon analysis methods were concentrations parametric ponds and in used A median test was conducted on the trararar arfinrfi however trararar dozen most common invertebrate taxa to and arfinrfi mercury concentration was below determine if their abundance in sewage ponds the acute value of 242.4 jugllaglju gl US environ- protection agency differed from that in industrial ponds for mental 1987 each taxa numbers of individuals collected in sewage ponds had higher nitrogen and each sample were used in analysis data were phosphorus concentrations than industrial and pooled over all ponds years and months with- radioactive ponds table 2 ammonia in each of the two groups sewage ponds and nhanh4 N concentrations in most ponds were industrial ponds pooling samples for years within the range found in unpolluted surface and ponds allowed ample sample size for com- water wetzel 1983 however nhanh4 N con- parison of gross invertebrate population differ- centcentrationsrations at ICPP sewage ponds were well ences between pond types A median test was above those usually found in eutrophic lakes also run on the total number of species collect- nitrite n02NO N concentrations indicated ed per pond during the entire sampling period high organic pollution at all sewage ponds to determine if species richness was greater at except NRFs which was the only sewage sewage ponds or industrial ponds A third pond where n02NO N concentrations did not median test was conducted to compare inver exceed those of industrial and radioactive

TABLE 1 selected heavy metal concentrations buglugl in wastewater ponds at INEL idaho august 1991 and EPA criteriaa1 pondapondbpond15 criteria metal anlie cppir2 trararar trail arfinrfi ctficefi tsfiresfir ugloglbugl arsenic 949.4 2 ld 1 3 5 2 igoe barium 71 100 100 100 100 100 100 50000 beryllium 5 10 10 10 10 10 10 535.3 boron 30 50 70 120 90 10 5000 lead 21212.1 3 3 3 2 3 2 3232f3.2 selenium 2 1 1 1 2 1 1 35 mercury 20 oiol010.1 02 oi01010.1 141.4 oloi010.1 oi01010.1 00128001290.01280.0129 concentrations at 01or below these levels should have no adverse effects on freshwater systems naval reactor facilities officials suggested the following clarifica- tion the criteria inin the last column havellave questionable applicability to the arfinrfi the EPA icummaximumimum contaminant level for mercury in public community drink- ing watelwaterwatey systems is 202 0 agljltglaal alinli argonne national laboratorylaboi atory west industrial waste pond cppir2 idaho chemical processing plant east percolation pond industrial and radioactive trai test reactor area warm waste pond radioactive trailtralltranTRAU test reactor area south cold waste pond industrial arfinrfi naval reactors facility industrial waste ditch ctficefi containment test facility disposal pond industrial tsfiresfir technical support facility disposal pond industrial and radioactive ANU water sample tested at envnodyneenvirodyne engineers st louis MO february 1988 d1 symbol means water sample contained less than the detection level which follows the symbol arsenicarseuic1 semeveme ililiitii111III111 fatat watelwater hardnessbaldnesshaidbaidbaldness ofot 100 eglmgl value is 131 3 at watelwater hardness of 50 eglmgl kmeicurygmercury 11II11 199511995 invertebrates IN wastewater PONDS 109

TABLE 2 nutrient concentrations in wastewater ponds at INEL idaho august 1991 a nitrogen phosphorus eglmgl as N eglmgl as P pond jhbphbph nhanh4NH nog n02 NOn03 NOn03 n03nh4n03 nhanh4 PO 3 sewage ponds anls2 902 019 017 046 029 150 120 cppslcpps1 752 1100 220 460 240 021 400 cpps2 723 1700 069 240 171 010 480 cpps3 733 1700 015 046 031 002 640 cpps4 743 1700 014 043 029 002 610 TRAs 687 041 013 510 497 1212 079 NRFs 990 040 002 014 012 030 300

Nonnonsewagesewage ponds annANU 742 097 009 074 065 067 140 cppir2 880 004 005 130 125 3049 001 trail 760 001 006 110 104 10400 007 trararar 843 015 001 027 026 173 001 arfinrfi 742 001 001 160 159 15900 040 ctfir 997 001 001 045 044 4400 009 tsfiresfir 975 004 002 011 009 217 012 samples weiewere collected between 0800 and 1400 hb mountain standard time bwaterawaterwater ph values fluctuate readily according to the INEL industrial waste management information system 1989 effluent ph ranges and numbers of months ph was sampled were as follows anlsl 78 989877 cppsicppsl 4 75 861286 12 trasi 2 71 801080 10 NRFs 74 iloiio11012110 12 trail 2 75 808066 trararar 636 3 686 8 2 arfinrfi 696 9 757 5 12 tsfiresfir 717 1 797 9 12 ponds nitrate n03 N concentrations were the water surface or shaken from emergent not noticeably different between sewage vegetation in the collection process in order ponds and industrialindustrialradioactiveradioactive ponds and of decreasing abundance the main taxa col- n03 N levels of all ponds were within ranges lected were rotifera daphnidaeDaphnidae ostracoda commonly found in unpolluted freshwater eucopepoda chydoridae corixidae chirono- wetzel 1983 midae oligochaeta baetidaeBaet idae psychodidae the n03 nnh4 N ratio is an indication acari dytiscidae and notonectidae the of organic pollution a lower number indicat- above taxa were also the most ubiquitous ing greater pollution wetzel 1983 the except chydoridae oligochaeta and psycho- n03 nnh4 N ratio was 1 at all sewage didae which were found in large numbers but ponds except anls2 and TRAs and 1 I1 at all in few samples industrial and radioactive ponds except aalianli the number of invertebrate taxa collected however only in ICPP sewage ponds were per pond ranged from 5 to 22 excluding ter- ratios small enough to be considered organi- rerestrialstrial taxa the number of aquatic taxa co- cally contaminated wetzel 1983 phosphorus llected ranged from 4 to 16 per pond concentrations at most sewage ponds were radioactive ponds were sampled only in july much higher than the concentration in the but the number of taxa collected was almost highest industrialindustrialradioactiveradioactive pond compared identical to july samples from radioactivenonnonradioactive with maximums in uncontaminated surface industrial ponds table 4 statistical analyses waters phosphorus concentrations in sewage were not performed on radioactive ponds ponds were 4 30 times greater but of the because only one activity trap sample was col- industrial and radioactive ponds only concen- lected industrial aaliannanliANU trailtralltranTRAH and 2 NRM trationstrations in ANU and arfinrfi were substantially and ctficefi and sewage ponds had similar P greater 7 and 2xax wetzel 1983 iiliA11.111 numbers of taxa per sample within most taxa the number of individu- invertebrate fauna als collected varied greatly from pond to pond forty nine taxa of invertebrates were col- table 5 A median test revealed that activity lected from waste ponds of which 30 were trap samples from sewage ponds contained aquatic table 3 most nonaquaticnonaquatic forms were more rotifera P oi0101.01 daphnidaeDaphnidae FP oi0101.01 found in small numbers collembola however and notonectidae P 0404.04 whereas industri- were found regularly and were probably on al ponds yielded more chydoridae P oi0101.01 110 GREAT BASIN naturalist volume 55

TABLE 3 invertebrate taxa and mean number collected order homoptera flomfrom 15 wastewater ponds at INEL idaho 1990 91gligia9111 family aphidae aphidsapbidsbaphids1111 0050.050oos05 family cercopidae spittlebugsbspittlebugs13 ooi0010 01 324x724hh family cicadellidae leafhoppersbleaftioppers11 0030 03 taxa n 96 family unidentified 11 0250 25 phylum rotifera 14711414711471.1414 order coleoptera beetles phylum toddmudamudd 0 05 family chrysomelidae leafbeetlesleaf beetles 0030 03 nematodaNema 005 1 phylum annelida family coccmellidaecoccinellidae ladybird beetlesbeetlesbbeetles1 ooi0010 01 class oligochaeta earthworms 6 32 family dytiscidae predaceous aquatic 632 ogs class hirudinea leeches diving beetles 0650.650 65 order rhynchobdellida family elmidaeelmidge riffle beetles ooi0010.010 01 family glossiphonidaeglossiphonndaeglossiphoniidae 0020.020 02 family gyrinidae whirligig beetles 001 phylum arthropoda family haliplidae crawling water beetles 0020.020 02 class crustacea family hydrophilidae water order cladocera water fleas scavenger beetles 0020 02 ptiliidae winged family Daphnidaphnidaedaphmdaedae 1351261351.261351 26 family feather beetles ooi0010.010 01 family chydondaechydoridae 10288102 88 family staphylinidae rove beetles 0020 02 family sididaemididae 0090 09 order trichoptera caddisfliescaddis flies olderorder eucopepoda copecopepodscopepodapods 15145 family leptoceridaeleptocendae 0050 05 lepidoptera butterflies and mothsbmoths1111 0.02000202 order ostracodaOstiaostraeodacoda seed scrimpsshshrimpsrimps 31717317 17 order moths 002 order amphipoda scadsscuds order diptera flies family ceratopogonidae biting midges 00.0101 family tahtndaetalitridae 0450 45 ooi001 class arachnoidea family psychodidae moth flies and sand flies 1681.681 68 olderorder acariaearlacan mites isilsi1511 51 15 family midges 11 52 order araneae spidersspiderspiders15sl 0040 04 chironomidae 1152 class insecta family tipulidae crane flies 0020 02 order collembola springspnngtailsspringtailstailstalis family unidentified adults 0800 80 family ogg99 family entomobryidaebentomobryidae11 0570.570 57 unidentified pupae 0990.990 family onychlurldaebonychiuridaeb 0300.300 30 order hymenoptera family formicidae antabantsbants11 0 03 olderorder ephemeroptera maymayfliesflies ants 0030.03 family platygasteridaebplatygastendae11 family baetidaeBaetidae 5715.715 71 ooi0010 01 braconidae11aebaeh11 family caenidaemaenidaeidae 0 01 family BraconidBraconidaebraconidaeb 0010 01 Caen ooi001 11 olderorder odonata family EncyrtidEncyrtidaeencyrtidaebencyrtidae11aeb ooi0010.010 01 pteromalidaebpteromalidae1111 suborderSuboider anisoptera dragonfliesdragon flies family PteromalidPteromalidaeaeb ooi0010.010 01 family scelionidaebsceliomdae1 01 family aeshnidaeaeshmdae ooi0010.010 01 ooi0010.010 wasps1717 suborder zygoptera damselfliesdamsel flies family sphecidae sphecid waspswaspsb ooi0010.010 01 family Coengoencoenagrionidaecoenagnonidaeagrionidae 0310.310 31 ainalnvertebratesinvertebrates were collected inin 383813 8 L activity traps suspended in the water order thysanoptera thripsbthnps11 column for 24 h one per pond per month collections were june october may 3 family thripidae common thnpsthripshsh 0 11 1990 and march 1991 for 12 ponds and july 1991 for radioactive thrip oil ponds family aeolothripidaeaeolothnpidae banded thnps1thripsb 0020.020 02 individuals111ndividuals found were mostly or exclusively terrestrial olderorder hemiptera true bugs family corixidae water boatmen 397639.7639 76 family notonectidae backswimmersback swimmers 0530 53 acariacarlaearlacan P oi0101.01 and baetidaeBaet idae FP oi0101.01 P 4545.45 and dytiscidae P 0707.07 were simi- numbers of oligochaeta P 44 eucope- lar between the pond with mercury and those poda P 5050.50 ostracoda P 09og09.09 corixidae without P 08os08.08 dytiscidae P 5454.54 and chirono- midae P 70 collected were not significant- discussion ly different between sewage and industrial ponds wastewater ponds at INEL were nutrient invertebrate numbers in pond arfinrfi which rich especially sewage ponds organic enrich- had a high mercury content were compared ment may be the cause of high abundance and to those in the remaining industrial ponds low number of invertebrate taxa found species where mercury was not detected samples richness at sewage ponds was similar to that at from arfinrfi contained more chironomidae P industrial ponds however species composi- 0202.02 and oligochaeta P oloi01.01 and fewer tion differed between sewage and industrial chydondaechydoridae P 03 and ostracoda P 0303.03 ponds differences were probably due to the than ponds annANUAN li trai and ctficefi numbers greater organic enrichment in sewage ponds of rotifera P loio10.10 daphnidaeDaphni dae P 10 activity trap samples from INEL ponds eucopepoda P loio10.10 acari P lsis15.15 baetibaebi contained fewer invertebrate taxa than compa- dae P ss5555.55 corixidae P 0707.07 notonectidae rable samples from natural waters gordon et al 199519951 invertebrates IN wastewater PONDS illiiiililillii111

TABLE 4 number of aquatic invertebrates per collec- eranee value from 0 lowest tolerance to tion activity trap set for 24 h from radioactive waste organic pollution to 10 highest eleven of the ponds at INEL idaho july 19911 families for which hilsenhoff 1988 presented cppir2bcpincppn trararar tsfiresfir tolerance values were found in INEL ponds taxa n 1 n 1 n 1 and only 2 had tolerance values of less than 4 daphnidaeDaphnidae 94 1 59 those 11 families and tolerance values are as chydoridae 0 0 129 follows aeshnidae and tipulidae 3 Baetbaetidaeidae eucopepoda 35 0 818 elmidaeElmelmidgeidae and leptoceridae 4 ceratopogon ostracoda 5 0 1620 amphipoda 0 0 1 idae 6 caenidaeCaenmaenidaeidae 7 chironomidae and baetidaeBaetidae 2 0 0 talitridae 8 coenagrionidaeCoenagrionidae 9 and psycho corixidae 1 5 0 didaedidace 10 the two families with a 3 tolerance dytiscidae 0 6 4 rating were represented by only single speci- chironomidae 7 0 18 mens in INEL wastewater ponds dataadataanata from radioactive waste ponds were not analyzed with those fromhrombrom sesewagewage and industrial ponds because only one sample was taken from radioaradicaradioactiveetive low invertebrate diversity in industrial ponds ponds may be caused by organic or chemical bcppir2cppn2 idaho chemical processing plant east percolation pond industrial and radioactive trararar test reactor area warmrm waste pond radioactive constituents although nutrients in industrial tsfiresfir technical support facility disposal pond industrial and radioactive waste ponds were within ranges found in nat- ural waters most industrial ponds at INEL 1990 neckles et al 1990 dominant taxa col- would be considered eutrophic wetzel 1983 lected from study ponds were similar to domi- additional organic enrichment in sewage ponds nant taxa collected in activity traps at natural did not affect species richness compared to wetlands in nebraska gordon et al 1990 and industrial ponds however species composi- manitoba neckles et al 1990 with the tion was different between the two pond exception of culicidae turbellaria neckles types metal and saline pollution has also been 1990 and gastropoda gordon et al 1990 found to decrease aquatic invertebrate diversi- neckles et al 1990 which were not collected ty savage and rabe 1973 seagle et al 1980 from wastewater ponds in our study fewer euliss 1989 taxa per sample were collected compared to in most instances the seven heavy metals activity trap samples from seasonal wetlands tested did not occur in concentrations great cowardin et al 1979 neckles et al 1990 enough to affect aquatic life only mercury was seasonal wetlands like organically enriched found at concentrations over chronic exposure systems of sewage ponds tend to have low levels at concentrations below chronic levels invertebrate taxa diversity wiggins et al freshwater organisms should show no chronic 1980 toxic effects US environmental protection the reduced number of taxa in wastewater agency 1987 chydoridae and ostracoda were ponds may be due to lack of emergent vegeta- scarcer and chironomidae and oligochaeta tion in most ponds odonateodonata families libelluli- more abundant in samples from pond arfinrfi dae and lestidaeLestleptidaeidae which were collected by wherein mercury was detected other toxins gordon et al 1990 but not from wastewater may occur in the water and no other ponds ponds are commonly associated with vascular with elevated mercury concentrations were hydrophytes merritt and cummins 1984 available for comparison therefore we do not vegetation has been found to be correlated know if mercury caused the difference detected with maemacmacroinvertebratemacromaeroroinverognveinvertebratertebraeb rate species richness although species richness of INEL ponds gilinsky 1984 was low comparison with natural wetlands another possible cause of low species rich- gordon et al 1990 neckles et al 1990 ness in wastewater ponds is high organic revealed that study ponds exhibited high waste content streams and wetlands receiv- invertebrate abundance of the taxa that waste- ing organic waste typically exhibit low inverte- water pond and nebraska wetland collections brate taxa diversity olive and dambach 1973 had in common wastewater pond samples con- brightman and fox 1976 kondratieff and tained higher densities of all except gyrinidae simmons 1982 kondratieff et al 1984 victor ceratopogonidae and hirudinea gordon et al and dickson 1985 pearson and penridge 1987 1990 gyrinidae and ceratopogonidae were hilsenhoff 1988 assigned arthropod families collected in almost identical amounts and from streams in the great lakes region a tol hirudinea were more abundant in nebraska 112 GREAT BASINz znaturalist volumeD 55

e 31991 fl 0 e reactors 1980-00 f a S TS hNRFs 05 CO otioiit t 1 e 3734 s 00 947 03 e 1 CO f fp mmamay 8 253 85169000 CO 73 10167 11 in 51 e 0 0 10 1 0 ohalg d i 640240 t & 01 K naval 3 10 10 1 1 i idaho n g0 00 0 0 ehl347 25 2 212 858.5 00 18 mm 181 U 00 01 a a1 01 0085 1 a z march 0 0 0 0 1 alternately

1 la and 00 95 f w arfinrfi arezINELH tat3 rt 8 18700 105lemwem 11 si- r 1 0 water 2 0 ponds 1990 2 arfinrfi 60 ra 27I1 2510 0 CD0 325 10 10 01 s n M 0 s0 & 05 05 6 0 0 0 2 L 0j i t3ta at aa CO aj s- a a S S S a 2a 0 0 0 0 0 0 0 4 0 0 0 alternately Q october S contained 1 ponds 0 industrial 0 CD ri s & ta 6 441 t3 water a august51 u i 1 trai 01 11 152 andtbas2 waste S n 0 3 g0 g0 2 10 g0 0 0 g0 g0 s a 1 1 aj0j s 0 0 0 0 0 0 0 0 0 0 0 0 c contained3 TBAs

01 1 1 8770 s trasl 1 1 1 711raz 1 frasl industrial 00 10 iaz 1991 455 0 g 8 00 35 141 10 t 1950 1 fchbli n3na ANU N 282 1- 1200 10 10 amsamymay 2 0 00 0 01 7 4 industrial 25 3 15 ponds n B1 aj0j 57tin 1 1 1 i s 0 01 0 0 10 01 0 0 1 0 5 0 of a s anda sewagepondsU C aprilwallm ca 5 af5f Ssewage a u ponds y and cu 0 CO 00 1 1 sewage 00 m 1 361501 1 1 ti 8 523001 1181 11 1 whoaarea 11 1 rhoa NRFs 10 6 1990 waste 1 1 1 191 rt CO 1 11 1 p 1 n 2- 7903 7 0 3 111 1 g0 1 0 ll 1 0j 21 1 aj i 1 a 2 0 0 0 0 0 01 0 1 0 0 0 s t frommocagoca reactor 2cold 2 2 tct C S S juueoctober 0 south 0 .5 1700 5 10 6007000 testhuml J h 011824 &i i T 00 10 ii c 24 CO 00 01 248 11 6 in if 700 500 ardandmrmJ 1- 11 300mee0 10 i i 10 b for TRAs 1315 1125zammM i7 S2 3 P97 fol 0CO ia od TBAs 355 1 v 10 05j10 03 1 s 2 CO 1 g 0 01 25 05 05 0 anls2 n 1 1 1 12 1 north girgysp- 1 0 1 1 01 1 H 5 setoolootS 0 1 0 0 0 0 1 c z r

ponds ardaareamrea 1991 & h CT trap2 0 a vi 5800 00 i 0 1 1 ponds 00 1218 1 5350 may & c 10 01 sewage1 reactor s 0 i7 I S is 8 ia 104500 S 1 3 13 activity p 1 11 izi cpps4 ua 4 10 g pondsa c a u5 16 1 2365 1 11 43 pi s10 1 1 ponds amzgez 465 industrial 161 CO g0 D 1 g muchpuchpuoh 3 05 gemal fool 0 0 g0 plant test n 300CO 05 A u cl 0 0 0 0 2 0 0 0 i 1

a andmcd industrial

1 industrial 0 w ielmoamom processing trai 1990 collection5 4799himm aqcq CO 1 jl 1 11 u 00 562meb Q 631CO t0ta 0800 0 twice 8 135010 1101 121 1 from CO f 0 cpps3 CD s co0 1& collected fromglobglom Q P 14 pond 111 pondsb 10 10 10 III 0 s 874 15 g0 10 4.5 g g a october iti 25 45 0 0 n 1 20 15 4 wm 6 U 00 1 cl0 oi 45 berbelperU 0 0 0 1 0 0 0 chemical collected & waste & andwam collected U 111 IIInumbers badbjd ililii ill june s g 1 sewage 3324 549 5100 nl 1 idaho s3sa 1 CO instances a 01 1 1 s 3 0 00 10 industrial i m 8 0 sii numbers 4300 bbsass ca 11 00 ohal fromgroegrom numbers invertebrates c1 cpps2 0CO cs2 10 10 8 g1ga233h sss & CO S 11 656eme 325 105 10 g U 10 g0 g0 oj 25 0 6 0 1 3 industrial n CO 0 s c5ca sixmir r iu U 01 and 0 s i 4 ggc05 west gac 0 2 swan 0 0 0 0 0 0 than ir emmazama Q q than onlyS s cppsi sl A F gfeaafea arfinrfi 05 0 00 c U pond g P 1 in frahrc 05 0942801 1794 SS 1 i&2 aquatics g 5376 0 ika vi 1 A pondsflalaboratorya CO 13 a vi 1 1 6300 s 3 airt w differentirt 00 17 910 0 0 fazAND P 8 10 5- 26S disposal S P D were CPPSI lmh1500 600700 oi 1 Q 1 10 10 tmt M lbsITS 8615 3785 1 050 S ba pi t 585 10 10 b3 higher SS of 00 g lower 25 0 1 4 ci 0 sewage 0 n 05 1 Cs 00 CO i notaezaga J 0ct 0 10 0 0 1 0 0 0 giiligialiNRFs ta national D t3 facility tlarmmrm mag are aremae mke minimums ga are number d e g west & 4f S S presented 01 iai5 S iii i ponds00t3ponds pondsgenesgeneo 3 test c 28302CO0 jaffcppsi I of a 0 raeeoeargonne 0 ajqj 00 ullmo SOI a c etric SO a 1 elw3141 sewage sewage 1 7 laboratory 1991 sewage i i i areeweewo anls2 n iralnalma zelgel107 lllgw 1 CO 15 1 1 1 1 1 1 containment anls1 r 8331 I 01 G G m 2 7 0 10 00 3 7 g0 1 a n so 5 68 0j nonparaarlelarlei 0 aj jnj21masxaymay 3 S 0 0 0 azuANU from from 0 0 mom 0 0 1 0 moa taxa 0 maximum S U lilfilsac month i liliiU national H aheare lii c b iai3 sjcmarchUTia t N ti pond collected i 00 llliicollected collected 47282 lluhlluhal S 142 8 datamalamata l cl l golpermoa 0 abundant anlsli 1 1 o a il 6 ugg andezeece g- g mcm I CO 49 CO elgu&g s and 3- 1 ilil S g g argonne 2 2 0 2 3 4 3 0 sewagesfathaiha i n 80 1 once 0 1 aj0j aj0j 00 ditch 3 0 0 0 0 0 1 0 0 0 0 0 becausejoosaagsste1990 numbers numbers a eliylils iylnumbers median mostaomaS waste 3 collected lil facility s a given 5 ae september s s anls2 3e shows shows 1 itjjl5051 a shows 01 nj 0 5 S 12 ctrct1 s a 0 zotnot CO ssiafiegindustrial t2ta were daemeemeo 0 C 2 kolu i 5 SM testzoma thehao 5 tat3 s andmcm reactors test test 1 a 1 chironomidae aleare 1 notonectidae 1 g 2 ti eucopepoda 0 g idae july aa oligochaeta t chydoridae TABLE daphnidae 0 dytiscidae w & ostracoda corixidae baetidae rotifera Daphni 0 C nl 0 csamples jc amples admedian emedian 1 dmedian emelian E i bedianwedian u odd ameantameans banlsl ei 199103 acarimoaga facility taxa S Baet lloilnaval 03 0 1 3 0 1 m y icestrai p Q sa cS icss 2 5 M 0 u Q U u m lliiljz2mfss 199519951 invertebrates IN wastewater PONDS 113 wetlands compared to our study ponds gordon ponds is limited due to a scarcity of published et al 1990 also in our study more cladocera papers porcella et al 1972 noted large popu- and ostracoda were collected compared to lations of daphnia in a reservoir fed mostly by activity trap samples from seasonal wetlands treated sanitary wastewater daphnidaeDaphni dae neckles et al 1990 which tend to have a rotiferarotiferal and notonectidae were more com- high invertebrate abundance wiggins et al mon in INEL sewage ponds than in industrial 1980 nutrient polluted natural waters also ponds all three species as well as oligochaeta have invertebrate communities containing eucopepoda ostracoda and corixidae sinclair many individuals of a few species brightman 1975 are common inhabitants of sanitary and fox 1976 lubini ferlin 1986 brightman wastewater oligochaeta eucopepoda ostra- and fox 1976 attribute this partially to a coda corixidae and chironomidae were reduction in competition from pollution intolantol abundant in sewage ponds but not more so erantarant forms than in industrial ponds cladocera euco- high invertebrate growth and abundance pepoda ostracoda corixidae and chironomi- have been associated with high algal produc- dae were also common in evaporation ponds tivity wallace and merritt 1980 richardson in california which contain salts and heavy 1984 which in turn has been associated with metals euliss et al 1991 high phosphorus and nitrogen concentrations invertebrate communities in INEL sewage liao and lean 1978 wetzel 1983 most INEL ponds differed from those in organically pol- wastewater ponds were eutrophic or highly luted streams however in making these com- eutrophic wetzel 1983 therefore wastewater parisonsparisons we note that our sampling methods ponds which are higher in nutrients than nat- did not target benthic organisms in nutrient ural wetlands would be expected to produce enriched stream reaches oligochaetes and more invertebrate biomass chironomids are dominant duda et al 1982 pearson the absence of fish in study ponds proba- and penridge 1987 crawford et al bly also contributed to high invertebrate den- 1992 but we found no difference in numbers sewage and industrial ponds some sities fish have been shown to decrease aquatic between kownacki 1977 and invertebrate densities gilinsky 1984 for most chironomid species oligochaete families 1986 charac- taxa collections from industrial ponds also had lewis are teteristic of clean waters and it is possible the more individuals than collections from natural species inhabiting sewage ponds differed from systems gordon et al 1990 neckles et al those in industrial ponds ostracoda have also 1990 even though industrial ponds were not been described as pollution tolerant kownacki as rich as sewage ponds nutrient 1977 but we found no difference in their in certain systems a large abundance of in- numbers at the 05os05.05 level of significance at the vertebrates has also been attributed to a paucity loio10.10 level sewage pond samples contained more of insect predators brightman and fox 1976 ostraostracodsostracodecods Baetbaetidaeidae may be either pollution williams 1985 dodson 1987 however sev- tolerant savage and rabe 1973 victor and eral predaceous taxa were collected from dickson 1985 or intolerant kownacki 1977 waste ponds most notably dytiscidae and depending upon the species we found more notonectidae because these taxa were col- baetidaeBaetidae in industrial ponds indicating they lected in greater numbers from wastewater as well as chydoridae and acari which were ponds than from natural wetlands gordon et also more abundant in industrial pond sam- al 1990 and because notonectidae were most ples may be less tolerant of low oxygen con- numerous in sewage ponds where many prey centrationscentrations than the other common taxa taxa were also most numerous we surmise the taxa found in greater abundance in sewage large number of invertebrates collected from ponds than in industrial ponds were those that waste ponds resulted mostly from a reduction could take advantage of the unique and difficult in competition from pollution intolerant taxa living conditions eutrophic waters typically high algal productivity and the absence of exhibit lower dissolved oxygen concentrations fish rather than from lack of invertebrate pre- and greater fluctuations in dissolved oxygen dation and ph than less organically enriched waters comparison of our results on water column some cladoceran species can form hemoglo- invertebrates with other studies of sewage bin when dissolved oxygen concentrations are 114 GREAT BASIN naturalist volume 55 low thus oxygen levels are rarely a limiting ing water body goulden 1976 in systems factor pennak 1989 the same is true of like some at INEL where water loss is rotirotifersrotiferafers certain genera are capable of with- through evaporation all waste processing standing anaerobic conditions for a short time occurs in the pond zooplankton are also and very low oxygen concentrations for important in waste elimination and transfer extended periods pennak 1989 since goulden 1976 patrick 1976 bogatovaBogatova and notonectidae breathe at the water surface yerofeyeva 1980 other aquatic invertebrates merritt and cummins 1984 they are unaf- that consume algae or bacteria or feed on zoo- fected by dissolved oxygen concentrations plankton and are then eaten by birds also most cladocera are less affected by ph fluctu- influence the reduction and transformation of ations than some taxa because they typically organic waste and its dissipation out of the occur over a wide ph range pennak 1989 if system ph levels are too high or too low cladocera and rotifera can withstand temporarily unfa- acknowledgments vorable environmental situations by producing resting eggs that are resistant to adverse we thank 0 D markham for suggestions chemical conditions under more favorable from initiation through project completion conditions cladocera and rotifera life cycles we appreciate the assistance of L knobel and allow them to respond quickly to improving R bartholomay of the US geological survey conditions pennak 1989 which provided water chemistry analysis we regarding the feeding habits of taxa that thank W L tucker experiment station statis- were more abundant in sewage ponds tician south dakota state university for pro- notonectidae were possibly taking advantage viding statistical advice and B mcdaniels of the reduced competition from other preda- and W G duffy of south dakota state uni- tors both rotirotifersrotiferafers and daphnia are omnivoomnivo- versity for assisting in invertebrate identifica- rous and feed on any suitable sized food parti- tion W G duffy 0 D markham and R C cle therefore food was abundant for them in morris reviewed the manuscript field and lab sewage ponds sinclair 1975 daphnia can assistance was provided by L maddison N alter their body structure in response to algal anderson PE saffel S alienallenailen and C birkelo concentrations which is thought to be a this research is a contribution from the INEL mechanism for surviving algal blooms pennak radioecologyRadioecology and ecology program and was 1989 thus while conditions in sewage ponds funded by the new production reactor are hostile to many species those that can tol- office idaho field office and the office of erate the conditions flourish due to an abun- health and environmental research US dant food supply and the absence offishof fishhishbish department of energy in summary wastewater ponds had low invertebrate diversity which we attribute to literature CITED lack of vegetation and inability of many BOGATOVA I1 B AND Z I1 yerofeyeva 1980 the use of condi- species to withstand the environmental container reared cultures of cladocera in polishing tions wastewater ponds also had high inverte- fish farm effluents hydrobiological journal 16 brate abundance which we attribute to reduc- 56 61 tion of competing taxa organic enrichment BORROR D J AND D M DELONG 1971 an introduction to the study of insects ard3rd edition holt rinehart and absence of vertebrate predators there and winston new york NY 812 appp was no indication that heavy metal concentra- BRIGHTMAN R S AND J L fox 1976 the response of tions were high enough to reduce water column benthic invertebrate populations to sewage addition invertebrate concentrations in most ponds pages 295 308 in third annual report on cypress wetlands florida university center for wetlands high invertebrate concentrations min INEL gainesville food wastewater ponds provided an abundant BRIX H 1993 wastewater treatment in constructed wet- source for many bird species migratory and lands system design removal processes and treat- resident which used INEL wastewater ponds ment performance pages 9 22 in G A moshinmoshiri bacteria protozoa and algae are important in editor constructed wetlands for water quality im-im they reduce the proprovementvement lewis publishers ann arbor MMII1 waste treatment because BROWN E M W SKOUGSTAD AND M J FISHMAN 1970 organic load of wastewater and convert waste methods for collection and analysis of water samples into a form useable by organisms in the receivreceive for dissolved minerals and gases techniques of 199511995 invertebrates IN wastewater PONDS 115

water resources investigations of the united states HALFORD D K AND J B MILLARD 1978 veltevertebrateVeitebrate geological survey book 5 chapter al USU S govern- fauna of a radioactive leaching pond complex in ment printing office washington DC 160 appp southeastern idaho great basin naturalist 38 CHADWICK J W S P CANTON AND R L DENT 1986 64 70 recovery of benthic invertebrate communities in hilsenhoff W L 1988 rapid field assessment of organ- silver bow creek montana following improved ic pollution with a family level biotic index journal metal mine wastewater treatment water air and of the north american Benthbenthologicalological society 7 soil pollution 28 427 438 65 68 CIEMINSKI K L 1993 wildlife use of wastewater ponds HOWE FE P AND L D FLAKE 1989 mourning dove use at the idaho national engineering laboratory of man made ponds in a cold desert ecosystem in unpublished master s thesis south dakota state idaho great basin naturalist 49 627 631 university brookings 311 appp KLOTZ L 1977 the effects of secondarily treated sewage CLAWSON K L G E START AND N R RICKS 1989 effluent on the willimanticshetucketwilhmanticshetucket riverbiver uni- climatography of the idaho national engineering versity of connecticut institute of water resources laboratory and2nd edition USU S department of com- storrs report 27 85 appp merce national oceanic and atmospheric adminis- kondratieff P F R A MAITHEWSMATTHEWS AND A L BUIKEMA trationtration idaho falls ID DOE ID 1211815512118 155 appp JR 1984 A stressed stream ecosystem macroinvermacromver COWARDIN L M V CARTER F C GOLET AND E T LAROELARGE tebratetebrake community integrity and microbial trophic 1979 classification of wetlands and deepwater habi- response hydrobiologia 111 81 91 tats of the united states USU S fish and wildlife kondratieff P F AND G M SIMMONS JR 1982 service office of biological services washington nutrient retention and macromacroinvertebrateinvertebrate community DC FWSOBS 7931 structure in a small stream receiving sewage efflu- CRAWFORD C G D J WANGSNESS AND J D MARTIN ent archivarcbivarchev fur hydrobiologie 94 83 98 1992 recovery of benthic invertebrate communities KOWNACKI A 1977 biocenosisbiocoenosis of a high mountain stream 4 fauna in the white river near indianapolis indiana USA under the influence of tourism the bottom following implementation of advanced treatment of of the stream rybi potok the high batratatra mts acta municipal wastewater archivarchev furfuir hydrobiologie hydrobiologicahydrobiological 19 293 312 126 67 84 LEWIS M A 1986 impact of a municipal wastewater effluent on quality DODSON S I1 1987 animal assemblages in temporary water periperlpenpenphytonperiphytonphyton and inverte- desert rock pools aspects of the ecology ofdasyheleaofdasyheleaDasy helea brates in the little miami river near xenia ohio subletsubletteisublettertei diptera ceratopogomdaeceratopogonidae journal of the ohio journal of science 86 2 8 north american Benthbenthologicalological society 6 65 71 LIAO C FHF H AND D R S LEAN 1978 nitrogen trans- lakes DUDA A M D R LENAT AND D L PENROSE 1982 formations within the trophogenic zone of journal of of 35 water quality in urban streams what we can expect fisheries research board canada journal of the watelwater pollution control federation 1102 1108 LUBINI FERLIN V 1986 sewage treat- 54113954 1139 1147 the influence of plant effluents on benthic invertebrates in EULISS N H JR 1989 assessment of drainwaterdramwaterbrainwaterdraindramwater evapo- ment in scheische ponds as waterfowl habitat in the san joaquin lake zurich switzerland SchweitzenschweltzerschweitzerischeschweitzenscheSchweitzer zeit ration in our valley california unpublished doctoral dissertation schrift fur hydrologicHydro logie 48 53 63 oregon state university corvallis MASON W T JR AND P P YEVICH 1967 the use of phlorinephloxmephloxinephloxinelne B and rose bengal stains to facilitate sorting EULISS N H R L JARVIS AND D S GILMER 1991 JR samples of the feeding ecology of waterfowl wintering on evapora- benthic transactions american microscopical society 86 221 223 tion ponds in california condor 93 582 590 mcbride R N R FRENCH A H DAHL AND J E FISHMANFlshwanSHMAN M J AND L C FRIEDMAN EDITORS 1989 shwas DETMER 1978 vegetation types and surface soils of methods for determination of substances inorganic the idaho national engineering laboratory in water and fluvial sediments ard3rd edition tech- site in IDO 12084 US department of eneigyenergy idaho of water resources investigations of the niques operations office idaho falls 29 appp united states geological survey book 5 chapter al MERRiMERRITTTr R W AND K W CUMMINS 1984 an introduc- USU S government printing office washington DC merrl tion to the aquatic insects of north america and2nd 545 appp edition kendallhuntKendall Hunt publishing co dubuque 10 GILINSKY E 1984 role of fish predation and spatial the 722 appp heterogeneity in determining benthic community MILLARD J B F W WHICKERWHICKFR AND D MARKHAM structure ecology 65 455 468 0 1990 radionuclideRadio nuclide uptake and growth ofbarnosbarnof barn swal- GLOYNA E F MALINA AND E M DAVIS EDI- E F J JR lows nesting by radioactive leaching ponds health ponds aeternaalter TORS 1976 as a wastewater treatment alternana- physics 58 429 439 tive for water resources center research in univer- MOSHIRI G A EDITOR 1993 constructed wetlands for sity of texas at austin 447 appp water quality improvement lewis publishers ann GORDON C FLAKE K HIGGINS C L D AND F 1990 arbor MIM 1 632 appp aquatic invertebrates in the rainwater basin area of NECKLES H A H R MURKIN AND J A COOPER 1990 nebraska prairie naturalist 22 191 200 influences of seasonal flooding on macroinverte GOULDEN C E 1976 biological species interactions and brate abundance in wetland habitats freshwater their significance in waste stabilization ponds pages biology 23 311 322 57 67 in E F gloyna J FE malina jr and E M OLIVE J H AND C A DAMBACH 1973 benthic macroinmaceoin davis editors ponds as a wastewater treatment vertebrates as indexes ofwaterof water quality in whetstone alternative center for research in water resources creek morrow county ohio scioto river basin university of texas at austin 447 appp ohio journal of science 73 129 149 116 GREAT BASIN naturalist volume 55

PAIRICKPATRICK R 1976 the effect of a stabilization pond on the strable biological benefits environmental manage- sabine estuary pages 33 55 in E F gloyna J FE ment 4 49 56 malina jr and E M davis editors ponds as a SINCLAIRNCLAIR R M 1975 freshwater biology and pollution wastewater treatment alternative center for ecology training manual EPA 4301 75 005 national research in water resources university oftexasof texas at technical information service springfield VA austin 447 appp TASKSK FORCE ON NATURAL SYSTEMS 1990 natural systems PEARSONPCARSONRR GANDLG AND L K PENRIDGE 1987 effects ofpolofoppolpol- for wastewater treatment water pollution control lution by organic sugar mill effluent on the macro federation alexandria VA 270 appp invertebrates of a stream in tropical queensland UNITEDNITED STATES environmental protection AGENCY australia journal of environmental management 1987 quality criteria for water 1986 EPA 4405864405 86 2420524 205 215 001 office ofwaterof water regulations and standards USU S PENNAKPLNNAK R W 1989 freshwaterfresh water invertebrates of the government printing office washington DC united states protozoa to mollusca ard3rd edition 19871302 m60645 john wiley & sons inc new york NY 628 appp VICTORCTOR R AND D T DICKSON 1985 macrobenthicMacrobenthic porcellapoboPOBC eliellLLI A D B P H MCGAUHEY AND G L DUGAN invertebrates of a perturbed stream in southern 1972 response to tertiary effluent in indian creek nigeria environmental pollution series A 38 reservoir journal of the water pollution control 99 107 federationfedeiation4444 2148 2161 WALLACE J B AND R W MERRITT 1980 filter feeding richardson J S 1984 effects of seston quality on the ecology of aquatic insects annual review of entom- growth of a lake outlet filter feeder oikosbikos 43 ology 25 103 132 386 390 WETZELETZEL R G 1983 limnology and2nd edition saunders ross L C M AND H R MURKIN 1989 invertebrates college publishing chicago IL 767 appp pages 35 38 in E J murkin and H R murkin edi- BIGGINSWIGGINS G B R J MACKAY AND I1 M SMITH 1980 tors marsh ecology searchleresearch program long term evolutionary and ecological strategies of animals in monitoring procedures manual delta waterfowl annual temporary pools archivarchev fur hydrobiologie wetlands research station technical bulletin 2 supplement 58 97 206 SAVAGE N L AND F W RABE 1973 the effects of mine WILLIAMS W D 1985 biotic adaptations in temporary and domestic wastes on macromacroinvertebrateinvertebrate commu- lentic waters with special reference to those in semi- nity structure in the coeur d alenedalene river northwest arid and and regions hydrobiologia 125 85 110 science 47 159 168 seagleSEACLESLAGLL H IIIT11 JR A C hennricksHENDRICKSHCNDRICKS AND J CAIRNS JR received 14 january 1994 1980 does impiimprovedoved waste treatment have demon accepted 7 september 1994 great basin naturalist 552 C 1995 appp 117 123

GROWTH AND reproduction IN AN ALPINE CUSHION PLANT astragalus kentrophyta VAR IMPLEXUS

wayne R owenibweniowen1

ABSTRACT A two year field experiment was conducted to investigate factors hypothesized to affect the reproduc- tive potential of astragalus kentrophyta var implexus and to test the importance of tradeoffstrade offs between growth and repro- duction in this species levels of mineral nutrients water herbivoreherbherbivoryivory and competition were manipulated seed output and growth of individuals in treatment groups were compared against control plants neither water nor mineral nutri- ents alone were shown to affect growth or reproduction herbivoreHerbherbivoryivory was shown to be similarly unimportant in affecting growth and reproduction competition with other species influenced growth but not reproduction no significant trade- offs between growth and reproduction were detected within years however there did appear to be a tradeofftrade off between these major fitness components when compared between years

key words astragalus alpine competition fecundity tradeofftrade off white mountains

the impact of resource availability on the limited experiments involving other organ- reproductive output of plants is well estab- isms from this habitat have shown that avail- lished harper 1977 schoener 1983 fowler ability of resources influences the competitive 1986 welden and slausenblausen 1986 plants may ability and distribution of species wright and experience resource limitation as a result of mooney 1965 mooney 1966 marchand 1973 competition inter or intraspecific or poor though this is not generally true of all alpine habitat quality resource limitations can also habitats korner 1989 second standing bio- occur when a portion of a plant s photosyn- mass and percent cover are substantially lower thetic organs are removed eg by herbivoreherbivoryherbivory on dolomitic soils than on adjacent sandstone damage which clearly interferes with the plant s and granite derived substrates suggesting that ability to provision its offspring marquis plants on the dolomite barrens might be rela- 1991 A number of authors cody 1966 tively resource limited mooney 1966 owen 1967 1977 macarthur and wilson harper 1991 A kentrophyta plants routinely grime third 1979 tilman 1982 weiner 1988 1990 abort the majority of flowers they produce have considered the ecological consequences each year owen 1991 a pattern that has been of resource limitation for individuals and pop- attributed to resource limitations in a broad ulatulationsions and have described various strategies spectrum of species and that plants might be expected to pursue to lovett doust lovett optimize the allocation of limited resources doustdoust19881988 this study tests whether the availability of an experiment was designed 1 to test resources limits the fecundity of astragalus whether there are resource constraints on the kentrophyta gray var implexus canby reproduction and growth of A kentrophyta barneby hereafter simply A kentrophyta and and 2 to assess the interactions between two to what extent tradeoffstrade offs between growth and major components of fitness ie growth and reproduction might influence patterns of reproduction under different regimes of reproduction observed in this species A ken resource availability to do this a factorial troptrophytahyta is an alpine cushion plant indigenous field experiment was established in which sep- to high elevations throughout the intermoun- arate groups of plants would receive either 1 tain west of north america barneby 1964 water or 2 nutrient supplements 3 protec- many lines of evidence suggest that repro- tion from herbivoreherbherbivoryivory or 4 relief from the duction in A kentrophyta might be resource potentially competitive influence of neighbors

IUDiversityuniversityiudiversity of california davis and white mountain research station university of california los angeles present address boise national forest 1750 front street boise ID 83702

117 118 GREAT BASIN naturalist volume 55

STUDY AREA as water 2 another 50 plants received sup- plementalplemental nutrients these plants were given the study was conducted on the alpine approximately 17 g of a balanced general pur dolomite barrens of sheep mountain pass pose fertilizer scott s all purpose builder above the patriarch grove bristlecone pine 121012 NPK providing each plant with 202.0 g forest in the white mountains of mono N in the form of ammoniacal nitrogen areasureas county CA elevations at the site range from and water soluble nitrogen 17ltit1.7 g P from 3535 m 11600 ft to 3660 m 12000 ft and phosphoric acid pao and 202.0 g K from sol- topographic relief of the site is minimal in the uble potash KO these quantities are equiv- white mountains A kentrophyta occurs only alent to application rates of 13813.8 11711.7 and on dolomitic soils lloyd and mitchell 1973 13813.8 kg ha 1 respectively A balanced fertilizer hall 1991 was chosen because experiments by chambers weather data were obtained from the white et al 1987 and shaver and chapin 1980 have mountain research station mt barcroft shown that plants in cold environments re- laboratory located 6 km north of the study spond most vigorously to resource augmenta- site at an elevation of 3800 m soils on the dolo- tion with fertilizer containing a balance of mite barrens have a high cation exchange essential nutrients the dry fertilizer was sca- capacity and are depauperate in nitrogen ttered in an approximately 2 cm wide ring phosphorus and potassium mooney et al around the perimeter of each test plant 1962 wright and mooney 1965 brayton and summer seasonal precipitation in 1989 was mooney 1966 mooney 1966 marchand 1973 apparently sufficient to solubilize the fertilizer 1974 the moisture holding capacity of and deliver it to the soil profile as the granules dolomite derived soils is equivalent to that of had completely disappeared from the surface adjacent granitic soils mooney et al 1962 in approximately one month this treatment wright and mooney 1965 marchand 1973 group will be referred to as fertilized 3 A vegetation of the white mountains is general- third treatment was designed to protect plants ly xerophytic this trend is especially prevalent from herbivoreherbivoryherbivory and predation on flowers and on the dolomite barrens lloyd and mitchell young fruits two locally common insects ha- 1973 bitubituallyally consume the reproductive parts of A kentrophyta the more common of these in- MATERIALS AND METHODS sects a darkling beetle tenebrionidae coleop- tera consumes flowers larvae of a locally com- in june 1989 195 healthy A kentrophyta mon lycinid butterfly species lycaenidae plants were selected randomly from within an lepidoptera occasionally consume immature area of approximately 020.20 2 ha decadent senes- A kentrophyta fruits tanglefoottangle foot brand cent plants were disqualified from inclusion sticky trap was applied in a circle around each in this experiment the specific location of the of 25 plants to exclude potential herbherbivoresivores site was chosen for its apparent homogeneity tanglefoot barriers were repaired as needed with respect to soil physical characteristics this treatment group will be called no vegetation and topographic profile plants predation 4 the fourth treatment sought to were randomly allocated to five treatment relieve a group of 20 A kentrophyta plants regimes 1 50 plants were provided with from neighborhood competition A 025 m three separate ILI111 L applications of water dur- radius circle around a central target A kentro ing the 1989 growing season plants were phytapayta plant was cleared of all other plants by watered during the driest part of the summer cutting them off at ground level this method 4 july 2 august and 19 august to maximize minimized ground surface disturbance the beneficial impact of the treatment water clearings were 020.2 m2ma in area the average was applied slowly to maximize infiltration in number of neighbors cametsrarametsmets removed was 63 a radius of 12512.512 5 cm around each plant this mostly tillers of poapoo rupicola covering an treatment supplied 616.16gi 1 cm of moisture to each average of 15 of the ground surface plant expected precipitation for the three excavations of A kentrophyta plants show that month growing season is 878.78 7 cm pace et al its roots grow straight downward into the soil 1968 the 1989 summer precipitation was 11iili1.11 1 with minimal lateral root spread owen 1991 cm this treatment group will be referred to roots of the target plants were therefore 199511995 astragalus GROWTH AND reproduction 119 thought to be well isolated from interactions production but previous experience owen with actively assimilating roots of other plants 1991 had shown that seed production is a sig- plants clipped in the cleared areas were nificantnificant function of flower production owen trimmed if they resprouted plants in this treat- 1991 flowers when aborted are dropped at ment group are referred to as the target a very early age owen 1991 and probably group 5 A final group of 50 unmanipulated represent a minimal per unit cost in resources plants was marked as a control group size to the plant bookman 1983 stephenson of the experimental groups was based on an 1984 therefore the cost of flowers should be analysis of expected variances in responses to proportional to a plantpiantplanes s seed output and can the treatments lower expected variances re- safely be disregarded for the purpose of this quire smaller necessary samples sokal and work fruits and seeds were cleaned and sepa- rohlfrohlfl9811981 rated in the laboratory counted and weighed plant sizes cushion area were measured and recorded on 23 june 1989 shortly after RESULTS initiation of growth for the season treatments were initially applied on 4 july 1989 in sep- weight of individual reproductive struc- tember 1989 all plants were remeasured and tures seeds and fruits was independent of the entire fruit and seed crop produced by total numbers of those items produced per each of the 195 plants was harvested since A plant in both years table 1 average seed and kentrophyta forms a tight cushion that never fruit weights were significantly correlated fi exceeds I1 cm in height and seeds are not 429429.429 in 1989 R 443443.443 in 1990 there were released from the plant before the end of the no significant differences between treatment growing season there was great confidence groups for the weight of individual seeds or that the entire seed crop of each individual fruits results not presented because seed was retrieved in early june 1990 I1 again mea- production is well correlated with other possi- sured the area of all plants just as they were ble measures of fitness in A kentrophyta and initiating growth for the season fertilized and weights of those seeds are independent of the water treatments were not repeated in 1990 numbers of reproductive structures produced so as to evaluate the potential for lags in the on a plant table 1 seed output was used as effectiveness of resource supplementation an index of total reproductive effort tanglefoot barriers were maintained during in a comparison of slopes of regression 1990 to test for interinterannualannual variation in the analyses growth was a significant function of effects of herbherbivoresivores and predators clear zones plant size in both 1989 and 1990 table 2 around target plants were maintained in 1990 though the relationship was weaker in 1990 all plants were allowed to grow through the the weight of individual seeds and fruits was season and in september 1990 all 195 plants independent of seasonal growth table 2 the were remeasured and all fruits and seeds har- amount of growth across years was significant- vested no attempt was made to quantify flower ly but poorly correlated

TABLE 1 correlation matrix for selected demographic traits values above the diagonal are correlation coefficients R based on 1990 data those below the diagonal are derived from 1989 data

seed seed fruit fruit reproductiverepi oductiveeductive seeds weight weight fruits weight weight weight produced average total produced average total total seeds produced 1 003 976 964 143 920 966 seed weight average 042 1 139 001 433 081 115 seed weight total 977 200 1 945 229 937 987987.98798711 fruits produced 963 024 033 1 106 963 968 fruit weight average 136 429 215 074 1 289 260 fruit weight total 943 120 949949.9499491111 952 284 1 981 total reproductive weight 973 1656 989 954 249 985 1 kendall rank correlation is significant at P 05 treatment differencedifferences noted with one anyvnyay ANOVA these differences do not affect the magnitude of significance oftleoftbeof the correlationcorrelations 120 GREAT BASIN naturalist volume 55

tailleTABLE 2 slopes of regressionsofregressions for selected demographic TABLE 3 result of an ANCOVA on seed production and traits on growth in 1989 and 1990 using the total data set growth by treatment group the covariatecovanate is plant size ie not partitioned by treatment where the overall the treatments are those listed in the text see also table 4 regressions are not significant there were also no treat- ment differences covariatecovanateCovanate treatment

growth in 1989 growth in 1990 F P F P 001 25 growth in 1990 168 1989 seed production 37164 1358 1990 production 1 plant size 340 iioilo110.110 seed 3981839 818 001 18541.854854 12 1989 growth 001 583 seed weight 038038.038 405054054 27207 0822 1990 growth 346 047 fruit weight 035035.035 036036.036 0893 2453 regressions aiedieateare significantly positive P 05 one way ANOVAs suggsuggestt clitlcienccsdif&rences between tieatmentticatinenttreatment groups ffirafir01 valesvalues of these traits P 05

A of simple linear was seed production square root transformed seriesserles regressions used to compare seed production with growth was a positive linear function of plant size to test for the presence of a tradeofftrade off between overall values of r2ra for regressions of seed these two components of fitness production on plant size were 206206.206 in 1989 primary when the data are corrected for the fact that and 182.182 in 1990 slopes of individual regres- larger plants are inherently more capable of sions for each treatment for seed production producing more flowers and fruits the analy- on plant size did not differ from the slope for sis finds no significant differences among control plants sis treatment groups by virtue of overlapping plant was a factor size minor but important 95 confidence intervals and therefore no both and reproduction influencing growth in tradeofftrade off between growth and reproduction A kentrophyta and indicates that size should within a given year was detected be considered as a covariate in an analysis of to compare tradeoffstrade offs across years the this variance of treatment effects in experi- ratio of 1990 to 1989 data was used table 5 ment analyses of covariance ANCOVA and this provides a number 101.0loio1 0 when 1990 data experimental results are presented in tables 3 values exceed 1989 values the converse is and 4 respectively plant size was a significant true when results are 101.0loio1 0 seed production covariate three of four analyses were in there was greater in 1990 than in 1989 regardless of no among groups differences treatment in treatment group in contrast growth in 1990 seed production reproduction for either year was less than that experienced 1989 with growth in did not differ among treatment groups the notable exception of target plants the in 1989 but there was a significant difference results can be interpreted as evidence for a between groups in 1990 FP 047047.047 A protect- tradeofftrade off between growth and reproduction ed leasleast t significant difference LSD test indi- they indicate that in general increased seed cates that growth in the target group was production is associated with decreased growth greater than that of individuals in other treat- furthermore plants may be relieved of trade- ment groups table 4 off constraints by removing competitors table 5 gives the results of two tailed t tests which should increase availability of mineral comparing mean reproduction and growth resources to the remaining target plant across years within treatment groups there were no significant differences for seed pro- discussion duction among treatment groups between 1989 and 1990 average size for plants in 1990 resource supplementation or alleviation of was consistently significantly greater than the resource competition did not significantly size of the same plants the previous year ie influence the reproductive output ofaofA kentro on average plants grew larger over the course phytapayta instead seed production was more close- of the experiment the no predation treat- ly related to the individual s past record of seed ment grew significantly less in 1990 than output tables 1 3 5 plants that produced 1989 whereas plants in the target group grew many seeds in 1989 tended to produce many significantly more in 1990 there were no sig- seeds in 1990 regardless of treatment growth nificantnificant differences in growth across years for while similarly unresponsive to the addition of plants in the control fertilized or water single resources increased significantly when groups potential competitors were removed tables 4 199519951 astragalus GROWTH AND reproduction 121

TABLE 4 treatment means SD in both 1989 and 1990 for important demographic traits controlcoicolatrolntrol nonolnoibugs fertilizedfertilized waterWaierkierater targetta rgetaget 1989 seed production 258 252 161 118 306 245 251 222 442 414 1990 seed production 322 322832 28 205 167 397 373 307 279 545 584

1989 plant size 59971 285128517 7 45946 187118718 8 68339 289272892289272892.7 7 63332 289142891289142891.4 4 76832 368383683368383683.8 8 1990 plant size 72473 312831288 8 55963 2156215662156.6 6 79340 324232426 6 74182 362743627362743627.4 4 83930 415941599 9

1989 growth 14784 13297132971329.71329 7 15300 98779877987.7987 7 17721 163421634163421634.2 2 17979 148691486148691486.9 9 15031 98869886988.6988 6 1990 growth 11561 15299152991529.91529 9 8084 1000100041000.4 4 15878 9044204420445 5 13950 1760176031760.3 3 24332 174901749174901749.0 0 growth inin 1990 varied significantly among treatments see table 3 the target groups grew more on average thinthan did plants inin inyany othelother treatment gioupgroup no other diffeiencesdifferences were significant

5 these results differ from those of wright 1989 is most sensitive to the proximity of its and mooney 1965 mooney 1966 and neighbors it is unclear however why repro- marchand 1973 which show that mineral duction among such species is rarely similarly nutrients were the primary factors limiting influenced as is the case with A kentrophyta other species that occur on dolomite in the the buffering of fitness components against white mountains artemisia tridentata two environmental stochasticity is characteristic of erigeron species and lupinus argenargenteusargenteousteus density vague demographics as described by respectively korner 1989 reports that the strong 1986 under density vague condi- effect of fertilization on the growth of species tions selection favors demographic functions from nutrient poor environments is often diffi- with indeterminate functional thresholds that cult to detect he does not cite studies that is current allocation decisions are only loosely address the relationship between growth and linked to current environmental conditions reproduction in nutrient supplementation strong 1986 experiments tradeoffstrade offs between growth and reproduc- the addition of mineral nutrients or water tion within years were not observed in this alone may have been insufficient stimuli for A experiment under any conditions A weak kentrophyta to increase either reproduction or tradeofftrade off between growth and reproduction growth if both factors were limiting multiple was identified in most treatment groups when limiting factors have been reported in a vari- data were compared across years table 5 it ety of species harper 1977 and are specifi- is of great interest that the target group alone cally predicted by tiiTiltilmansmarsmais 1980 1982 mod- experienced an increase in both seed produc- els of optimal resource consumption that tion and growth in 1990 compared to 1989 val- there may be multiple resource limits to A ues and thus did not experience a tradeofftradeoffstrade offofftoffs kentrophyta growth and reproduction is sup- the absence of well defined tradeoffstrade offs between ported by the response of A kentrophyta to primary components of fitness could be due to the removal of competitors in this study one of several reasons lack of a discernible tanglefoot barriers were very effective at tradeofftrade off would be noted if resources were not excluding ground moving herbherbivoresivores and truly limiting it may also be that growth and predators this was evidenced by the lack of reproduction are not co limiting for this foliar damage or partially eaten fruit and the species in this environment if this were true capture of many insects in the traps flowers factors that influence growth and reproduction of A kentrophyta are produced in sufficient are likely to be independent eg one fitness excess to buffer individuals against the levels component might be canalizedcanalized and the other of flower and fruit predation observed in this dependent on environmental conditions population finally a tradeofftrade off between growth and repro- growth in A kentrophyta as has been re- duction would not be detected if a resource ported for a number of species from andaridarld regions other than one provided in this experiment throughout the world fonteyn and mahall were limiting 1981 robberecht et al 1983 ehleringer adult A kentrophyta mortality at the sheep 1984 parker and salzman 1985 shaw 1987 mountain study site is low juvenile mortality manning and barbour 1988 and chapin et al is extremely high even though germination 122 GREAT BASIN naturalist volume 55

tailleTABLE 5 cross year comparisons of fitness components 1990 values represented as a fraction of 1989 trait values values ofoftt and the associated probabilities P represent results of two tailed t tests for differences in values between years refer to table 4 for raw data control no bugs fertilized water target

seed production 9089 125 1161.16ilg1 16 132 1151.151 15 1181.181 18 t 141 171 180 139 071 P 17 10 08 17 49 plant size t 706 502 490 505 350 P 01 01 01 01 01 growth 9089 085 098 056 086 207 t 1131.131 13 2502.502 50 0400 40 142 2122.122 12 P 26 02 70 16 05

malmaiValvilucssaltiesvaltiesmaitlestiesiles listed represent the ratio olof199001 1990 traitirali values to those of 1989

tests under controlled conditions show seed literature CITED viability of greater than 95 and recruitment is low owen 1991 these demographic attri- BARNEBY R C 1964 atlas of north american astragalus memoirs of the new york botanical garden 13 butes would certainly favor a strategy that 1 1187 routes resources away from the risky business BOOKMAN S S 1983 costs and benefits of flower abscis- of reproduction toward growth the small but sion and fruit abortion in asclepias speciosospeciosaspeciosa ecology consistent portion of A kentrophyta s annual 6426464 264 273 BRAYTON MOONEY population accumulation of biomass allocated to repro- R AND H A 1966 vari- ability of Cercocercocarpmcercocarpuscarpus in the white mountains of cali- duction guarantees that each plant will proba- fornia as related to habitat evolution 20 383 391 bly produce at least a few seeds each year CHAMBERS J C J A MACMAHON AND R W BROWN while being able to dedicate most of each sea- 1987 response of an early seraiseral dominant alpine grass and a late seraseraiseral dominant alpine forb to N and P sols s accumulated resources to growth and son availability reclamation and revegetation research survival that the allocation of resources to 6 219 234 reproduction but not growth in this species is CHAPIN FE S J B MCGRAW AND G R SHAVER 1989 constant over a broad range of resource avail- competition causes regular spacing of alder in abilities is consistent with a bet hedging life alaskan shrub tundra oecologia 79 412 416 CODY M L 1966 A general theory of clutch size history strategy kozlowski and steamsstearns 1989 evolution 20 174 184 philippi and seger 1989 steamsstearns 1989 ehleringer J R 1984 intraspecific competitive effects resource limitations on organisms are rarely on water relations growth and reproduction in simple or solitary while fruit and flower pre- encehafarinosaencelia fannorafannosa oecologia 63 153 158 dation can be an important limit on fecundity FONTEYN P J AND B E MAHALL 1981 an experimental analysis of in desert such an effect was not noted here similarly structure in a plant community journal of ecology 69 883 896 reproductive output of plants growing on the FOWLER N 1986 the role of competition in plant commun- the sheep mountain dolomite barrens would ities in and and semiarid regions annual review of appear to be resource limited although single ecology and systematics 17 443 464 resource augmentation had no direct effect on GRIME J P 1979 plant strategies and vegetation process- seed production in combination however es john wiley and sons new york NY resources can influence the amount of realized HALL C A EDITOR 1991 natural history of the white inyo range university of california press berkeley growthcrowth will affect 9rowth that in subsequent years HARPER J L 1977 population biology of plants academic reproduction press new york NY KORNER C 1989 the nutritional status of plants from acknowledgments higher altitudes oecologia 81 379 391 KOZLOWSKI J AND S C STEARNS 1989 hypotheses for the production of excess zygotes models of bet I1 would like to thank the white mountain hedging and selective abortion evolution 43 research station for providing logistic and 1369 1377 financial support for this project especially LLOYD R M AND R S MITCHELL 1973 A flora of the the crew at the mt barcroft laboratory T white mountains california and nevada university of california press berkeley holmes E nagy A and two anony- fitter LOVETT DOUST J AND L LOVETT DOUST 1988 plant mous reviewers made significant improve- reproductive ecology oxford university press new ments on earlier drafts of this manuscript yorknyyorkonyyork NY 199519951 astragalus GROWTH AND reproduction 123

MACARTHUR R H AND E 0 WILSON 1967 the theory SCHOENER T W 1983 field experiments on interspecific of island biogeography princeton university press competition american naturalist 122 240 285 princeton NJ SHAVER G R AND FE S CHAPIN 1980 response to fertil- MANNING S J AND M G BARBOUR 1988 root systems ization by various plant growth forms in an alaskan spatial patterns and competition foiforholbol soil moisture tundra nutrient accumulation and growth ecology between desert subshrubssub shrubs american journal of 6166261 662 675 botany 75 885 893 SHAW R G 1987 density dependence in salvia lyratelyrata MARCHAND D E 1973 edaphic control of plant distribu- experimental alterations of densities of established tion in the white mountains eastern california plants journal of ecology 75 1049 1063 ecology 54 233 250 SOKAL R R AND FE J ROHLF 1981 biometry WH 1974 chemical weathering soil development and freeman and company new york NY geochemical fractionation in a part of the white STEARNS S C 1989 tradeoffstrade offs in life history evolution mountains mono and inyo counties california functional ecology 3 259 268 USGS professional paper 352 J stephenson A G 1984 the cost of over initiating fruit MARQUIS R J 1991 evolution of resistance in plants to american midland naturalist 112 379 386 herbivoresherbivores evolutionary trends in plants 5 23 29 STRONG D R 1986 density vagueness abiding the varivarlvari- MOONEY H A 1966 influence of soil type on the distribu- ance in the demography of real populations pages tion of two closely related species of erigeron ecology 257 268 in J diamond and T J case editors 4795047 950 958 community ecology harper and row publishers MOONEY H A G ST ANDRE AND R D WRIGHTWKIGHT 1962 new york NY alpine and subalpine vegetation patterns in the TILMAN D 1980 resources a graphicalgrap cicalhical mechanisticmeehanmechan is tictletie white mountains of california american midland approach to competition and predation american naturalist 68 257 273 naturalist 116 362 393 OWEN W R 1991 the reproductive ecology of an alpine 1982 resource competition and community struc- legume A kentrophyta var implexus unpublished ture princeton university press princeton NJ dissertation university of california davis 226 appp WEINER J 1988 variation in the performance of individ- PACE N D W KIEPERT AND E M NISSEN 1968 climat- uals in plant populations pages 59 81 in A J davey ologicalto data summary for the crooked creek labora- M J hutchings and A R watkinson editors plant tory 1949 1967 and the barcroft laboratory 1953 population ecology blackwell scientific publications 1967 university of california white mountain london research station publication berkeley 1990 resource competition and community struc- PARKER M A AND A G SALZMAN 1985 herbivore ture trends in evolution and ecology 5 360 364 exclosure and competitor removal effects on juve- WELDEN C W AND W L SLAUSEN 1986 the intensity nile survivorship and growth in the shrub of competition versus its importance an overlooked Gutiergutierreziagutierrezbarezia microcephalmicrocephalamicrocephalya journal of ecology 73 distinction and some implications quarterly review 903 913 of biology 61 23 44 PHILIPPIphilippitandjT AND J SEGER 1989 hedging one s evolution- WRIGHT R D AND H A MOONEY 1965 substrate ori ary bets revisited trends in ecology and evolution entedanted distribution of bristleconebnstlecone pine in the white 4 41 44 mountains of california american midland naturalist robberecht R B E MAHALL AND P S NOBEL 1983 7325773 257 284 experimental removal of intraspecific competitorss effects on water relations and productivity of a received 21 january 1994 desert bunchbunchgrassbuncbgrassgrass hilaria ngidarigidabrigida oecologia 60 accepted 28 october 1994 21 24 great basin naturalist 552 0 1995 appp 124 134 calileuctra A NEW GENUS AND TWO NEW SPECIES OF stonefliesSTONE FLIES FROM california plecoptera leuctridae

W D SheparShepardshepardlshepard1shepardddl1 and R W Baumann2

ABSTRACT calileuctracaltleuctra is proposed as a new genus in the family leuctndaeleuctridae with calileuctracahleuctra ephemera designated as the type species all stages of calileuctracahleuctra ephemera are described calileuctracahleuctra dobryidobayi is described in the male and female stages both species inhabit the mediterranean climatic region of california A phylogenetic analysis of the gen- era in the family leuctndaeleuctridae is given which places cahleuctracalileuctra near the genus Perloperlomyiamyia

key words insecta plecoptera leuctridaeleuctndae calileuctra description distribution phylogeny

both of us have been collecting stonestonefliesflies tionseions all from mountains surrounding the los from streams all across california several years angeles basin ago one of us WDS collected a small and poorly sclerotized stonstoneflyefly nymph from an inter- calileuctra new genus napa mittent valley stream the male adult TYPE SPECIESPECIESS calileuctra ephemera new that was reared from the nymph could not be species determined using existing keys by WDS the ADULTS body brownish weakly sclero- specimen was then given to RWB for identifi- tized setation sparse except for abundant tiny cation his identification kept us collecting at setae clothing hairs figs 1 10 wings the same site for nine years the single male macropterous or brachypterous venation as specimen was first thought to be a new species illustrated fig 3 prosternum with prester- in the asian genus rhopalopsole however num separate furcasternumfurcasternum fused to base of recent work indicates that the male represents triangular basisternumbasisternum meso and metaster- a new genus in the family leuctridae despite num similar except basisternumbasi sternum rectangular extensive searching in surrounding areas only fig 2 the napa valley population has been found MALE tergum IX with posterior border A few years after discovery of the first new heavily sclerotized and irregularly serrate or species RWB found in the natural history dentate tergum X with posterolateral corners museum of los angeles county a small series each with one or two elongate horns project- of an interesting new leuctrid from the san ing posteriorly figs 4 11 sternum IX pro- gabriel mountains later two additional fe- jecting posteriorly to cover base of paraprocts males of this species were collected in the with vesicle broadening posteriorly figs 6 santa ana mountains however we decided 13 paraprocts fused into a complex T that fresh male specimens were needed before shaped subanal probe with two ventromedial a description could be undertaken projections off subanal probe figs 8 9 13 keith dobry who was doing fieldwork in FEMALE with weak abdominal sclerotisclerote the los angeles area was encouraged to look zationbation sternum vilVII completely sclerotized for additional specimens of this leuctrid species sternum villVIII largely membranous sternum he was successful in locating two additional IX completely sclerotized subgenital plate populations one in the san gabriel moun- poorly produced sternum X incompletely tains the other in the santa monica mountains sclerotized figs 7 14 cerci one segmented this species is known from only four popula elongate in male poorly sclerotized on sides

ldepartinentcpaitment of entomology california academy of sciences golden gate park san francisco CA 94118 mailing address 6824 linda sue way fair oaks CA 95628 2departmeutepaitment of zoology monte L bean life science museum brigham young university provo UT 84602

124 199511995 calileuctra NEW GENUS 125 apically flat and membranous figs 4 11 one half length of cereal segments simple in female figs 7 14 calileuctra NYMPH mature nymph weakly sclero- tized body elongate se tation scarce harper and stewart 1984 adult key abdominal segments I1 VII with membranous 585658 56 in hind wing culgulguicu not forked calileuctra pleural fold mesosternal Y ridge with double 58 in hind wing culcuicu forked 58a stem arms meeting furcal pits at posterior 58a 58 in hindwinghindwing m cu joining culcuicu beyond fork ends barapparaproctsparaproetsroots fused basally with no visible of culcui 59 suture sparse setation cerealcercalgereal segments each ssa in hindwinghindwing mmeucu joining culcuicu before fork with apical fringe of 10 15 setae setae approx- of cuiculguigul 60 imately one half length of cercalcereal segments distribution napa valley and los stewart and stark 1988 nymphal key angeles basin CA 5 pronotum with no long marginal setae fig DIAGNOSDIAGNOSISIS Mmalesmaiesalesaies are best characterized 814a8 14a paraprocts of both sexes fused by their elongate flat basally with no distinct medial line of unique topped cerci separation 5aaa females are characterized by sternum villVIII being incomplete and the lack of a posteriorly fronoPronopronotumturn with 2 4 long hairs on anterior projecting subgenital plate nymphs are char- and posterior margins fig 82a8 2aaa 812a8 12a paraprocts of both sexes fused acterized by abdominal segments I1 VII hav- with distinct medial line of separation ing a membranous pleural fold the subanal or slightly separated medially fig lobes having basal fusion but no distinct fusion 8 ahi2hi821118.2111 8 12hi812h1 6 line and the cerealcercal segment setae being one half length of the cerealcercal segment 5aaa abdominal segments 1 6 divided by ventro- lateral membrane ENA and SW ETYMOLOGY prefix was select- cali 1 the I zealeuctra ed to denote california the origin of the spec- imens the suffix leuctraleuctra was selected to indi- 5aaa abdominal segments 1 7 divided by ventro- cate placement of the genus in the family lateral membrane WNA calileuctra gender the is leuctridae of name neuter key to adults of calileuctra KEY modification modifications are with given for the following identification keys for la wings macropterous male epiepiproctproct bifurcate dorsally tergite IX posteriorlyposte noily emarginate with nearctic leuctrid genera harper and stewart two large heavily sclerotized teeth cercicerel with api- 1984 nymphal and adult keys stewart and cal tooth tergite X with one tooth on each postero- stark 1988 nymphal key wording style lateral corner sternite IX with posteriorpostenor projection and figure citations are as presented in the broadly rounded female sternite VII broad with a posteriorly projecting rectangular lobe sternite original keys VIII membranous C dobryidobayi

harper and stewart 1984 nymphal key ib wings brachypterous male with epiepiproctproct with one dorsal hook tergite IX with single sclerotized pos- length 383738 37 body robust less than 8 times width terior plate bearing numerous teeth cerci without body conspicuously clothed with hairs apical tooth tergite X with two teeth on each pos- about one fifth the length of middle ab terolateralterolateral corner sternite IX with posterior pro- segment subanal lobes of mature male a jection broadly angulate female sternite VII elon- fused strongly keeled plate much pro- gate with lateral constrictions sternite VIII with duced with no posterior notch fig two arcuate sclerotized plates C ephemera 134413 44 megaleuctra calileuctra ephemera species 38 body more elongate fine hair pile inconincon- new spicuous appearing naked subanal lobes figs 1 9 of mature male fused one half to two MALE general color brown dark brown thirds length leaving a notch at tip 38a pattern as illustrated fig 1 length of body 38a 38 subanal lobes fused but with complete 454.5 mm brachypterous length of forewing suture apical setae on cerealcercal segments 252.5 303.0 mm wings light brown venation simi- usually less than one half length of cer- lar to the genus perlomyiaPerlomyia fig 3 Prothprothoracicoracie ealcal segments Perloperlomyia myia basisternumbasisternum triangular in shape fig 2 38a subanal lobes basally fused no suture inm abdominal tergum IX with posterior border basal half apical setae on cercalcereal segments complete projecting and serrate tergum X 126 GREAT BASIN naturalist volume 55

N

X

fig 1 cahleuctracalileuctra ephemera habitus

incomplete medially posteroposterolaterallylaterally with two membranous and sclerotized expanded near elongate projections figs 4 5 sternum IX apex figs 8 9 with basal pear shaped vesicle posterior bor- FEMALE general color and wing vena- der extending to base of subanal probe fig tion similar to male brachypterous length of 6 cerci extending beyond genitalia with api- forewing 353.5 404.0 mm sternum vilVII constrict- cal membranous area expanded and flattened ed laterally projecting slightly over sternum figs 4 5 6 epiproctEpiproct small and hook shaped VIIIv111 sternum villviliVIII reduced to 2 small arcuate fig 5 subanal probe large elongate both sclerotized plates fig 7 199511995 calmeucmcalileuctra NEW GENUS 127

3

figs 2 3 calileuctracaliteuctra ephemera 2 ventral view of thorax 3 wings

NYMPH body lightly sclerotized light in a posteriorly directed U shape mesosternum color setation sparse except on labrum legs narrowest anteriorly widest by coxaecomae mem- and cerci size small 727.2 minmm long head branous except for weak sclerotization of the slightly broader than thorax color pattern faint furcal pits and the Y ridge Y ridge with faintly mouthpartsMouthparts of the herbivorousdetritivorous sclerotized double stem arms connecting to type type I1 stewart and stark 1988 labrum posterior ends of furcal pits transverse ridge and clypeus with numerous long setae man- connecting anterior ends of furcal pits dible typical for leuctridae 4 dorsal cusps 2 metasternum similar to mesosternum scleromclero only transversely rectangular large 2 small and I1 small ventral cusp on tizationtization in a area side of first large dorsal cusp bowl shaped limited by the furcal pits a transverse ridge the ends of the furcal molar region with transverse ridges in the connecting anterior pits and the area between the pits and the ridge bowl and with a pectinate scraping ridge on all legs similar but increasing in size posteri- the ventromedial edge maxillary palpi 5 seg orly setation consists of abundant very small palpi 3 segmented glossae dentedmented labial flossae setae clothing hairs and sparse longer paraglossaeparaglossate and subequal size and short in setae tibiae and femora with setal fringes paraglossaeparaglossate slightly larger pronotum quad- apex of tibiae with a pair of spines tarsi 3 seg ranrangulargular transverse anterior and posterior dentedmented first segment short and conical sec- sclerotized bands median longitudinal suture ond very short ringlike with apex cleft third unsclerotized color pattern weak mesonotum elongate and cylindrical suture between first with two scleritisscleritessclerites anterior sclerite transverse and second tarsomerestarsomeres very narrow and hard and roughly trapezoidal posterior sclerite to see tarsomerestarsomeres with ventral pad of numer- roughly U shaped metanotum like mesono- ous fine setae tarsal claws slender abdominal tum wing pads three or more times as long as terga very weakly sclerotized setation sparse wide posterior wing pads a little shorter than except on end of tenth segment abdominal anterior wing pads longitudinal axes of wing fold present on segments I1 VII subanal lobes pads almost parallel to axis of body ProsteProsterprostermimprostemummimmum incompletely fused cercalcereal segments with api- naked and membranous except for two small cal fringe of 10 15 setae setae about one half scleritisscleritessclerites between the coxaecomae scleritisscleritessclerites forming length of the segments 128 GREAT BASIN naturalist volume 55

2

y

R

el

A

17 7

51

figs 4 7 calileuctra ephemera 4 male terminalia dorsal view 5 male terminalia lateral view 6 male terminalia ventral view 7 female terminalia ventral view 199511995 calimucmcalileuctra NEW GENUS 129

this intermittent stream has water present mam1 only a few months each year some years there is no wwaterater ie 1987 and 1990 when water is present it flows down a small steep canyon across a grassy flat under hwybwy 128 and down a short cliff into capell creek the stream course appears to be less than 350 in 1000 ft long all specimens have been collected in the grassy flat or just downstream the stream course has a substrate of either bedrock or rocks on a clay soil there is no obvious hyporheichyporheic zone detrital input is usually leaves from trees mainly live oak grass and star thistle TYPE SPECIMENS HOLOTYPE male type locality 19 II11 1983 WDS A 160 reared from nymph to be deposited in the entomology collection at the california academy of sciences san francisco CA ALLOTYPE female type locality 25 II11 1984 WDS A 240 to be deposited with the holotype PARATYPES 1 male type locality 18 II11 1984 WDS A 234 reared from nymph deposited at monte L bean life science museum brigham young university provo UT 3 females type locality 18 II11 1984 WDS A 234 deposited with male paratype additional specimen I1 nymph typetype locality 27 111988II11 1988 WDS A 527 deposit- ed with the holotype and allotype ETYMOLOGY the trivial name was select- ed to indicate the temporary nature of the stream at the type locality and also to indicate the difficulty encountered when trying to col- lect specimens BIOLOGY all specimens were collected during the last two weeks of february when the stream was flowing streams in this area of the coastal mountains experience a mediter- ranean climate with a december to february rainy season local intermittent streams usually have surface flow only from january through march first all figs 8 9 calileuctra ephemera 8 male subanal probe when collected specimens were right lateral view 9 male subanal probe ventral view either late instar nymphs 3 or adults 4 all field collected adults 4 females were swept from vegetation overhanging the stream two EGG shape oval size uniform 01330.1330 133 mm of the three nymphs collected were held in in length 00950.0950 095ogs mm in width surface coarsely styrofoam containers until they boltedmolted to the rugose large coarse punctures present inm an adult stage both individuals were males irregular distribution the bowl shaped molar region of the man- TYPE LOCALITY california napa co dibles is similar to molar modifications found 3363.363 36 kinkm gl91212.12 1 mi N on hwybwy 128 from the inter- in beetle larvae that feed on fungal tissues in section of hwybwy 128 and hwybwy 121 unnamed general and fungal spores in particular tributary to capell creek ca 300 in 275 ft lawrence 1977 lawrence and hlavac 1979 elevation lawrence and newton 1980 since fungal 130 GREAT BASIN naturalist volume 55 tissues are high in protein martin 1987 use headwater tributary of elsmere creek it has of them as a food would aid the fast growth an extremely steep gradient and is hard to and development of nymphs access thus the habitat has been preserved present information suggests that cakleuccalileuc more than the surrounding drainage area tra ephemera has a facultatively long egg dia- TYPE SPECIMENS HOLOTYPE male type pause very fast nymphal development and locality 22 IV 1991 K F dobry to be short stadium for both nymphs and adults deposited in the entomology collection at the the high protein content of fungal tissues california academy of sciences san francisco martin 1987 may aid in the fast growth and CA ALLOTYPE female same data as holotype development of nymphs of this unique species to be deposited with holotype PARATYPES I1 these characteristics are similar to those of male and 1 female same data as holotype 2 zealeuctra snellen and stewart 1979 an males and I1 female CA los angeles co eastern north american genus and another santa monica mountains east fork arroyo inhabitant of intermittent streams sequitdequit 5 mi NW pacific coast highway off mulholland highway 28 II11199211 1992 K E dobry calileuctracahleuctra dobryidobayi new species 2 females CA orange co santa ana moun- figs 10 14 tains trabucotrabucco canyon 1300 111198811 I1 1988 R W B MALE general color brown dark pattern baumann B J sargent C kondratieff and go C R 3 males and I1 female CA as illustrated fig 10 length of body 404.04 0 606.06 0 nelson los san gabriel canyon 23 IV 1960 mm macropterous length forewingofofforewing 454.54 5 555.55ss 5 angeles co gibbo mm wings light brown venation similar to the D LACM remaining paraparatypestypes to be genus perlomyiaperlomyzaPerlomytamyza tergum IX with membra- deposited at monte L bean life science myiamyla provo nous median band dividing tergum into two museum brigham young university sclerotized halves each half bearing a small UT nipplenippienipplelikempplehkelikeilke projection and a large earlike pos- ETYMOLOGY the trivial name honors terior projection tergum X also divided into keith FE dobry los angeles CA who collect- two halves each half with a gently rounded ed many of the specimens knoblike lobe and an enlarged lateral posteri- BIOLOGY specimens were collected as or lobe which ends in a sclerotized prong that adults between january and april all popula- extends about one third the way up the cercus tions are from the los angeles basin and figs 11 12 sternum IX broadly rounded experience a mediterranean climate posteriorly extending only to base of subanal probe large vesicle present at median anterior PHYLOGENY malginmargin vesicle with truncate apex fig 13 cerci enlarged elongate extending beyond leuctrid phylogeny has been examined genitalia posteriorly sclerotized on lateral from a cladisticcladistic point of view only two times margins apex roundedlounded ending in a sclerotized the two studies ricker and ross 1969 nelson lateral prong figs 11 12 13 epiproctEpiproct with and hanson 1973 are somewhat contradictory narrow bifurcate apex fig 11 subanal probe however examination of the analysis given in large elongate broadest medially apex pointed both studies shows calileuctra to possess many fig 13 character states that are termed primitive or FEMALE general color and wing vena- ancestral following nelson and hanson s tion similar to male length of body 505.0so5 0 606.06go 0 more comprehensive analysis the character mm macropterous length forewingofforewingof 505.05 0 606.06go 0 states present in calileuctra are as follows 101 0 mm abdominal sternum VII enlarged 202 0 303 0 404 0 505 0 606 0 717 1 808 0 909 0 10210 2 11011 0 expanded slightly over VIII posteromedial 12012 0 13013 0 14214 2 15015 0 16016 0 17017 0 18019018 019 0 area formed into a narrow medially rounded 20020 0 21121 1 22022 0 23023 0 24124 1 25125 1 26126 1 27127 1 lobe sternum VIII small and only lightly scle- 28128 1 29029 0 30130 1 and 31231 2 first number rotized fig 14 character second number character state NYMPH unknown see nelson and hanson 1973 for a key to the EGG unknown characters and character states character TYPE LOCALITY california los angeles states for calileuctra and those cited in nelson co south fork elsmere canyon san gabriel and hanson 1973 for other leuctrid genera mountains the type locality is a very small were run through the PAUP 31131.1 program 199519951 calileuctra NEW GENUS 131

AV

31

fig 10 cahleuctracalileuctra dobryidobayi habitus 132 GREAT BASIN naturalist volume 55

6

A 0

12

j 6 il ff

13 14

figs 11 14 calileuctra dobryidobayi 11 male terminalia dorsal view 12 male termterminaliamaliamallamaila lateral view 13 male tennterminaliamaliamallamaila ventral view 15 female terminalia ventral view 199519951 calileucaucalileuctra NEW GENUS 133

ancestor moseliajoselia leuctra pachyleuctra LC r despaxiadespaniaDespaxia paraleuctra zealeuctra rhopalopsole Perloperlomyiamyia calileuctra tyrrhenoleuctra megaleuctra

15

fig 15 phylogeny of the leuctridae using the branch and bound algorithm with all ningrung phylogeny programs for us and for his characters unordered this analysis found one many helpful comments C riley nelson uni- minimum length tree fig 15 with a length of versity of texas austin provided a review and 68 a consistency index of 0820.82 and a retention made valuable suggestions boris C kondratieff index of 0800.80 colorado state university also offered many this new tree is not considerably different helpful suggestions as well as helped collect from that given by nelson and hanson 1973 specimens the late charles L hogue kindly it differs only in the collapse of the sister loaned specimens from the los angeles county group relationship between rhopalopsole and museum LACM keith F dobry helped zealeuctra and the exclusion of euleuctra and greatly in the collection of additional speci- leuctraLeuctra divisadavisa from consideration the sta- mens jean A stanger made the many excel- bility of this tree with calileuctra added is taken lent illustrations as evidence of the consistency of this data set and the overall stability of this new classifica- literature CITED tion it is heartening to find the cladogram of nelson and hanson 1973 stable despite the HARPER P P AND K W STEWART 1984 chapter 13 previous extinctiorextinction absence of calileuc plecoptera in R W merritt and K W cummins ie editors an introduction to the aquatic insects of tra in this particular case an extinct taxon did north america kendallhuntKendall Hunt publishing co not particularly influence the overall topology dubuque 10 722 appp of the dogramcladogramcla hence there is hope in our LAWRENCE J F 1977 the family pterogeniidaepterogenndae with search for relationships among living taxa notes on the phylogeny of the heteromera the bulletin 31 25 26 despite known extinction events coleoptenstscoleopterists LAWRENCE J F AND T F HLAVAC 1979 review of the in this tree calileuctra is a the sister taxon derodontidae coleoptera polyphaga with new of the group containing ferlomyiaperlomyiaPerloFerlomylamyia rhopalop species from north america and chile the sole zealeuctra paraleuctra despaxiadespaniaDespaxia pachy coleoptenstscoleopterists bulletin 33 369 414 leuctraleuctra leuctraLeuctra and moseliajoseliaMoselia in leuctrid phy- LAWRENCE J F AND A FE NEWTON 1980 coleoptera associated with the fruiting bodies of slime molds logeny a basal calileuctra occupies near posi- myxomycetes the coleoptenstscoleopterists bulletin 34 tion and as such gives an important addition to 129 143 our knowledge of the group MARTIN M M 1987 invertebrate microbial interactions cornell university press ithaca NY 148 appp acknowledgments NELSON C H AND J F HANSON 1973 the genus per lodyialomylalomyia plecoptera leuctndaeleuctridae journal of the kansas entomological society 46 187 199 many thanks go to charles H nelson uni- RICKER W E AND H H ross 1969 the genus versity of tennessee at chattanooga for run zealeuctra and its position in the family leuctndaeleuctridae 134 GREAT BASIN naturalist volume 55

plecoptera leuctndaeleuctridae canadian journal of zoology STEWART K W AND B P STARK 1988 nymphs of north 47111347 1113 1127 american stonstoneflyefly genera plecoptera thomas say SNELLENSNCLLEN R K AND K W STEWART 1979 the life cycle foundation entomological society of america 12 and drumming behavior of zealeuctra claasseniclaassensclaas seni 1 460 prisonfrison and zealeuctra hiteihidei ricker and ross plecoptera leuctndaeleuctridae in texas USA aquatic received 2277 september 1994 insects 1651 65 89 accepted 17 january17january 1995 great basin naturalist 552 C 1995 appp 135 141

CARBON ISOTOPE discrimination IN THE c4ca4 SHRUB ATRIPLEX confertifoliaCONFERTI FOLIA ALONG A SALINITY GRADIENT

darren R SandquSandquististl1 and james R Ehlerehleringerlehlennger1ingerlingeri

ABSTRACT carbon isotope discrimination A was measured for leaves ofatriplex conferticonfertifohaconfertifoliafolia along a salinity gradi- ent in northern utah over this gradient the variation of A values was high for a c4ca4 species and the A values were posi- tively correlated with salinity in both years of the study of the possible explanations for this pattern the A results are consistent with the notion that salinity induces an increase in the bundle sheath leakinessleakmess of these c4ca4 plants

key words carbon isotope ratio salt stress bundle sheath leakiness halophyte desert ecology

the analysis of carbon isotope ratios and carbon isotope discrimination in c4ca4 plants 13c12CCQ has become a useful tool for under- as standing various integrated aspects of plant 1 metabolism including numerous investiga- A a bab44 b300 acicaaccicaa clcyalcy tions of plantpiant environment interactions the impact of environmental factors on carbon iso- where a 0404404404.40 is discrimination against the tope discrimination A by plants with c3ca3 pho- heavier 13cocog2 molecule relative to the lighter totosynthesis has been well studied however 12co2gogcog based on differential rates of diffusion only a limited number of studies have exam- bab3&3 22029029o is the discrimination due to a ined variation of A values in c4ca4 plants greater affinity for 12co2CO relative to 13co2CO by oleary 1988 farquhar et al 1989 peisker riburibuloserivuloselose phosphatebisphosphatebis carboxylase lubiscorubiscoRubisco and henderson 1992 in part this disparity and bab4b typically 5205.20522 is discrimination stems from c4ca4 plants having much smaller based on the steps leading to and including A variation of values than c3ca3 plants c02CO fixation by phosphoenolphospho enol pyruvate car- additionally A values in c3ca3 plants have been boxylaseboxy lase pepoPEPQPEPC after atmospheric 602coggog correlated with water use efficiency and this enters the leaf the bab4b term varies slightly as a has lead to an emphasis on applying carbon function of temperature and is negative isotope analyses to breeding programs greater proportion of 13co2CO due to fractiona- farquhar et al 1989 ehleringer et al 1993 tion associated with the hydration of c02CO to however a few recent studies have demon- hc03 mook et al 1974 the discrimination strated that variation of A values in c4ca4 plants terms of equation I1 a bab3b and bab4b are con- may reflect environmental influences on phys- stants for the most part and thus differences iological function bowman et al 1989 among A values are the result of changes in 0 meinzer et al 1994 in this study we exam- andor cicacc during c02CO assimilation ined variation of A values in a c4ca4 perennial in c4ca4 plants cog is initially fixed by PEPC shrub atriplex confertifoliaconfertifolia torr & frem in the mesophyll cells transported and decar wats and its relationship to natural condi- boxylatedboxylated in the bundle sheath cells and then tions ofsoilof soilsoli salinity refixedprefixed by lubiscorubiscoRubisco however before the assimi- the A value of a c4ca4 plant integrates two lation by rubiscolubisco a fraction of the c02CO may factors that can impact productivity 1 the diffuse out through apoapoplasticplastic portions of the ratio of intercellular to ambient coo concen- bundle sheath cells this is known as leaki- tration cilcilcacalcaccca which can reduce photosynthe- ness and is thought to be reduced by suber- tic activity when low and 2 bundle sheath ization of bundle sheath surfaces farquhar leakiness 0 which reduces photosynthetic 1983 this leakiness however may be in- efficiency when high farquhar 1983 mod- creased by environmental stresses such as eled the relationship between these factors salinity bowman et al 1989 and an increase

department of biology university of Uutahtahtab saltlakesaltsait lake city ut84112UT 84112

135 136 GREAT BASIN naturalist volume 55

in leakiness represents an energetic cost to the bury mountain range the four sites range in plant as a result of incomplete carbon assimi- elevation from 1366 in to 1286 in fig 1 site I1 lation or overovercyclingcycling ehleringer and pearcy 1366 in is dominated by sagebrush artemisia 1983 jenkins et al 1989 henderson et al tridentata with low densities of atriplex con 1992 fertifoliafertifolia juniperus osteospennaosteosperma and tetrady leakiness affects A because it causes the mia spinosa weedy grasses and annual species bundle sheath cell to become an open system of the chenopodiaceae are also found within and therefore allows expression of discrimina- disturbed areas of this and all other sites tion by lubiscorubisco bab3b the proportion of cog greasewood sarcobatus vermiculatus is the that leaks out of the bundle sheath cell 0 dominant species at sites 2 1317 in and 3 modifies the degree to which bab3b is expressed 1294 in with A confertifoliaconfeilifoliaconferticonjertifolia co occurring in and thereby determines the relationship low frequency site 4 1286 in along the mar- between A and cicacc eq 1 at low 0 values gins of the salt flats is a heterogeneous site the relationship between A and cicacc is nega- with a mixed community of salt tolerant tive at high 0 the relationship is positive and species S vermiculatus is the dominant at 0 0320.32 A is constant at 44c44o444440 regardless of species with moderate densities of allenrofeaallenrolfea cica equation I1 also predicts that for any occidentoccidentalisoccidentalistalis atriplex gardnerigardneri A confertifoliaconfertifolia given ciccica an increase in 0 results in an chrysothamnus viscidiflorusviscidiflorus kochia ameri- increase in A given these relationships varia- cana and suaeda tortorreyanareyana tion of A values in c4ca4 plants can provide an weather data for this transect are taken indication of bundle sheath leakiness and its from the grantsvillegrantsville weather station grants relationship to environmental stresses ville thoeletooele county UT 1307 in located 17317.3 to date much work investigating variation km E and 828.2 km S from the center of our of A in c4ca4 plants has come from either labora- study transect evans al tory gasexchangegas exchange studies et 1986 leaf and soil samples bowman et al 1989 henderson et al 1992 or theoretical models peisker 1982 farquhar leaves of atriplex confertifoliaconfertifolia and soil 1983 peisker and henderson 1992 there is samples were collected from each of the four little direct information on environmental transect sites in october 1991 and 1992 with stresses that influence A in natural popula- the help of the 1991 and 1992 plant ecology classes from the university of utah recently tions of c4ca4 plants except see walker and sinclair 1992 here we report on changes in matured leaves ofaofA confertifoliaconfertifolia were collect- ed from five to eight individuals per site in A values for the c4ca4 species atriplex confertifo lia found along a natural salinity gradient in 1991 and three per site in 1992 leaf samples utah the purpose of this study was to deter- were oven dried 70c70 C 7 d ground with mine if A values changed in relation to soil mortar and pestle and analyzed for carbon salinity under field conditions and if these isotopic composition windy ike delta S mass changes corresponded to variation in 0 values ratio spectrometer finnigan MAT san jose pee two previous laboratory studies have shown CA relative to the dee belemnite stan- that higher soil salinity does increase A values dard analyses were done at the stable isotope facility for environmental in c4ca4 plants and that this change is a result of ratio greater 0 bowman et al 1989 meinzer et al research SIRFER university of utah salt 1994 for A confertifoliaconfertifolia we hypothesized lake city UT carbon isotope ratio values 8 that the same trend would be found over a were transformed to discrimination A values transect of naturally increasing soil salinity as A aa8a 8 2 METHODS 5plp5p1 p 8 is the study sites where p measured carbon isotope ratio of the plintpiantplant and 5aaa8 is the carbon isotope ratio four study sites of increasing salinity were of cobCO in the atmosphere 008oos008.008 or 80 chosen along a south to north transect in the farquhar et al 1989 the standard per mil northern end of skull valley thoeletooele county 0 o notation is used throughout for ease of UT flanking the western slope of the stans presentation and the overall longtermlong term error 19951 CARBON ISOTOPE discrimination IN ATRIPLEX 137

1400

site 1 1375 E C 135- 00 16 site 2 1325 mu sltesslte3site 3 sltesslte4site 4 1300 salt flat

1275 j 0 5 10 15 20 25 30 35 40 transect distance from site 1 km

fig 1 study transect in cross section shown is the topography over the transect and localities of each study site based on the approximate linear distance from site 1 associated with carbon isotope determination ratio extracts are highly correlated with soil isis011o011o0110 paste ECs for soils within and near our tran- soil samples were collected from two depths sect D G williams unpublished data 15 20 cm and 40 60 cm in two to six excava- electrical conductivity is reported in zohoszmhosjumhos 1 tion pits at each site approximately 200 g of cm 1 I1 chosmhosumhos cm 1 010.1oi ms m 1 05020.502 freshly extracted soil from each hole and depth mm nacl and the data were log transformed was placed immediately into soil canisters for statistical analyses Interinterannualannual compar- sealed and kept cool until analysis in the labo- isons of means for each soil trait were done by ratory in the lab one subsample per canister t tests and correlations between soil trait and was removed for salinity analyses the remain- plant carbon isotope discrimination means ing soil was used for gravimetric water content were determined by pearson product moment determination based on the difference between correlation soil fresh wet weight and dry weight ie water content relative to the soil dry weight RESULTS soils were dried at 70c70 C for 7 d in 1991 the soil salinity analysis was based transect characterization on electrical conductivity EQEC of an aqueous salinity increased across the gradient in solution extracted from a soildeionized 12 both the 1991 and 1992 samples electrical water and 1992 from a soil mixture in 15 conductivity increased by two orders of mag- deionized water was no evi- mixture there evievl nitude over the entire transect table 1 site I1 dence that the 12 mixture was ion saturated was the least saline and salinity progressively thus to standardize these ratios the ECs of increased toward the highly saline site 4 samples using a 12 solution were extrapolated there were few differences between years to EC based on a 15 ratio assuming a linear in soil electrical conductivity significant dif- dilution relationship tests confirmed that this ferencesferences were found at only two sites and at extrapolation was valid even for EC values only one depth per site furthermore sites higher than those found in actual field samples gave opposite results soils of site 3 at the although a more standard procedure for 15 20 cm depth had greater conductivity in salinity determination is the soil paste 1991 than 1992 t 4334.33 P oi0101.01 and soils method the 15 ratio method we used is rec- from site I1 at the 40 60 cm depth had higher ommendedommended as a simpler technique to deter- conductivity in 1992 than in 1991 t 4604.60 FP mine relative salinity contents rhoades 1982 oi0101.01 and is suitable for the purposes of this study gravimetric water content also increased ie standardized comparison of relative salin over the transect from site I1 to site 4 table 1 itiesaties among sites additionally the ECs of 15 soil water content was somewhat greater in 138 GREAT BASIN naturalist volume 55

TABLE 1 soil properties at two depths for sites 1 4 along the study transect n number of pits one sample for each depth per pit soil water content was measured as gravimetric water content and electrical conductivity is of an aque- ous extract from 151 5 stilwatersoilwatersoil water mixture extrapolated for 1991 from 121 2 ratio see text electrical electrical soil water soil water conductivity conductivity content content amhoscmmhoscmmhoscmsem mhoscmmhoumhoscmscmsem 152015 20 cm 406040 60 cm 152015 20 cm 406040 60 cm mean SE n mean SE n mean SE n mean SE n

OCTOBEROCTOBLR 1991 site 1 466 0300 4 534 0234 4 89 157 4 70 32 4 site 2 415 0687 4 723 0360 4 91 76 4 324 811 4 site 3 1179 1446 4 1724 0892 4 2309 1141 4 2066 6577 4 site 4 2484 7578 6 3941 7841 6 3596 5876 6 3382 5307 6 OCTOBERoc IOBLK 1992 sitelaitelsite 1 289 0454 2 381 0402 3 84 63 3 93 37 3 site 2 479 0226 2 556 0499 3 144 292 3 324 1110 3 site 3 246 0270 2 1026 3672 3 546 4593 3 984 8585 3 site 4 1066 0950 2 NA 1640 14400 2 3250 3500 2

1991 than in 1992 but significant differences discussion at both depths were found only at site 1 15 20 cm depth t 3343.34 P 05os05.05 40 60 cm variation in carbon isotope discrimination depth t 3523.52 P 05os05.05 rainfall over the 10 values of c4ca4 plants is inm part dependent upon wk period prior to sampling in 1991 was much the proportion of 602CO that is initially fixed by ultimately diffuses of greater than that of 1992 82582582.5 minmm vs 18818.8 PEPC and out the bun- mm which likely accounts for the trend of dle sheath cells without being refixedprefixed iei e the greater water content in the soils during the leakinessleakmess 0 leakiness might be influenced 1991 sample period by environmental stresses such as salinity bowman et al 1989 meinzer et al 1994 carbon isotope discrimination because such stresses could disrupt mem- along the transect the carbon isotope dis- brane properties or the biochemical coordina- cycles crimination for atriplex confertifoliaconfertifolia ranged tion between the c4ca4 and c3ca3 operating from a low of 4744.74 0960 at site I1 in 1992 in the mesophyll and bundle sheath cells 096o peisker to a high of 6556.55 011c011oolloolio0110 at site 3 in 1991 respectively and henderson 1992 the other component influencing variation of fig 2 this range of nearly 20 is high for c4ca4 A claca 3 illustrates plants farquhar et al 1989 the mean A in c4ca4 plants is cic figure how value was always greater than 44o444440 and for the relationship between A and cdcclaca depends only a single sample was the individual shrub upon the value of 0 from eq 1 and provides value less than 444440 these high A values a model for how changes in 0 and cocclaca can 44o A indicate that the mean 0 values were always account for the changes in values we greater than 0320.32 eq 1 observed with respect to the environmental parame- we found that A values of A confetticonferticonfertifoliaconfertzfoliafolianoila ters examined along the transect mean leaf A increased by 2o20ao inm concordance with increas- was not significantly correlated with water ing salinity fig 2 these A values were content during any observation but was pos- always greater than 44o4444404 4o40ao therefore the 0 val- itively correlated with log EC fig 2 ues must be greater than 0320.320 32 cf fig 3 A inclusion of the notably low A value of site 4 2c2o20ac increase in A values at 0 0320.320 32 cannot in 1991 resulted in a nonsignificant positive be explained solely by changes in ccclaca given trend but when excluded A was significantly the typical range of clacacc values for c4ca4 plants correlated with log EC in 1991 at the deeper under ambient conditions 0200200.200 20 0400.400 40 pearcy soil depth R 101.0iolo P oi0101.01 in 1992 there and Ehlerehleringermger 1984 to do so would require was a highly significant positive correlation of either extreme leakinessleakmess values 0 06 or an A and log EC for both the shallow soils R increase of cicaclaca with increasing salinity since 978978.978 P 05os05.05 and deeper soil depths R A and cilcacalcacc are positively related when 0 999999.999 P ooi001001.001 fig 2 0320.320 32 leakinessLeakmess values greater than 060.60og 6 have 199511995 CARBON ISOTOPE discrimination IN ATRIPLEX 139

go 909.0 1 soil depth 15 20 cm 0 0 6 z 65 T 80 005 656.5 T 808.0 h ti0 i I1 70 I1 L 0 0 1i 0 040.4 60 555.5 TT 0 0 505.0 0ook00101 0 0 4.040 40 0 03 303.0 A 0 202.0 35 a 353.5 0 02 soil depth 40 60 cm 10 4060 0 01 65 1 I 656.5 T 4 0 X 1 0 T ilhl 0 02 04 06 08 1 1L L I1 0 1 T cica 555.5 h 04CH L fig 3 model for the relationship between carbon isoiso- tope discrimination A and cica ratio of intercellular to 0 ambient 602coggog based on equation 1 and for 0 values rang- 454.5 ing from 010 1 to 060.60og 6 dashed and solid lines represent the range of A values for each 0 value depicted based on a 488480 high leaf temperature 34c34oc where bab44 48o4 80 solid line and a lower leaf temperature gsg25c25 C where bab4b 5705 70 dashed line 353.5 10 100 1000 10000 log soil electrical conductivity gihosumhosgmhos cm 1 tion could be due simply to the high degree of edaphic variability at site 4 this location had fig 2 relationship between carbon isotope discrimina- the greatest topographic variability highest tion A of atriplex conferticonfertifoliaconfertzfohafolianoila leaves and log electrical species diversity and greatest overall variance conductivity log EQEC of soil at two depths 15 20 cm and for soil conductivity and water content table 40 60 cm for sites 1 4 along the transect closed symbols 1 site 4 was also extremely wet in 1991 near 0 are study site means for 1991 and open symbols 0 are those for 1992 error bars are lseISE 40 water content at 40 60 cm depth which may have diluted the salinity of these soils thereby reducing the salinity experienced by never been reported and the latter explana- the plants without a more detailed study how- tion is unlikely since salt stress typically ever this deviation remains unexplained decreases or does not change cicacc long and previous studies have found contrasting baker 1986 flanagan and Jefjefferesjeffenesjefferiesfenes 1988 A patterns of the relationship between A and simpler explanation for the change in A values salinity in a laboratory study with 11 c4ca4 is that 0 increases with higher salinity A 2o20ao2 species henderson et al 1992 found that 0 increase based on changes in 0 values can be values were invariable and low remaining at 0 0.21 thereby resulting in a negative rela- easily accommodated within the limits of cccca 021 tiontionshipship between A and fig 3 the found for c4ca4 plants fig 3 thus changes in cicacc small variation they observed in A values was A values for A conferticonfertifohaconfertifoliafolianolia are more likely attributed to changes in cc values however due to an increase of 0 associated with a cica in an earlier study with the c4ca4 monocotsmono cots zea change in salinity consequently the presence mays and andropogon glomeratusglomeratus bowman et of a significant relationship between A values al 1989 found that A values of salt stressed and EC fig 2 plants were more dramatically influenced by the trend of increasing A values with changes in elcacicacc than were control plants the increasing salinity held in all but one site in increase of A values with salinity was ex- the two year study site 4 in 1991 this devia plained by a changing relationship between A 140 GREAT BASIN naturalist volume 55

and cilcacalca due to increasing 0 values as the ability among previous studies of carbon iso- waterastatus of salt stressed plants declined tope discrimination in atriplex conferticonfertifoliafolia through the day bowman et al 1989 mean A values range from 44o444440 marino et al recently meinzer et al 1994 also observed 1992 to 6906.90 troughton et al 1974 yet that increasing salinity resulted in increases of each of these observations is consistent with A values using two sugarcanesugarcane cultcultivarsivars they the notion that 0 values exceed 0320.32 and are showed that change in A value could be therefore high compared to halophyticnonnonhalophytic c4ca4 ascribed to greatelgreater 0 values as salinity in-in species henderson et al 1992 creased and that variability of cicaaa had much in the present study we have shown that less impact on the increase of A values in salinity may be one factor that significantly contrast walker and sinclair 1992 reported influences variation of A values in c4ca4 plants that A values of two australian c4ca4 atriplex most likely through an effect on bundle sheath species decreased at sites with increased salin- leakiness while variation in A values of c4ca4 ity the A values of these australian atnplexatriplex plants may provide new insights into plant leaves were greater than 44o4444404 4o40ao which could salinity dynamics along environmental gradi- have been achieved only with a bundle sheath ents results also suggest that caution is neces- leakage greater than 0320.320 32 fig 3 since the sary when using A values of c4ca4 plants to inter- relationship between A and aacica is positive at pret historical changes in atmospheric cog 0 0320.320 32 fig 3 the walker and sinclair data concentrations and 13cC values as has been suggest that salinity affected a decrease oficofcicaofcc proposed by marino et al 1992 and therefore a decrease of A our findings of a positive correlation be- acknowledgments tween A values of atnplexatriplex conferticonfertifohaconfertifoliafolia and salinity are in contrast to findings of walker we thank university of utah students in and sinclair 1992 our observations like 1991 and 1992 plant ecology classes for assis- those of bowman et al 1989 and meinzer et al tance in sample collection craig cook for 1994 suggest that changes in leaf carbon iso-iso assistance in carbon isotope analyses and dr tope discrimination result from an increased david williams for salinity analyses compar- bundle sheath leakage when plants are exposed isons dr williams and two anonymous to a salinity stress the mechanism of change reviewers also provided helpful comments on in 0 values is likely to be associated with phys- a previous version of this manuscript ical changes in the bundle sheath permeability to cog to biochemical or hc03 andor literature CITED changes in the coupling of lubiscorubisco and PEPC activity such biochemical changes due to BOWMAN W D K T HUBICK S VON CAEMMERER AND salinity have been previously found guy and G D FARQUHAR 1989 short term changes in leaf reid 1986 have shown that salinity may carbon isotope discrimination in salt and water stressed ca grasses plant physiology 90 162 166 reduce lubiscorubisco activity in co plants without a c44 in c3ca ehleringer J AND R W PEARCY 1983 variation in in concomitant decrease in PEPC activity quantum yield for 602COgog uptake among c3ca3 and c4ca4 increased salinity naci has also been shown plants plant physiology 73 555 559 ehleringer J R A E HALL AND G D FARQUHAR to increase PEPC activity in some c4ca4 halo phytespaytes shomer ilan et al 1985 any such 1993 stable isotopes and plant carbon water rela- tions academic press san diego CA 555 appp in of carboxylation increase in the activities c4ca4 EVANS J R T D SHARKEY J A BERRY AND G D enzymes relative to those of c3ca3 carboxylation FARQUHAR 1986 carbon isotope discrimination enzymes in c4ca4 plants should increase 0 values measured concurrently with gas exchange to investi- peisker and henderson 1992 thus under gate 602coggog diffusion in leaves of higher plants natural conditions it appears that salinity australian journal of plant physiology 13 281 292 FARQUHAR G D 1983 on the nature of carbon isotope could increase A values of A confertifoliaconfertifolia by discrimination in c4ca4 species australian journal of influencing an increase in 0 values plant physiology 10 205 226 the relationship between salt stress and 0 FARQUHAR G D J R ehleringer AND K T HUBICK 1989 of c4ca4 plants may be species specific or even carbon isotope discrimination and photosyn- population specific and may account for dis- thesis annual review of plant physiology and molecular biology 40 503 crepancies among different studies of A values 537 FLANAGAN L B AND R L JEFFERIES 1988 stomatal in c4ca4 plants for example there is high varivarlvan limitation of photosynthesis and reduced growth of 199519951 CARBON ISOTOPE discrimination IN ATRIPLEX 141

the halophyte plantago dantimamaritinamantimamaritimamarimaylmayitima L at high salinity OTEARYLEARY M H 1988 carbon isotopes in photosynthesis plant cell and environment 11 239 245 biosciencebioscience3838 325 336 GUY R D AND D M REID 1986 photosynthesis and the PEARCYdarcySARCY R W AND J ehleringer 1984 comparative plant influence of coiCO enrichment on SC valuesvalnesvaines in a c3ca3 physiologyecoecophysiology of c3ca3 and c4ca4 plants cell and halophyte plant cell and environment 9 65 72 environment 7 1 13 HENDERSON S A S VON CAEMMERER AND G D PEISKERolskerSISKER M 1982 the effect of 602coggog leakage from bundle FARQUHAR 1992 short term measurements of car- sheath cells on carbon isotope discrimination in c4ca4 bon isotope discrimination in several c4ca4 species plants Photosyntheticphotosyntheticaa 16 533 541 australian journal of plant physiology 19 263 285 PEISKERdisker M AND S A HENDERSON 1992 carbon terres- JENKINS C L D R T FURBANKFURRANK AND M D HATCH trial c4ca4 plants plant cell and environment 15 1989 mechanism of c4ca4 photosynthesis A model 987 1004 describing the inorganic carbon pool in bundle RHOADESdoadesHOADES J D 1982 soluble salts pages 167 179 in sheath cells plant physiology 91 1372 1381 methods of soil analysis part 2 chemical and micro- LONG S P AND N R BAKER 1986 saline terrestrial biological properties ASASSSAASA SSSA madison WI environments pages 63 102 in N R baker and S P SHOMERlomerIOMERIOMERILANllanILAN A D MOUALEM BENO AND Y WAISEL long editors photosynthesis in contrasting enviedvienvi- 1985 effects of nacl on the properties of phospho ronronmentsments elsevier scientific publishers new york enolenolpyruvatepyruvate carboxylase from suaeda monnicamonoica and NY chionschloris gayana physiologiapbysiologiaphysiologic Planplantarumplantariumtarum 65 72 78 MARINO B D M B MCELROY R J SALAWITCH AND TROUGHTONKUGHTON J H P V WELLS AND H A MOONEY 1974 W G SPAULDING 1992 glacial to interglacial variavarlavaria- photosynthetic mechanisms and paleoecology from tions in the carbon isotopic composition of atmos- carbon isotope ratios in ancient specimens of c4ca4 and pheric cog nature 357 461 466 CAM plants science 185 610 612 MEINZER FE C Z PLAUT AND N Z SALIENDRA 1994 WALKERalgerALKER C D AND R SINCLAIR 1992 soil salinity is cor- carbon isotope discrimination gas exchange and related with a decline in 13cC discrimination in leaves growth of sugarcanesugarcane cultcultivarsivars under salinity plant of Atriplex species australian journal of ecology 17 physiology 104 521 526 83 88 MOOK W G J C BOMMERSON AND W H STAVERMAN 1974 carbon isotope fractionation between dis- received 20 may 1994 solved bicarbonate and gaseous carbon dioxide accepted 16 august 1994 earth and planetary science letters 22 169 176 cleatcreatgleatgreat basin naturalist 552 C 1995 appp 142 150

demography OF astragalus SCAPHOIDscaphoidesES AND EFFECTS OF HERBIVORY ON population GROWTH

peter lesicallesica1vesical

ABSTRACT losses in fecundity due to predispersal herbivoreherbberbherbivoryberbivoryivory can be large howeverhoweveivel the effects of this loss on long- term population viability have rarely been investigated I1 conducted a demographic study of astragalus scaphoides fabaceae a long lived perennial endemic to east central idaho and adjacent montana by following mapped individu- als at two sites from 1986 to 1993 astragalus scaphoidscaphoideses suffers losses of dispersalpredispersalpre fecundity averaging nearly 50 fromblom insect seed piepiedationpredationdation and inflorescence predation by insects and livestock cattle reduced fecundity by 0 85 nonetheless estimates from matrix projection models indicate that both sample populations had positive growth in most years elasticity analyses revealed that population growth occurred in spite of relatively small contributions by recruitment compared to growth and survival of nonreproductive plants results suggest that populations of this long lived perennial depend little on reproduction and recruitment for growth and can persist in association with seasonal i otationrotationdotation livestock grazing

key words demography herbivoreherbherbivoryivory livestock grazing predation matrix projection models elasticity analysis astragalus rare plant

the importance of herbivoreherbherbivoryivory in determining predation particularly by exotic species plant population dynamics and composition of has often been cited as a threat to endangered vegetation has long been debated ehrlichehrilch and plant populations greig smith and sagar 1981 birch 1967 slobodkinSlobod km et al 1967 belsky parsons and browne 1982 willoughby 1987 1986 A great deal of evidence suggests a neg- norton 1991 pavlik et al 1993 negative im- ative impact of herbivoreherbherbivoryivory on the host plant pacts of herbherbivoresivores were shown but a causal harper 1977 crawley 1983 dirzoderzo 1984 link to declining population size has rarely been however researchers have recently presented demonstrated evidence for positive interactions mcnaughton astragalus scaphoidscaphoideses jones redbrydb is 1986 paige and whitham 1987 endemic to a small area of east central idaho A plant s life history plays an important role and adjacent montana barneby 1964 it was formerly a candidate for listing in determining the effects of herbivoreherbivoryherbivory loss of as a threatened reproductive output from seed predators can or endangered species by the US fish and be disastrous for an annual or biennial but wildlife service category 3cac USDI FWS 1993 and is currently listed as sensitive in may have little effect on a long lived perenni- idaho moseley and groves 1990 and montana al furthermore effects of herbivoreherbherbivory will ivory lesica and shelly 1991 most populations of depend on the age or stage ege g seeds adults A scaphoidscaphoideses occur on public lands subject to at which it occurs 1984 most studies derzodirzo livestock grazing high levels of inflorescence have focused on the effects of herbivoresherbivores on and seed predation have been observed in particular components of fitness over relatively some populations lesica and elliott 1987a short spans is time this is unfortunate because here I1 report the results of an eight year it is the longtermlong term effect on population growth demographic study ofA scaphoidscaphoideses at two sites that determines the importance of herbivoreherbherbivoryivory the purpose of the study is to document levels to population viability few studies have inte- of herbivoreherbherbivoryivory and to assess its importance to grated the effects of herbivoreherbherbivoryivory on population population growth using stage based transi- dynamics and growth harper 1977 but see tion matrix models and elasticity analysis de louda 1982 1983 kroon et al 1986 caswell 1989

division of biological science universityuniverunivei sity of montana missoula MT 59812 and conservationconser vation biology research 929 locust missoula MT 59802

142 199519951 demography AND HERBIVORY IN astragalus 143

METHODS fied as weweevilsweevilyevils small beetles in the family curculionidae weevil larvae feed on maturing species studied seeds and leave the developing or mature astragalus scaphoidscaphoideses is a caulescent peren- legume by creating a small hole in the outer nial with a taproot surmounted by a branched wall seed predation by weevil larvae was caudex reproductive individuals are 20 50 inferred from the presence of fecula andor an cm high with a cluster of pinnately compound exit hole in the legume basal leaves and 3 10 leaves at intervals along the erect stem the inflorescence is composed study sites of 1 4 racelesracemes arising from the aeilsaxils of the the sheep corral gulch population occurs upper leaves each raceme is composed of a in southern beaverhead county MT on a gen- naked peduncle 5 15 cm long surmounted tle south facing slope at 1920 inm tst8s r12w by a tight cluster of 10 30 flowers that sig mean july and january temperatures at expands in fruit nonreproductive individuals dillon 32 kinkm NW and 275 in lower are 19019.0igo generally have 1 4 basal leaves and may have and 666.6gg C respectively mean annual precipi- a sterile stem less than 15 cm tall with 1 5 tation is 241 mm vegetation is dominated by leaves the branching caueaucaudicesdices of reproduc- artemisia tridentata and agropyron spicatumspicatum tive plants may bear up to four stems and aster scopulorum and phlox hoodiihoodei are com- more than a dozen racelesracemes barneby 1964 mon forbs livestock were managed on a rest unpublished lesica data rotation system by which grazing occurred in flowers astragalus scaphoidscaphoideses generally dur- different seasons in most consecutive years ing the first three weeks of june the most evidence of heavy spring grazing by livestock conspicuous form of herbivoreherbivoryherbivory of these plants was observed in 1989 1990 and 1993 is the removal of inflorescences during flower- the haynes creek population is in central principal ing inflorescence predation has two leahilemhi county ID approximately 48 km W of and livestock subfamily sources insects ants sheep corral gulch it occurs on a moderate formicinae and moth larvae melacosomamalacosoma sppapp southeast facing slope at 1555 in j19nt19n r23er28e family lasiocampidae were observed remov- s2sa mean july and january temperatures at ing inflorescences at a site near haynes creek salmon 24 km NW and 365 in lower are in idaho Pedpedunclespeduncleduncles below the flowers were 16216.2 and 676.7 C respectively mean annual girdled and withered inflorescences were precipitation is 252 mm vegetation is domi- often still of the present near the base plant nated by artemisia dentatatritridentatatridentate agropyron spica by livestock also inflorescence predation tum and bromus tectoriumtectorumtectorum this site was not occurred but differed from insect predation in grazed by livestock before early july during that pedpedunclespeduncleduncles were all removed at the same the course of the study height and severed inflorescences were not found below the plants in eithereltherleither case the field methods cluster of basal leaves was usually left intact it was possible to assign primary responsibility two permanent monitoring transects were for inflorescence herbivoreherbherbivoryivory at a site in a partic- established at each of the study sites in early ular year to either insects or ungulaungulatesungulatedtes based july 1986 following methods outlined in lesica on the appearance of damaged plants and the 1987 transects were located subjectively to presence or absence of droppings hoof prints represent the populations and were read in or trampled vegetation however it was not early july because fruits were mature or near- possible to unambiguously assign each case of ly so but seed dispersal had not yet begun at herbivoreherbherbivoryivory to one or the other source inflor- each site the transects were parallel to each escence predation by insects was observed at other and the slope each transect consisted of both study sites in all years that inflorescences 50 adjacent im2ima1 m2ma mapping quadratsquadquadransrats placed were produced but ungulate predation was along the transect line the position of each A common only at sheep corral gulch scaphoidscaphoideses plant encountered in the quadratsquadquadransrats dispersalpredispersalPre seed predation occurred at was mapped and classified for three traits 1 both sites in most years larvae were collected size 2 inflorescence production and 3 from developing legumes in 1gsg9861986 and identididenti fecundity using the following classification 144 GREAT BASIN naturalist volume 55

1 size classes the pods counted intact seeds and recorded D dormant no aboveground parts evidence of insect predation observed analysis S small nonreproductives data 131 3 leaves stage structured transition matrix projec- L large nonreproductives tion models summarize the way in which sur- 4 leaves vival growth and reproduction at various life R reproductive history stages interact to determine population 2 inflorescence production growth van groenendael et al 1988 caswell A inflorescence produced no fruit 1989 matrix projections assume fixed transi- P inflorescence was removed due tion probabilities between stages in a popula- to predation tion through time lefkovitch 1965 menges I1 inflorescence produced at least 1990 they also assume density independent one mature fruit population growth and thus do not give an 3 fecundity total number of mature accurate projection of longtermlong term population fruits future nonetheless they can be used to sum- when stems weiewelewere removed below the point of marize short term population dynamics inflorescence articulation I1 made a conserva- caswell 1989 oneyearone year transition probabili- tive estimate of the number of inflorescences ties were estimated as the number of plants in removed based on the size of the remaining life stage class i moving into classyclasselass j over the plant evidence of livestock and native anguungu course of one year divided by the number of lates ege g droppings hoof prints trampled plants in stage i at the beginning of the year vegetation was noted along each transect and this method assumes that an individual s tran- for the site as a whole sition depends only on its life stage class at I1 found that some plants would go unde- the beginning of the period and is indepen- tected for one to several years but reappear in dent of its transition the previous year the subsequent years lesica and steele 1994 equilibrium growth rate Xi is the dominant these dormant plants may have produced eigenvalue of the transition matrix lefkovitch small leaves that had senessenescedced and disap- 1965 caswell 1989 Xi 101.0loio indicates popula- peared by early july however my observa- tion increase while X 101.0io indicates tions in may and june suggest that most of decrease X integrates the effects of survival them produced no vegetation on the years in growth and fecundity of the different life his question the presence of dormant plants can tory stages into a single parameter there are be inferred by comparing transect maps from two ways in which a reproductive plant can the full sequence of years the proportion of undergo a transition 1 the plant itself moves dormant plants ranged from 1 to 23 with a into a different class or stays the same and 2 mean of 10 in 1987 1991 plants have dis- the plant produces progeny in one or more appeared for as many as five years before re- classes these two probabilities recruit reprocepro appearing however in 1986 1992 at the two are presented separately in the matrices but sites 71 of dormant plants reappeared after must be added together to solve for XL details one year and 88 reappeared after two years on the construction and use of matrix popula- lesica and steele 1994 As a result ca 10 tion models can be found in caswell 1989 of the plants were undetected inm the first and and menges 1990 X was calculated using last years of the study and ca 3 were unde- ramasstageRAMAS stage ferson 1991 tected in the second and second from last elasticity measures the relative change in years thus I1 have chosen to eliminate the the value of X in response to changes in the first and last years 1986 1993 of the study value of a transition matrix element elasticity from demographic analysis recognizing that a matrices allow comparison of the relative con- small ca 3 error still remains in mortality tributions of various life history transitions to and recruitment estimates in 1987 and 1992 population growth and fitness de kroon et al on years when fruit production was ade- 1986 elasticities sum to unity and regions of quate I1 collected 50 randomly selected the matrix may be summed to compare the im- mature fruits from at least 25 plants I1 opened portance of growth and survival to recruitment 199519951 demography AND HERBIVORY IN astragalus 145

caswell 1989 elasticities for nonreproductive predation plants are sums from the small S and large inflorescence predation attributable to unguangu L classes elasticities were calculated using lates was virtually absent from the haynes ramasstageRAMAS stage person 1991 ferson creek population droppings and hoof prints the of seeds pass directly when majority of cattle were the only signs of ungulatesungulatedtes at from production to germination in less than ungula sheep corral gulch droppings occurred in one year seeds should not appear as a sepa- 3 9 of the mapping quadratsquadquadransrats during the rate stage in matrix models caswell 1989 study inflorescence predation by Silvertown al 1993 of insects silvertown et seeds astragalus occurred at both sites in all years scaphoideses germinate readily without stratifica- scaphoid A significant number of inflorescences were tion and elliott 1987b that lesica suggesting produced in six of eight years at haynes creek most seeds germinate the same year they are and inflorescence predation accounted for produced nonetheless A scaphoides form scaphoides may fecundity losses of 14 50 over the course of a seed bank not including a seed bank in the the study fig 3 most of this herbivoreherbivoryherbivory was matrix model may affect the value of X kalisz attributable to insect damage at sheep corral and mcpeek 1992 especially when it is io10101.0 gulch reproductive plants were common in however it will have little effect on analyses only four of eight years inflorescence predation based elasticities Silver al 1 on silvertowntown et 1993 I resulted in fecundity losses of 19 90 and calculated separate elasticities for reproduc- the proportion of inflorescences lost to preda- tive transitions and recruitment by dividing tion was highest in 1989 1990 and 1993 the reproductive recruitment elasticities years in which predation was due mainly to proportionately between the two components livestock fig 3 losses to predation were estimated from seed predation occurred at both sites in the number of inflorescences lost using the nearly every year in which significant fruiting calculated means for seedsfruitseeds fruit and fruits occurred fig 3 overall loss of seeds to wee- inflorescence cumulative fecundity losses vil predation ranged from 0 to 33 with a were calculated by multiplying the propor- mean of 18 insect seed predation was gen- tions of inflorescences and seeds remaining erally higher at sheep corral gulch than at after predation and subtracting from one haynes creek fig 3 losses of fecundity due to the combined RESULTS effects of inflorescence and seed predation were 19 90 in 1986 1993 with means of population growth the number of astragalus scaphoidscaphoideses plants in the transects at both sites increased 250 by about one third between 1986 and 1993 A fig 1 equilibrium population growth rate 225 0 sheep corrolcorralcorrotrr 21 was loio101.0 at both sites over the course of v hoyneshaynes the study and was 25252.5 at sheep corral gulch 200 in 1988 89 and 1990 91 at no time during 175 the study was kX 080.8 at either site table 1 e survivorship 150

between 40 and 50 of the astragalus E 125 scaphoidscaphoideses plants observed at the start of the V study in 1986 were still alive in 1993 fig 2 loo100 approximately 50 of the 1989 cohort the first large cohort recruited during the study 75 survived for more than 3 4 years taken 50 together these results indicate that A 87 88 89 90 91 92 scaphoidscaphoideses is a long lived perennial with ca year 50 mortality occurring in the first 3 4 years but a large proportion of plants living to be fig 1 density of astragalusofastragalus scaphoidscaphoideses plants in the 1010yearsyears two sample populations 1987 1992 146 GREAT BASIN naturalist volume 55

tableTABLL 1 stage based transition matrices for astragalus scaphoidscaphoideses at two sites in 1987 1992 four stages are recog- nized dormant D small nonreproductive S large nonreproductive L and reproductive R the reproductive and recruitmenti re columns must be added together before solving for X the dominant eigenvalue see methods

sheep corral gulchi 1987 88 1990 91 fromprom from to D S L R re to D S L R re D 67 18 20 0 0 D 14 06 0 0 0 S 11 55 24 0 0 S 21 23 06 0 986 L 22 06 36 0 0 L 50 42 26 29 242 R 0 0 03 10 0 R 14 12 57 10 14 X 1181 18 A 2692 69

1988 89 1991 92 from from to D S L R re to D S L R re D 23 04 02 0 0 D 70 24 21 25 0 S 17 27 05 0 40 S 30 27 37 33 20 L 43 53 45 0 70 L 0 0 14 22 0 R 17 08 43 100 25 R 0 0 0 0 0 k 2512 51 X 0830 83

1989 90 from to D S L R re D 80 17 14 06 0 S 10 73 37 22 91 L 10 02 35 56 03 R 0 02 01 16 0 Xi097097 haynes creek 1987 88 1990 91 flomfrom from to D S L R re to D S L R re D 50 03 04 0 0 D 21 03 0 0 0 S 40 45 04 0 5 S 21 34 03 0 95 L 0 24 37 0 16 L 50 31 21 18 27 R 10 06 52 60 20 R 07 19 66 64 0 k 1881.881 88 1311 31

1988 89 1991 92 from from to D S L R re to D S L R re D 57 13 04 0 0 D 75 03 06 05 0 S 14 42 24 05 110 S 0 44 30 15 30 L 14 16 28 14 14 L 25 08 36 53 03 R 14 04 32 67 05 R 0 0 11 20 0 tX 1131 13 X 0830 83 1989 90 flomfromprom to D S L R re D 64 12 05 0 0 S 27 38 14 04 23 L 0 20 48 27 04 R 09 0 33 58 0 X1097vogtl0970970.97 199511995 demography AND HERBIVORY IN astragalus 147

loo100 sheep corralcorrol gulch 10 90 inflorescence 172g ESSems seed 2 08 80 HI combined 0 70 V 060 6

60 040 4 0 1 95 a U CL 137 x L 50 136 a 020 2 x B 0 9 sheep corralcorrol m x 40 Mn 1 x v haynes 00oo fl 86 89 91 93 30 yearyeanyeon 20 86 87 88 89 90 91 92 93 year haynes creek 10 fig 2 depletion curves for the 1986 sample popula- p7pa inflorescence tions of astragalusofastragalus scaphoidscaphoideses at the two study sites 0 08 seed 21 combined 6 51 and 44 at and sheep Z 06 haynes creek 108 corral gulch respectively fig 3 80 0 04 elasticity analysis 70 138 elasticity gives the proportional impor- 02 130 tance of demographic transitions to population five 00 growth elasticity matrices for years of 86 88 89 90 91 93 transitions for the two study sites are given in year table 2 elasticities were summed into four life history transition categories 1 recruit- fig 3 proportion of astragalusofastragalus scaphoidscaphoideses reproduc- ment and survival and growth of 2 dormant tive output lost to inflorescence predation dispersalpieprepredispersalpiedispersal 3 nonreproductive and 4 reproductive seed predation and the combination of the two in those years when significant flowering occurred two study plants fig 4 growth and survival of conrenonre at sites numbers of inflorescences in samples aieare given productproductivesproductizesives was consistently important at both above bars sites with mean elasticities of 42 and 36 at haynes creek and sheep corral gulch respectively survival of dormant plants was accounting for an average of less than 17 of important in two years at sheep corral gulch population growth at both sites high levels of and one year at haynes creek with mean inflorescence and seed herbivoreherbherbivoryivory are undoubt- elasticities of 19 and 29 survival of reprocepro edly one of the main reasons for the low con- ducductivestives had mean elasticities of 23 and 20 tributiontribution of recruitment to X in this species for the two sites and mean elasticities for nonetheless both sample populations became recruitment were 16 and 17 larger during the study furthermore popula- tion growth rate was S 101.0iolo in four of five years discussion at both sites and never 08 growth and sur- vival of dormant and nonreproductive plants loss of astragalus scaphoidscaphoideses fecundity contributed 60 to population growth at due to inflorescence and dispersalprepredispersal seed both sites these results suggest that popula- predation was high at both sites ranging from tions of A scaphoidscaphoideses can persist and even ca 20 to 90 further losses in reproduc- grow larger in spite of heavy losses in repro- tive output due to ants or rodents may have ductive output and low recruitment occurred following dispersal recruitment was large reductions in fecundity due to cerbiherbi the least important stage transition in the life vores have been documented for astragalus history of A scaphoidscaphoideses during my study species green and Palmpalmbaldbald 1975 as well as 148 GREAT BASIN naturalist volume 55

TABLE 2 elasticities for astragalus scaphoidscaphoideses stage transition matrices at two sites for 1987 1992 the left three columns D S L represent nonreproductive growth and survival the reproductive R column represents growth and survival of reproductives the recruitment column re represents recruitment from seed sheep corralcorraicorralaral gulch 1987 88 1990 91 D S L R re D S L R re D 048 022 015 0 0 D 001 005 0 0 0 S 004 032 009 0 024 S 001 018 002 0 185 L 032 015 055 0 077 L 002 082 023 014 118 R 0 0 101 568 0 R 003 099 215 204 029

1988 89 1991 92 D S L R re D S L R re D 001 003 001 0 0 D 686 126 0 0 0 S 001 017 002 0 139 S 126 061 0 0 0 L 002 081 049 0 143 L 0 0 0 0 0 R 003 057 222 224 056 R 0 0 0 0 0

1989 90 D S L R re D 413 074 015 001 0 S 048 292 037 002 011 L 042 007 030 005 001 R 0 017 002 004 0

haynesis creek 1987 88 1990 91 D S L R re D S L R re D 002 003 002 0 0 D 001 005 0 0 0 S 001 045 002 0 141 S 001 058 004 0 160 L 0 082 058 0 153 L 004 080 038 046 068 R 003 059 231 164 055 R 001 079 193 264 0

1988 89 1991 92 D S L R re D S L R re D 055 048 006 0 0 D 617 020 041 006 0 S 007 086 020 005 113 S 0 042 030 003 005 L 015 064 044 028 028 L 068 018 082 021 001 R 031 034 109 287 021 R 0 0 036 011 0

1989 90 D S L R re D 081 028 013 0 0 S 023 061 024 007 039 L 0 065 170 093 014 R 018 0 134 230 0

many other plants janzen 1971 hendrix 1988 environments jordan and nobel 1979 signi- louda 1989 louda 1982 1983 has shown ficant reductions in a single reproductive bout that seed predation can lead to lowered could greatly increase chances of population recruitment however reductions in seed out- extirpation for these sorts of species on the put will not necessarily lead to reduced other hand many populations of long lived recruitment if germination safe sites are limit- plants will have more stable populations ing harper 1977 analysis of the matrix pro- whose persistence is more dependent on the jection models suggests that recruitment is not growth and survival of established plants limiting population growth ofaofA scaphoidscaphoideses silvertownSilvertown et al 1993 survivorship curves recruitment from seed is likely to be imim- indicate that astragalus scaphoidscaphoideses is a long portant to population growth for short lived lived species and elasticity analysis suggests species and is essential for semelparoussemelparous ones that recruitment is indeed less important to furthermore successful reproductive episodes population persistence than growth and sur- are rare for some perennial species in rigorous vival of nonreproductive plants 199519951 demography AND HERBIVORY IN astragalus 149

sheep corral gulch dation upper portions of plants are most 10 dormantormant accessible to livestock and newer growth is I1 non reproductive generally selected by livestock arnold and 080 8 reqL reproductive dudzinski 1978 valentine 1990 furthermore JPSPSJ recruitment sugars such as found in flower nectar also 2 060 6 increase palatability arnold and dudzinski 1978 valentine 1990 thus livestock often ej 04 remove only the upper portions of broad Xx x leaved plants predation that mainly affects Xx X X X 020 2 N fecundity is likely to endanger populations i X X sxaxxi X 1 only when grazing removes most inflores 00 1 L L J j bencescences consistently for many years because 88 89 90 91 92 population growth is not likely to be limited year by recruitment on the other hand grazing that lowers growth and survival eg high density stocking during periods of growth will haynes creek have a much more detrimental effect on popu- 10 1 F I1 M dormant lation viability UI1 non reproductive 08 eegeggLN reproductive lssj recruitment acknowledgments 060 6 6 I1 am grateful to joe elliott anne garde 0 r and lou hagener for help in the field james ui 040 4 liebherr of the comstock museum ithaca s x NY and will lanier of the entomology 020 2 X X research lab bozeman MT identified x insects kimball harper and an anonymous alxl s 00ooi001 il ii 0 88 89 90 91 92 reviewer gave helpful comments on the manu- year script funding was provided by the idaho and montana bureau of land management fig 4 elasticities summed into four life history transi- and the montana natural heritage program tion categories recruitment and survival and growth of dormant nonreproductive and reproductive plants for astragalus scaphoidscaphoideses at two study sites 1987 1992 literature CITED

ARNOLD G W AND M L DUDZINSKI 1978 ethology of inflorescence predation of astragalus scaaca free ranging domestic animals elsevier amsterdam 198 appp phoidesphotideshoidesboides livestock P was greatest in years when BELSKY A J 1986 does herbivoreherbherbivoryivory benefit plants A were present in 1993 inflorescence predation review of the evidence american naturalist 127 was greater than 85 and A scaphoidscaphoideses was 870 892 grazed in preference to the highly palatable BARNEBY R C 1964 atlas of north american astragalus 1 york grass agropyron spicatumspicatum PE lesica personal parts I and 2 memoirs of the new botanical gardenl3gardenly13 1 1188 observations that garden observation these suggest CASWELL H 1989 matrix population models sinauer livestock could nearly eliminate reproductive associates sunderland MA 328 appp output under high stocking rates and repeated CRAWLEY M J 1983 herbivoreherbivoryHerbivory the dynamics of animal heavy spring grazing if carried on over a long plant interactions university of california press enough period of time however results of my berkeley 437 appp DIRZO R 1984 Herbherbivory a phytocentricphytocentnc study suggest that A scaphoideses populations herbivoreivory viewpoint scaphoid pages 141 165 inm R dirzoderzo and J sarukiansarukhanSarukhan editors can persist if predation is moderate at least in perspectives on plant population ecology sinauer some years rotation grazing systems in which and associates sunderlandSunderlandeilandelland MA spring grazing occurs only one in three years EHRLICH P R AND L C BIRCH 1967 the balance of appear to be compatible with the longtermlong term nature and population control american naturalist persistence of A scaphoideses populations loi10197101 97 108 ofa scaphoid FERSONPERSON S 1991 ramasstageRAMAS stage generalized stage based these results have implications for other modeling for population dynamics applied bio- long lived perennials exposed to livestock pre mathematics setauket NY 150 GREAT BASIN naturalist volume 55

GREENGRLLN T W AND I1 G PALMBALD 1975 effects of insect 1983 seed predation and seedling mortality in seed pipredatorseclitoi s on astragalus cibacibanuscibariuscibarioustiusrius and astragalus the recruitment of a shrub haplopappus renetusvenetus utahensis legummosaeleguminosacleguminosae ecology 56 1435 1440 asteraceae along a climatic gradient ecology 62 cricrlgriigGRIgraig K SMHIJANDGsmrrj J AND G R SAGAR 1981 biological causes 511 521 of rarity in carlina vulgaris pages 389 399 in H 1989 predation in the dynamics of seed regener- synge editor biological aspects of rare plant conser- ation pages 25 51 in M A leekleck V T parker and vation john wiley and sons chichester england R L simpson editors ecology of soil seed banks VAN grolnlndaigrolnendai LI1 J M 11 DLDE khoonKKOONKROON AND H CASWELL academic press new york NY 1988 projectionpi ejection matrices in population biology mcnaughton S J 1986 on plants and herbherbivoresivores trends in ecology and evolution 3 264 269 american naturalist 128 765 770 HARPERHARPLR J L 1977 population biology of plants aca- MENCESMENGES E S 1990 population viability analysis for an demic pipressess london 892 appp endangered plant conservation biology 4 52 62 HENDRIXHLNDRIX S D 1988 herbivoreheibivoiyherbivoryHerbivory and its impact on plant MOSELEY R AND C GROVES 1990 rare threatened and reproductioni epi oductionobduction pages 246 263 in J lovett doust and endangered plants and animals of idaho idaho L lovett doust editorsediedltoismols plant leproductivereproductive ecology natural heritage program boise 33 appp oxford university press new york NY NORTON D A 1991 tnlepideatrilepidea adamsiiadamski an obituary for a janenJANLNJANZEN D H 1971 seed predation annual review of species conservation biology 5 52 57 ecology and systematics 2 465 492 PAIGE K N AND T G WHITHAM 1987 overcompensation JORDAN P W AND P S NOBEL 1979 infrequent estab- in response to mammalian herbivoreherbivoryherbivory the advantage lishmentlishment of seedlings of agave desertideberti agavaceae in of being eaten american naturalist 129 407 416 the northwestern sonoran desert americanamerlean journal PARSONS R FE AND J H BROWNE 1982 causes of plant of botany 66 1079 1084 species rarity in semisemiaridarid southern australia bio- KALISZ S AND M A MCPEEKM pekkPEEK 1992 demography of an logical conservation 24 183 192 age stiuctuiedstructured annual lere sampled projection matri- PAVLIK B M N FERGUSON AND M NELSON 1993 ces elasticity analyses and seed bank effects assessing limitations on the growth of endangered ecology 73 1082 1093 plant populations II11 seed production and seed bank DL KROON H A plaiserPLAISLRPLAISFR J M VAN croengroenendaelGROEN ENDALL AND dynamics of erysimum capitatum sspasp angustatumangustatum H CASWLICASWELL L 1986 elasticity the relative contribu- and oenothera deltoidesdelt oides sspasp howelhowellehowelliihowellnhowellisliililiii biological tion of demographic parameters to population conservation 65 267 278 giowthiategrowth rate ecology 67 1427 14314311 sieverskeverSILVERsllvertownsilvertownTOWN J M FRANCO I1 PlplsantyPISANTYSANTY AND A MENDOZA llli11 FKOVITCH1 KO Vircil L P 1965 the study of population growth 1993 comparative plant demography relative in organi01 organisinsgamsmssins grouped by stage biometrics 21 1 18 importance of life cycle components to the finite llli11 sacas1casl A P 1987 A technique foiforboibol monitoringmonitoiing nonrhizomanomhizoma rate of increase in woody and herbaceous perenni- tous perennial plant species in permanent belt tran- als journal of ecology 81 465 476 sects natural areas journal 7 65 68 SLOBODKIN L B F E SMITH AND N G HAIRSTON 1967 lisicall11 shaSKA P AND J C ellionELLIOTT 1987a distribution age regulation in terrestrial ecosystems and the implied structure and predationpiedation of bittenbitterrootoot milkvetchmilkvetch pop- balance of nature american naturalist 10loi1011 109 124 ulationsulations in lemhilcmhilemdi county idaho report submitted USDI FISH AND WILDLIFE SERVICE 1993 endangered to the bureau of land management boise ID and threatened wildlife and plants review of plant 1987b 1987 monitoring study of bitterroot milk taxa for listing as endangered or threatened species vetch populations in leahilemhi county idaho report notice of revienreview federal register 58 511445119051144 51190 submitted to the bureau of land management VALENTINE J FE 1990 grazing management academic boise ID press san diego CA 533 appp leshLLSKLESICA A P AND J S SHELLY 1991 sensitive threatened and willoughby J W 1987 effects of livestock grazing on endangered vascularvascu lailalial plants of montana montana two rare plant species in the red hills tuolumne natural heritage program occasional publication 1 county california pages 199 208 in T S elias editor helena 88 appp conservation and management of rare and endan- lesuLLSKLESICA A P AND B M SILLLLSTEELE 1994 prolonged dormancy gered plants california native plant society in vascuvascularlailalial plants and implications for monitoring sacramento studies natural areas journal 14 209 212 LOUDA S M 1982 distribution ecology variation in received I1 april 1994 plant recruitment over a gigiadientgradientadient in i elationrelation to accepted 7 septemberSeptembei 1994 insect seed predation ecological monographs 52 25 41 great basin naturalist 552 0 1995 appp 151 157 LAHONTAN SAGEBRUSH ARTEMISIA ARBUSCULA SSPSSE LONGIloncicaulislongicaulisCAULIS A NEW TAXON

alma H Winwarwinwardlwmwaicandedl and E durant McArthurmcarthur22

ABSTRACT A new subspecies of artemisia arbuscula is described A arbuscula sspasp longicaulislongicaulis winward & mcarthur sspasp nov this taxon is a landscape dominant in portionspoitportionstons of northwestern nevada and adjacent california and oregon at elevations from 1050 to 2000 rn on shallow or argillic clayey soils it differs from A arbuscula sspasp arbuscula in its long floral stalks and large leaves morphological chemical ecological and cytological data suggest that it is of hybrid origin it is hexaploid ax6x we hypothesize that 2xax A arbuscula sspasp arbuscula and 4xax A tntridentata sspasp byominwyominwyomm gensis are its parents

key words nevada taxonomy chemotaxonomychemotaxonomy allopolyploid hybrid tndentataetridentataeTridentatae

in preliminary reports we winward et al sagebrush A nova was best treated as a sub- 1986 1991 provided a brief description of a species of A arbuscula but beetle 1960 new taxon of artemisia found in northwestern restored it to neinelsonneisonNelsorssots s 1900 original species nevada and adjacent california and oregon status beetle 1960 recognized two sub- we suggested referencing it by the common species of A arbuscula arbuscula and ther name lahontan sagebrush pending a formal copolamopola his treatment has been generally description this paper provides that formal accepted winward and tisdale 1977 mcarthur description and details concerning its taxono- et al 1981 shultz 1986 although winward my distribution general ecology and origin 1980 has observed an unusual variant of A arbuscula in eastern oregon that reaches a TAXONOMY height of I1 in he suggested that further taxo- nomic treatment of A arbuscula would be new taxon is a member of subgenus the appropriate tridentataeTridentatae of artemisia the true sagebrushessagebrushersagebrushes during the past few decades fieldworkers beetle 1960 mcarthur et al 1981 shultz in western nevada have observed a sagebrush 1986 we suggested winward et al 1986 that in that does not fit the existing artemisia taxo- this taxon may have originated as a hybrid with nomic keys 1972 termed this sage- parental lines consisting of low and big sage- brunner brush wide lobe with the comment brush A arbuscula and A dentatatridentatatridentatdtridentatedtri because dr feels this ofA of its general morphology and ecology we sug- beetle may be an ecotype triden gested that it be considered a subspecies of A tata sspasp wyomingensis I1 concur others have arbuscula further studies indicate that this new referred to it as wonder sagebrush junk taxon is in fact best treated as a subspecies of sagebrush or N sagebrush winward et al A arbuscula 1986 accessions of two populations trough the type specimen of A arbuscula came springs humboldt county NV cultures ul from a collection along the andaridarld plains of the and uss and leonard creek humboldt lewis now known as the snake river nuttall county NV culture u55 of this taxon were 1841 subsequent workers have submerged the established in common gardens of the forest species as a subspecies of big sagebrush A service s shrubland biology and restoration tridentata sspasp arbuscula hall and clements research work unit at several locations 1923 or in contrast recognized a number of around central utah there they were treated races and subspecies within the species ward as an ecotype ofA tritrldentata sspasp wyomingensis 1953 beetle 1960 ward proposed that black following beetle and brunner brunner 1972

range and watershed management intermountaininteimountam region forest serviceSe ivice USU S department of agriculture ogden UT 84401 hibibhiiib sciences laboratory intermountain research station forestfoiest service USU S department of agriculture provo UT 84606

151 152 GREAT BASIN naturalist volume 55 eg mcarthur and plummer 1978 welch and 0 nevada pershing co 646.4 km west of mcarthur 1979 1981 1986 mcarthur et al toulon mcarthur & mcarthur 1683 1981 1985 mcarthur and welch 1982 welch SSLP two sheets et al 1986 1987 the new taxon is a landscape nevada washoe co mustang dominant over much of its range winward et al mcarthur & mcarthur 1684 SSLP 1986 and both domestic and wild animals 0 oregon lake co 32 kinkm east ofadellocadellof adelladeliadeil feed extensively on it brunner 1972 welch and sanderson & mcarthur 1590 SSLP mcarthur 1986 winward et al 1986 welch 0 oregon malteurmalheur co near mcdermitt et al 1987 nevada winward sn 31 october 1986 OGDF two sheets SSLP description artemisia arbuscula sspasp longicaulislongicaulis win- distribution AND ECOLOGY ward & mcarthur sspasp nov similis A arbuscula artemisia arbuscula sspasp longicaulislongicaulis occurs sspasp arbuscula sed ramis floralibusfloralibus bultomulto lon on several hundred thousand hectares in gioribusibus fohisfodisfoliis magnioribus differt similar gior et northwestern nevada and in adjacent areas of to A arbuscula sspasp arbuscula except flower california and oregon at elevations from stalks are much longer and leaves are larger about 1050 to 2000 in fig 1 it often occurs the longer flower stalks and larger leaves in pure stands it may also share dominance also differentiate sspasp longicaulislongicaulis from sspasp with other sagebrush taxa such as big sage- thennopolathenthedthermopolathermonopolapola which differs from sspasp arbuscula brush A dentatatritridentatatridentate sspasp dentatatritridentatatridentate and wyo and longicaulislongicaulis by having deeply trifid leaves minmingensisgensis low sagebrush A arbuscula sspasp beetle 1959 arbuscula and black sagebrush A nova at we chose the common name lahontan sage- lower elevations it is interspersed with salt brush because the old shorelines of pleisto- desert shrub species such as shadscaleshad scale cene lake lahontan are one of the centers of atriplex confertifoliaconfertifolia bailey greasewood sar- its current distribution and may have provided cobatus baileyibailebaileyyyi mormon tea ephedra sppapp the ecological setting for the taxon s origin and budsagebuddage artemisia spinescentspinescensspinescens shockley s development winward et al 1986 199iggi19911 desert thorn lycium shockleyishockleyi and horse type toulon pershing county nevada brush tetradymia sppapp except for artemisia USA 1053 in S C sanderson and E D our taxonomy follows welsh et al 1993 and mcarthur 1593 21 august 1986 holotype mozingo 1987 the most common grass BRY isotopesisotypesIso types OGDF RENO SSLP and understory species at upper elevation lahontan UTC other specimens examined sagebrush sites is bluebunchblue bunch wheatwheatgrassgrass nevada douglas co topaz lake elymus spicatusspicatus at lower elevations thurber sanderson & mcarthur 1594 SSLP and desert needleneedlegrassesgrasses stipa thurberianathurberiana four sheets and S speciosaspeciosospeciosa and indian ricricegrassegrass stipa nevada humboldt co golconda hymenoideshymenoides bottlebrushbottlebrush squirreltail elymus plummer ssnn 1985 SSSLPS LP elymoideselymoides and sandberg bluegrass poa nevada humboldt co leonard secunda are more common areas supporting creek plummer & mcarthur sn 3 A arbuscula sspasp longilongicauliscaulis receive between october 1975 culture uss SSLP 175 and 350 mm of precipitation annually with nevada humboldt co trough most as winter precipitation the frost free springs jackson mountains plummer season ranges from 90 to 110 days lahontan brunner & mcarthur sn 3 october sagebrush grows most commonly on aridisolsAridisols 1975 culture ul SSLP but at upper elevations it also occurs on nevada humboldt co trough mollisolsMollisols soil conservation service US de- springs jackson mountains mcarthur partmentpartment of agriculture personnel have located 1532 culture ul SSLP A arbuscula sspasp longilongicauliscaulis on at least 17 soil nevada humboldt co trout creek series generally these soils have low available basin jackson mountains mcarthur water holding capacities and a shallow depth 1501 SSLP two sheets to an argillic horizon andor bedrock these nevada lyon co dayton sanderson soils are similar to those of low sagebrush A & mcarthur 1595 SSLP two sheets arbuscula sspasp arbuscula communities 199511995 LAHONTAN SAGEBRUSH A NEW TAXON 153

OREGON 61 mcdermitt

alturasalburas

LL0

winnemucca

UL

Susanville

reno austin

NEVADA

fig 1 extent of the known distribution of Artemisia arbuscula sspasp longicaulislongicaulis 154 GREAT BASIN naturalist volume 55

fosberg and hironaka 1964 zamora and mcarthur et al 1981 1988 the following tueller 1973 G K brackley and C A taxa are thought to have originated as hybrids plummer personal communication 0 A argiargillosallosa A cana sspasp viscivisciduladula X A general distributions of the three sub- longilobalongiloba beetle 1959 species of A arbuscula are as follows sspasp 0 A tridentata sspasp spiciformspiciformisis A arbuscula western wyoming and eastern tridentata sspasp vastyanavasevaseyanayana X A cana sspasp utah to eastern washington and northeastern viscivisciduladula beetle 1959 goodrich et al california sspasp thenthennopolathermopolathermonopolapola western wyoming 1985 mcarthur and goodrich 1986 and adjacent idaho and northern utah to 0 A tridentata sspasp xericensis A tridentata northern nevada and eastern oregon sspasp sspasp dentatatritridentatatridentate X A dentatatritridentatatridentate sspasp longicaulislongicauhslongicaulis western nevada extending into vastyanavaseyanavaseyana winward 1970 rosentreter adjacent california and oregon and kelsey 1991 0 A tridentata sspasp wyomingensis A supporting DATA AND discussion dentatatritridentatatridentate sspasp dentatatritridentatatridentate X A dentatatritridentatatridentate sspasp vastyanavaseyanavaseyana with perhaps some morphological chemical and cytological involvement with A nova as well data are consistent with the hypothesis that A beetle and young 1965 winward 1975 arbuscula sspasp longicaulislongicauhslongicaulis is of hybrid origin mcarthur 1983 with A arbuscula sspasp arbuscula as one parent figure 2 shows a polygonal representation of and A dentatattltrltritridentatatridentatetrzdentata sspasp wyomingensis as the morphological features of the new subspecies other parent hybridization and introgression and its putative parents including permanent are thought to have been important in the evo- leaf width length and lengthwidthlength width ratio and lutiolutionarynary development and differentiation of flower and vegetative stalk lengths values are tndentataetridentataeTridentatae taxa ward 1953 beetle 1960 shown in table 1 morphological differences

A arbuscula sspasp longilongicaulislongicauhscaulis

A arbuscula sspasp arbuscula A A tntritrldentata sspasp wyomingensis

B

A E C

D

fig 2 polygonal graph comparing morphological features of Artemisia arbuscula sspasp longicaulislongicauhslongicaulis and its putative par- ents A arbuscula sspasp arbuscula and A ttltntrltridentata sspasp wyomingensiswyormngensis data from table 1 A flower stalk length B vegetative stalk length C leaf width D leafleaflw1waw ratio E leaf length 199511995 LAHONTAN SAGEBRUSH A NEW TAXON 155

TABLE 1 morphological measurements means SD ofaof A arbuscula sspasp arbuscula araraARAR1 A arbuscula sspasp longi caulis ARARARARI1 and A tridentata sspasp wyomingensis artawaartrwaARTR a arararar81ARAB arabARARARARI ARTRARTRW flower stalk length cm 16416 4 17 24524 5 343.4 10410 4 42424.2 vegetative stalk length cm 393 9 080.8os 57 12121.2 474 7 24242.4 leaf length mm 59 11 737 3 iglg101.0 838 3 24242.4 leaf width mm 303 0 040.4 34340404 272708080.8 leaf1waw ratio 202.0 22 31 data foiforhorborogi arara irelreaieare fromhrombrom the following collections california lassen co shaffer mountain sinsandersonsandeisonSanderiondeisondervon & mcarthurMcArt huihur 1591 SSLP nevada eureka co tuscaroratu icaioia mountains 40 kmkrakna north ofofcarlincarlincariln frischknecht 210 SSLP nevada humboldt co 13 km northwest of paradise valley town holmgrenhoimHolm giengren 128 SSLP nevada lander co bade creek toiyabe national forest goodrich 8868 OGDF nevada landetlander co toiyabetoiyahe range toiyahe national forest goodrich 9966 OGDF nevada nye co toiyabe range toiyabe national forest goodrich 12201 OGDF nevada pershing co 40 km northeast Winnemuccaofwinnemuccaof winward s n 25 october 1841984 OGDF nevada washoe co buffalo hills 40 km northwestnoinol thwestthweat oiof gerlach sanderson & mcarthur 1592 SSLP data forfoirormol arabARARARABI aiealeare from collections nevada humboldt co golcondaColgolgoiconda plummer sns n 1985 SSLP nevada humboldt co trough springs mcarthurmeartMcArthuthui 1532 SSLP nevada humboldt co trout creek basin mcarthurmcauthurMcArthuihur 1501 SSLPSSLF nevada pershing co toulon sanderson & mcarthur 1593 SSLP nevada pershing co 646 4 km west of toulonoftoulon mcarthurMcAithur & mcarthurMcArthuihur 1683 SSLP nevada washoe co mustang meartmcarthurMcArthuihul & mcarthurMcArthulhui 1684 SSLP oiegonbiegonoregon lake co 32 kmkrakna east ofofadellocadelladelladeliadeil sanderson & mcarthurMcAithur 1590 SSLP oregon malteurmalheur co near mederMcDermcdermittraitt nevada winward sns n 31 october 1986 OGDF means foifolfor each tasataxa are based on N 8 differences aieare significant by one way analysis ofvananceof variancevanance woolf 1969 for flower stalk length P 01 vegetative stalk length FP 05 and icaflengthleaf length FP 01 between arabarabaararaARAR1 and ARARARABI data foifor artr1artraartrw aieare fromblom winward 1970 means from ARTR weiewere not statistically compared with the othelotherothet taxa may be summarized as follows 1 flower A arbuscula sspasp longicaulislongicaulis is hexaploid stalk length averages 50 longer for A arbus- 6xax 2nan 54 mcarthur et al 1981 reported cula sspasp longicaulislongicaulis than for A arbuscula sspasp this taxon as A dentatatritridentatatridentate sspasp wyomingensis arbuscula and over twice as long as A triden wide lobe and gave chromosome counts as tata sspasp wyomingensis 2 A tridentata sspasp 2nan 36 54 for two transplanted population wyomingensis has longer narrower leaves than samples growing in a common garden we the other taxa 3 A arbuscula sspasp longilongicauliscaulis now suspect both bona fide A tridentata sspasp has larger leaves and a longer vegetative stalk wyomingensis and A arbuscula sspasp longi than A arbuscula sspasp arbuscula caulis were growing in the accessional rows wilt et al 1992 compared 34 phenolic A dentatatritridentatatridentate sspasp wyomingensis is uniform- compounds from six taxa of tridentataeTridentatae in- ly tetraploid 2nan 36 for the ca 70 popula- cluding A dentatatritridentatatridentate sspasp wyomingensis A tions that have been examined and A arbus- arbuscula sspasp longicaulislongicaulis undescribed at the cula sspasp longicaulislongicaulis is uniformly hexaploid n time and called by them lahontan sage- 27 for seven populations including trout brush and A arbuscula sspasp arbuscula their creek mcarthur 1501 adell sanderson and work demonstrated that typical high pressure mcarthur 1590 toulon sanderson and liquid chromatography patterns for A arbuscula mcarthur 1593 topaz lake sanderson and sspasp longicaulislongicaulis and A dentatatritridentatatridentate sspasp wyo mcarthur 1594 dayton sanderson and minmingensisgensis are remarkably similar their fig 2 mcarthur 1595 and mustang mcarthur and representative chromatograms of those two mcarthur 1684 populations cited earlier in taxa are the most similar of the six taxa studied this report mcarthur et al 1981 mcarthur A arbuscula sspasp arbuscula A nova A triden and sanderson in review tata sspasp tridentata A t sspasp vastyanavaseyanavaseyana A t sspasp morphological chemical and cytological wyomingensis and lahontan sagebrush wilt evidence reveals affinities of A arbuscula sspasp et al 1992 we analyzed the wilt et al 1992 longicaulislongicaulis for both A a sspasp arbuscula and A frequency of detection of individual phenphenolicsolics dentatatritridentatatridentate sspasp wyomingensis in a series of by assigning percent similarity values for each common garden experiments hanks et al shared phenolic table 2 those values are 1973 mcarthur and plummer 1978 welch not dramatically different for A arbuscula sspasp and mcarthur 1979 1981 1986 nelson and longilongicauliscaulis in respect to the other five taxa but krebill 1981 mcarthur et al 1981 mcarthur A arbuscula sspasp arbuscula and A dentatatritridentatatridentate and welch 1982 welch et al 1987 lahontan sspasp wyomingensis are the two most similar at sagebrush accessions ul and uss trough 54 to A arbuscula sspasp longicaulislongicaulis for total springs NV and uss leonard creek NV phenphenolicsolics the wilt et al 1992 data are very were treated as wyoming big sagebrush they close for A arbuscula sspasp longilongicauliscaulis and A t fit with that group in those experiments sspasp wyomingensis closer than for any other sagebrush taxa hybridize naturally mcarthur two taxa examined et al 1988 we propose that A arbuscula sspasp 156 GREAT BASIN naturalist volume 55

TABLE 2 percent frequency of shared phenphenolicsolics for A nova ARNO A ttltntrltridentata sspasp tntritrldentata ARTR A tndentatridentatridental ta sspasp vastyanavaseyanavaseyana ARTARTRRv A tntritrlfridentata sspasp wyomingensiswyormngensis ARTR A arbuscula sspasp arbuscula araraararaARAR8 and A arbus- cula sspasp longiiongilongicaulislongicauhscaulis ARARARARI1 data from wilt et al 1992 ARNO artrtartr1artret ARTRV ARTRW araraARAR arar1ARABIarary ARNO 52 49 57 59 52 artrtartra 52 76 48 52 ARTRV 48 58 53 ARTRW 52 54 ARARARAB 1 54

longicaulislongicaulis is an allopolyploid derivative from FOSBERG M A AND M HIRONAKA 1964 soil properties diploid n 9 A arbuscula sspasp arbuscula and affecting the distribution of big and low sagebrush in southern idaho forage plant physi- 18 A communities in tetraploid n dentatatritridentatatridentate sspasp byominwyomin ology and soil vegetation relationships special gensis A arbuscula sspasp arbuscula occurs in publication of the american society of agronomy 5 both diploid and tetraploid populations in the 230 236 northwestern nevada area tetraploid A triden GOODRICH S E D mcarthur AND A H WINWARD tata sspasp wyomingensis also occurs there 1985 A new combination and a new variety in artem- ttltntrltridentata great basin naturalist 45 99 104 and sanderson in popula- isia mcarthur review HALL H M AND F E CLEMENTS 1923 the phylogenet- tion dynamics of sagebrush populations migrat- ic method in taxonomy the north american species ing in response to climatic change during the of artemisia chrysothamnus and atriplex publica- various phases of ancient lake lahontan and tions of the carnegie institute of washington 326 other pluvial lakes of the lahontan basin 1 355 HANKS D L E D MCARTHUR R STEVENS AND A P morrisonmornson 1965 could have provided the PLUMMER 1973 chromatographic characteristics opportunity for the origin and establishment and phylogenetic relationships of artemisiaofartemisia section ofaofA arbuscula sspasp longicaulislongicaulis morphological tndentataetridentataeTridentatae USDA forest service research paper cytological and chemical data are consistent INT 141 US department of agriculture forest range with the hybrid origin hypothesis service intermountain forest and experi- ment station ogden UT 24 appp MCARTHUR E D 1983 taxonomy origin and distribu- acknowledgments tion ofbigosbigof big sagebrush artemisia ttltntrltridentata and allies subgenus tndentataetridentataeTridentatae pages 3 13 in K L johnson we thank gary brackley jim brunner chu editor proceedings of the first utah shrub ecology gelinge lin sherel goodrich craig plummer workshop utah state university logan 50 appp marty mcarthur E D AND S GOODRICH 1986 artemisia tntritrl stewart sanderson wilt and the late dentata sspasp spiciformspicifarmisspiciformis distribution and taxonomic plummer is perry for their help in various phas- placement pages 55 57 in E D mcarthur and B L es of this study we thank field personnel of welch compilers proceedings symposium of the the soil conservation service US department biology of artemisiaofartemisia and chrysothamnus USDA of agriculture nevada offices who provided forest service general technical report INT 200 intermountain research station ogden UT 398 much of the information for figure 1 we also PP appreciate assistance provided by the MCARTHUR E D AND A P PLUMMER 1978 biogeography pittman robertson w82r wildlife habitat and management of important western shrubs a project and US department of agriculture case study section dentatetridentateTn of artemisiaofartemisia great cooperative state research service grant 91 basin naturalist memoirs 2 229 243 MCARTHUR E D AND S C SANDERSON in review 38300615738300 6157 additional chromosome counts in subgenus tndentatridentatridental tae of artemisiaofartemisia with implications literature CITED mcarthurMCARTHUK E D AND B L WELCH 1982 growth rate differences among big sagebrush artemisia tndentatridentatridental BEETLE A A 1959 new names within the section ta subspecies journal of range management 35 Tridenttndentataetridentataeatae of Artemisia rhodora 61 82 85 396 401 1960 A study of sagebrush the section tiidentatndenta MGARTHuRMCARTHUR E D C L POPE AND D C FREEMAN 1981 tae of artemisiaofartemisia wyoming agricultural experiment chromosomal studies of subgenus tndentataetridentataeTridentatae of station bulletin 368 1 83 artemisia evidence for polyploidyautoautopolyploidy american BEETLE A A AND A YOUNG 1965 A third subspecies in journal of botany 68 589 605 the artemisia tridentata complex rhodora 67 MCARTHUR E D B L WELCH AND D L NELSON 405 406 1985 developing improved cultcultivarsivars of sagesagebrushessagebrusherbrushes BRUNNER J R 1972 observations on artemisia in nevada and other composite shrubs pages 188 196 in J R journal of range management 25 205 208 carlson and E D mcarthur editors symposium 199519951 LAHONTAN SAGEBRUSH A NEW TAONTAXON 157

range plant improvement proceedings selected a superior selection of low elevation mountain big papers presented at the 38th annual meeting society sagebrush USDA forest service research paper for range management denver CO 220 appp INT 370 intermountain research station ogden MCARTHUR E D B L WELCH AND S C SANDERSON UT 10 appp 1988 natural and artificial hybridization between WELCH B L E D mcarthurMCARTHUB AND R L RODRIGUEZ big sagebrush artemisia tntridentata subspecies 1987 variation in utilization of big sagebrush acces- journal of heredity 79 268 276 sions by wintering sheep journal of range manage- MORRISON R B 1965 quaternary geology of the great ment 40 113 115 basin pages 265 285 inm H E wright jr and D G WELSH S L N D ATWOOD S GOODRICH AND L C frey editors the quaternary of the united states HIGGINS 1993 A utah flora and2nd edition revised princeton university press princeton NJ 922 appp brigham young university print services provo MOZINGO H N 1987 shrubs of the great basin univer- UT 986 appp sity of nevada press reno 342 appp WILT F M J D GEDDES R V TAMMA G C MILLER NELSON A 1900 new plants from wyoming XII bul- AND R L everetiEVEREEVERETTTr 1992 interspecific variation of letin of the torrey botanical club 27 258 274 phenolic concentrations in persistent leaves among NELSON D L AND R G KREBILL 1981 A sagebrush wilt six taxa from subgenus tridentataetndentataeTrident atae of artemisiaartemisioofartemisia disease of unknown cause great basin naturalist 4411 asteraceae biochemical systematics and ecology 184 191 204120 41 52 nutNurNUTTALLrALL T 1841 descriptions of new species and genera WINWARD A H 1970 taxonomic and ecological relation- of plants in the natural order of the compositae ships of the big sagebrush complex in idaho unpub- transactions of the american philosophical society lished dissertation university of idaho moscow 80 ii7iia11 7 357 453 PP rosentreter R AND R G KELSEY 1991 xericxene big 1975 evolutionary development of artemisiaofartemisia tntritrl sagebrush a new subspecies in the artemisia triden dentata taxa page 163 in H C stutz editor tata complex journal of range management 44 proceedings of the symposia and workshop wild 330 335 land shrubs brigham young university provo UT SHULTZ L M 1986 taxonomic and geographic limits of 168 appp

artemisia subgenus tridentataeTridentatae beetle mcarthur 1 1980 taxonomy and ecology of sagebrush in asteraceae anthemideae pages 20 28 in E D oregon oregon agricultural experiment station mcarthur and B L welch compilers proceed- bulletin 642 1 15 ings symposium of the biology of artemisia and WINWARD A H AND E W TISDALE 1977 taxonomy of chrysothamnus USDA forest service general tech- the artemisia tntritrldentata complex in idaho univer- nical report INT 200 intermountain research sity of idaho college of agriculture idaho agricul- station ogden UT 398 appp tural experiment station bulletin 19 1 15 WARD G H 1953 artemisia section senphidiumseriphidium in WINWARD A H E D MCARTHUR AND G K BRACKLEY north america a cytotaxonomic study contributions 1991 lahontan sagebrush another sagebrush in of the dudley herbarium 4 155 205 nevada page 13 in abstracts 44th44tb annual meeting WELCH B L AND E D MCARTHUR 1979 variations in society for range management denver CO 45 appp winter levels of crude protein among artemisia tntritrl WINWARD A H E D MCARTHUR D A KAFFER C A dentata subspecies grown in a uniform garden journal PLUMMER AND G K BRACKLEY 1986 another sage- of range management 32 467 469 brush in nevada technical note range nevada 44 1981 variation of monoterpenoid content among USDA soil conservation service reno NV 2 appp subspecies and accessions of artemisia tntritrldentata WOOLF C M 1969 principles of biometry D van grown in a uniform garden journal of range manage- nostrand co inc princeton NJ 359 appp ment 34 380 384 ZAMORA B AND P T TUELLER 1973 artemisia arbuscula 1986 wintering mule deer preference for 21 acces- A longilobalongiloba and A nova habitat types in northern sions of big sagebrush great basin naturalist 46 nevada great basin naturalist 33 225 232 281 286 WELCH B L E D MCARTHUR D L NELSON J C received I1 march 1994 PEDERSON AND J N DAVIS 1986 hobble creek accepted 11 august 1994 cleatgleatgreat basin naturalist 552 C 1995 appp 158 163

DOUGLAS FIR TUSSOCK MOTH ORGYIAORCYIA pseudotsugataPSEUDOTSUGATA mcdunnough ON SUBALPINE FIR IN NORTHERN UTAH

E matthew hansellhansenlhansen1

ABSABSIRACtraunTRAUr1 douglas hirbirfir tussock moth orgyia pseudotsugapseudotsugatata mcdunnough defoliation was detected by aerial sur- vey on three areas of the wasatch cache national forest in 1990 and 1991 these are the fustfirst documented tussock moth outbreaksoutbioutby eaks in utah ground surveys revealed that subalpine fir abies lasiocarpalasiocampa hook nutt was heavily defoli- ated duringdining the outbreakoutbieakbleak douglas firhirbir pseudotsugamenziesiipseudotsuga menziesmenziem birbmirb franco though a minor component in the affected aleasareas had noticeably less defoliation and mortality adjacent stands of douglas fir had little or no visible tussock moth activity defoliation on subalpine fir was typically found evenly distributed throughout the crown rather than concen- trated at the top ninety four percent of subalpine fir with defoliation ratings of 90 or more were killed top kill occurred on neailynearlyneally one half of subalpine firs defoliated 25 89 heavily defoliated trees tended to occur in pockets bounded by areas of light defoliation after three consecutive years of defoliation tussock moth populations collapsed no life stages weiewelewere detected in 1993 from visual inspections of foliage or in pheromone traps

key words tussock moth subalpine fir defoliadefohatorsdefoliatorsforstors douglas fir utahutahforestsforests forest insects

the douglas fir tussock moth orgyia interestingly the first record of tussock moth pseudotsugapseudotsugatata mcdunnough is a significant defoliation in the united states was on sub- defoliator of douglas fir and true firs through- alpine fir at Jarjarbidgeharbidgebidge NV in 1927 balch 1930 out its host range in western north america affected areas were of subalpine fir occurring tussock moth outbreaks on the wasatch either purely or in mixture with limber pine cache national forest UT from 1990 to 1992 pinus flexflexilisilis james these two conifers and were unique because subalpine fir abies lasioalasio quaking aspen populus tremultremuloidesoides michamichx carpa hook nutt experienced greater defo- formed virtually the entire forest at Jarjarbidgeharbidgebidge liation and mortality than douglas fir pseudo- balch 1932 in contrast to the utah sites that tsuga menziemenziesiimenziesnmenziesiasiisllslisn birbmirb franco these tussock contain or are surrounded by substantial moth infestations are the first documented douglas fir balch s records are the only litera- subalpine fir outbreaks in utah ollieu 1978 tunnock et al ture indicating to be a principal 1985 host while more recent authors consider the previous tussock moth outbreaks indicate species to be secondary wickman et al 1981 berryman 1988 three hosts depending on location in primary objective of this study docu- british columbia and northern washington the was to ment the effects of these outbreaks on sub- douglas fir is preferred southern washing-washing in alpine fir information will be used to predict ton oregon and idaho douglas fir white fir future impacts of tussock moth defoliation on abies concolor cordgord and glend 1 and grand subalpine fir resource this report summarizes fir A grandis dougldougi 1 lindl are preferred stand conditions following infestation and in california nevada arizona and new attempts to characterize the effects of defolia- mexico white fir is preferred wickman et al tion on individual trees 1981 in these areas subalpine fir and other members of the pine family are typically defoli- METHODS ated after the preferred host is consumed the wasatch cache national forest outbreaks aerial detection surveys delineated tussock experienced light defoliation of douglas fir moth activity near the baxter sawmill site within stands of completely defoliated sub- ogden bangerranger district wasatch cache nation- alpine fir al forest UT in 1990 additional infestations

USDA forestfoiest serviceSc ivice foiestforest pest management 4746 south 1900 east ogden UT 84403

158 199519951 TUSSOCK MOTH ON SUBALPINE FIR 159 were detected at blind hollow and amazon ratings from 1992 and 1993 were used to de- hollow logan ranger district wasatch cache termine defoliation effects on tree condition national forest UT in 1991 subsequent ground surveys confirmed tussock moth popu- RESULTS lations at these locations A total of 35 pairs of 20 basal area factor blind hollow variable plots and 1300 ac seedlingsaplingseedling sapling prior to the outbreak composition of all fixed plots were established in july 1992 at the live trees greater than 5 inches DBH was 69 three areas plots were installed along a tran- subalpine fir 28 aspen and 3 douglas fir sect at 100loom m intervals starting from a refer- total pre outbreak live basal area was 1789178.9 sqaq ence point in areas with visible defoliation no ftacatac by 1993 live basal area was reduced to other attempt was made to randomize plot 1668166.8 sqaq ftacatac site elevations range from 7800 locations or to cover the entire affected area to 8100 ft aspect ranges from southwest to tree data collected include species diameter south to east on slopes varying from 10 to at breast height DBH height age 5 and 10 50 year radial growth insectinsectdiseasedisease damage and subalpine fir seedlings and saplings 0 494.9 an ocular estimate of percent defoliation inches DBH were significantly affected by additionally the distal 18 inches of three ran- tussock moth sixty stems per acre in this size domly selected lower crown branches on all class representing 25 of the stocking were host species were examined for pupae or egg killed table 1 pole sized subalpine fir 5 898.9sg masses inches DBH were more frequently defoliated plots were remeasured in july 1993 to eval- than larger diameter classes fifty eight per- uate changes in insect related tree injury and cent of stems 5 898.9sg inches DBH 34 of stems mortality percent defoliation and presence of 9 11911.9 inches DBH and 39 of stems 12 tussock moth life stages one pair of plots at inches DBH were defoliated stems exhibiting baxter sawmill was cut and lost during salvage top kill increased proportionately with per- operations to maximize data collection con- cent defoliation four percent of subalpine fir sistencysistency the same individual conducted defoli- stems over 5 inches DBH were killed by tus- ation estimates on 32 of 35 plots for plot estab- sock moth lishmentlishment and remeasurement in the 12 inches diameter class none of INDIDS the insect and disease damage 737.3 douglas fir per acre were visibly defoliat- survey program bousfield et al 1985 was ed table 1 among subalpine fir in this class used to calculate per acre average stand char- 3 of 65165.1 per acre were defoliator killed acteristicsacteristics individual tree defoliation ratings twenty eight percent survived defoliation were classified into the five categories of while 69 were not visibly defoliated weatherby et al 1992 1 undamaged 2 western balsam bark beetle dryocoetes con lightly defoliated 1 24 of total needle com- fusus swaine killed 494.9 subalpine fir stems per plement defoliated 3 moderately defoliated acre these trees were attacked in 1991 coin- 25 74 4 heavily defoliated 75 89 and ciding with the peak of the tussock moth out- 5 very heavily defoliated 90 defoliation break

TABLE 1 trees per acre condition summary of subalpine fir and douglas fir following a douglas fir tussock moth out- break blind hollow wasatch cache national forest july 1993 summary calculated from 10 variablefixedvariable fixed plot pairs SAF subalpine fir DF douglas fir defoliation class diameter undamaged light moderate heavy very heavy mortality class SAF DF SAF DF SAF DF SAF DF SAF DF SAF DF 0 49 300 00 1200 00 300 00 00 00 00 00 600 00 5 89 390 00 160 00 185 00 60 00 62 00 60 00 9 119iigailga1191 259 29 70 00 60 00 00 00 00 00 00 00 12 40040 0 73 13413.413 4 00 48 00 00 00 00 00 21 00 160 GREAT BASIN naturalist volume 55

the 1992 survey found an average of 313.1 the 1992 survey found an average of 484.8 pupae and 050.5os egg masses per three branch pupae and 121.2 egg masses per three branches samples the 1993 survey found no current sampled no current life stages were found in life stages on any sample tree and no life 1993 on the plots or in the area additionally no stages were visible in the area tussock moths were caught in pheromone traps placed in the baxter sawmill area in 1993 baxter sawmill amazon hollow prior to the outbreak composition for all live trees greater than 5 inches DBH was 65 prior to the outbreak composition of all subalpine fir 25 aspen and 10 douglas fir live trees greater than 5 inches DBH was 73 total live basal area was 1761176.1 sqaq ftacatac at the subalpine fir 24 aspen 2 douglas fir and onset of the outbreak live basal area in 1993 1 lodgepole pine total live basal area was was 1128112.8 sqaq ftacatac site elevations range from 1255125.5 sqaq ftacatac at the onset of the outbreak 7400 to 7900 ft aspect is south southwest live basal area in 1993 was 72272.2 sqaq ftacatac site west and northwest on slopes varying from 10 elevations range from 7500 to 7800 ft aspect to 30 is east on slopes varying from 10 to 25 subalpine fir seedlings and saplings had one hundred subalpine fir seedlings and considerable defoliator damage more than saplings per acre or 10 of stocking in that 250 seedlings and saplings per acre or 55 of class were killed table 3 mortality in the stocking in this size class died table 2 most three size classes greater than 5 inches DBH surviving seedlings and saplings were only ranged from 50 to 62 top kill was common lightly defoliated forty nine percent of sub- for all defoliation intensities of the surviving alpine fir stems 505.0so 11911.9 inches DBH were defoliated subalpine fir 5 inches DBH killed by tussock moth trees with top kill 60 had top kill including 63 of stems clas- increased proportionately with percent defoli- sified as lightly defoliated ation only 3 of subalpine fir stems in the in the 12 inches size class 28 of 434.3 lightly defoliated category experienced top douglas fir per acre were defoliator killed kill compared to 92 of surviving trees in the with another 16 defoliated but surviving heavily and very heavily defoliated classes table 3 among 29729.7 subalpine fir per acre in in the 12 inches diameter class douglas that class 50 were defoliator killed and fir had 10 of 22622.6 trees per acre defoliator another 31 were defoliated but survived killed fifty seven percent were not defoliated western balsam bark beetle killed 262.6 sub- with another 33 defoliated but surviving alpine fir per acre annosus root disease was table 2 among 38538.5 subalpine fir per acre in found on 424.2 trees per acre this size class 7 were defoliator killed and the 1992 survey found 202.0 pupae and 060.6og egg 77 were defoliated but survived masses per three branch samples the 1993 western balsam bark beetle has also been survey failed to detect any current life stages active at sawmill killing 38238.2 subalpine baxter sample tree summary fir per acre mostly in 1990 or 1991 annosus root disease heterobasidion annosum fr arefbref two hundred ninety one host sample trees was found on 464.6 subalpine fir per acre were rated for defoliation and monitored for

TABLE 2 trees peipelper acre condition summary of subalpine fir and douglas fir following a douglas fir tussock moth out- break baxter sawmill wasatch cache national forest july 1993 summary calculated from 13 variablefixedvariable fixed plot pairs SAF subalpine fir DF douglas fir defoliation class diameter undamaged light moderate heavy very heavy mortality class SAF DF SAF DF SAF DF SAF DF SAF DF lafIAFSAF DF 0 49 231 00 1385 00 231 00 23123.123 1 000 0 00 00 2535253.5253 5 000.00oo 0 5 89 129 00 00 00 00 00 00 00 00 00 293 00 9 119119111.91 25 00 78 00 25 00 21 00 00 00 250 00 121211 34 12812.812 8 17917 9 65 27 10 00 00 13 00 27 22 199519951 TUSSOCK MOTH ON SUBALPINE FIR 161

TABLE 3 trees per acre condition summary of subalpine fir and douglas fir following a douglas fir tussock moth out- break amazon hollow wasatch cache national forest july 1993 summary calculated from 12 variablefixedvariable fixed plot pairs SAF subalpine fir DF douglas fir defoliation class diameter undamaged light moderate heavy very heavy mortality class SAF DF SAF DF SAF DF SAF DF SAF DF SAF DF

0 49 5000 00 2750 00 75075 0 000.00oo 0 00 00 25025 0 000.00oo 0 loo1000100.0100 0 25025 0 5 89 68 00 134 00 43 00 00 00 00 00 404 00 9 119119111.91 50 00 95 00 31 00 00 00 00 00 230 00 12 57 24 55 04 07 03 12 00 18 00 14814 8 13

survival tables 4 5 defoliator caused mor- over douglas fir all three study sites are in tality was found to increase with the degree of close proximity to stands where douglas fir is defoliation in the very heavily defoliated class the primary overstory component these doug 94 ofsubalpineofsubalpine firs and 100 of douglas firs las fir stands experienced little or no visible were killed none of the sample trees in the defoliation this contrasts to balchbaich s Jarjarbidgeharbidgebidge lightly defoliated class were killed incidence NV site where subalpine fir limber pine and of top kill also increased with degree of defoli- quaking aspen form practically the whole of ation although trees in the heavily and very the foresthorest balch 1932 heavily defoliated classes were more likely to another exception to the tussock moffmoth s pref- succumb than exhibit top kill this parallels erence for douglas fir white fir or grand fir other tussock moth study results where has been observed in urban areas along the degree and incidence of top kill is related to colorado front range in these cases blue severity of defoliation wickman 1978 spruce picea pungentpungens engelm has been the surviving defoliated trees began to recover preferred host over white fir and douglas fir by 1993 tables 4 5 average defoliation rat- D Leatherman 2 personal communication ing for lightly defoliated subalpine fir in 1992 in colorado s native forests douglas fir is the was 777.7 in 1993 the same trees had an aver- principal host age rating of 393.9 with no visible defoliation the defoliation pattern seen on the wasatch of that year s needles the other defoliation cache national forest outbreaks differed great- classes for subalpine fir and douglas fir had ly from previously recorded patterns such as similar recoveries some of the most dramatic in oregon s blue mountains wickman 1978 recoveries however can be partially attrib- recommends estimating defoliation according uted to the most heavily defoliated trees of to the percent of crown totally defoliated from their respective classes succumbing and there- the top down that technique was abandoned fore not being rated in 1993 for this study because most needle loss was distributed evenly throughout the crown rather discussion than concentrated at the top application of wiekwickmanWickmanss method would have misrepresent- although douglas fir tussock moth had been ed many trees with significant defoliation by previously captured in pheromone traps in having them rated at 10 defoliation in utah the wasatch cache outbreaks are the other words the wasatch cache national first to be documented in the state tunnock et forest outbreaks did not fit the top down al 1985 more significantly a literature review defoliation pattern observed in other outbreaks revealed the wasatch cache outbreaks to be J Weatherbyweatherby33 personal communication unique in that subalpine fir is apparently the this study indicates that subalpine fir may preferred host type baichbalch s 1930 1932 stud- be locally more susceptible to tussock moth ies are the only that list subalpine fir as a pri- mortality than either grand fir or douglas fir mary host more recent literature indicates sub- despite the difference in percent defoliation alpine fir to be secondary to douglas fir white fir or grand fir wickman et al 1981 johnson entomologist2entomologist colorado state forest service and lyon 1988 at the wasatch cache out- entomologist3entomologist USDA forest service forest pest management boise breaks subalpine fir appears to be preferred idbadbidaho 162 GREAT BASIN naturalist volume 55

tabletailleIABLL 4 condition otof subalpmesubalpine fir sample trees within tussock moth monitoring plots at blind hollow baxter sawmill and amazon hollow wasatch cache national forest july 1993 average 1992 average 1993 class limits defoliation defoliationdefoliation22 total no top kill mortality defoliation classi defoliation defoliation defoliation oftreesof trees undamaged 00 00 00 51 0 0 0 0 light 1 24 77 39 71 7 10 0 0 moderatemodel ate 25 74 395 363 31 14 45 3 10 heavy 75 89 775 775 9 2 22 5 55 very heavy 90 913 725 63 2 3 59 94 itreesidrees1 filesFILLS issincdassigned defoliationdclidalideltdeit itionaition elassclass1 isstvs based on 1992 defoliation ratings 2s111vivingmvivingmviving trees floifrolfromtrom 1992 defoliationdctolidotoli itionaition classcl issibs

TABLEfabirabi L 5 condition of douglas fir sample trees within tussock moth monitoring plots at blind hollow baxter sawmill and amazon hollow wasatch cache national forest july 1993 average 1992 averageverage 1993 class limits defoliation defoliationdefoliation22 total no top kill mortality defoliation classelass defoliation defoliation defoliation of trees undamaged 00 00 00 17 0 0 0 0 light 1 24 65 20 20 0 0 0 0 moderatemodel ate 25 74 400 217 3 0 0 0 0 heavy 75 89 800 650 2 0 0 1 50 very heavy 90 950 3 0 0 3 100

1 nicesnieesileeliceileo s feignedassigned defoliationdcfoh itionaition classcl iss based1 isadiscd on 1992 defoliation r itingsicingsratings inviting2s111vivinginviving trees flomfrom 1992 defoliationdefoil itionaition clasiclasselass estimation techniques the wasatch cache virtually all host type was killed many of these results can be compared to those of wickman areas are bounded by stands of similar compo- 1978 at the 90 defoliation level wickman sition and density that were only lightly defoli- found 24 grand fir mortality and 30 douglas ated in a study of five case histories in oregon fir mortality 90 defoliation in wiekwickmanWick marsmais s and california wickman et al 1973 found study means complete defoliation in the top almost one half of tree mortality occurring in 90 of the live crown at the wasatch cache patches coinciding with high moth population outbreaks 57 of subalpine fir defoliated at centers 90 were killed 90 defoliation using the douglas fir tussock moth outbreaks typical- methodology of this study means that 90 of ly span two to four years moth populations the estimated total needle complement was develop rapidly and then abruptly subside consumed at the 99 defoliation level wick- after only one to two years of outbreak popula- man found that grand fir died at 53 and tions wickman et al 1981 the wasatch douglas fir at 46 this compares to 96 cache outbreaks have followed this pattern moitamoltamoitalitymortalitylity on wasatch cache subalpine fir moderate to heavy defoliation at baxter sawmill rated at 95 defoliation was first detected from aerial survey in 1990 within the infested study areas the degree defoliation was very heavy in 1991 in 1992 of damage varies greatly from one plot to the moth activity dramatically declined and in next one plot at amazon hollow had all host 1993 no life stages were discovered by either type defoliator killed while a plot 100 in dis- visual inspection or pheromone trapping tant was only lightly defoliated although the while it is beyond this study s scope to iden- very heavily defoliated areas are restricted in tify causal agents that initiated the wasatch size usually less than 5 ac the amount of mor- cache outbreaks it should be noted that a tality in these pockets is substantial an area prolonged drought coincided with the infesta- not sampled at baxter sawmill due to salvage tion most damage occurred on drier sites logging operations included over 20 ac where such as ridge tops and southerly facing slopes 199511995 TUSSOCK MOTH ON SUBALPINE FIR 163 this corresponds to patterns seen in other dawn hansen cindy hampton john guyon outbreaks bergstrom 1980 the affected and bent olsen all forest pest management trees were apparently drought stressed at the intermountain region helped with data pro- time of defoliation the sudden moth popula- cescessingsing table preparation and editing irene tion collapse mimics that of other outbreaks voit intermountain research station assisted where a nuclear polyhedrosis virus appears to with the literature search be the major mortality factor wickman et al 1973 literature CITED conclusion BALCH R E 1930 the fir tussock moth reveals ability to cause serious damage forest worker 621762 17 18 1932 the fir tussock moth hemerocampa pseudo fir although uncommon douglas tussock tsubatatsugata mcd journal of economic entomology 25 moth can cause considerable damage to sub- 1143 1148 alpine fir while damage in the three study BERGSTROMBERCSTBOM D 1980 new lessons from old tussock moth areas was variable pockets of heavy defoliation outbreaks USDA forest service pacific northwest had substantial subalpine fir mortality research station 4 appp larger BERRYMAN A A 1988 dynamics of forest insect popula- diameter trees are apparently less susceptible tions plenum press new york NY to mortality except in these pockets where vir- BOUSFIELD W R EDER AND D BENNETT 1985 user s tually all host type was killed although a minor guide and documentation for insect and disease component in the heavily defoliated areas local- damage survey INDIDS rlrissig85 19 USDA forest service northern region missoula MT ly douglas fir appears to be less preferred JOHNSON W T AND H H LYON 1988 insects that feed host material all study areas are in close prox- on trees and shrubs 2ndand edition cornell university imity to douglas fir stands that exhibited little press ithaca NY or no tussock moth activity western balsam OLLIEU M 1978 detection of douglas fir tussock moth in the intermountain region using baited sticky bark beetle and annosus root disease con- traps USDA forest service intermountain region tritributed to subalpine fir mortality though visi- ogden UT 7 appp bly minor relative to defoliator impacts while TUNNOCK S M OLLIEU AND R W THIER 1985 history forecasting losses in volume would be difficult of douglas fir tussock moth and related suppression efforts in the and rocky based on this study the fate of individual trees in intermountain northern mountain regions 1927 through 1984 USDA forest can be reasonably predicted given degree of service report 851385 13 intermountaininterintel mountain and northern defoliation regions missoula MT WEATHERBY J C K A KNAPP B R GARDNER J ROBERTS acknowledgments AND P MOCETTINI 1992 A biological evaluation of the douglas fir tussock moth outbreak in southern idaho 1991 USDA forest service report 11492114 92 I1 am grateful for the many people who 01 intermountain region ogden UT helped with this project david leatherman WICKMAN B E 1978 how to estimate defoliation and colorado state forest service julie weather- predict tree damage USDA agriculture handbook by steve munson and john anhold all forest 550 pest WICKMAN B E R R MASON AND C G THOMPSON management intermountain region ppro-ro 1973 major outbreaks of the douglas fir tussock vided critical review of the manuscript alan moth in oregon and california USDA forest dymerski john guyon dawn hansen john service general technical report PNW 5 anhold valerie deblander all forest pest WICKMAN B E R R MASON AND G C TROSTLE 1981 management jill douglas fir tussock moth USDA forest service intermountain region forest insect and disease leaflet 86 ansted craig yanase and lisa robinson all utah department of agriculture assisted with received 21 april 1994 data collection julie weatherby and john accepted 14 november 1994 anhold provided input for the survey design great basin naturalist 552 0 1995 appp 164 168

SEASONAL NUTRIENT CYCLING IN potamogeton pectinatusPECTINATUS OF THE LOWER PROVO RIVER

C mel lytlellyttellytle1 and bruce N smithl2smithly

ABSTKAC 1 A common submersed aquatic plant of great basin wetland and riverine systems potamogeton pectinapechina tus L sago pondweed is a key waterfowlwater fowlbowl food nutritional qualities of submersed aquatics in the great basin are little understoodundeidundei stood the purpose of this study was to determine the seasonal element cycling and nutritional qualities of P pectinatuspectinatus drupelet leaf and root tissues from the lower provo river leaf tissue protein was 27 dry weight in july but declined to 15 by december drupelet protein content was 9 in july and 656 5 in october lignocellulose in leaf tissue was lowest in july at 34 and increased as the season progressed percent fat was highest in leaf tissue at 12 in july sugars were highest in P pectinpectinatusatus leaf tissues in december and july calcium and magnesium concentrations increased in P pectinpectinatusatus tissues over the entireeatnentn e season leaf tissue zinczine was 329 ppm dry weight in october leaf iron concentration was highest in september at 1184 ppm while root tissue iron was 7166 ppm manganese content in leaf tissue peaked in october at 4990 ppm copper concentrations in leaves and roots were variable high protein in leaf tis- sue would benefit local nesting and brooding waterfowl populations that feed on this aquatic trace metal concentrations in leaf and lootroot tissues from possible anthropogenic activities appear very high during fall migratory months metal bioaccumulation by this species in other great basin wetlands and possible metal toxicity in waterfowl warrant further study

key words sago pondweed potamogeton pectinatuspectinatus nutritional qualities trace element cycling metal bioaccumulation waterfowl

A common submersed aquatic plant of the condition and nutritional requirements are great basin potamogeton pectinpectinatusatus is a key based on studies from other areas of north primary producer in fresh and saline wetlands america yet energy and sustenance required kantrud 1990 waterfowl feed on all plant by waterfowl species that frequent the great parts including drupelet leaf and root tissues basin are largely provided by resident aquatic anderson and ohmart 1988 korschgen et al plants of these P pectinatuspectinatus ruppia mariti 1988 sherwood 1960 noted that whistling ma L widgeon grass scirpus marmaritimusitimus L swans olorolor columbianuscolumbianus fed heavily on tubers alkali bulrush scirpus pungentpungens L olney and drupedrupeletslets during fall migration in the bear three square scirpus acutisacutus L hardsterhardstemhard stem river migratory bird refuge and ogden bay bulrush and zannichellia papalustrislustris L horned refuge other waterfowl species canada pondweed are common plant species man- geese branta canadensis mallards anas platy aged in national refuges and waterfowl man- rhynrhynchosrhynchopschos pintails anas acutdabutaacutaacute gadgadwallswalls anas agement areas potamogeton pectinatuspectinatus is con- strepera canvascanvasbacksbacks aytha vallisneria and sidered the most important of these species redheads aytha americanaamerlamericand also fed on P for diving and dabbling ducks kadlec and pectinpectinatusatus leaf and root tissues localized inter- smith 1989 the purpose of this study was to mountain trumpeter swan cygnus buccinator determine the seasonal element concentra- populations are also largely dependent on sub- tions and nutritional qualities of P pectinatuspectinatus mersed aquatic plants as food anderson et al from a local great basin river drainage 1986 henson and cooper 1993 little is known concerning nutrient dynam- METHODS ics and seasonal element cycling of P pectinapechina tus from great basin wetlands kadlec and plant harvests were conducted monthly smith 1989 consequently how this aquatic from three locations within the lower provo species may affect waterfowl nutrition is poorly river drainage from july 1991 to december understood most assumptions concerning body 1991 1 just below deer creek dam 4024n

deparI1 departinenttmentament of botany and range science 401 WIDB brigham young university provo UT 84602 uthaiuthoi to whomwhonn correspondconespondencccorrespondeneceneeenecence should behe addressedaddi essed

164 199511995 SEASONAL NUTRIENT CYCLING IN P pectinatusPECTIN ATUS 165

TABLE 1 A range of measured water column and sedi- TABLE 2 mean exchangeable fe and mn from lower ment characteristics ph and electrical conductivity EQEC provo river drainage sediments ppm dry weight SES E from the lower provo river drainage n 3 means sharing the same letter are not significantly different P 05 water sediment depth cm fe mn clarity clear opaque velocity isecmsec 0 040.40 4 0 7 61661.661gig 6 07a0 7aaa 19219.219 2 12a1 2aaa depth cm 5 60 120 7 15 56514a56556.5 14a 11411.411408b08b temperature C 3 14 3 12 15 22 61361.361 3 17a1 7aaa 949.49 4 2 lb ph 74 69 22 30 57157 1 08a0 8aaa 12012 0 I17b1 ab7b EC mhoscm3kmboscm3 425 1570 m31w11131w elev 1603 m 2 near the sundance tionseions and intervals as plant samples turnoffturnoff4040 22n 11134w elev 1560 m 3 sediments were air dried and extracted for 200 m from the mouth of the provo river exchangeable iron fe and manganese mn near utah lake 4014n llew11144w elev with didiethylenetriaminepentaethylene triaminthiamin eventaepenta acetic acid 1347 m water column and sedimentlllwcharac- DTPA and detected by atomic absorption teristicsteristics measured in the lower provo river spectroscopy water samples were analyzed are found in table 1 sediment conditions for ph electrical conductivity mhoscm3ymhoscm3 ranged from stony with gravelly patches to and available fe and mn with an orion micro- silty clay mud stands of P pectinatuspectinatus were processor ion analyzer901 ph meter a wheat- most abundant on muddy sediments stone bridge and by atomic absorption spec- whole plants leaf and root tissues of P pec trostroscopycopy tinatustenatus were sampled in replicate from each mean concentrations and standard errors location Drupedrupeletslets shoot stems and leaves SE were determined for each plant sedi- tissues and belowbelowgroundground root rhirhizomeszomes and ment and water sample to determine if sig- turions tissues were separated from plant litter nificantnificant variation in plant tissue nutrient and and sediments invertebrates were removed element concentrations existed between the from samples when rinsed in warm water different months we used analysis of variance 38c38 C cleaned samples were rinsed in ANOVA where month was considered the deionized water and dried in a forced air oven fixed effect and sample site the experimental at 70c70 C plant sediment and water samples unit in a repeated measures design if signifi- were analyzed at brigham young university cance P 05os05.05 was found tukey s multiple department of agronomy and horticulture comparison procedures were used to separate plant and soil analysis laboratory dry plant means tissue samples were weighed and ground inm a wiley mill to pass a 40 mesh screen and 0250.25 RESULTS AND discussion g samples were digested in folin wu tubes with 5 ml of concentrated hn03HNO samples available fe and mn concentrations in water were left covered for 16 h before digestion in samples were 0060.06 oolooi0010.01 and 00010.001 ppm an aluminum block for I1 h at 100cloo100 C three ml sediment exchangeable fe and mn contents of 70 hc104 was added and samples were were found between the normal soil range of refluxed at 200c200 C until the solution cleared 5 65 ppm yet under anoxic conditions that approxapprox 30 min samples were then brought to are common in sediments fe and mn may be- somi50 ml volume with deionized water orson et come more available for root uptake spencer al 1992 element contents were detected by and brewer 1971 tisdale et al 1985 table 2 direct aspiration into a perkin elmer model significant differences in sediment exchange- 5000 atomic absorption spectrophotometer able mn were found between surface sedi- all blanks and standards were run with the ments 0 7 cm and the rest of the sampled same procedures percent total nitrogen and profile table 2 phosphorus were determined using a kjeldahl element concentrations and forage quali- digestion followed by analysis with an ALP- ties were determined for P pectinatuspectinatus tissues KEM rapid flow analyzer from july to december leaf and root tissue sediment 0 30 cm and water 1000 ml dry matter as a percentage of fresh weight samples were obtained from the same loca remained constant at 6 7 with the highest 166 GREAT BASIN naturalist volume 55 dry matter content observed in october percent nitrogen N and phosphorus P in throughout the season P pectinatuspectinpectmatusatus element leaf tissue reached peak concentrations in july and forage composition varied with growth but were significantly lower by december F stage significant variation in leaf tissue pro- 233723.37 df 414 P ooiool001.001 TF 793079.30 df tein was found F 216921.6921 69 dfd f 414 P 414 P oolooi001.001 table 4 vermaakvermaas et al 1983 001 between july september october and stated that P pectinpectinatusatus played an important december table 3 drupelet protein content role in P cycling in aquatic systems cultured was higher in july than in october in all P pectinatuspectinatus grown in water relatively high in months sampled leaf tissue protein was high- phosphate p04 P 03030.3 ppm accumulatedbioaccumulatedbio er than drupelet protein percent protein in p32 toQ 4734738 times the amount found in the leaf tissue was higher than values reported in water column nitrogen and P content in P othelother studies linn et al 1975 kantrud 1990 pectinatuspectinatus can be well above that required for acid detergent fiber ADF analysis revealed plant growth this would indicate luxury con- that leaf tissue was lowest in lignocellulose sumption of these elements jupp and spencer fiber in july but significant differences F 1977 ho 1979 madsen 1986 significant 3033.033 03 dfd f 414 P 07 in fiber content concentrations of calcium ca and magnesium were not observed as the year progressed mg accumulated F 291229.12 df 414 P linn et al 1975 found P pectinatuspectmatuspectinatus leaf fiber ooi001001.001 F 27871278.71 df 414 P ooi001001.001 in content of 33 that is similar to values leaf tissue between july and december this observed in this study increased fiber content may indicate abiotic deposition though no vis- would decrease the overall forage quality of ible crustationincrustationencrustationen on exterior leaf or stem sur- leaf tissue significant variation did exist F faces was observed hutchinson 1975 report- 17740177.40177 40 dfd f 414 P 001 in leaf tissue fat ed that P pectinatuspectinatus leaves were higher in ca content and was highest in july total non- fe K mg na and several micronutrientsmicronutrients than structural carbohydrates sugars in leaf tissues other aquatic plants yet no mention of time were highest in december and differed from sampled was given for these mineral concen- all other months F 421942.1942 19 dfd f 414 P trationstrations therefore no knowledge of seasonal 001 by october drupelet fat and sugar con- accumulation was determined potassium K tent were both higher than values found in content was highest in september and differed july significantly from percent K content in july F

TABITABLE L 3 mean protein fiber fat and sugar content in P pectinpectinatusatus drupelet and leaf tissue over five months forage quality constituents expressed as dry weight SES E n 3 means sharing the same letter are not significantly differ- ent FP 05 month tissue protein adfaadf1adaa fat TMC oc ddryry wt july leaf 27427.427 4 03a0 3aaa 34234.234 2 oga0 9aaa 12212 2 01aoia0 la 838.38 3 oia0 la aug leaf 24924 9 03ab0 aab3ab 35635 6 28a2 8aaa 656.56 5 02b0 ab2b 818 1 04a0 4aaa sept leaf 21421.421 4 03b0 ab3b 39739.739 7 04a0 4aaa 686 8 02b0 ab2b 797.97 9 02a0 2aaa oct leaf 20320 3 14b1 ab4b 37937.937 9 13a1 3aaa ti717.17 1 lib1 lb 868 6 oia0 la dec leaf lsiisi15115 1 03c0 3cac 38138 1 osa0 5aaa 595 9 I1 lc 110liolloiio11.011001b01b july drupelet 90900505 33433.433 4 060 6 6107616.1gi 070.7 12012.012 0 030.30 3 oct druDiudrupeletpelet 65650808 36336 3 131 3 740874 080.8 16316.316 3 121.2lbib1 2 acid detergentclott ijcnt tillifiberhiberbiber ADF a measure of percent lignocellulose 01or fiber 11rotal11 rotalnotalpotal nonstructirralnonstikonsti ultiii il carbohydrate inqrnornqFNC a measure ofofiiiigaissugars

TABLE 4 mean mineral element concentration in P pectinatuspectinatus leaf tissue over five months element content expressed as dry weight SES E n 3 means sharing the same letter are dotnot significantly different P 05 month tissue N P K ca mg S dry wt july leaf 4407a44 07a 060.6og0601a01aoia 191.9lgig1902a02a 131.31301a01aoia 030.303004a004a 1201a121.2 oia aug leaf 2810a28 ioaloa 050501aoia 3501b35 0 lb 1201a121.2 oia 0505001b001b sept leaf 3401a34 oia 050.50502a02a 3701b37 01b 1400414004ab141.4 004abab 06002c06 002c 1802b18 02b oct leaf 3302a333.3 oga 050501a01aoia 313101b01b 1701b171.7 0 ib 06001c060.6og 001c dec leaf 242401b01b 020201b01b 303.03003b03b 232301c01c 070.707002c002c 060.6og0601c01coieole 199511995 SEASONAL NUTRIENT CYCLING IN P pectinatusPECTIN ATUS 167

TABLE 5 mean trace element concentration in P pectinatuspectinatus leaf tissue over five months element content expressed as ppm dry weight SES E n 3 means sharing the same letter are not significantly different FP 05 month tissue znan fe mn cu

ppm idry wt july leaf 213 14a 633 67a 122 6aaa 214a21 4aaa aug leaf 185 ioaloaioa 1097 58b 1744 colblolb101b 10lb10 ab1b sept leaf 211 la 118475b1184 75b 38613861117c117c 10lb10 ib oct leaf 329 ab4b 963 73b 4990 48d llabll0b11 ob dec leaf 295 13b 1038 63b 2130 65b 8sob80bob

264026.40 df 414 P oolooi001.001 percent sulfur in the summer months when P pectinatuspectinatus was decreased between july and december F growing rapidly by fall and early winter protein 134113.41 df 210 FP 0303.03 table 4 content decreased but was still higher than zinc anzn concentration in leaf tissue was concentrations found in drupedrupeletslets apparently significantly higher F 365636.56 df 414 FP protein content in P pectinpectinatusatus leaf tissue ooiool001.001 in october and december than in all from the lower provo river was higher than other months table 5 mean fe content was concentrations reported elsewhere high pro- higher in august leaf tissue than in july F tein content in leaves and stems in the sum- 125912.59 df 414 FP 001001.001 after which fe mer months would greatly benefit nesting and content remained fairly constant throughout brooding waterfowl that feed on this aquatic the remainder of the sample period leaf tissue species drupelet fat and sugar content was mn content increased through the season and higher than that for leaf or root tissues in was highest in october F 58738587.38 df october this would tend to confirm why dru 414 FP oolooi001.001 table 5 dudkin et al 1976 beletspelets are so eagerly sought after by staging found that FP pectinpectinatusatus growing in polluted and migrating waterfowl trace metal fe and coastal waters of the black sea accumulated mn contents in leaf and root tissues accumu- mn to 050.5 dry weight this mn concentra- lated over the season and were very high by tion corresponds to values found in this study fall however water and sediment concentra- yet mn concentrations in water and sediment tions appear normal should be determined provo it from the lower river appear normal whether the trace metal concentrations copper cu in leaf tissue varied significantly observed are of natural or anthropogenic ori- ooi001001.001 with high F 444844.48 df 414 FP gin future research should develop a greater concentrations in july followed by lows in understanding of heavy metal accumulation in august through december table 5 this and other key great basin aquatic plant root rhirhizomeszomes and turions of root tissues species FP pectinpectinatusatus were not separated for analysis mean root tissue forage qualities compared to leaf tissues were lower in percent protein but higher in fat content table 6 phosphorus was the only mineral element with a concen- TABLE 6 forage quality mineral and trace element con- P rhizome and than in leaf centrationcent ration of pectinatuspectinatus root tissue root tration higher in root tissues tissues turions averaged over five months forage quality con- mineral N P K ca and mg contents of root stituents and mineral content expressed as and ppm drsdry tissues in this study were similar to contents weight SES E n 3 studies and wallwaliwailwaii found in other kollman protein ADF fat tncatnc11anca 1976 van sierssenvierssenViers sen 1982 high trace metal dry wt concentrations were also found in root tissues 130 10 nabndbnd1 109log10 9 30 11911.911ilg 9 13 and mn concentra- like leaf tissues mean fe N P K ca mg tions in root tissues appear inordinately high dry wt 210221 020.2 0aaa0401010.1oloi 292.92902020.2 150215 02 030103 01 FUTURE RESEARCH conclusions AND znan fe mn cu apppppm dry wt seasonal variation did exist in forage quali- 167 25 7166 1438 2051 570 14814.814 8 333.33 3 P pechina ties and nutrient concentrations in pectina total nonstmeturalnon&tructui al carbohydrate TNQTNC a measure of hiigaisugessugms s tus protein content in leaf tissue was highest intnot deterinineddetel mined 168 GREAT BASIN naturalist volume 55

acknowledgments tionseions of potamagetonpotamogeton pectinatuspectinatus communities archivarchev fuer hydrobiologie Supplementsupplementbandband 50 439 472 funding and materials for this study were KORSCHGEN C E L S GEORGE AND W L GREEN 1988 feeding ecology of canvascanvasbacksbacks staging on pool 7 of provided by the department of botany and the upper mississippi river waterfowl in winter range science at brigham young university university of minnesota press minneapolis 624 appp and the utah chapter of the wildlife society LINN J G E J STABASTARA R D GOORICH J C MEISKE AND D E OTTERBY 1975 nutritive value of dried or ensiledenfiled aquatic plants I1 chemical composition CITED literature journal of animal science 41 601 609 MADSEN J D 1986 the production and physiological ANDERSONANDEHSON B W AND R D OHMART 1988 structure of ecology of the submerged aquatic microphytemacrophyte com- the winter duck community on the lower colorado munity in badfishbaddish creek wisconsin unpublished river patterns and processes waterfowl in winter doctoral dissertation university of wisconsin press university of minnesota minneapolis 624 appp madison 449 appp ANDERSONANDLRSON HERRON AND B REISWIG 1986 D R R C ORSON R A R L SIMPSON AND R E GOOD 1992 A estimates of annual survival rates of trumpeter swans mechanism for the accumulation and retention of banded 1949 82 near red rock lakes national wild- heavy metals in tidal freshwater marshes of the life refuge montana journal of wildlifeofwildlife management upper delaware river estuary estuarineEstuanne coastal 5021850 218 221 and shelf science 34 171 186 DUDKIN M S I1 V ARESHIDZEARESHIDZL AND G D LUKINA 1976 SHERWOOD G A 1960 the whistling swan in the west chemical composition of seaweed in the coastal with particular reference to great salt lake valley waters of the black sea ukrainian SSR USSR utah condor 62 370 377 resursy rastitelRastitel nye 12 133 137 SPENCER D W AND P G BREWER 1971 vertical advec- HENSONHLNSON P AND J A COOPER 1993 trumpeter swan tion diffusion and redox potentials as controls on the incubation in areas of differing food quality journal distribution of manganese and other trace metals of wildlife management 57 709 716 dissolved in water of the black sea journal of geo- ho Y B 1979 inorganic mineral nutrient level studies in physical research 76 5877 5892 potamogeton pectinatuspectinatus L and enteromorpha proliferaprohferaprolproiifera TISDALE S L W L NELSON AND J D BEATON 1985 min forfar loch scotland hydrobiologia 62 7 15 soil fertility and fertilizers fth4th4tb edition macmillan hutchinsonhu rcillnson G E 1975 A treatise on limnology volume publishing new york NY 753 appp 111 sons iliIII limnological botany john wiley & new VAN VIERSSEN W 1982 the ecology of communities york NY 660 appp dominated by zannichellia taxa in western europe JUPP B P AND D H SPENCER 1977 limitations on macro III111 chemical ecology aquatic botany 14 259 294 lake 1 phytespaytes in a eutieutrophicophic loch leven I effects of VERMAAK J F J H SWANEPOEL AND H J SCHOONBEE phytoplankton journal of ecology 65 175 186 1983 the phosphorus cycle in hermistonGergermistonmiston lake with KADLECKADLLC J A AND L M SMITH 1989 the great basin special reference to the role of potamogeton pectinatuspectinatus pedersen marshes in L M smith R L and R M L pages 317 321 in proceedings oftheodtheof the international for kaminski editors habitat management migrating symposium onOD aquatic macrophytes nijmegannijmegadNijm egaDegan and wintering waterfowl in north americaamenea texas netherlands tech university press lubbock 651 appp KANTRUD H A 1990 sago pondweed potamogeton received 31 may 1994 pectinatuspectinatus L a literature review USU S fish wildlife accepted 3 january 1995 service resource publication 176 89 appp KOLLMAN A L AND M K WALIWALL 1976 Intraintraseasonalseasonal nationsvariationsva in environmental and productivity rela great basin naturalist 552 C 1995 appp 169 173

FACTORS influencing FISH assemblages OF A HIGH ELEVATION DESERT STREAM SYSTEM IN WYOMING

bernard carteri and wayne A huberti

ABSTRACT seven fish species were found in the bitter creek drainage of southwest wyoming but only speckled dace rhinichthys osculus flannelmouth sucker catostomus latiiatilatipinnislatipinmspinnis and mountain sucker catostomus platyrhynchusplatyrhynchous were indigenous no relationships were found between fish standing stocks and habitat features but species richness was related to elevation and stream width no fish were found above an elevation of 2192 m only the most downstream study reach had more than three species present two indigenous species speckled dace and mountain sucker and a nonnativenormative species fathead minnow pimephales kromelaspropromelasmelas weiewelewere predominant fishes in the drainage these three species withstand intermittent stream flows that are common in the drainage

key wordsfishwords fish streams desert wyoming habitat distribution

fish communities in streams become more system in southwestern wyoming and 2 complex as habitat diversity increases along determine the factors that influence fish abun- the length of a stream variation in fish com- dance and community structure within the munity structure within a stream system can fol- drainage low patterns of zonation or addition specific fish communities can be associated with zones STUDY AREA defined by water temperature or geomorphogeomorphy the study was conducted in an intermittent logic features or community complexity can drainage bitter creek a tributary to the green increase with progression downstream by river in the red desert of southwest wyo- addition of species moyle and nichols 1973 ming fig 1 the study area consists of bitter guillory 1982 mcneely 1986 hughes and creek and four tributatributariesries little bitter salt gammon 1987 platania 1991 rahel and wells bean springs and gap creeks frequent- hubert 1991 however such patterns may ly no measurable surface flow occurs in bitter differ in andaridarld drainages of the western united creek at bitter creek WY during midsum- states with depauperate ichthyofauna cross mer and midwinter flow data available in the 1985 water resources data system at the wyoming little is known about the fish communities water resources center university of wyo- in high desert stream systems in southwestern ming laramie bitter creek at salt wells wyoming annual precipitation over most of WY generally has no measurable surface flow these drainages is 16 cm with much of it as from july to february salt wells creek has snow in headwater areas during late winter and more persistent flows near its mouth but thunderstorms during late summer discharge records of no measurable flows occur in mid- is highest during spring runoff and streams summer and midwinter when no measurable frequently become intermittent during sum- flow occurs in these streams isolated pools of mer and winter because these systems in standing water can be found in the stream wyoming are at high elevations 1800 m channels elevation of the study area ranges above mean sea level water temperatures are from 1800 to 2400 m cool compared with other desert streams the streams in the bitter creek drainage typi- climate in these areas typically consists of dry cally are downcut by at least 151.5 m with steep moderately warm summers with long cold clay banks having no vegetation riparian vege- winters tation consists of grasses and sagebrush artem- the purpose of this study was to 1 describe isia sppapp upland vegetation is primarily the fish species present in a high desert stream latter

lWywyomingoming cooperative fish and wildlife research unit university of wyomingofwyoming laramie WY 82071316682071 3166 the unit isis jointly supported by the university of wyomingofwyoming wyoming game and fish department and national biological survey

169 170 GREAT BASIN naturalist volume 55

5 saltsait wellsweilswelis wyoming

rock springs 4 wyoming bitter creek 00 wyoming

6 3 2

5 P

IS 4 CD Q 3 2 CD

ioos 0 N nyoWYO ING 2

20 km 1 Researchasearch location

fig 1 map of the bitter creek drainage WY showing the location of the 16 study reaches

baxter and simon 1970 reported four fish velocity was determined within each reach species in collections at two sites in bitter using the dye flow method binns 1982 stream creek speckled dace rhinichthysrhimchthys osesculusosculusculus discharge at time of sampling was computed fathead minnow pimephales kromelaspropromelasmelas and from width depth and velocity mountain sucker catostomus platyrhynchusplatyrhynchous alkalinity hardness and ph were mea- were reported from a site about 10 km upstream sured at the time of sampling alkalinity and from the mouth blueheadbluebeadBluehead sucker catostomus hardness were determined with field test kits discobolus was the only species reported from hach model al 36dt ph with an electron- a site near salt wells ic meter mean elevation and channel slope at each study reach were estimated from 757.5 METHODS minute topographic maps fish were sampled in each 100loom m reach by sixteen 100 m long study reaches were electroelectrofishingfishing small mesh 64 mm block nets selected to represent variation in stream size were placed at each end and two or three elec and habitat in the drainage during summer trofishing passes were made over the entire 1993 wetted width mean depth and sub- reach three pass depletion estimates of species strate were determined across transects at 10 abundance were made in most reaches two inm intervals dominant substrate at each tran- pass depletion estimates were used when sect was visually determined following bainbarnbarmbamm et 80 offish captured by the first two passes al 1985 sand silt 2 mm diameter gravel were captured during the first pass fish 2 16 mm pebble 17 64 mm cobble abundance was computed using the zippin 65 256 mm and boulder 256 mm water method platts et al 1983 all fish were 199511995 DESERT FISHES 171 weighed to enable computation of standing As study reaches declined in elevation and as stock estimates width water velocity and discharge increased standing stocks of individual species total the number of species increased standing stock of all species and number of because the most downstream reach on species in a reach were evaluated for their bitter creek had twice as many species as any relation to nine habitat variables using simple other reach and flow at the reach was en- linear and multiple regression analyses hanced by discharge from a sewage treatment independent variables were included in plant we assessed relations with the omission regression models if they were significant at of that reach again no relationships were P 0505.05 we further limited inclusion of de- found between any of the habitat variables and pendent variables in multiple regression mod- standing stocks of fish but the number of els to ones that were not correlated at FP f 0505.05 species NS was significantly related to eleva- computations were performed using statistixstatisticStatistix tion E and water velocity V 404.0 analytical software 1992 NS 159515 95 0 006800068 E P 0014 r2ra 55 and RESULTS NS 3003 00 10 11 V FP 0018 r2ra 51

seven fish species were collected speckled among the 15 study reaches with a maximum dace fathead minnow utah chub gila atatraridarldrafiafaniafanna of three species present the number of species bonneville redsidebedside shiner richardsoniusRichardrichards oniussonius increased with decline in elevation and water balbalteatusteatus hydrohydrophloxphlox mountain sucker white velocity sucker catostomus commetcommersonicommercommersoncomcommersomoommersomsoni and flannel- mouth sucker C latipinnislatipinnis abundance varied discussion substantially among study reaches table 1 mean total standing stock of all species was of the seven fish species in the bitter creek 303.0 gm3gme and ranged from 0 to 21321.3 gm3gme no drainage only three speckled dace flannel- fish were found in the four reaches above mouth sucker and mountain sucker are 2192 m indigenous baxter and simon 1970 absence habitat features varied among the 16 study of fish above 2192 m is probably due to a cli- reaches table 2 flow was measurable at all mate that is too cold for barmwaterwarmwaterwarmwater fishes reaches stream width water velocity and dis- additionally no trout occur naturally or have charge increased downstream sand silt sub- become naturalized inm the watershed strate occurred over 90 of almost all study the number of species increased with pro- reaches alkalinity ph and hardness also gressiongres sion from headwater to downstream increased downstream reaches table 1 with the exception of the no significant relations were found most downstream reach on bitter creek no between any of the nine habitat variables and more than three species specked dace standing stocks of individual species or total mountain sucker and fathead minnow were standing stock of all species however there found in any of the study reaches the high were significant relations between the number elevation reaches with fish tended to have of species and four habitat variables predominantly or exclusively speckled dace and mountain sucker NS 208820.8820 88 0 0091000910.0091 E P 0003 r2fi2ra gi61gl61.61 much of the increase in species richness NS 0 13 08120.8120 812 W P 0010 r2ra 52 with downstream progression was due to the NS 3403.403 40 1100811.00811 008oos V P 029 r2ra 33 and most downstream reach on bitter creek NS 0570.570 57 3124531 245 F P 022 r2ra 32 where six species were found table 1 four of six species were not natives fathead min-min where NS number of species E eleva- now white sucker utah chub and bonneville tion in meters W mean wetted width in redsidebedside shiner mountain sucker was not found meters V water velocity in meters per sec- in this reach but it was common throughout ond and F flow in cubic meters per second most of the bitter creek drainage while this the best two variable model was study reach was lowest in elevation among the 16 study reaches it also was downstream from NS 143614 36 0 006500065 E 0530.530 53 W P 0001 r2fi2ra 80 the outfall of the wastewater treatment facility tqaq CA 172 GREAT BASINtz naturalistz volumeCD 55

r 1 11 CO OS 10 1 N CO CO ll 003 10IO 2092.09 CO CD 10IO 02 05 07 01 ebl021 ellelz011 809209 2.2 4.6 2 2 13 0 05 0 22mm wm46 95 0 0 0 v 0 0 0 0 co co68 he os ft &fl3 gaagapCT creeku eapgapco creeku url oal U CO u co co 00oo 0 1 1 1 1 I I 1 575.7 CO i i CO OO i 57mg 15610 213 eab012 014 eeb002 218 lo10 00 1 in 1 13 0 21 222.2 mm353.5 78 mi010.1 il 0 0 0 CO CO bbbmCO 35CO

OT bab0 M &0 c c 3 co 10 co os 1 beanscd spring beans spring r 00OO elm012i I 085025CO 002 2292.29CO CO r 08 0 141.4 069869 222.2 343.4f 808.00 1 wg1 pw 1 I eg lw CO Nbb82 mhCO 8000OO CQ 73a 0 0 0 0

1 10 0 1 1 009 067087027CO 005 00OO 689229M 00OO D OS 001 M 2.1 0 767.6 2 2 ml21co 28co co 38co 66gm 79 wells wells 0 0 0 f- S 4 i creek creek U

10 saltS saltlo co co 10 00oo C i ca 1 i c3 T co elm013 032co 005 00oo 238co 00oo 1 111 12 ele 0 28 24 28 85io 0 0 0 co co co co 00oo co 1993OS 05

i 1 1 00 10 0 td i I1 IT 0.6 CO 0180.18 005 001 196OS CO 10 t 1 04 02 06 10 13 ham 0 0 04 33 lo 777.7 5 06 1 5 CO 145 771 summers 00 0 0 0 0 0 ti 3

M CO 1 l i oo 0 CO 1 00 o 1 1 fl 18 f i 0090.09 013ezeewe 002 2048042.04 OS 00 10 04 l 04 26 0.7 duringC 4 1 M 4 21 0 eg07 0 29 48 85 0 0 co 0 0 0 070 co co 00oo 3 S 1993OT OS tat3 bitter bitter U s S CQ tj 10 M 01 ura l i co to S 00 m ira li 1 1 sampled creek ol w creek co 0200.20CO0 0120.12 006 00OO 806206606 CD tc i ai 02 58 05 65 2 24f 012 0 08 0 262.6 46 81 3 in D 3 co co cobmmm 00oo 0 0 summer co littleB little5 0 0 0 0 s 3 3 i WY bc oi i co co 05 0090.09 CO 0030.03 t 1 f I1 CO CD ool01l 29 g CO 14 0 027 0003 11 212mab 030 52 767.6 co 1 2 2 I1 I om lu duringc i CO CO 10 be1 M 3 0 0 0 nj tat3 drainagea a tat3 1l 00 CO CO CO i I1 1 1 r 0080.08 013 eeb002ehm0.02 i 2332.33CO to 0 sampled& tle mpr 1 1 2002.0 0 0 21 020 mm262.6 700 beCOme CO CO CO 66CO t- AU 0 0 0 creekS

u sewVVY

1 1 1 1 S i I1 cl 0.1 f CO CO 00 01 01 24 22 01 01 47 1 1 bitter g1ga N f a CO 0140.14i 0130.13 0130.13i i 1881.8800 00 LO I1 0 01ea 60 72 013 01 38 mm959.5 mg878.7 t 6 0 0 0 i 95 fflfal v v v v M f 0 0 0 0 co os 00oo u drainage thezaezao5 S inC Vs fi 10 0 00 00oo 015i f 006tem0.06 001 1981.98ammas co co in 05 04 100 10 18 0 0 01 27f 32 72 5 5 i I1 I1 s i CO CO t 00 creek 0 0 0 0 reaches co U

1 1 1 1 M i 0.1 a 4 1 1 CO LI 01 01 L CO CO CO 0 I1 bitter 081021 007een 0020.02 600800200 10IO creekm creek 12 0 0 02 0 25 212.1 800 4 010 0 1 V V 4 i I CO CO blCO 00OO study 0 0 0 0 2 u aia i u theameamo ID 161 B i bitter bitter oi D CO 00 g 1 0 CO 080.8 23 31 in CO 0260.26emmCO 0090.09 006 202 D 0 atco m 08tm CO m 27 0 0 01 0 262.6 202.00 797.9 3 0 c1ca w 3 co co cobmmm 20co irana D 0 0 0 0 J fishes3 reaches y2ya 1CO 1 i 1 i aqcq 1.2 2 CO CO D oiioilt 1 1 11 01 12 l 1 wo1 1 I CO OO CO r l lo1 CO OS 013 016 004 206bem2.06mem 00 of cl rl 0 2 19 0 03 0 28 171.7t 93 0 2 V i co co lg os study 0 0 0 0 m2ma 2 lil i melmolperL t1 CO t- i16t ta n 1 1 1 1 in 1 CO 0 CO 0 1 17 42 014taw 010 0020.02 610210 to oi to a 25 i 13 0 02 26 191.9 66 1 1 1 I am g N at 0 0 0 0 CO CO 49 0 i a stocks 0 features s iro bab0 g suckerS 0 3 standingo sa s3 iai3 habitat S bedsideredside co Scu Q u sucker 2 c 0 lee100 100 minnow 333 ca dacemaca 3 fo S 0 c1 g s 5 slope 0 velocity 1000 sucker m m i wetted x x CO valervalorwater 1 2 m3second chub M msecond Ss flannelmouth 1 a ll i l S il I x g mgl mgl TABLE zelszeis bonneville shiner TABLE S S width depth discharge g gx M mountain H elevation sall alkalinity hardness speckledssll n c & ax t bagbcg G 111fathead channel aumamm species habitat g m eglg egl feature s a s e e a saleoraleewhite S mean aomamean 0 water zoosc- p utah 1 ai u o 5 s total1 5 a a o a & ija mumumm PH rbt 2 fal ivaiv3 icseScscseSE ij1ij ffl s s Q d a 199511995 DESERT FISHES 173 for rock springs WY and was only 13 kinkm fish department for their assistance and H upstream from the confluence of bitter creek li T patton FE rahel R wiley and an anony- and the green river the more permanent mous referee for review of the manuscript flows due to the wastewater treatment facility the project was supported by the wyoming may have enabled fish not adapted to inter- game and fish department mittent flows to persist in this reach also the relatively short distance to the green river literature CITED may enable upstream migration of fish to this reach contributing to higher species diversity analytical SOFTWARE 1992 statistixstatisticStati stix version 4044.00 user s paul repeated invasion of nonnativenormative species from manual analytical software st MN BAIN M B J T FINN AND H E BOOKEBOOKL 1985 downstream reservoirs maintains species quantifying stream substrate for habitat analysis stud- diversity in the virgin river UT cross 1985 ies north american journal of fisheries manage- also human disturbances have been found to ment 5 499 500 create environmental conditions favorable to BAXTER G T AND J R SIMON 1970 wyoming fishes bulletin 4 wyoming game and fish department nonnativenormative fish in california moyle and nichols cheyenne 1973 1974 therefore enhanced flows due to BINNS N A 1982 habitat quality index procedures manu- the wastewater treatment facility and invasion al wyoming game and fish department cheyenne of nonnative species from the green river CROSS J N 1985 distribution offishof fishhishbish in the virgin river a tributary of the lower colorado probably contribute to the diversity of fish river environ- in mental biology of fishes 12 13 21 the downstream portion of bitter creek GUILLORYCUILLORY V 1982 longitudinal gradients of fishes in during summer 1993 flowing water thompson creek louisiana southwestern naturalist occurred at all study reaches when they were 2710727 107 115 sampled precipitation in spring and summer HUGHES R M AND J R GAMMON 1987 longitudinal changes in fish assemblages and water quality in the 1993 was substantially greater than normal willamette river oregon transactions of the enabling measurable surface flows to persist american fisheries society 116 196 209 during summer however study reaches up- mcneeley D L 1986 longitudinal patterns in fish stream from the outfall of the rock springs assemblages of an ozark stream southwestern naturalist 31 375 380 WY wastewater treatment facility are frequent- MOYLE P B AND R D NICHOLS 1973 ecology of some ly intermittent during late summer fathead native and introduced fishes of the sierra nevada minnow has been described previously as a foothills in central california copeiacopela 1973 478 490 species associated with intermittent streams 1974 decline of native fish fauna of the sierra baxter and simon 1970 pflieger 1975 our nevada foothills central california american mid- land naturalist 92 72 83 observations indicate that two indigenous PFLIEGER W L 1975 the fishes of missouri missouri speciespeciess speckled dace and mountain suck- department of conservation columbia er and one introduced species fathead PLATANPLATANIA IA S P 1991 fishes of the rio chama and upper minnow can survive in the frequently inter- rio grande new mexico with preliminary comments on their longitudinal distribution southwestern mittent streams consequently these three naturalist 36 186 193 fishes are the only species occurring over most PLATTS W W W F MEGAHAN AND G W MINSHALL of the bitter creek drainage but fathead min- 1983 methods for evaluating stream riparian and nows tend to be limited to lower elevations biotic conditions USDAUS DA forest service general report 138 70 than the two native species it is not known technical INT appp RAHEL E AND A HUBERT 1991 assemblages how by F J W fish the invasion fathead minnow may and habitat gradients in a rocky mountain great affect the native speckled dace and mountain plains stream biotic zonation and additive patterns sucker in this desert stream system of community change transactions of the american fisheries society 120 319 332

acknowledgments received 21 april 1994 accepted 3 october 1994 we thank M fowden K johnson W wengert and R wiley wyoming game and great basin naturalist 552 0 1995 appp 174 176

speciation BY aneuploidy AND polyploidy IN MIMULUS scrophulariaceaelscrophulariaceae1SCROPHULARIACEAE1

robert K vickeryjr2vickery jr 2

key words mimulus speciation evolution aneuploidy polyploidy

speciation by aneuploid and polyploid and simiolusSimiolus 14 15 16 30 base numbers of changes in chromosome numbers is so common the sections suggest extensive evolution by in flowering plants as to be almost a character- both aneuploidy and polyploidy for the genus istic of the angiospermsangiosperms elegant examples of as a whole the base number appears to be x these patterns of evolution are exhibited by 8 inasmuch as the other plausible base num- monkey flowers of the genus mimulus scro- ber x 7 is found only in one apparently de- phulariaceae rived desert species M mohavensis lemmon the genus mimulus contains some 150 table 1 species occurring in western north and south next let us consider the chromosome num- america with a few outlying species in eastern bers by individual species all species counted north america japan vietnam the himalayas thus far are the same in each of several sections new zealand australia and south africa the specifically in mimulastrum erythranthe center of diversity is california with a second- benoeoenoe and placusDidiplacusdipsacus the other sections are ary center in chile some species are annuals of polymorphic for their species chromosome deserts grasslands or forests some are bienbiennialsnials numbers and frequently exhibit speciation by of marshy places some are herbaceous peren- aneuploidy andor polyploidy often in com- nials from springs streamstreamsidessides or lake shore plex combinations for example the various habitats and others are woody shrubs of the species of section mimulusEueumimulus exhibit n 8 dry california chaparral the species form clus- 11 and 12 species of section euranuseunanus exhibit ters reflecting these various life forms there n 8 10 and 16 species of section daradanparadan are 8 10 such clustersciucluelusteis commonly recognized thus exhibit n 8 9 16 17 18 and 30 and as sections of the genus mimulus grant 1924 species of section simiolusSimiolus exhibit n 13 14 pennell 1951 chuang and heckard personal 15 16 24 28 30 31 32 46 and 48 table 1 communication section simiolusSimiolus which shows by far the chromosome numbers of over 50 species most speciation by aneuploidy andor poly- table 1 that is approximately one third of ploidy of all sections of the genus consists of the mimulus species have been ascertained six species groups that is complexes of related by vickery and his workerscoworkersco vickery 1978 species and varieties first is the M guttatusgustatusguttatus vickery chu et al 1981 vickery simpson et al complex centered in california it has as its 1981 vickery et al 1982 1985 1986 1990 un- base number x 14 with aneuploid forms at published and by chuang and heckard per- n 13 and n 15 table 1 as well as tetra- sonal communication chromosome numbers ploid forms with n 28 second is the alpine reveal intriguing patterns of evolution by aneu- western united states M tilingtilingiiii complex ploidy and polyploidy with its base number ofofaofxx 14 and aneuploid first let us consider the base chromosome forms at n 15 n 16 and an unusual poly- numbers of the eight mainmaln sections of the ploid form at n 24 the third species group genus section mimulastrum has a base num- is the M dentilobusdentilobus complex of southwestern ber of x 7 euranuseunanus and erythranthe have united states and northwestern mexico with base numbers ofaofxof x 8 Paradparadanthusanthus 8 9 10 its base number of x 16 and an aneuploid mimuluseumimulusEu 8 11 12 benoeoenoe 9 Diplacus 10 form at n 15 fourth is the M luteusgluteus complex

I1IAA talk presented 4 septemberSeptem bei 1993 as part of the symposium plant evolution at the national institute of genetics mishima japan 2bioloiydpaitmlnt2ffiolgy dopailmerit univerityuniversityUniumanumvnveritysitssity oiof utah salt lake city UT 84112 USA

174 199519951 NOTES 175

TABLE 1 chromosome numbers in the genus mimulus by from the central and southern andes of south sections counts by chuang and heckard and by vickery america its base number is x 30 but there and coworkersco see text for references workers are n 31 and n 32 forms as well fifth taxon n there is the M glabratusglabratus complex that ranges mimulastrum gray x 7 from canada to patagonia its varieties in cen- M mohavensismohavensts lemmon 7 tral north america exhibit the base number of Eumimulus gray x 8 11 12 the complex x 15 in the rio grande M alatas altonaiton 11 M gracilisgractlis R biby 8 drainage we find tetraploidstetraploidy with n 30 M ringens L 812 from northern mexico to southern colombia euranuseunanus gray x 8 we find the aneuploid tetraploid n 31 vari- M bolanboianbolandenderiderl gidygrdygray 8 of the complex ecuador south to M layneselayneae greenegregreeneGieene jepson 8 eties from M brevipesbrevipenbrevipes bentham 8 southern argentina and including the juan M cusiccusickcusickiicusicktikiikliti greene piper 8 fernandez islands off the coast of chile we M nanus hook & amarn 8 M torresitorreyi gray 10 find the aneuploid hexaploid species and vari- M biglovbigloviibtglovitii gigrayay 16 eties with n 46 chromosomeschromosomes apparently Paradparadanthusanthus clantgiantglantgrant x 8 9 10 each change in chromosome number facilitat- M bicolor hartweg ex bentham 8 ed adaptive radiation further south is M filiazilfilicaulisfihcaiihscaulis watson 8 an last M brewen greene coville 16 the M wiensiiwienski complex of the mountains of M florifloribundusbundus douglas 16 western mexico with its base number ofaofxof x M moschatusmoschnwschatitsatus douglas 16 M latidenslatidens gray greene 16 16 and three apparent new species that are M arenariusarenanusareavenariusnariusnafiusnanus grant 16 morphologically distinct and reproductively M primulaidespnmuloidesprimulaidesoidesoldes redbrydb 91718 M repensrapens R bibr 10 isolated vickery et al unpublished one has M nepalensis bentham 1630 n 16 chromosomeschromo somes one has n 32 chromo erythranthe greene x 8 somes and the third has two forms one with M cardinaliscardicarddnabsnahs douglas 8 32 chromosomeschromosomes and the other with 48 M eastwoodiae redbrydb 8 n n M lewish pursh 8 1 4 chromosomeschromo somes incipient aneuploidy M nelsonelsonnnelsonunelsoniinii grant 8 does speciation by aneuploidy and M ruperupestrisrupestnsstris gleenegreene 8 how M verbenaceusverbenaverbenaceousceus greene 8 polyploidy occur we carefully examined meiosis benoeoenoe glaygray x 9 in M glabratusglabratus var utahensis and M gladraglabra M dictuspictus cuicurranGUI ran gray 9 tus var fremontiifremontiatii two of the widespread M tricolor lindllindi 9 fremon M pygmaeus grant 9 or 10 diploid varieties of the M glaglabratusbratus complex M piloselluspdoselluspilosellusseilusselius greene 9 and their intervarietal F hybrids first of Diplacus gray i 10 1317 cells examined in diakinesis or meta- M alidusaridus abrams grant 10 of first meiosis and vickery 1970 M aurantiacusamantawantawanttacustacus curt 10 phase tai M calycinuscalyctnuscalycinnscinus eastweasaw io 1972 1090 exhibited regular 15 bivalent M clevelanddededevedevelandudhuelanduhanduii branagbrandg 10 chromosomeschromo somes another 23 cells or 171.7 had M fasiculatus pennell mcminn 10 M longifloruslongiflorus nutt grant io aneuploid numbers of chromosome pairs rang- M puniceuspunipuniceousceus nutt steudstaud 10 ing from only 6 to as many as 13 plus 4 18 sitsimiolussirmolusstrSirSimimolusolus greene x 14 15 16 uniunivalentsvalents these cells presumably could pro- M gutatusgustatus fischer ex DC 14152814 15 28 M lacinialaciniatuslacimatustacilacimatustus gray 14 duce aneuploid galetesgametesgametes at least in some cases M nasutus greene 1314 A sizeable minority 204 cells exhibited 14 II11 M glaucescentglaucescens greene 14 2 1 13 11 and 1 IV or complement frac- M platycalyxplatycalyx pennell 15 and I or II I M tilingtilingiiii regel 14152414152428 28 tionationtio nation with its uneven groupings of chro M gemnigemniparusgemmparusparus weber 16 mosonesmomosomessomes these cells might produce aneu- M dentidentilobuslobus rob & fern 1516 M wiensiiwienskiwwnszt vickery 16 ploid galetesgametes also second of 782 additional M glabratusglabratus HBK 153031 cells observed in anaphase I1 294 37537537.5 M andiandicoluscolus HBK 46 M pilosiusculusptlosiusculus HBK 46 exhibited unequal disjunction laggard chromo M externus skottsscottsSkotts skottsscotts 46 somes or chromatin bridges these cells also M luthus L 3031323031 32 some M cuprouscupreus dombrain 31 could result in aneuploid galetesgagametesmetes 47 of M undescribed these abnormalities occurred in glabratusglabratus n sp A 16 var fremonfremontiifremontiatii only 18 occurred in M glabratusglabratus n sp B 32 var utahensis but most 229 occurred in the in- n sp C 324832 48 141 44 tertervarietal hybrids thus varieties differ in their potential for producing aneuploid galetesgametesgametes 176 GREAT BASIN naturalist volume 55 and intervarietal hybrids are particularly nificancenificance in the mimulus glabratusglabratus complex scrophu- prone to do so this suggests to me that natur- lariaceaelanaceaelamiaceaeae american journal of botany 59 488 493 VICKERY R K JR 1978 case al hybridization probably plays a significant studies in the evolution of species complexes in mimulus evolutionary biology role in evolution in monkey flowers finding 1140411 404 506 occasional plants in various populations with VICKERY R K JR Y E CHU K flnemanFlFINEMANNEMAN AND S aneuploid chromosome numbers indicates PURCELL 1981 chromosome number reports on the that aneuploid gametes not only are produced scrophulanaceaescrophulariaceae in IOPB chromosome number galetes reports LXX presented by askell love taxon 30 68 but actually function of 95 cells exam- third VICKERY R K JR M SIMPSON AND M nellesteinNELL ESTEIN ined inm anaphase II11 22 were polyploid and 1981 chromosome number reports on the scrophu- could presumably lead to polyploid galetesgametesgametes lanaceaelariaceaelamiaceaeae in IOPB chromosome number reports thus we see significant numbers of the very LXX presented by askel love taxon 30 68 69 cytological abnormalities VICKERY R K JR S A WERNER AND E D MACARTHUR in the basic diploid 1982 chromosome number reports on the scrophu- varieties that could lead to evolution by aneu- lariaceaelanaceaelamiaceaeae in IOPB chromosome number reports ploidy and polyploidy that is to the very pat- LXXV presented by askell love taxon 31 360 terns of evolution that we actually see in the VICKERY R K JR S A WERNER D R PHILLIPS AND M glabratusglabratus complex S R PACK 1985 chromosome counts in section simiolusSimiolus of the genus mimulus X the M glabratusglabratus complex madrono 32 91 94 literature CITED VICKERY R K JR B Y KANG T K MAC S R PACK AND D A PHILLIPS 1986 chromosome counts in mimulus GRANT A L 1924 A monograph of the genus mimulus sect erythranthe scrophulariaceae III111 madronoMadrofiogio annals of the missouri botanical garden 11 99 389 3326433 264 270 PENNELLPLNNLLL F W 1951 mimulus pages 688 731 in L VICKERY R K JR FE RAHMEN S R PACK AND T MAC abrams illustrated flora of the pacific states 1990 chromosome counts in section simiolusSimiolus of the volume 3 stanford university press stanford CA genus mimulus scrophulariareaescrophulanareaescrophulariaceae XI M glabratusglabratus tafTAI W AND R K VICKERYVICKCRY JR 1970 cytogenetic rela- complex contcont Madromadronofioflo 37 141 144 tiontionshipsships of key diploid members of the mimulus glabratusglabratus complex scrophulariaceae evolution 24 received 6 july 1994 670 679 accepted 24 september 1994 1972 unusual cytological patterns in microsporomicrosporon genesis and pollen development of evolutionary sig great basin naturalist 552 0 1995 appp 177 180

speciation IN MIMULUS OR CAN A SIMPLE FLOWER COLOR MUTANT LEAD TO SPECIES divergencedivergencesdivergence11

robert K vickery jrjrejr22

key words mimulus erythranthe speciation reproductive isolation flower color mutations pollinapollinatorstors bumble- bees hummingbirds

the general pattern of speciation in nature versus 4 5 cm corollasCorollas ofmof M eastwoodiae has been clear for a long time the diver- rydberg and M rupestrisrupestris greene the two gence of portions of a population usually small cliff dwelling species are partially reflexed levin 1993 usually in geographic isolation red and typically hummingbird pollinated mayr 1976 and the accumulation of genetic also and last flowers of the rocky mountain changes by selection andor genetic drift crow variety ofofmM lewisii pursh are magenta pink and kimura 1970 that produce reproductive with all five corolla lobes gently recurved isolation and normally character divergence as rather than reflexed thus forming a beelandbee land well the critical step is reproductive isolation ing platform flowers of the sierra nevada and yet that step except for polyploid forma- variety of M lewisii are lavender pink rarely tion which in itself is not always effective white with corolla lobes thrust forward both dewet 1980 has rarely been observed varieties of M lewisii are bee pollinated actually happening in nature A promising mimulus lewisii flowers and those ofmof M east group in which to study speciation events in woodinewoodiae and M rupestrisrupestris produce only modest progress is section erythranthe of the genus amounts of nectar whereas the other species mimulus vickery 1978 produce abundant nectar table 1 thus the the six species of monkey flowers compris- species differ markedly in flower shape flower ing section erythranthe are moisture requiring color nectar production and consequently in herbaceous perennials 1 10 dindm in height with pollinapollinatorstors servicing the flowers in the forma- variously shaped opposite leaves and bilabiate tion of the six species evolution appears to flowers that have four stamens one style with have responded to selection imposed by polli- a bilobed sensitive stigma and five corolla nator preferences and ecological opportunities lobes that range in color from orange to red the result is that members of the complex rarely yellow and from lavender pink to have radiated into a wide variety of different magenta pink rarely white see grant 1924 habitats and niches for further details when considered species A bright yellow flowered mutant has by species rollascocorollas of M cardinalis douglas appeared on the scene in this setting of polli vary from orange to red rarely yellow and nator driven ecologically opportunistic evolu- are sharply and fully reflexed hummingbird tion in two populations ofmof M cardinalis bright pollinatedpollinated flowers corollasCorollas ofmof M verbenaverbenaceusverbenaceousceus yellow flowered morphsmorphe have become well greene are partially reflexed that is the established one population is in the siskiyouSiskiyou upper two corolla lobes are reflexed whereas mountains of oregon which is the northern the lower three are gently recurved flowers limit of the range ofofmM cardinalis grant 1924 are orange red to red rarely yellow and the other population is on cedros island baja also are hummingbird pollinated Cocorollasrollas of california and is at the southern limit of the M nelsoniinelsonianelsonii grant are partially reflexedre flexed also species range As mayr 1976 suggests new and have orange red to red flowers which are forms often evolve from isolated populations longer than those ofmof M verbenaceusverbenaverbenaceousceus 6 7 cm such as these on the periphery of a species

theithe opening talk in the symposium mechanisms of speciation inin higher plants given 1 september 1993 at the XV international botanical congressCongiessgress yokohama japan biology department university of utah salt lake city UT 84112 USA

177 178 GREAT BASIN naturalist volume 55

IABLLTABLE 1 nectar production in the species of section nate to test these two questions I1 used the fact erythranthe measured at 0800 h in the wild vickery and that M cardinalis flowers are borne in pairs I1 sutherland 1994 averages are based on 20 or more mea grew plants of red and of yellow flowered M suiesuiementssuresurementsments from a population representative of each species or variety cardinalis from cedros island in the green- house of the biology department university species volume ininuiA sugar of utah the greenhouse is free of pollinatorspollinators M cardcardinalisindis 39 115 I1 carefully hand pollinated one flower of each M vcrbenaceusverbenaceusverbenaverbenaceousceus 67 58 of ten pairs of red flowers and of ten pairs of M rupestrisrupestnsrupe stris 09 19019.019igo 0 M eastwoodiae 15 13713 7 yellow flowers the hand pollinated flowers of M ndsonnnelsoniinelsonelsonianiinil 18318 3 19219 2 both the red flowered and yellow flowered M lewisii plants set moderate numbers of seeds per cap- rocky mountains 05 05 sule 50 150 while the unpollinated flowers sierra nevada 07 113 set no seeds at all this finding corroborates my earlier observations on the cedros island range adjacent to new ecological opportuni- M cardinalis vickery 1990 that flowers do not ties A bright yellow flowered morph of M self pollinate and that pollinatorspollinators are required verbenaceusverbenaverbenaceousceus has appeared also and become for seed set well established in a population growing in an are the rewards for pollinatorspollinators the same in isolated spring area vasey s paradise at the yellow flowers as in red that is do yellow bottom of the grand canyon of the colorado flowers and red flowers produce equal volumes river AZ that species northwestern limit of nectar with the same concentrations of sug- flower colors in section erythranthe are ars red flowers of the cedros island M car- due to various combinations of six anthocyanin dinalis produced an average based on flowers pigments three pelargonidinspelargonidms apricot pink from 30 greenhouse grown plants volume of and three cyanidinscyamdmscyanidine lavendellavender pink and at 959.5 iiifitgl of nectar with 18218.2 sugar yellow flow- least one carotene pigment pollock et al 1967 ers produced an average based on measure- the lavender to magenta flowers of M lewisii ments of flowers from 40 greenhouse grown are due to variousvailous combinations of the pelar- plants of 10910.9log allui of nectar with 23023.0 sugar gonidin and cyanidin anthocyanin pigments there is so much variation that these values without the yellow carotene flowers of the red are not significantly different flowered species have all or most of the six finally the key question do pollinatorspollinators show anthocyanin pigments plus the carotene pig- a preference for red or yellow flowers to study ment red color results from a visual blend of this question I1 placed 24 red flowered and 24 pink pigments and yellow pigment yellow yellow flowered plants in a random arrange- flowered plants have a pair of recessive genes ment in a meadow in the red butte canyon at one locus that suppresses anthocyanin pro- natural area in the wasatch mountains be- duction pink pigments leaving just the yel- hind the university of utah and observed pol low carotene pigment showing so a single linatorslinatorelinators that visited this experimental popula- mutation when homozygous changes flower tion Pollinapollinatorstors that came were humming- colorfromcolor from red to yellow birds and bumblebees with rare visits from if the change from red to yellow flowers flies but no hawkmothshawkmoths or honey bees leads to a change in pollinatorspollinators for example Pollinapollinatorstors were observed for three 50sominmin from hummingbirds to bumblebees or hawk periods on each of 5 d on 28 july 1987 there moths then the first major step in reproduc- were 55 hummingbird visits to the 39 red tive isolation has been established by a single flowers present and 20 visits to the 35 yellow gene change when homozygous I1 once repro- flowers chi square 1437914.379 p ooi001001.001 ductive isolation has been established by color which indicates a significant preference for differences presumably selection would fine red flowers that day there were 10 bumble- tune it eg by favoring more tubular flowers bee visits to red flowers and 12 to yellow flow- for hummingbird pollinated flowers and by ers chi square 018180.1818 no significant pref- favoring a landing platform morphology and erence on 31 july there were 176 humming- nectar guides for bee pollinatedpolhnated flowers bird visits to the 42 red flowers in bloom that are pollinatorspollinators lequirrequireded for seed set in day in the population and 40 visits to the 21 mimulus cardinaliscardinally or do the flowers self polli yellow flowers chi square 7024670.246 p ooi001001.001 199511995 NOTES 179 which indicates a significant preference for for daily details of numbers and chi square red that day there were six bumblebee visits calculations on average bumblebees visited to red and one to yellow there were too few them 24 56 and 128 times respectively and bumblebee visits for a meaningful x2xa value to hummingbirds 43 98 and 52 times respective- be calculated the same pattern of three ly vickery 1990 bumblebees significantly observation periods was continued on 2 4 eschewed red and orange flowers and prefer- august but once again there were too few pol- entiaentiallylly visited yellow flowers hummingbirds linator visits to obtain meaningful x2xa values significantly preferred orange visited red apparently most hummingbirds had migrated flowers in proportion to their frequency in the south and there were few bumblebees all sea- population and significantly eschewed yellow son that year on the first day of the experiment flowers results for M verbenaceusverbenaverbenaceousceus are much when the plants had just been placed in the clearer than those for M cardinalis there is a meadow all pollinatorspollinators would be naive for both definite preference for yellow by bumblebees red and yellow flowered M cardinalis plants and a clear avoidance of yellow by humming- inasmuch as red butte canyon is hundreds of birds thus this color change has lead to sig- miles from the nearest M cardinalis popula- nificantnificant partial isolation between the normal tion in northern arizona therefore the highly orange and red flowered morphsmorphe and the yel significant preference for red appears to be low flowered mutant morph under the condi- real and not the result of learned behavior tions of this experiment apparently hummingbirds strongly preferred would M cardinalis react like M verbe the red flowers but also visited the yellow flow- nacelusnaceus in the better experimental locality at ers to some extent the few bumblebee visits the mouth of parley s canyon to probe this did not suggest a preference question I1 added red orange and yellow results show that the change in flower color flowered morphsmorphe of M cardinalis to the M from red to yellow did affect the frequencies verbenaverbenaceusverbenaceousceus red orange and yellow flow of pollinator visits but not in an all or none ered morphsmorphe of the previous experiment the way that would immediately establish repro- new experiment was run 8 17 august 1988 ductive isolation however the change would with the population being observed for 15 probably be enough to initiate partial incipi- periods of 1 h each at different times of day ent reproductive isolation on average there were 61 red 57 orange and would M verbenaceusverbenaverbenaceousceus with its normal red 22 yellow flowers ofmof M cardinalis see vickery morph and mutant yellow morph produce the 1990 for day to day numbers and chi square same reactions in pollinatorspollinators the flowers ofmof M calculations on average bumblebees visited verbenaceusverbenaverbenaceousceus differ from those of M cardinalis them 28 30 and 29 times respectively and in that only the upper two corolla lobes are hummingbirds 59 60 and 6 times respectively reflexed whereas all five of those of M cardi- bumblebees eschewed red and orange flowers nalis are reflexed both species sometimes and significantly preferred yellow flowers have wild populations with orange red flowers despite their low numbers in the population instead of the typical red flowers hummingbirds significantly eschewed yellow for the M verbenaceusverbenaverbenaceousceus experiment plants of flowers and preferentially visited orange flow- red flowered and yellow flowered individuals ers M verbenaceusverbenaverbenaceousceus plants were run again at from vasey s paradise in the grand canyon this time with M cardinalis plants and exhibit- plus plants of an orange red flowered popula- ed the same attractiveness or lack of attrac- tion from yecora sonora mexico were placed tivenesstiveness to the pollinapollinatorstors as before the pres- on a lawn by clumps of native gambel oak at ence ofmof M cardinalis flowers did not alter pol- the mouth of parley s canyon salt lake city linator response to M verbenaverbenaceusverbenaceousceus flowers UT this location had an abundance of pollina the color shift from red or orange to yellow tors in contrast to the paucity of pollinapollinatorstors in leads to marked partial reproductive isolation the red butte canyon meadow used previ- in M verbenaverbenaceusverbenaceousceus as well as in M cardinalis ously the test population was observed for 15 how effective is the partial reproductive periods of 1 h each at different times of day isolation to test this I1 placed 198 plants of from 26 july through 8 august 1988 on aver- M verbenaceusverbenaverbenaceousceus one sixth yellow flowered age there were 73 red flowers 87 orange flow- and five sixthssixtus red flowered to simulate a popu- ers and 136 yellow flowers see vickery 1990 lation with a well established mutant in four 180 GREAT BASIN naturalist volume 55 experimental areas the experimental garden instead of the 86 red flowered and 128 yellow on the university of utah campus red butte flowered seedlings actually observed this is a canyon natural area the mouth of parley s highly significant difference xax2 146730146.730 canyon and at silver fork big cottonwood p 0001 and greatly strengthens the point canyon salt lake county UT I1 harvested of pollinator faithfulness clearly pollinator seeds of each plant and planted seeds harvest- preference for yellow and faithfulness to yel- ed from 20 yellow flowered plants and grew low are having a large effect though not an them to flowering if pollinatorspollinators were visiting all or none effect we are seeing strong incipi- the flowers at random then they should pick ent reproductive isolation due to color change up and carry pollen from red flowers five times in different areas with different conditions more often than pollen from yellow flowers and different guilds of pollinatorspollinators the effect pollen loads and resulting seed sets were well might be less or might be stronger even lead- below the 500 1500 seeds per capsule that ing eventually to effective reproductive isola- may occur in M verbenaceusverbenaverbenaceousceus so results were tion and speciation not skewed by saturation of the stigma also assuming all else to be neutral such as relative acknowledgments growth rates of yellow and red pollen tubes speed of flowering of red and yellow flow I1 appreciate the financial support of the US ered plants randomness of placement of red national science foundation grant BSR and yellow flowered plants and sample size of 8306997 I1 thank dr stephen sutherland for red and yellow flowered plants then the nectar measurements and for carrying out the expected five to one visitation rate should red butte canyon experiment on M cardinalis hold inasmuch as red is genetically dominant to yellow then five sixthssixtus of the seedlings literature CITED should be red flowered and one sixth yellow CROW E AND M KIMURA 1970 an to flowered that is of the 214 seedlings grown J F introduction population genetics theory harper & row new 178 should be red flowered and 36 yellow york NY 591 appp flowered in fact there were 86 red flowered dewettdewetjDEWET J M J 1980 origins of polyploidspolyploidy pages 3 15 in seedlings and 128 yellow flowered seedlings W H lewis ed polyploidy plenum press oxford the ratio is 2 red to 3 yellow flowers which is GRANT A L 1924 A monograph of the genus mimulus annals of the missouri botanical gardens 11 99 389 far from the expected ratio of 5 red flowers to LEVIN D A 1993 local speciation in plants the rule not I1 yellow flower this suggests considerable the exception systematic botany 18 197 208 pollinator faithfulness to one color or the MAYR E 1976 evolution and the diversity of life other however in addition to pollinator faith- harvard university press cambridge MA 721 appp fulness could be self pollination POLLOCK H G R K VICKERY JR AND K G WILSON there 1967 flavonoidFlavonoid in mimulus cardinalis and does self pigments in mimulus cardinalis not pollinate but its related species I1 Anthocyananthocyaninsanthocyamnsins american journal M verbenaverbenaceusverbenaceousceus does at the average rate of 10 of botany 54 695 707011 seeds per capsule average normal seed set is VICKERY R K JR 1978 case studies in the evolution of 110 seeds per capsule therefore self pollina species complexes in mimulus evolutionary biology 11 404 506 tion would account for 9 of the yellow flow 1990 pollination experiments in the mimulus car- ered seedlings ie 9 of the 214 seedlings or dinalisdanalis M lewishlemishlewisii complex great basin naturalist 50 19 seedlings would be expected to be yellow 153 159 flowered as a result of self pollination of the VICKERY R K JR AND D sutherland 1994 variance and remaining 195 seedlings five sixthssixtus or 162 replenishment of nectar in wild and greenhouse populations of mimulus great basin naturalist 54 would be expected to be red and one sixth or 212 227 33 would be expected to be yellow therefore I1 should expect to observe 162 red flowered received 6 july 1994 seedlings and 52 ie 33 19 the results of accepted 27 september 1994 self pollination yellow flowered seedlings great basin naturalist 552 0 1995 appp 181 182 FALL LAMB production BY A california BIGHORN SHEEP matthew mccoylmccoy1mccoye walt bodie2bodied and elroy tadlootaylootaylor3

key words parturition california bighorn sheep ovis canadensis idaho

parturition is timed to maximize survival of big jacks creek during february and march offspring thompson and turner 1982 1988 four ewes were fitted with radio collars parturition occurring outside an optimum time and periodically located to monitor their period lowers reproductive fitness and there- movements and status three radio collared fore should be selected against timing of ewes were observed with lambs in may 1988 parturition in bighorn sheep ovis canadensis A fourth collared ewe 34 was observed with a has been related to resource abundance geist lamb A less than two weeks old based on size 1974 bunnell 1982 thompson and turner and behavior on 26 october 1988 an average 1982 risenhoover and bailey 1988 and climat- gestation period of 174 days for bighorn sheep ic conditions stewart 1982 parturition varies shackleton et al 1984 indicated conception by latitude between subspecies thompson occurred about 25 april 1988 ewe 34 and and turner 1982 and by elevation within sub- lamb A were located monthly through march species risenhoover and bailey 1988 peak 1989 during 1987 1990 onset of parturition lambing periods occur in march for desert occurred from 11 april to 3 may and mating bighorn sheep 0 c nelsoni hanson 1960 activity was observed between october and sandoval 1980 witham 1983 may for cali- december in an adjacent drainage the birth fornia bighorn sheep 0 c californianacalifornianscaliforniana jones of lamb A occurred approximately six months 1950 and early june for rocky mountain out of cycle ewe 34 was observed 4 january bighorn sheep 0 c canadensis bunnell 1982 1990 with a lamb B that appeared to have thompson and turner 1982 unusual lambing been born during the normal lambing period periods such as january for desert bighorn april june 1989 lamb B was conceived be- russo 1956 welles and welles 1961 and july tween october and december 1988 while ewe for rocky mountain bighorn stewart 1982 34 was nursing lamb A were attributed to extremes in climatic condi- ewe 34 may not have bred in 1987 or stress tions and elevations related to transplanting may have caused her vegetation in the big jacks creek drainage to abort stress can affect any aspect of repro- owaheeowyhee county IDTD is dominated by sage- duction deforge 1976 contact with rams brush artemisia sppapp bluebunchbluebunch wheatgrasswheatgrass during march and april 1988 may have caused pseudoroegneriapsezidoroegneria spicataspicatespicata and sandberg blue- ewe 34 to come into estrus presence of males grass poa sandbergsandbergiiii climatic conditions are has been found to induce estrus in female characterized by warm dry summers and cool merino sheep watson and radford 1960 and winters total precipitation from november feral goats coblentz 1980 recurrent estrus 1988 through march 1989 was equal to the 10 was observed in a cow elk cervus claphuselaphus year average however most precipitation that was associated with bulls but not bred occurred as rain in november and march ten during previous estrus periods morrison ewes one ewe lamb and three ram lambs 1960 from chilcotinChilcotin BC and two rams from east lamb survival has been related to forage fork owaheeowyhee river ID were transplanted to quality wehausen et al 1987 festa bianchet

lldaho11daho departmentdepaitment of fish and game 3101 S perlinePPowerlineeiline road nampa ID 83686 present address bureau of land management 3948 development avenue boise ID 83705 bahodaho department of fish and game 3101 S PoPowerimeeddmewerline road nampa ID 83686 3bureaubureau of land management 3948 development avenue boise ID 83705

181 182 GREAT BASIN naturalist volume 55

1988a precipitation patterns as they affect 1988b age specific reproduction of bighorn ewes plant growth douglas and leslie 1986 popu- in alberta canada journal of mammalogy 69 157 160 lation density and 1986 douglas leslie and GEIST V 1974 on the relationship of ecology and behav- mother s age festa bianchet 1988a cheatgrassCheatgrass lorior in the evolution of ungulaungulatesungulatedtes pages 235 246 in V brome bromus tectoriumtectorumtectorum seedlings were avail- geist and F walthers editors the behavior of anguungu able in november and sandberg bluegrass lates and its relation to management international greenup was observed in january south facinbacinfacing union conservation nature publication 9 HANSONHANSOM G 1960 lamb survival on the desert game slopes were generally free of snow soon after range desert bighorn council transactions 4 60 61 storms cattle grazing occurred in riparian JONES FE L 1950 A survey of the sierra nevada bighorn areas and on plateaus adjacent to drainages pages 29 76 in sierra club bulletin 1950 areas that received limited use by bighornsbighorns MORRISONMORBISON J A 1960 characteristics of estrus in captive elk behaviour 16 84 92 during summer and lambing periods bibighornb0rn 9 risenhoover K L AND J A BAILEY 1988 growth and mule deer odocoileus hemionus popula- rates and birbirthingberthingthing period of bighorn sheep in low tions were at low densities competition for elevation environments in colorado journal of forage was probably not a limiting factor mammalogy 6959269 592 597 festa bianchetblanchet 1988b reported that lambs RUSSO J P 1956 the desert bighorn sheep in arizona arizona game and fish department wildlife born to ewes four to nine years old had signifi- bulletin 1 cantly higher survival rates than those born to SANDOVAL A V 1980 preferred habitat of desert bighorn two to three year old ewes ewe 34 was esti- sheep in the san andres mountains new mexico mated to be five years old in 1988 mild cli- unpublished thesis colorado state university fort matic conditions availability of green forage collins 282 appp shackletonSHACKLE TON D M R G PETERSON J HAYWOOD AND A during weaning limited competition for for- BOTTRELLBOTTBELL 1984 gestation period in ovis canadeocanaden age and probable previous lambing experi- sis journal of mammalogy 65 337 338 ence allowed ewe 34 to raise an out of season STEWART S T 1982 late parturition in bighorn sheep lamb and survive concurrent fallwinterfall winter lacta- journal of mammalogy 63 154 155 THOMPSON R W AND J C TURNER 1982 temporal geo- tion and gestation periods this observation graphic variation in the lambing season of bighorn suggests that under favorable conditions sheep canadian journal of zoology 60 1781 1793 bighorn sheep may be able to successfully WATSON R H AND H M RADFORDRADFOBD 1960 influence of reproduce outside generally observed repro- rams on the onset of oestrous in merino ewes in the ductive periods spring australian journal of Agricultural research 2 65 71 WEHAUSEN J D V C BLEICH B BLONG AND T L literature CITED russi 1987 recruitment dynamics in a southern california mountain sheep population journal of BUNNELLBUNNLLL FE L 1982 the lambing period of mountain wildlife management 51 86 98 sheep synthesis hypotheses and tests canadian WELLES R E AND F B WELLES 1961 the bighorn of journaljouinal of zoology 560 1 14 death valley fauna of the national parks of the COBLENIZCOBLENTZ B E 1980 A unique ungulate breeding pat- united states fauna series 6 242 appp tern journal of wildlife management 44 929 933 WITHAM J B 1983 desert bighorn sheep in southwest- deforedeporeDEFORCE J R 1976 stress Is it limiting bighorn ern arizona unpublished dissertation colorado desert bixhornbighorn council transactions 19 30 31 state university fort collins 93 appp DOUGLAS C L AND D M LESLIE 1986 influence of weather and density on lamb survival of desert received 22 november 1993 mountain sheep journal of wildlife management accepted 20 june 1994 5015350 153 156 fesiaFLSIA BIANLIIEIBIANCIIET M 1988a nursing behavior of bighorn sheep concorrelateselates of ewe age parasitism lamb age birthbirthratebiithdatebirthdatedate and sex animal behavior 36144536 14451454144536144514541454 great basin naturalist 552 0 1995 appp 183 187 AGE GROWTH AND reproduction OF leatherside CHUB cliaclingilaGILAGZLA COPEI

jerald B Johnsonjohnsonl2johnson1212 mark C buobelkiBOO and dennis K ShiozawaShiozawal1

key words gila copei leatherside chub life history reproduction age growth

the leatherside chub gila copei is a small iratelyimately 1700 m main creek flows directly cyprinid native to eastern and southern areas into deer creek reservoir an impoundment of the bonneville basin of utah idaho and of provo river and was sampled in july 1993 wyoming to wood river idaho and to regions n 11 500 m upstream from the reservoir of the snake river idaho and wyoming 4024n40 24n lllsw128wellswI1 I1 at an elevation of 1650 m above shoshone falls baxter and simon 1970 chubs used for determining reproductive pat- simpson and wallace 1982 sigler and sigler terns below were collected from the latter 1987 gila copei is currently listed as a can- site in 1978 79 creeks at both locations flow didate for federal protection under the slowly at low gradient through meadows the endangered species act main creek site is downstream from beaver conservation efforts for G copei would dams tall grasses and small trees grow along benefit from accurate life history data yet the banks collections for both creeks were made life history of G copei is not well known this from vegetated pools separated by shallow species was thought to live less than five years riffles stream substrate is silt gravel and sigler and sigler 1987 based on bright col- boulders oration of males and abdominal distension in females sigler and miller 1963 concluded G MATERIALS AND METHODS copei spawns between june and august using similar coloration criteria on males from because cilacliagilaglia copei is a species of special sulphur creek WY baxter and simon 1970 concern our permit was limited to 40 speci- suggested breeding occurred in late summer mens and care was taken to collect the entire simon 1951 found females distended with size range following collection by elec eggs in early august other than these few shockingtrotroshocking fish were placed on ice and trans- observations no data on age growth or repro- ported to brigham young university BYU duction are available where they were stored frozen individuals we present data on age growth and repro- were then thawed rinsed in water blotted dry duction of leatherside chub from central utah and weighed 00010.001 g on a denver instrument these data were generated as a first step to XD 1200do1200d electronic balance standard understanding and protecting this potentially length SL was determined ooiool0010.01 mm using threatened endemic fish species fowler ultra cal lililillielileIII1111118 electronic calipers ages were determined by grinding otolithsotolithus STUDY SITE lapilus to a thin section and counting opaque bands under a leica dissecting microscope age and growth data were obtained from 40x opaque bands were validated as annuli 36 CG copei collected from thistle and main using a marginal increment analysis because creeks both tributatributariesries to larger rivers that juvenile ages 1 2 and adult fish ages 3 8 flow into utah lake thistle creek a tributary demonstrated distinct growth rates they were to spanish fork river was sampled in may evaluated separately identification of annuli september and october 1993 n 25 was facilitated by generating digitized images 3952n 11132w at an elevation ofofapproxapprox of otolithsotolithus on a video monitor using a hitachi

department of zoology brigham young university provo UT 84602 2address2addlesswaddress coicoreor i espondenceesponderespondencecorrespondencence to this author

183 184 GREAT BASIN naturalist volume 55 cctv6CCTV camera fitted to a Heerheerbruggbrugg wildwildo minnow traps hand nets and electroelectrofishingfishing dissecting microscope annual growth incre- gear and preserved in formalin daily temper- ments along the longest axis of the otolith ature was recorded from september 1978 to were then measured 000100010.001 mm using video july 1979 image analysis software mocha release 101.0iolo standard length was measured mm and jandel scientific rundel 1993 which reduces preserved wet mass ooi0010010.010 01 g was recorded for measurement errors introduced when reading each specimen monadsgonads from all n 176 otolithsotolithus directly under a microscope mcgowan individuals 50 minmm SL were removed and et al 1987 weighed 00010.001 g no fish 50 mm SL had en- size at age was back calculated from otolith larged monadsgonads A gonadosomatic index GSI measurements using a modified fraserleefraser lee was generated for each fish using the follow- formula campana 1990 ing formula andreasen and barnes 1975

L lo10L lcL lorxcorxLR RRrorcborc roR GSI gonad weight body weight X 100 where lxalxL is estimated SL at age x lclg is length mean monthly GSI values were used to deter- at capture rxarxR is otolith radius at age x and rerg mine onset and duration of spawning ova is otolith radius at capture lolp10 is estimated counts were made on nine fish collected in length at swim up estimated at 4 minmm from may 1979 the relationship between number data on gila atrariaatratwariaaria varley and livesay 1976 of ova present and SL was evaluated by linear and roR is otolith radius at swim up estimated regression from otolithsotolithus at ooi0010.01 mm there was no significant difference in back RESULTS calculated lengths at age I1 between main and thistle creek chubs main creek n 11 opaque bands on leatherside chub otolithsotolithus thistle creek n 25 T 1961.96 df 34 P appear to be valid annuli as demonstrated by 06og06.06 numbers of age II11 n 2 and age illIII111 an increase in the marginal growth increment n 3 fish from main creek precluded statis- throughout the growing season for both adult tical comparisons however back calculated and juvenile fish fig 1 lengths at age II11 and age iliillIII111 for main creek fish were within the range of comparably aged fish from thistle creek hence growth data for the two populations were combined an age 025u y growth curve was generated for the combined E ages 121 2 samples by averaging back calculated sizes at E 0 ages 383 8 age fE 020 chub collected in 1993 t 1 j leatherside were U sexed by dissection and examination of monadsgonads E 015 individuals lacking mature monadsgonads were classi- 1 fied .9 as juveniles immature testes were trans- 9 olo0100 10 lucent and threadlike while mature testes 750 9.9c were opaque white or pinkish and firm 20 reproductive states of ovaries were deter- roco 00505 mined according to criteria in holden and S berry 1983 immature ovaries were small 000 1 translucent and lacked yolked ova mature ovaries were larger and contained both imma- ture ova and firm yolked ova 4 5 6 7 8 9 10 11 reproductive data were obtained from a collection of 176 adult leatherside chubs month archived in the monte L bean museum at 1 mean marginal widths S BYU 5592 5619 5629 5686 5688 5775 fig I increment 2 SEE mea- sured from otolithsotolithus in cliagila copei n 36 immature age monthly collections from main creek august classes 1 2 and mature age classes 3 8 plotted sepa- 1978 to september 1979 were made using rately 199519951 NOTES 185

ages of 36 G copei collected in 1993 ranged 1979 and ranged from 938 in a 67 mm SL from one to eight years with SL of 38 110 59599 g female to 2573 in a 92 mm SL 146 g mm table 1 chubs grew rapidly to 58 mm female average count for leatherside chubs SL at about age II11 fig 2 table 1 from age collected in may 1979 was 1813 significant II11 on annual growth was slower and fairly correlations existed between SL and fecundity uniform mean GSI values for males and rar2 8282.82 P 05os05.05 n 9 and weight and females fig 3aaa were highest for both sexes in fecundity rar2 7272.72 P 0050.05 n 9 spring with maxima in may female GSI 12312.3 male GSI 272.7 increasing water tem- discussion peraperaturestures from january through may fig ab3b were associated with increased GSI values for A maximal age of eight years in our sample both sexes average water temperature in may of G copei indicates a life span much longer corresponding to GSI maxima was 949.4 C than previously thought sigler and sigler fecundity as measured by ovum counts in- 1987 longevity in G copei may be a life his- creased with SL for females collected in may tory trait that has evolved in response to living

TABLE 1 capture and back calculated standard lengths SL ofofgilagila copei from thistle and main creeks central utah

slatSL at capture mean 1 ackback calculatedcalciilatedalated SL at annulusis age N mean range 1 2 3 4 5 6 7 8

1 8 44 38 49 32 2 9 76 65 85 41 65 3 2 87 71 104 42 68 82 4 1 85 51 65 77 83 5 1 97 38 53 70 87 96 6 7 92 83 110 35 52 63 73 82 89 7 7 94 88 105 36 54 66 73 80 86 91 8 1 96 31 46 54 62 67 76 86 93 overall means 37 58 67 74 81 87 90 93

loo100

80

E E 60

C W 72 40 W

CD 20

0 0 1 2 3 4 5 6 7 8 age

fig 2 mean back calculated standard lengths at age 2 SE for cilacliagilaglia copei n 36 in central utah shaded block indicates estimated age at first reproduction 186 GREAT BASIN naturalist volume 55

a 18 coupled with the facts that the smallest fish with developed monadsgonads collected in 1993 was 16 19 female 65265.2gsg minmm SL and no fish in the museum collec- X 14 E male 50 mm had enlarged gonads sug- W tion min SL monads 12 gests that first reproduction inm G copei occurs C0 at age II11 10 cu0 high GSI in may followed by decreased 0r 8 GSI in june and minimal values in july and D aa 6 august fig 3a indicates that peak spawning occurred in may with some activity possibly 4 extending into early june gila copei appar- 2 ently follows a pattern of reproduction common to various cyprincyprinidsids living in temperate climates 0 munro et al 1990 this pattern is character- J A S 0 N D J F MMAA M J J A ized by the onset of spawning in late spring month followed by a period of gonadal recrudescence and inactivity size of monadsgonads begins to increase in autumn and continues through winter with D 1610 in b final 14 maturation occurring in early spring temperature influences the onset of 12 if aD 0 1 spawning differences in temperature as a 10 41431 0 function of latitude between main creek this 2CU 8 study and southwestern wyoming simon 0 0 0CL 6 1951 could explain the discrepancy between E Eu 4 onset of chub spawning at these locations may F 2 vs august A more detailed investigation of CG copei will be required to resolve questions 0 of differences in reproductive and life history 2 h characteristics among populations A J A S 0 N D J F M A M J J A literature CITED month ANDREASEN J D AND J R BARNES 1975 reproductive life history of catostomus ardensaddens and C discobolus in fig 3a3daa mean gonadosomatic indices 2 SES E for male the weberwebel river utah copeiacopela 1975 643 648 and gilagoia female cilaclia copei n 176 b mean monthly temper BAXTER G T AND J R SIMON 1970 wyoming fishes atuiesaturesagures 2 SES E from august 1978 to july 1979 in main wyoming game and fish department cheyenne creek wasatch county UT 168 appp CAMPANA S E 1990 how reliable are growth back calcacalcu lations based on otohthsotolithsotolithus canadian journal of fish- erieserles and aquatic science 47 2219 2227 in an environment where annual precipitation eries in HOLDEN M A AND C R BERRY 1983 vitellogenesisVitellogenesis in and stream flow vary considerably successful the utah chub gila atratrariaadranaatranaariaarla and its use in evaluating chub reproduction and recruitment may be reproduction in a transferred population Encyencycliaencyclicclia uncertainunceitdin in any given year an extended life 60 32 42 span would increase the likelihood that appro- MCGOWAN W FE E D PRINCE AND D W LEE 1987 an increase inexpensive microcomputer based system for making priate environmental conditions for reproduc- lapidrapid and precise counts and measurements ofzonaof zona tive success would be met at some time in an tion in video displayed skeletal structures in fish individual s life thus longevity may be a bet pages 385 395 in R C summerfelt and G E hall hedging strategy steamsstearns 1976 for living in editors age and growth of fishfisbfisaeishelsh iowa state univer- in press ames unpredictable conditions sity MUNRO A D A P SCOTT AND T J LAM 1990 repro- the growth pattern of G copei is typical of ductive seasonality in teleoststeleosts environmental influ- other fishes in which rapid juvenile growth ences CRC press inc boca raton FL 254 appp decreases at the onset of sexual maturity as ROFF D A 1984 the evolution of life history parameters in finite energy are allocated to both teleoststele osts canadian journal of fisheries and aquatic resources science 41 989 1000 growth and reproduction roff 1984 the RINDELRUNDEL R 1993 mocha image analysis software user s inflection point in the growth curve fig 1 manual jandel scientific san rafael CA 189 appp 199519951 NOTES 187

SIGLER W F AND R R MILLER 1963 fishes of utah STEARNS S C 1976 life history tactics a review of the utah department of fish and game salt lake city ideas quarterly review of biology 51 3 47 203 appp VARLEY J D AND J C LIVESAY 1976 utah ecology and SIGLEBSIGLER W FE AND J W SIGLER 1987 fishes of the great life history of the utah chub cilacliagila attatratrariaadranaatranaariaafiaafla in flaming basin a natural history university of nevada press gorge reservoir utah wyoming utah division of reno 425 appp wildlife resources publication 761676 16 salt lake SIMON J R 1951 wyoming fishes wyoming game and city 29 appp fish department cheyenne 129 appp SIMPSON J C AND R L WALLACE 1982 fishes of idaho received 15 june 1994 university press of idaho moscow 238 appp accepted 7 september 1994 great basin naturalist 552 0 1995 appp 188 191

consumption OF A TOXIC PLANT ZIGADENUS paniculatus BY MULE DEER

william S Longlandlonglandl2longland1212 and charlie ClementClementssl1

key words death camascomas Zigazigadenuszygadenusdenus mule deer poisonous plants coevolution

the abundance of green vegetation in nature dixon 1934 second in most cases of native can yield false impressions of the availability ungulaungulatesungulatedtes eating a plant species that is toxic to of food resources to herbherbivoresivores because many domestic animals the plant does not produce plants have evolved anti herbivore defenses noticeable toxic effects in the former indicat- defensive mechanisms commonly include ing that native herbherbivoresivores may possess detoxi- incorporation of distasteful or toxic secondary ficationfication mechanisms for some plant toxins lay- chemical compounds into plant tissues effects cock 1978 thus deer consume without adverse of different compounds on consumers range effects a variety of plants poisonous to live- from mild unpalatable to severe illness or stock stoddart and rasmussen 1945 dean death from poisoning Herbherbivoresivores have conse- and winward 1974 reciprocal examples in quently evolved a host of means for coping with which native plants are toxic to native cerbiherbi defensive compounds resulting in an evolu- vores but benign to domestic animals are tionary arms race between plants and cerbiherbi lacking in the literature vores freeland and janzen 1974 although herein we report on four years of observa- evidence of plantherbivoreplant herbivore coevolution can tions of an eastern sierra nevada mule deer be found for herbherbivoresivores ranging from phyto- herd feeding on substantial quantities of foot- phagous insects to mega vertebrates we con- hill death camas Zigazigadenuszygadenusdenus paniculatus a lilia- centratecentrate specifically on mule deer odocoileus ceous bulb plant that is toxic to domestic sheep hemoniushemonius feeding on toxic plants cattle and horses fleming et al 1921 kings- because domestic grazing animals lack a bury 1964 james et al 1980 panter et al 1987 coevolutionary history with the plant commu- the genus Zigazigadenuszygadenusdenus includes several species nities in which they forage they are often all containing toxic sterosteroidalasteroidalidal alkaloids james affected by toxic secondary compounds to a et al 1980 death camas emerges earlier than greater degree than native herbherbivoresivores this most plants making it particularly hazardous has significant economic impact on the range for spring grazing of livestock panter and livestock industry due to direct losses such as james 1989 these plants have been variously death reduced fecundity or reduced weight described as the most important poisonous gain and to indirect costs of minimizing such plants in the western US kingsbury 1964 losses nielsen et al 1988 james et al 1992 and the most dangerous poisonous plants in historical familiarity with local plant assem- north america clarke and clarke 1975 blages has provided herbherbivoresivores foraging in foothill death camas has been described as their native ranges with two advantages over one of the more toxic zigadenusZigazygadenusdenus species introduced domestic counterparts freeland kingsbury 1964 james et al 1980 and janzen 1974 laycock 1978 laycock et al our study site is located at t20n rise 1988 first native mammals often avoid eating s36 just west of reno NV on an alluvial fan at toxic plant species that are eaten by domestic the southern base of leavinepeavine mountain woody glazersgrazersgrazers for example toxic plants eaten by vegetation is dominated by basin big sage- livestock such as azalea azalea sppapp and lark- brush artemisia tridentata tritrldentata and bit spur delphinium sppapp are avoided by mule terterbrushterbruschbrush purshia dentatatritridentatatridentate death camas deer even when these plants are abundant emerges at this site in mid march flowers in

1usdaSDA agricultural researchRe seaichsearch service 920 valley road reno NV 89512 2addresswaddressaddi ess correspondence to thisthih author

188 199511995 NOTES 189 april and remains green into may A herd of ends of leaves alternately deer may occasion- mule deer usually numbering 20 25 animals ally sample plants in their environment free- has foraged extensively in this area from land and janzen 1974 and removal of short october to may since we began making obser- leaf segments may represent cautious sam- vations in fall 1988 pling of a plant deer find undesirable the lat- we first noticed deer consuming death ter possibility sampling seems less likely camas on 28 march 1989 before plants flow- than the former selectivity because we have ered and confirmed this with additional obser- observed individual deer feeding on several vations in all subsequent years examination death camas plants consecutively furthermore of death camas foliage immediately after deer total numbers of plants consumed on our tran- left the foraging patches consistently revealed sects were several orders of magnitude greater fresh herbivore damage we found that deer than the number of deer foraging in the study herbivoreherbherbivoryivory left a characteristic leaf damage pat- area and it seems unlikely that deer would tern with most or all leaves of a foraged plant have to sample repeatedly so many plants to cleanly bitten off perpendicular to their long discover they are undesirable axes in addition to direct observations of deer we found significant annual variation in the consuming death camas fresh deer pellet frequency of death camas consumption rang- groups were found in patches of plants ex- ing from 383.8 to 18918.9 of total plants counted hibitinghibiting this characteristic damage pattern showing evidence of deer herbivoreherbherbivoryivory G during all five springs 1989 1993 during 23289328232.8 df 3 P 0001.0001 table 1 maximum observation periods we found no evidence of and minimum percentages of plants eaten deer exhibiting toxic effects from death camas table 1 illustrate that frequency of herbivoreherbherbivoryivory consumption and neither we nor personnel also varied spatially in each of the four years from the nevada department of wildlife we sampled there was significant variation which surveys deer in the area by air have among transects in numbers of plants eaten found any fresh deer carcasses in the vicinity P 001001.001 for all years while the minority of each year from 1990 through 1993 we plants in the local death camas population walked 10 12 permanently located parallel were eaten the values in table I1 also represent transects and categorized all death camas a surprisingly high frequency of herbivoreherbherbivoryivory on plants seen as either eaten or uneaten by deer a plant species with such a notorious reputation transects were 500 m long 20 m wide ie we the relatively low proportions of damaged generally saw all plants occurring 10 m from plants indicate that deer may be selective for the transect lines and spaced 30 m apart particular death camas plants this is support- usually deer removed only the distal 2 5 cm ed by the fact that deer generally ate only a of leaves but on several occasions we found few non neighboring plants from large patches plants eaten to within 2 cm of ground level of death camas rarely did the majority of plants were considered eaten regardless of the plants within a patch show evidence of her amount of leaf removed we tested these data bijorybivory the apparently selective use of indi- for temporal differences in frequency of death vidual death camas plants significant tempo- camas consumption by comparing numbers of ral and spatial variation in death camas use eaten versus uneaten plants among the four and infrequent extensive herbivoreherbherbivoryivory on small years of the study using a G test of indepen- patches of plants could be due to variation dence we similarly tested for spatial effects among plants or patches in toxicity or to dif- on consumption by comparing eaten versus fering availabilities of superior foods leading uneaten plant counts among individual tran- to variation in the use of toxic foods sect lines within years our observations suggest that death camas is there are at least two potential explana- more palatable to deer than to domestic cattle tions for the partial consumption of leaves that or sheep domestic animals must be force fed we noted perhaps ends of leaves are less toxic death camas in captivity experiments fleming than leaf bases and deer preferentially con- 1918 fleming et al 1921 panter et al 1987 sume less toxic plant parts kingsbury 1964 and must be stressed or left with few alterna- suggests that death camas bulbs are the most tive foods in nature before they consume it toxic part of the plants and a gradient of panter et al 1987 mule deer at our study site decreasing toxicity could occur from bulbs to however occur at a low density and consume 190 GREAT BASIN naturalist volume 55

TABLETABLL 1 numbers and percentages of foothill death camas plants consumed by mule deer along 500soom m transects 1990 1993 at peavlepeavinepeavme mountain washoe county NV

number of plants 1 plantpiantplants eaten per transecttransactect number of yeaiyealyear transects total eaten maximum minimum XXSDSD 1990 12 2646 501 290 73 18618.618 6 10610.610log 6 1991 12 2726 259 447 26 16416 4 13413.413 4 1992 10 3073 118 326 16 83 96 1993 10 3799 202 154 25 80 40 includes combined datad it i koinflom illtiltiiallaliail transectstiantran sects death camas each spring although alternative because even limited past exposure of a plants are available because bitterness is a herbivore to a particular toxin can result in general property of alkaloids laycock 1978 reduced toxic effects selection for detoxifyingdetoxifying death camas is quite bitter most herbherbivoresivores rumen microflora may also account for intra- apparently find bitterness distasteful laycock specific variation in toxicity among individuals 1978 however bitterbrushbitterbrush purshia tntHdentata of a domestic species such individual varia- which is named for its bitterness is a pre- tion in susceptibility to death camas toxicity ferred browse plant of mule deer although has been reported in force feeding experi- bitterbitterbrushbrush is also consumed by domestic ments with domestic sheep fleming et al ungulaungulatesungulatedtes it is not highly preferred by them 1921 kingsbury 1964 perhaps it is possible to perhaps because bitterness is a greater feed- utilize this individual variation in selectively ing deterrent to domestic animals than to deer breeding for reduced vulnerability to particu- native herbherbivoresivores have been observed con- lar toxins currently most domestic grazing suming a variety of plant species known to be animals are products of artificial selection for toxic to domestic herbherbivoresivores laycock 1978 productivity rather than for resistance to envi- including an anecdotal report of mule deer in ronronmentalmental challenges utah consuming death camas and several other another avenue for applied research con- toxic plants stoddart and rasmussen 1945 cerns the possibility of ameliorating effects of recent work stimulates the interesting possi- toxic plants through the transfer of rumen bility that herbherbivoresivores consume specific toxic innocula from animals resistant to specific tox- plants to rid themselves of gut parasites ins to those that are susceptible jones 1985 barbosa et al 1991 gauld and gaston 1992 reported that transfer of rumen cultures from however this hypothesis only addresses why goats that were resistant to poisoning by leu- toxic plants are consumed rather than why the caena leucocephala to susceptible goats and consumers are physiologically able to tolerate steers eliminated adverse effects of leucaena the toxins although we can only speculate consumption in the previously susceptible ani- about reasons mule deer are less affected by mals this example suggests that even inter- death camas toxicity than domestic ruminants specific transfer of rumen fluids may effective- a likely explanation is that deer possess rumen ly reduce toxic effects in some cases microflora that have acquired the ability deer herbivoreherbherbivoryivory we witnessed on leavinepeavine through natural selection to detoxify this plant mountain may affect the demography of the freeland and janzen 1974 laycock 1978 local death camas population defoliation exper- such selection is perhaps to be expected for iments indicate that death camas probably suf- native ruminants because the microflora com- fers reduced reproductive output after her munity has seen prolonged exposure to native bijorybivory tepedino 1982 knapp 1986 while toxic plants it is certainly possible however plants adapted to herbivoreherbherbivoryivory may compensate that deer are able to detoxify death camas by for loss of biomass by allocating additional some other mechanism for example since energy to growth andor reproduction highly deer are browsers their diets include large toxic species instead employ an evolutionary amounts of tanninsfannins cooper and owen smith strategy of defense against herbivoreherbherbivoryivory and thus 1985 robbinsbobbins et al 1987 that may precipitate may not exhibit compensation catesgatescatesgates 1975 the alkaloids in death camas into a harmless laycock 1978 when such defenses are cir- tannate freeland and janzen 1974 cumcumventedvented by herbherbivoresivores with detoxification 199511995 NOTES 191 mechanisms toxic plants should experience ous to livestock in the western states USDAUS DA SEA reduced fitness agricultural information bulletin 415 JAMES L F D B NIELSEN AND K E PANTERPANTEB 1992 impact of poisonous plants on the livestock industry acknowledgments journal of range management 45 3 8 JONES R J 1985 leucaena toxicity and the ruminal we thank dr jeanne chambers dr kip degradation of mosinemimositemimosinemi pages 111 119 in A A panter and two anonym for seawright M PE hegarty L F james and R FE anonymousousons reviewers keeler editors plant toxicology proceedings of the thoughtful reviews of the manuscript this australiaaustralla USU S poisonous plant symposium brisbane paper is a contribution of the USDAUS DA agricul- queensland department of primary industries tural research service conservation biology yeerongpilly of rangelands unit reno NV KINGSBURY J M 1964 poisonous plants of the united states and canada prentice hall englewood cliffs NJ literature CITED KNAPP A K 1986 physiologyEcoecophysiology of zigadenusZigazygadenusdenus nuttalnettnuttnuttallnnuttalliinuttalliaallnliiiliill a toxic spring ephemeral in a warm season grassland BARBOSA P P GROSS AND J KEMPER 1991 influence of oecologiaoecologia7171 69 74 plant allelochemicalsallelochernicalsallelo chemicals on the tobacco hornhornwornhornworkworn and LAYCOCK W A 1978 coevolution of poisonous plants and its parasitoid cotesiacohesia congregatacongregatecongregata ecology 72 large herbherbivoresivores on rangelands journal of range 1567 1575 management 31 335 342 CATESGATES R G 1975 the interface between slugs and wild LAYCOCK W A J A YOUNG AND D N UECKERT 1988 ginger some evolutionary aspects ecology 56 ecological status of poisonous plants on rangelands 391 400 pages 27 42 in L F james M H ralphs and D B CLARKE E G C AND M L CLARKE 1975 veterinary nielsen editors the ecology and economic impact toxicology macmillan publishing new york NY of poisonous plants on livestock production COOPER S M AND N OWENSMITHOWEN SMITH 1985 condensed westviewWestview press boulder CO tanninsfannins deter feeding by browsing ruminants in a NIELSEN D B N R RIMBEY AND L F JAMES 1988 south africanafi leanican savanna occologiaOccooecologia67logia 67 142 146 economic considerations of poisonous plants on live- DEAN R E AND A H WINWARD 1974 an investigation stock pages 5 15 in L FE james M H ralphs and into the possibility of tansy ragwort poisoning of D B nielsen editors the ecology and economic blacktailedblackblackmailedtailed deer journal of wildlife disease 10 impact of poisonous plants on livestock production 166 169 westviewWest view press boulder CO DIXON J S 1934 A study of the life history and food PANTER K E AND L FE JAMES 1989 death camas early habits of mule deer in california part 2 food habits grazing can be hazardous rangelands 11 147 149 california fish and game 20 315 354 PANTER K E M H RALPHS R A SMART AND B DUELKE FLEMING C E 1918 range plants poisonous to sheep 1987 death camas poisoning in sheep a case report and cattle in nevada nevada agricultural experi- veterinary and human toxicology 29 45 48 ment station bulletin 95 ROBBINSBOBBINS C T S MOLE A E HAGERMAN AND T A FLEMING C E N FE PETERSON M R MILLER AND L H HANLEY 1987 role of tanninsfannins in defending plants WRIGHT 1921 death camas plants poisonous to against ruminants reduction in dry matter digestibil- sheep cattle nevada agricultural experiment ity ecology 68 1606 1615 station bulletin 101 STODDART L A AND D I1 RASMUSSEN 1945 deer man- FREELAND W J AND D H JANZEN 1974 strategies in agement and range livestock production utah agri- herbivoreherbherbivoryivory by mammals the role of plant secondary cultural experiment station circular 121 17 appp compounds american naturalist 108 269 289 TEPEDINO V J 1982 effects of defoliation on reproduction GAULD I1 D AND K J GASTON 1992 plant allelochemi of a toxic range plant zigadenuszigadennsZigazygadenusdenusdenns paniculatuspamculatus great cals tntritrltrophic interactions and the anomalous diver- basin naturalist 42 524 528 sity of tropical parasitoids the nasty host hypothe- sis oikosbikos 65 353 357 received 12 october 1993 JAMES L F R F KEELER A E JOHNSON M C WILLIAMS accepted 30 august 1994 E H CRONIN AND J D OLSEN 1980 plants poison great basin naturalist 552 0 1995 appp 192

USE OF AN UNUSUAL FOOD SOURCE BY ROCK WRENS troglodytidae

polly K Phillipsphillips1phillipse1 and alienallenailen E Sanborn2

keykeywordswords rock wrens food source foraging salpinctes obsoletusobsoletes troglodytidae feeding behavior

on 12 july 1993 we observed an interesting one possible explanation for our observations exchange between an adult rock wren is that the parent was tutoring the offspring salpinctessalpmctes obsoletesobsoletus and two juveniles while about the availability of food in vehicle grill- at toroweapTorow eap point on the north rim of the work tutoring and observational learning have grand canyon we observed an adult wren been documented in laboratory experiments in accompanied by two juveniles near our vehi- blackbirds mason et al 1984 tits sherry and cle all three birds walked beneath the vehicle galef 1984 1990 and pigeons palameta and by the rear wheel but the adult moved immeimme- lefebvre 1985 and have also been document- diately to the front end whereupon it hopped ed in the wild in other birds using usual food onto the front bumper and began to inspect the sources schaadt and rymon 1982 we have grill the adult found and ate an insect that no way of knowing whether the adult we had been trapped in the grillwork while stand- observed was attempting to teach what we ing on the bumper the adult began to vocalize believe were its offspring about an unusual after consuming the insect the juveniles food source we hope this observation will appeared to show a positive phototacticphonotacticphonotactic stimulate further study of feeding in fledgling response to these calls stopped foraging under birds with the possibility of discoveries in the rear of the vehicle and moved to the front social learning after the juveniles arrived at the front of the vehicle the adult continued collecting insects literature CITED from the grill the adult ate none of these insects but merely held them in its beak while FISHER J AND R A HINDE 1949 the opening of milk bottles by birds british birds 42 347 357 walking back and forth across the bumper buds the MASON J R A H ARZT AND R FE REIDINGER 1984 adult continued to vocalize periodically paus- comparative assessment of food preferences and ing to face the juveniles then it continued for- aversions acquired by blackbirds viavla observational aging in the grill it appeared to us that the learning auk 101 796 803 adult was showing the insects to the young PALAMETA B AND L LEFEBVRE 1985 the social trans- of a food finding technique in what the mission in pigeons neither of the young birds joined adult on is learned animal behaviour 33 892 896 the bumper however and within a few minmin- SCHAADT C P AND L M RYMON 1982 innate fishing utes the adult and juveniles flew off not to behavior of ospreys raptor research 16 61 62 return that afternoon As far as we have been SHERRY D FE AND B G GALEF JR 1984 cultural trans- mission without imitation milk bottle opening by able to determine this sort of acquired or birds animal behaviour 32 937 938 derived behavior has not been reported previ- 1990 social learning without imitation more about ously for rock wrens nor for any member of milk bottle opening by birds animal behaviour 40 the family troglodytidae 987 990 other observers have noticed birds taking received 13 june 1994 advantage of unusual food sources such as the accepted 16 november 1994 opening of milk bottles fisher and hinde 1949 there is generally a question however as to whether the behavior was by chance or learned

volosylolosy department miami dade community college north campus 11380 NWN W 27th avenue miami FL 33167349533167 3495 school Naturalofnaturalotof and health sciences banybarry university 11300 NEN E second avenue miami shores FL 33161669533161 6695 address reprint requests to this author

192 information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit cal natural history of western north america high quality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication coucouragedraged references IN THE TEXT are cited by author and SUBMIT manuscripts to richard W baumann date eg martin 1989 or martin 1989 multiple editor great basin naturalist 290 MLBM PO box citations should be separated by commas and listed 20200 brigham young university provo UT in chronological order use et al after name of 846024602020084602020084602 0200 A cover letter accompanying the man- first author for citations having more than two uscript must include phone numbers of the author authors submitting the manuscript and FAX number and acknowledgments under a centered main emailE mail address when applicable the letter must heading include special publication numbers when also provide information describing the extent to appropriate which data text or illustrations have been used in literature CITED also under a centered main other papers or books that are published in press heading lists references alphabetically in the fol- submitted or soon to be submitted elsewhere lowing formats authors should adhere to the following guidelines manuscripts not so prepared may be returned for mack G D and L D flake 1980 habitat rela- revision tiontionshipsships of waterfowl broods on south dakota manuscript preparation consult vol 51 no stock ponds journal of wildlife management 2 of this journal for specific instructions on style 44695 700 and format these instructions guidelines FOR sousa W PE 1985 disturbance and patch dynamics SUBMITTED GREAT BASIN manuscripts TO THE on rocky intertidal shores pages 101 124 in supply greater detail than those pre- naturalist S T A pickett and PE S white eds the ecolo- sented here form guidelines in condensed the are gy of natural disturbance and patch dynamics printed at the back of the issue additional copies academic press new york are available from theeditorthe editor upon request also coulson R N and J A witter 1984 forest ento- check the most of the great basin recent issue mology ecology and management john wiley for changes and refer to the CBE style naturalist and sons new york manual ath5th edition council of biology editors inc 669 appp one illinois center suite 200 111 east wacker TABLES are double spaced on separate sheets and drive chicago IL 60601429860601 4298 USA PHONE 312 designed to fit the width of either a single column 6160800616 0800 FAX 312 6160226616 0226 24 or a page use lowercase letters to indicate foot- TYPE AND DOUBLE SPACE all materials including literature cited table headings and figure legends notes FIGURES initially avoid hyphenated words at the righthandright hand margins photocopies OF are submitted underline words to be printed in italics use stan- with the manuscript editors may suggest changes dard bond 2222x28x 28 cm leaving 25 cm margimarglmarginsns on lettering on figures should be large enough to all sides withstand reduction to one or two column width originals must be no larger than 22 X 28 cm SUBMIT 3 COPIES oftheodtheof the manuscript and the oriorlorigi-91 22x28 nal on a 5255.25 or 35 inch disk utilizing WordwordperfectPerfect NOTES if the manuscript would be more appro- 424.2 or above number all pages and assemble each priate as a short communication or note follow the copy separately title page abstract and key words above instructions but do not include an abstract text acknowledgments literature cited appendices A CHARGE of 50 per page is made for articles tables figure legends figures published the rate for individual subscribers will TITLE PAGE includes an informative title no longer be 35 per page however manuscripts with com- than 15 words names and addresses of authors a plex tables andor numerous halfbalf tones will be running head of fewer than 40 letters and spaces assessed an additional charge reprints may be pur- footnotes to indicate change of address and author chased at the time of publication an order form is to whom correspondence should be addressed if sent with the proofs other than the first author FINAL CHECK ABSTRACT states the purpose methods results cover letter explaining any duplication of and conclusions of the research it is followed by information and providing phone numbers 6 12 key words listed in order of decreasing FAX number and emailE mail address importance to be used for indexing 3 copies of the manuscript and WordwordperfectPerfect TEXT has centered main headings printed inallindinallallailali diskette capital letters second level headings are centered conformity with instructions in upper and lowercase letters thirdthird4evellevellevei head- photocopies of illustrations ings begin paragraphs ISSNissn0017001736140017 3614 GREAT BASIN naturalist vovolvoi 55 no 2 april 1995 CONTENTS articles diets of young colorado squawfish and other small fish in backwaters of the green river colorado and utah robert T muth and darrel E snyder 95 invertebrate fauna ofofwastewaterwastewater ponds in southeastern idaho karen L cieminski and lester D flake 105 growth and reproduction in an alpine cushion plant astragalus kentrophyta var implexus wayne R owen 117 calileuctra a new genus and two new species of stonestonefliesflies from california plecoptera leuctridae W D shepard and R W baumann 124 carbon isotope discrimination in the c4ca4 shrub atriplex conferticonfertifoliafolia along a salinity gradient darren R sandquist and james R ehleringer 135 demography of astragalus scaphoidscaphoideses and effects of herbivoreherbherbivoryivory on population growth peter lesica 142 lahontan sagebrush artemisia arbuscula sspasp longicaulislongicaulis a new taxon alma H winward and E durant mcarthur 151 douglas fir tussock moth orgyia pseudotsugapseudotsugatata mcdunnough on subalpine fir in northern utah E matthew hansen 158 seasonal nutrient cycling in potamogeton pectinatuspectinatus of the lower provo river C mel lytle and bruce N smith 164 factors influencing fish assemblages of a high elevation desert stream system in wyoming bernard carter and wayne A hubert 169 notes speciation by aneuploidy and polyploidy in mimulus scrophulariaceae

I1 robert K vickery jr 174 speciation in mimulus or can a simple flower color mutant lead to species divergence robert K vickery jr 1771 77 fall lamb production by a california bighorn sheep matthew mccoy walt bodie and elroy taylor 181 age growth and reproduction of leatherside chub gila copei jerald B johnson mark C belk and dennis K shiozawa 183 consumption of a toxic plant Zigazigadenuszygadenusdenus paniculatus by mule deer william S longland and charlie clements 188 use of an unusual food source by rock wrens troglodytidae polly K phillips and alienallenailen FE sanborn 192