Full Issue, Vol. 50 No. 1

Great Basin Naturalist

Volume 50 Number 1 Article 16

3-31-1990

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

Recommended Citation (1990) "Full Issue, Vol. 50 No. 1," Great Basin Naturalist: Vol. 50 : No. 1 , Article 16. Available at: https://scholarsarchive.byu.edu/gbn/vol50/iss1/16

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]. dbwdbadk

T H E

GREATE AT BASINB A NMAUST

knemoe

aaa& A

VOLUME 50 N0 I1 MARCH 1990

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

associate editors

BRIAN A MAURER JIMMIE R PARRISH department of zoology department of zoology brigham young university brigham young university provo utah 84602 provo utah 84602 other associate editors are in the process of being selected

editorial board richard W baumann chairman zoology H duane smith zoology clayton M white zoology berranjerran T flinders botany and range science william hess botany and range science all are at brigham young university ex officio editorial board members include clayton S huber dean college of biological and agricultural sciences norman A darais university editor university publications james R barnes 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication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 1990 are 25 for individual subscribers 15 for student and emeritus subscriptions and 40 for institutions outside the united states 30 20 and 45 respectively 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 business should be directed to the editor great basin naturalist 290 MLBM brigham young university provo UT 84602 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian harold B lee library brigham young university provo UT 84602

EDITORIAL production STAFF joanne abel technical editor carolyn backman assistant to the editor heidi larsen production assistant

ISSN 017 01736143614 3909065045221390650452213 90 650630 45221 GREAT BASIN naturalist MEMOIRS SERIES

the great basin naturalist memoirs series was established in 1976 for scholarly works in biological natural history of greater length than can be accommodated in the great basin naturalist the memoirs series appears irregularly approval for the production is the respon- sibisibilitylity of the great basin naturalist editorial board information concerning the production of the memoirs series can be obtained from james R barnes editor great basin naturalist 290 MLBM brigham young university provo UT 84602

GREAT BASIN naturalist MEMOIRS all memoirs are available for purchase direct inquiries to editorial office attnanttn carolyn backman 290 MLBM brigham young university provo UT 84602 only checks or money orders drawn on USU S banks or US currency are accepted please include 2 per copy for mailing and handling charges

no I1 the birds of utah 10 no 2 intermountain biogeography a symposium 15 no 3 the endangered species a symposium 6 no 4 soil plant animal relationships bearing on revegetation and land reclamation in nevada deserts 6 no 5 utah lake monograph 8 no 6 the bark and ambrosia beetles of north and central america coleoptera scolytidae a taxonomic monograph 60 no 7 biology of desert rodents 8 no 8 the black footed ferret 10 no 9 A utah flora 40 no 10 areclassificationofthegeneraofscolytidaeA reclassification of the genera of scolytidae coleoptera 10 no 11 A catalog of scolytidae and platypodidaePlatypod idae coleoptera part 1 bibliography 30 no 12 research in the auchenorrhyncha homoptera a tribute to paul W oman 30 information FOR AUTHORS

the great basin naturalist welcomes previously references IN THE TEXT are cited by author and unpublished manuscripts pertaining to the biologi- date eg martin 1989 or martin 1989 multiple cal natural history ofwestern north america pref- citations should be separated by commas and listed erence will be given to concise manuscripts ofup to in chronological order use et al after name of 1200012 000ooo words first author for citations having more than two au- manuscripts SHOULD BE SUBMITTED to james R thors barnes editor great basin naturalist 290 acknowledgments under a centered main MLBM brigham young university provo utah heading include special publication numbers 84602 A cover letter accompanying the manu- when appropriate script must include phone numbers of the author literature CITED also under a centered main submitting the manuscript it must also provide heading lists references alphabetically in the fol- information describing the extent to which data lowing formats text or illustrations have been used in other papers in mack G D andland L D flake 1980 habitat rela- or books that are published in press submitted or tiontionshipsships of waterfowl broods on south soon to be submittesubmittersubmittedsubmitted elsewhere authors should dakota stock ponds journal of wildlife man- adhere to the following guidelines manuscripts not so prepared may be returned for revision agement 4469544 695 700 sousa W P 1985 disturbance and patch dynam- CONSULT THE MOST RECENT ISSUE of this journal acsonics on rocky pages for general style and format also refer to the CBE icson intertidal shores 101 124 S A pieketpickettpiekettpickettandtand S in T Picket and P eds 1 style manual ath5th edition council of biology edi- white I the tors 9650 rockville pike bethesda MD 20814 ecology of natural disturbance and patch dy- USA 24 namics academic press new york TYPE AND DOUBLE SPACE all materials including coulson R N and J A witter 1984 forest literature cited table headings and figure legends entomology ecology and management john avoid hyphenated words at righthandright hand margins wiley and sons inc new york 669 appp underline words be printed italics I1 to in use stan- TABLES are double spaced on separate sheetsheetss and dard bond 22 x 28 cm leaving 25 cm margins on designed to fit the width of either a single column all sides or a page use lowercaselower case letters to indicate foot- SUBMIT 3 COPIES of the manuscript number all notes pages and assemble each copy separately title photocopies OF FIGURES are submitted ini- page abstract and key words text acknowledg- tially with the manuscript editors may suggest ments literature cited appendices tables figure changes lettering on figures should be large legends figures enough to withstand reduction to one or two TITLE PAGE includes an informative title no lon- column width originals must be no larger than ger than 15 words names and addresses ofauthors 22 X 28 cm a running head offewer than 40 letters and spaces NOTES if your manuscript would be more footnotes to indicate change of address and author ap- propriatepropriate as a short to whom correspondence should be addressed if communication or note follow other than the first author the above instructions but do not include an abstract ABSTRACT states the purpose methods results A CHARGE of 45 per page is made for articles and conclusions of the research it is followed by published the for will 6 12 key words listed in order of decreasing im- rate subscribers be 40 per portance to be used for indexing page however manuscripts with complex tables andor numerous half tones will assessed TEXT has centered main headings printed in alallailali be an additional capital letters second level headings are centered charge reprints may be purchased at in upper and lowercaselower case letters third level head- the time of publication an order form is sent with ings begin paragraphs the proofs VOUCHER SPECIMENS authors are encouraged to FINAL CHECK designate properly prepare label and deposit cover letterietterlettietter explains any duplication of infor- high quality voucher specimens and cultures docu- mation and provides phone numbers menting their research in an established perma- 3 copiescoples36opies ofthe manuscript nent collection and to cite the repository in publi- conformity with instructions cation photocopies of illustrations ISSN 0173614017 3614 GREAT BASIN naturalist volvoi 50 no i1 march 1990 CONTENTS articles A plasma protein marker for population genetic studies of the desert tortoise xerobates agassiagassizizi james L glenn richard C straight and lacklaekjackjaek W sites jr 1 effects of nitrogen availability on growth and photosynthesis of artemisia tridentata sspasp wyomingensis paul S doescher richard F miller jianguojiangoo wang and jeff rose 9 form and dispersion of mima mounds in relation to slope steepness and aspect on the

columbia plateau I1 george W cox 21 esox lucius esocidae and stizostedion vitreum percidae in the green river basin colorado and utah harold M tyus and james M beard 33 influence of soil frost on infiltration of shrub coppice dune and dune interspace soils in southeastern nevada wilbert H blackburn and M karl wood 41 seed production and seedling establishment of a southwest riparian tree arizona walnut juglans major juliet C stromberg and duncan T patten 47 forage quality rillsealeofrillscaleof rillsealescale atriplex sticklesucklesuckiesuckleyistickleyiyi grown on amended bentonite mine spoil

I1 marguerite E voorhees 57 summer food habits of coyotes in idaho s river of no return wilderness area chcharleschariesarlesaries L elliott and richard guetig 63 infection ofyoung douglas firs by dwarf mistletoe in the southwest robert L mathiasen carleton B edminster and frank G hawksworth 67 new mexico grass types and a selected bibliography of new meximexicoco grass taxonomy kellykeilykellawkellywW allred 73 notes noteworthy mammal distribution records for the nevada test site philip A medica 83 formation of pisolithus tincttinctoriustinctoriousorius ectomycorrhizae on california white fir in an eastern sierra nevada mineminesoilsoil R F walker 8586 bone chewing by rocky mountain bighorn sheep K A keating 89 distribution of limber pine dwarf mistletoe in nevada robert L mathiasen and frank G hawkswohawksworthrth 91 sorex prebleiprebles in the northern great basin mark A ports and sarah B george 93 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 50 31 MARCH 1990 nolnoino I1

A PLASMA PROTEIN MARKER FOR population GENETIC STUDIES OF THE DESERT TORTOISE XEROBATESxerobatesagassizjAGASSIZI

james L clennglenngiennglenn1glenna 122 richard C straight 1 and jack W sites jr 3

ABSTRACT fifty seven individual plasma samples from desert tortoises xerobates agassiagassizizi representing 10 separate populations were analyzed by polyacrylamide gel electrophoresis using alkaline buffers an albumin like protein was found to be polymorphic for two electromorphs in northern populations inhabiting the mohave desert province while sonoran desert populations to the south were monomorphic the genetic divergence demonstrated in this survey is similar to earlier studies and provides evidence for the colorado river as a potential barrier to gene flow among tortoise populations these data suggest that tortoise plasma examined by various electrophoretic methods may provide a nondestructive means ofdetermining the broad regional origin ofdesert tortoises

desert tortoises xerobates agassiagassizizi pres- addressed is the question of genetic compati- ently inhabit two regions of southwestern bility of utah s tortoise populations especially utah separated east and west by the beaver if any of the tortoises east of the beaver dam dam mountains the population dynamics mountains are transtranslocatedlocated to the western slope of the beaver dam slope tortoises west of few data are available comparing the the mountains have been severely impacted physiology or morphology of utah s separate by both human and animal activities during tortoise populations rainboth et al 1989 the past several decades despite both fed- electrophoreticelectrophoreticallyally analyzed whole blood eral and state protective regulations tortoise homogenates of 146 desert tortoises from two numbers in the slope region are probably at an separate localities in california for allozymeallozyme all time low mike coffeen UDWR personal expression at 23 loci the two populations communication two stable populations near were quite similar as each locality contained st george east of the beaver dam moun- unique elements only when allozymeallozyme combi- tains face new human development projects nations were used and there was a consider- that threaten the future of these previously able degree ofoverlap in allozymeallozyme frequency isolated populations the eastern populations lamb et al 1989 included five tortoises from are found in paradise and city creek canyons paradise canyon in an analysis of phylogephylone and relocation of some tortoises from these agraphicographic patterns in mitochondrial DNA populations is presently under consideration mednamtdna of the gopher tortoise complex by the utah division of wildlife resources their data indicated that these individuals fit however several relocation issues remain within an eastern mohave clone represent- unresolved including methods of collection ing the northern arizona and eastern nevada conditioning transport sex ratios and speci- region also jennings 1985 included five men numbers involved one problem to be tortoises from the beaver dam slope region

benomvenomenom research laboratory veterans administration medical center salt lake city utah 84148 ogie2hogleogle zoological cardensgardengardens salt lake city utah 84148 department3department ofzoology brigham young university provo utah 84602 2 J L GLENNGLENNETALET AL volume 50

TABLE 1 genotypes and variability estimates for the polymorphic GP 1 locus resolved in 10 localities xerobatesofxerobatesof agassiagassiziagassiztzizt

number of GP 1 localityLO actyahty N genotypes A H 1 pima co AZ 3 3aaaaa 10 00 2 tucson AZ 6 gaa 10 00 3 finalfinaipinal co AZ 4 4aaaaa 10 00 4 maricolamaricopamarlMarimancopacopa co AZ 4 4aaaaa 10 00 5 kingman AZ 3 2aa1bb2aaaaa ibb 13 00 6 beaver dam slope AZ 12 9aa2abibbgaa gab ibb 15 0083 7 paradise canyon UT 12 3aa6ab3bb3aaaaa gab abb3bb 151.5isls1 5 0208 8 lincoln co NV 2 laa1aajabiablab 151.5isls1 5 0250 9 riverside co CA 4 laa1aa3abaab3ab 151.5lsis1 5 0375 10 san Bernardbernardinomo co CA 7 3aaaab3aaaaa aab4ab 15 0214 mean number ofalleleilocufofalleleslocus mean heterozygosity direct count in biogeographic investigations comparing in heparinized tubes 3 ml by venipuncturevenavenapuncturepuncture blood and tissue enzymes using horizontal jugular vein or antebrachial sinus from 15 starch gel electrophoresis his study included tortoises from the beaver dam slope AZ and no specimens from east of the beaver dam paradise canyon UT additional plasma sam- mountains in utah but showed that a north ples were donated by colleagues involved to south variation was evident and that the in desert tortoise projects in other parts of arizona beaver dam slope specimens fit AZ and in CA and NV localities and sample within the northern mohaveanMohavean group sizes for the populations sampled are listed in mitochondrial DNA and isozyme studies table 1 and their geographic locations are are costly and may involve traumatic to the plotted in fig 1 tortoise and labor intensive biopsies of inter- nal tissue or the sacrificing of specimens for Electrophoresiselectrophoresis necropsy the purpose of this study was to plasma samples were lyophilized and stored determine whether general proteins in tor- at 4 C polyacrylamide gel electrophoresis toise plasma could be used to detect geo- PAGE was run in a bio rad vertical slab graphical differences in tortoise populations electrophoresis cell model 220 the gel was our primary interest was to compare plasma 151.5isls mm thick sample wells 10 mm long were proteins among utah s aforementioned tor- formed with canalcocanaleo stacking gel 25252.5 acry toise populations using alkaline polyacry lamide the 7 acrylacrylamideamide separating gel lamidebamide gel electrophoresis however as elec- was 80 mm in length samples were elec trophoretic profiles of plasma or serum from trophoresed toward the anode at 20 ma xerobates agassiagassizizi have not been previously constant current and were stopped 10 mm reported we examined plasma profiles from from the bottom ofthe gel the electrophore- tortoises from arizona california and ne- sis buffer used was 000250.025025 M trisatris0tris0192Tris 0192192 M vada as well this report follows the taxo- glycine ph 838.3 bromophenol blue was used nomic grouping of gopher tortoises by bram- as tracking dye the gel was fixed in 10 ble 1982 and the nomenclature revision of acetic acid40acid 40 isopropanol50 water for 60 bour and dubois 1984 applying xerobates min stained with coomassiecoomassie blue 005oos0050.05 in as the genus for the desert tortoise 10 acetic acid10acid 10 isopropanol80 water for 60 min and destained in 10 acetic acid MATERIALS AND METHODS 10 isopropanol80 water using three or four changes of solution over a 24 hr period study area and sampling only qualitative differences were exam- A total of 68 desert tortoises were collected ined in this study and only the faster migrat- from 11 localities in arizona AZ california ing proteins were used since alkaline PAGE CA nevada NV and utah UT repre- is not the preferred method for resolving senting most of the range of this species in the differences in basic globulin like proteins united states plasma samples were collected due to their short migration distances the 1990 DESERT TORTOISE GENETIC STUDIES 3

N V 7 UT & AZ AB

1 I

CA n

1 1 1

fig 1 distribution ofcollectionofcollection localities oftortoise genotypes AA BB and AB listed in table 1 numbers indicate localities electrophoretic nobilitiesmobilitiesmobilities of rattlesnake zygosityzygosity H direct count and the mean num- crotalus atrolatrox plasma albumin and bovine ber of alleles per locus A the genetic dis- serum albumin were used as markers and tance and similarity coefficients of neinelnet 1972 compared with the albumin like proteins of 1978 and rogers 1972 were calculated for all the desert tortoise samples both albumin pairwise combinations of samples corrected markers migrated at faster rates than the for small samples sizes as described by tortoise albumin like proteins analyzed in levene 1949 and all such matrices were this study clustered by the UPGMA algorithm of sneath and sokal 1973 genotype from genetic analysis ratios the largest samples localities 6 and 7 n 12 for two protein loci identified in order of de- both were tested for conformance to hardy creasing anodal mobility were designated weinberg proportions by the Vx2xa goodness of general proteins GP I1 and 2 allelic data at fit option of BIOSYS again corrected for small both loci were recorded as individual geno- sample sizes levene 1949 types for analysis with the BIOSYS 1 program ofswoffordofswofford and selander 1981 measures of RESULTS genetic variability computed for each popula- sixty eight individual samples were ana- tion sample included average locus hetero lyzed from four states including 11 localities 4 J L GLENN ET AL volume 50

2 3

GP 2 GP 11

AA BB AB A BB AB

fig 2 alkaline polyacrylamide gel illustrating GPIGP I1 and GP 2 loci enclosed in dotted rectangle with allelic variation at GPIGP I1 locus for plasma samples from three individuals run in duplicate animals 1 2 and 3 were consistently scored as genotypes AA BB and AB respectively

TABLE 2 matrix ofgenetic distance coefficients ofseiofneiof neinel 1978 above diagonal and rogers 1972 below diagonal for all pairwise combinations of xerobates agassiagassizizi localities locality numbers are as listed in table I1 and D values are rounded off to two decimals

locality 1 2 3 4 5 6 7 8 9 10 I1 0000.00ooo 000 0000.00ooo 0030.03 ooi0010.01 0170.17 0000.00ooo 005 0060.06 2 0000.00ooo 0000.00ooo 0000.00ooo 0030.03 0010.01 0170.17 0000.00ooo 005oos0.05 0060.06 3 0000.00ooo 0000.00ooo 0000.00ooo 0030.03 001 0170.17 0000.00ooo 005oos0.05 0060.06 4 0000.00ooo 0000.00ooo 0000.00ooo 0030.03 ooi0010.01 0170.17 0000.00 005oos0.05 0060.06 5 0170.17 0170.17 0170.17 0170.17 0000.00ooo 0000.00ooo 0000.00ooo 000 0000.00ooo 6 008oos0.08 008oos0.08 008 0080.08 0080.08 008 0000.00ooo 000 ooi0010.01 7 0270.27 0270.27 0270.27 0270.27 0100.10olo 0190.19olg 001 0000.00ooo 0000.00ooo 8 0130.13 0130.13 013 00130.1313 0040.04 0040.04 015 0000.00ooo 0000.00ooo0 00 9 0190.19 0190.19 019 019olg0.19 0020.02 olo0100.10 0080.08 0060.06 0000.00ooo 10 0180.18 0180.18 018 0180.18 0010.01 0100.10olo 009 0050.05 ooi0010.01

considered as separate populations the pro- df I1 in both cases suggesting that this is a tein profiles present in 7 of 9 plasma samples simple mendelian dominantcodominantco system with from the desert tortoise natural area CA two alleles segregating in some populations and 2 samples from utah were unique but table 2 summarizes pairwise comparisons were very likely artifacts due to their badly of two genetic distance coefficients rogers hemolyzed condition these 11 samples were 1972 nei 1978 and shows that between excluded from the data analysis this reduced sample divergence was minimal neinelnetsnerss D val- the number of samples to 57 and the number ues for example range from 0000.00ooo to 0170.170 17 of localities to 10 table 1 general protein I1 four of the arizona samples maricolamaricopaMaricopa co was polymorphic for two electromorphs des- finalpinal co pima co and tucson are identical ignated A and B in several samples fig 2 nelneineinet s D 0000.00ooo five localities having the B table I1 summarizes the ratios of GPIGP I1 geno- allele at GPIGP I1 formed a distinct separate clus- types across these 10 localities and the esti- ter albeit the total degree of divergence from mates of variability across both loci geno- the monomorphic populations was slight D type ratios at localities 6 and 7 conformed to 0050.05oos0 05 within this group the arizona and hardy weinberg expectations xax2 27892.789 california populations were nearly identical xa P 095095.095 x2X 503503.503 P 478478.478 respectively while the paradise canyon UT samples were 1990 DESERT TORTOISE GENETIC STUDIES 5

141 4 5 9 LE1 10 6 8 7

1818.18 iglg16.16 1414.14 1212.12 loio10.10 0808.08 06og06.06 0404.04 0202.02 00oo00.00 rogers distance

fig 3 dendrogram based on rogers 1972 genetic distance values for 10 samples xerobatesofxerobatesof agassizi see table 1 fig 1 clustering was by the UPCMAUPGMA algorithm of sneath and sokal 1973 and the cophenecophenetictic correlation value was 08260.826

the most divergent these relationships are ern sonoran populations were monomorphic also visually displayed in the UPGMA den at the GP 1 locus an east west mohave dif- drogramgrogram presented in figure 3 using the ference was observed due to the eastern isola- statistical analysis method of rogers 1972 tion of the BB genotype table 1 fig 2 in all other UPGMA dendrograms gave identi- populations from the eastern mohave region cal or nearly identical topologies data not of utah and northwestern arizona fig 1 shown the B allele was not present in any of the central and southern arizona samples those discussion samples expressing the B allele may differ in frequency dessauer 1970 reported that while some the of this allele as suggested by blood proteins in reptiles are relatively the differentiation between paradise canyon conservative others are quite polymorphic and the beaver dam slope but the present many species can be readily distinguished by sample sizes are too small for accurate deter- the electrophoretic nobilitiesmobilimobilitiesties of blood pro- minationmination despite the small sample sizes the teins and certain subspecies and populations heterozygosity estimates are more similar for can also be distinguished by differences in the paradise canyon population and the three plasma albumin like proteins dessauer and mohaveanMohavean populations 8 9 and 10 than for fox 1958 masat and dessauer 1968 masat the paradise canyon and beaver dam slope and dessauer 1968 also found that the populations table 1 if the paradise canyon albumin like proteins of the testudinesTestudines have tortoises differ from the beaver dam slope slower migration rates in alkaline buffers populations in frequency of the B allele and than do these same proteins in most other are in fact more similar to california tortoises reptiles and mammals we also found that this could reflect a a divergence of allele under alkaline conditions the desert tortoise frequencies between the slope and paradise albumin like proteins migrate at slower rates populations in allopatry with allele frequen- than rattlesnake plasma albumin and bovine cies at paradise simply drifting to values serum albumin similar to california populations similarity the results of this investigation suggest by convergence b transport of tortoises geographical differences in genetic variability from california or nevada to st george and of the albumin like protein GP 1 of desert dumping into paradise canyon but not at tortoises the northern mohaveanMohavean popula- the slope ie human induced gene flow tions were polymorphic whereas the south between california nevada and paradise or 6 JjlglennetalL GLENN ET AL volume 50

c transport and release of arizona tortoises duceddeuced status is based on three general obser- most with AA genotypes at the GP- I locus on vationsvations one is the fact that some tortoises the beaver dam slope but not paradise found in the st george region obviously have canyon which would cause the A allele at the been captive specimens exhibiting rope or slope to increase in frequency at the expense chain holes in their shells or having painted of the B allele and push the slope population areas on their shells second residents of away from paradise canyon and california washington county often keep tortoises as allele frequencies see genotype differences in pets and some have escaped or were pur- table 1 these are not mutually exclusive posely released third the beaver dam and hypotheses since a certain amount of mixing virgin mountains presently form an east west may have occurred at all localities of califor- barrier between natural assemblages and a nia nevada beaver dam slope and paradise few reptiles found west of the mountains are canyon populations not present east of them ege g crotalus scutuscurtu the long history of human collection and latus phyllorhynchus decurtisdecurtusdecurtus dipsosaurus translocation of desert tortoises between dorsalis the possibility still remains that states constitutes a variable that could influ- some of utah s tortoises found east of the ence the genetic structure of desert tortoise mountains may be derived from ancestral populations especially when comparing al- stock of naturally occurring tortoise popula- lele frequencies between populations well tions that have since mixed with captive re- known as captive release sites hundreds leased specimens support for the natural perhaps thousands of tortoises have been population relies on the fact that the mohave picked up along roadways and released in dif- desert province extends into this region and ferent regions over the past several decades many other sonoran life zone animals found mike coffeen and eric coombs UDWR on the western slope are also found east of the personal communication this logistical dis- mountains and are natural assemblages like placement of tortoises continues at present the desert tortoise some of the reptiles are certain localities have been popular release life zone specific and occur on both sides for sites eg regions of southern california example the banded gecko coleonyxColeonyx varie desert tortoise natural area arizona gatus gila monster heloderma suspectumsuspectum mcdowell mountain region near phoenix and sidewinder crotalus cerastes nevada near las vegas and recreation areas the geographical differences observed in and utah st george the most common this investigation are similar to those found in avenues of translocation by motorists crossing the allozymeallozyme survey by jennings 1985 the the mohave desert are east to west and vice mtdnamedna survey by lamb et al 1989 and the versa favorable habitats for tortoises exist morphometricmorpho metric analysis of tortoise remains by from washington county in southwestern weinstein and berry 1987 specifically the utah to southern california a region heavily present study supports the earlier molecular traveled over a major interstate highway for investigations of lamb et al 1989 which decades in addition to the release of tortoises showed divergence between tortoise popula- in this region by motorists and local citizenry tions north and west ofthe colorado river and utah s beaver dam slope populations have those to the south and east their report pro- been the site of approximately 200 captive vided good evidence that tortoise populations release tortoises regulated by the utah divi- now isolated on opposite sides of the colorado sion of wildlife resources since 1970 mike river have likely been separated from each coffeen UDWR personal communication other for several million years the mtdnamedna although utah s tortoise populations lo- lineages from central and southern arizona cated east of the beaver dam mountains are formed a single haplotype that differed from often regarded as captive escapees in a the northern haplotypes in CA NV UT and indigenousnonnonindigenous setting there are no scientific extreme northwestern AZ by a minimum of 17 data confirming this view the hypothesis restriction site changes see fig 2 in lamb seems to have originated from several et al 1989 this is one of the highest levels sources eg magazine and newspaper arti- of intraspecific genetic divergence reported cles and the opinions of a few naturalistsnaturalists for any animal species and exceeds that and herpetologists support for the intro reported for many interspecific comparisons 1990 DESERT TORTOISE GENETIC STUDIES 7

the exception in our study was the small ble exception of locality 5 noted above very sample from near kingman AZ locality 5 in similar to the findings of others in that the fig 1 which genotypically grouped with the broad regional genotypes in desert tortoises mohaveanMohavean populations north and west of the are approximately concordant the PAGE colorado river consequently the genotypic screening of the plasma protein marker may composition for population 5 must be inter- provide one inexpensive method of objec- preted with caution the single BB homo- tively determining the regional origin of zygote in a sample of three individuals would tortoises if allele frequency data are to be not be expected unless the B allele was segre- used additional specimens from paradise gating at a high frequency these results may canyon and beaver dam slope populations be due to several factors eg the transloca- are needed to determine the significance of tion of this specimen by humans a sampling the allele frequency differences between error for a low frequency allele or a degrada- these two localities additional samples from tional artifact in this sample if future sam- throughout nevada and california would also pling verifies the presence of a high be required also some variations observed frequency B allele at this locality it could in the slower migrating proteins could be ex- represent an ancestral polymorphism shared amined with more high resolution techniques with mohaveanMohavean populations north ofthe colo- ege g isoelectric focusing two dimensional rado river this anomalous result under- electrophoresis combined with bio image an- scores the need for statistically adequate sam- alyticalalytical instrumentation that can quickly and ple sizes in all future genetic studies of the accurately scan and record qualitative differ- desert tortoise therefore identifying the ences in electrophoretic profiles morpho specific origin of any individual tortoise on metric data from live tortoises should be col- the basis of nuclear gene markers may be lected to compare paradise canyon and difficult weinstein and berry 1987 suggest western slope populations in conjunction with using a combination ofphysiological and mor future molecular analyses in fact external phometricphometric screening methods to designate re- morphology such as shell shape could be a gional types they analyzed shell morphology more functional criterion for the survivability of adult 180 mm desert tortoises of both of paradise canyon tortoises on the beaver sexes by using morphometricmorphometric data gathered dam slope see weinstein and berry 1987 by the bureau of land management at 31 since genetic differences between these pop- different localities these measurements were ulatulationsions appear to be slight collected from tortoise remains by several persons over a 48 year period these authors acknowledgments noted that live tortoises were not used in their analysis and that shell morphology of tortoise we greatly appreciate the cooperation of remains does incur some shrinkage over time the following individuals for their efforts in following death they recommend further supplying plasma samples used in this study studies comparing live tortoises none of trip lamb savannah river ecology labora- these authors suggested that the differences tory aiken south carolina randy jennings observed in their investigations justify sub- university of new mexico albuquerque jim specific designation for any of the regional jarchow neglected fauna international populations and thus xerobates agassiagassizizi re- tucson arizona brian henen ken nagy mains a monotypic species and charles peterson laboratory of bio- alkaline PAGE was useful in examining the medical and environmental sciences uni- albumin like proteins in tortoise plasma but versity of california los angeles and paula does not resolve slight differences in the elec- acer kingman animal hospital kingman trotrophoreticlphoretic mobility among the majority of arizona trip lamb and ken nagy also the plasma proteins however this method reviewed this manuscript and offered valu- did detect the polymorphic nature ofthe albu able comments for which we are grateful min like GP 1 protein and this protein may michael coffeen utah division of wildlife be one marker that could be used for desig- resources was instrumental in facilitating the nation of broad regional types since the re- fieldwork involved in this study we also sults of this investigation are with the possi thank martha wolfe veterans administration 8 J L GLENN ETALET AL volume 50

medical center salt lake city utah for her graphic patterns in mitochondrial DNA of the technical skills and appreciate the coopera- desert tortoise xerobates agassiagassiziagasstzizi and evolution of cecil schwalbe and terry johnson tionary relationships among the north american game gopher tortoises evolution 43 76 87 arizona and fish department and LEVENE H 1949 on a matching problem arising in randy radant and tim provan utah division genetics annals of mathematical statistics 20 of wildlife resources this research was sup- 91 94 ported in part by the utah division ofwildlife MAsMASAT R J AND H C DESSAUERDESSAUEB 1968 plasma albuminsalbu mins of reptiles comparative biochemistry and physi- resources and hogle zoological gardens ology 25 119 128 salt lake city utah NEI M 1972 genetic distance between populations american naturalist 106 283 292 1978 estimation of average heterozygosity and literature CITED genetic distance from a small number of individuals genetics 89 583 590 BOUR R AND A DUBOIS 1984 xerobates agassiz 1857 RAINBOTH W J D G BOTHBUTH AND F B TURNER 1989 synonymesynonymysyno nyme plus ancien de scaptochelys bramble allozymeAllozyme variation in mojave populations of 1982 reptiliaReptthaiha shelomicheloniichelomiCheloniilomi testudinidae bulletin the desert tortoise gopherusGopherus agassizizt copela mensuelmansuel de la societe lmneennelinn6enne de lyon 53 19891151989 115 123 30 32 ROGERS J S 1972 measures of genetic similarity and BRAMBLE D M 1982 scaptochelys generic revision genetic distance studies in genetics university and evolution of gopher tortoises copela 1982 of texas publications 72 145 153 852 867 SISEATHSNEATH P H A AND R R SOKAL 1973 numerical tax- DESSAUER H C 1970 blood chemistry ofreptiles physi- onomy W H freeman san francisco evolutionary ological and evolutional y aspects appp 1 54 in C SWOFFORD D L ANDANDRR B SELANDER 1 gans 1981 BIOSYS cans and T S parsons eds biology of the rept- a FORTRAN program for the comprehensive anal- press ilia vol 3 academic new york ysis of electrophoretic data in population genetics dessauer H C anawannwAN 1 W fox 1958 geographic variation and systematics journal of heredity 72 281 283 plasma in protein patterns of snakes proceedings WEINSTEIN M N ANDANDKK H BERRY 1987 morphometncmorphometricMorphometricmetrie of the society of experimental biology and analysis of deseideseldesert t tortoise populations bureau of medicine 98 loi101101105losios105 land management report ca950 ctrct7 003 riv- JENNINGS R D 1985 biochemical variationvanation ofthe desert erside california 39 appp tortoise gopherusGopherus agassiagassizizi unpublished thesis university ofnew mexico albuquerque 72 appp received 5 november 1989 LAMB T J C AVISE ANDJAND J W GIBBONS 1989 phylogeo accepted 16 january 1990 great basin naturalist soi501 1990 appp 9 19 EFFECTS OF NITROGEN availability ON GROWTH AND photosynthesis OF ARTEMISAARTEMISIA tridentatatridentateDENTATATRI SSP wyomingensis

paul S doescherdoescher11 richard F millermiller2millera2 jianguojiangoo wang 2 and jeff rose2rosea

ABSTRACT this study examined the effects of alterations in soil nitrogen on the growth ofaof rtemisiaArtemisia tntritrldentata sspasp wyomingensis nutt soil nitrogen content was altered by applying sugar 45 grnarngm2gme nitrate 45454.54 5 gm2gme or ammonium 45454.54 5 gm2gme and the results were compared with a control treatment no soil amendments addition of either form of nitrogen significantly increased leaf nitrogen content mean maximum length of ephemeral leaves number of ephemeral leaves per terminal shoot and current year s vegetative stem length over the control and sugar treatments both soil water and predawn xylem potentials during active growth were lower in the nitrogen treated plots the higher growth activity and greater leafmassleaf mass ofaof A tntritrldentata in the nitrogen treatments may have been responsible for this result higher photosynthetic rates observed in the nitrogen treatments during an early june sampling period also lend support to this observation this study suggests A dentatatntritrltndentatatridentatatridentate sspasp wyomingensis would opportunistically take advantage of increasedinci eased availability of soil nitrogen the ability of this species to respond positively to increased soil nitrogen may enhance its competitiveness over associated perennial species

artemisia dentatatritridentatatridentate nutt is a semidecidu volume much larger than that of perennial ous perennial shrub occupying 44844.8 million grasses sturges 1977 ha in the western intermountain sagebrush relatively little research has examined the steppe it is the most abundant shrub in this response of this species to soil nutrients such ecosystem during the past century in- as nitrogen limited work however indi- creases of A dentatatritridentatatridentate have been attributed cates A tridentata to be an effective competi- to overgrazing of perennial grasses by domes- tor for soil nutrients caldwell et al 1985 tic livestock cultivation of lands too andaridarld to demonstrated that this species successfully produce crops and alterations in fire fre- competes for soil phosphorus with the native quency hironaka and tisdale 1963 tisdale perennial grass agropyron spicatumspicatum pursh et al 1969 tisdale and hironaka 1981 As scribn & smith the accumulation of nutri- A tridentata has increased in the great ents and higher soil nitrogen mineralization basin both production and diversity of herba- rates in surface soils beneath A dentatatritridentatatridentate ceous understory species have declined nu- canopiescanopies may also convey an ecological advan- merous physiological and morphological char- tage to plants during active growth periods acteristicsacteristics ofaofA dentatatritridentatatridentate have been shown to charley and west 1975 1977 doescher enhance its effectiveness as a competitor with et al 1984 few studies however have eval- native bunchbunchgrassesbuncligrassesgrasses especially for soil mois- uated the response of A tridentata to in- ture depuit and caldwell 1973 eissenstat creased or decreased amounts of available soil and caldwell 1988 miller and shultz 1987 nitrogen carpenter 1972 working in the miller 1988 among these the ability of A colorado plateau reported that 134 kg nha dentatatritridentatatridentate to maximize leaf area early in the applied to A tridentata yielded an 81 in- growing season by overwinteringoverwintering one third of crease in total leafy material compared with a its leafleafbiomassbiomass and by developing ephemeral nontreated control however carpenter and leaves early in the spring strongly enhances west 1987 found little response to nitrogen its ability to photosynthesize during favorable in A dentatatritridentatatridentate grown on mine spoils the growth periods depuit and caldwell 1973 form of nitrogen whether nhanh4NH or no3n03noa may miller and shultz 1987 miller 1988 A deep also be an important factor in the mineral well developed root system also allows A tri nutrition of fridlandaridland shrubs wallace et al dentata to capture soil moisture from a soil 1978

department ofRangeland resources oregon state university corvallis oregon 97331 heastern2easterneastern oregon agricultural research center burns oregon 97720

9 10 PSP S doescheretalDOESCHER ETALET AL volume 50

our experiment was designed to determine lated sugar 45 gmgm2gme 3 ammonium how depletion or addition of different forms of nh4is04nh42s04 nitrogen 45 gm2gme and 4 nitrogen affects A dentatatritridentatatridentate growth and car nitrate hn03 nitrogen 45 gmgm2gme2 sugar bon assimilation rates our hypothesis was addition was assumed to increase the CNC N that A tridentata responds favorably to in- ratio to decrease availability of soil nitrogen creases in soil nitrogen baathetalbaathBaatbaachhetalet al 1978 both ammonium and sugar were broadcast onto the 5 X 5 m plots ni- MATERIALS AND METHODS trate was diluted in water 1 I1 part hn03 to 5 parts water and applied with a backpack the study was conducted at the squaw sprayer all herbaceous plants were dormant butte experimental range in southeastern at the time of application soil and plant oregon 11943w longitude 4329n lati- growth measurements were recorded during tude 67 km west of bumsburns on the northern the following 1987 growing season fringe of the great basin the 37 year mean soils were analyzed for ammonium and ni-ni annual precipitation for this area is 284 mm trate concentration in the A and B horizons in precipitation during the 1987 crop year sep- five plots per treatment on 14 april 26 may tember august was 296 mm the squaw and 25 july soil analysis was performed using butte experimental range typically receives a kcfkc1KCI extracting technique horneck et al in most of its moisture between october and press june generally as snow with little precipita- soil water content was measured from tion received in july and august the mean 1 april to 15 september once every two weeks temperature in winter is 060.6og C with the in each of the A and B horizons one soil daily minimum averaging 484.8 C and the sample was collected for each of the two mean temperature in summer is 17617.6 C with depths within each plot for all treatments soil the daily maximum averaging 26826.8 C the water was measured gravimetrically and soil study site is located in an artemisia tridentata water release curves were developed for each sppapp wyomingensisstipawyomingensislstipa thurberianathurberiana habitat depth to define soil water potential type at an elevation of 1372 m doescher et ephemeral leaf number and maximum al 1984 this site has been excluded from length and vegetative stem elongation were grazing by domestic herbivoresherbivores for the past 40 measured on five randomly selected terminal years soil texture is gravelly fine sandy loam branchletsbranchbranchlesslets of the single artemisiaArtemista located and classified as xerollicXerollic Durothdurothidsids lentz within each plot leaf measurements were and simonson 1986 soils vary in depth from recorded on three dates during initiation and 35 to 45 cm and are underlain by an induratedindurated expansion of ephemeral leaves 15 april to duripandurigan 5 20 cm thick which is underlain by 5 may vegetative stem elongation was mea- unweathered basalt A detailed description of sured on five dates from initiation to termina- soil nutrient levels is provided by doescher tion of growth leaf nitrogen content was et al 1984 measured on current year s leaves both ephemeral and collected from experimental procedures persistent vegetative stems on 15 and 21 april I1 june and A completely randomized plot design was I1 august on all plots collections represented used with 10 replications of each treatment three phenological stages initial leaf elonga- plots 5 X 5 m were laid out with an A triden tion rapid leaf and stem growth and early tata located in the center of each plot to flowering the semimicro kjeldahl method maximize uniformity we selected plots that was used to determine total leafnitrogen con- had vigorous appearing A dentatatritridentatatridentate plants of tent bremmer 1965 specific leaf weight g similar growth form and size plant measure- mm2ma was obtained by measuring leaf area on ments were recorded on the center A triden current year s green leaves on 12 dates during tata plants and soil measurements were col- the growing season leaves were removed lected within 151.5 m of the stem base the from one randomly selected terminal branch remainder of the plot was used as a buffer in each plot and placed in a damp cooler treatments were applied both in march several hours later leaf area was measured on and late november of 1986 treatments were a leafarea meter and weight was determined 1 control no amendments added 2 franugranu by oven drying the leaves at 60 C for 48 hr 1990 EFFECTS OF NITROGEN availability 11

TABLETABLEI 1 soil nitrate and ammonium content ppm at soil depths ofoof 0200 20cm20 cm and 20 40cm40 cm treatment sampling date control sugar nitrate ammonium n03NO 0 20 cm april 14 050 048 124212 42b 120 a may 26 i1 16 1 14 684 116ilg1 16 july 25 182a1 82 3283.28328a 11 6464b 5502au02ab 20 40 cmern april 14 007474 084a0 84 2060b20 6011 1 64 may 26 1 56 1 10ioa 846846b 1 46 a july 25 1621.621 62 1 90goa 90290211b 3463.46346alA nhi 0 20 cmcraeraern a aprilprii 14 632 4184.184 18 5145 143a 778 may 26 492 482 7227.227 22a 492 july 25 080 007070 330000a oto07007010771 20 40 cmem april 14 788 682 10 183 1232a may 26 896 768 110411.0411 04 119611 96a a july 25 1041.041 04a liolloiloiio1101.101 W 284 930 numbers followed by the same letters are not significantly different P 05 between treatments for each soil depth and date

xylem potentials scholander et al 1965 fied using analysis of variance procedures waring and cleary 1967 on current year s time was set as a variable in addition to treat- vegetative branchbranchletsbranchlesslets were measured during ment least significant differences LSD the 1987 growing season with a pressure P 05 were calculated only when the F chamber PMS corporation corvallis ore- value was significant P 05 steel and tor- gon predawn 19 may 3 june and 21 july riene 1980 the general linear model proce- and midday 15 april and 3 june measure- dure of SAS was used to evaluate treatment ments were recorded between 0430 and 0630 differences for specific leaf weight and photo- 1130 and 1230 hr respectively five branch synthesis SAS 1988 this procedure permit- lets were measured in each treatment sam- ted statistical analysis of unbalanced data sets ples were selected at random removed from both interactions and main effect means were the shrub and immediately measured in the separated using fischer s least significant pressure chamber difference test only statistically significant photosynthesis was measured on one ran- results are reported in the results and dis- domly selected vegetative branchlet of an cussion artemisia plant in 5 of the 10 plots for each treatment on eight dates from mid april RESULTS through early august measurements were recorded between 1200 and 1300 hr using a soil nitrogen LI 6000 LICORLI COR lincoln nebraska por- the addition of nitrate increased nitrate table photosynthesis meter with a quarter levels 25 and 28 fold in the upper and lower liter chamber to attain an adequate amount soil depths respectively at the beginning of of leaf area in the chamber we recorded the growing season table 1 nitrate levels measurements on both previous and current remained high compared with the other three season vegetative branchletsbranchbranchlesslets initial photo- treatments at both depths during the growing synthesis values were used and corrected season the shallow character of the soil prob- using the formula developed by LICORLI COR ably limited nitrate losses to leaching maxi- mcdermitt 1987 mizing the amount available for plant uptake statistically significant treatment effects for the addition of ammonium increased ammo- variables measured on artemisia were identididenti niumgium 188 in the B horizon the increase in 12 P S doescheretalDOESCHER ETETALAL volume 50

171 el control 11 1 9 sugar E 20 0 E E 0 nitrate 111 A A ammonium C C 15

E 3 T E 10 T

0 5

0 april 15 april 20 april 25 april 5030 may 5 date

fig I1 ephemeral leafleafmaximummaximum length ofArtemisia tntritrldentata sspasp wyomingensiswyomingensts during the early growing season standard errors are presented for each mean

the A horizon was not significant the sugar appeared to increase at a greater rate in early treatment did not appear to change soil nitro- may than in sugar and control plots gen levels when compared with the control the addition of nitrogen increased current possibly indicating that carbon was not limit- year s vegetative stem length compared with ing decomposer microflora mcgarity 1961 no nitrogen addition throughout the growing season 3 stem length growth fig was similar between both nitrogen forms in april and may in the early spring both mean maximum but continued at a more rapid rate in the length of ephemeral leaves and number of nitrate treatment in june at the termination ephemeral leaves per terminal shoot were of vegetative stem elongation stems in the greater in the ammonium and nitrate treat- nitrate and ammonium treatments were 175 ments than in the sugar and control treat- and 140 longer respectively than shoots in ments figs 1 2 ephemeral leaf lengths the control treatment stem elongation in the during april in the nitrogen treated plots did control and sugar treatments was similar not differ from one another but were greater the addition of either form of nitrogen did than in the nonnitrogennonnitrogen treated plants con- not increase specific leaf weight averaged trol and nitrate treated plants did not differ across the growing season as compared with from one another in leaf length for the may the control treatment table 2 the addition sampling period analysis of main effect of sugar however reduced specific leaf means for ephemeral leaf numbers revealed weight across dates on the average by 12 that plants in nitrogen treated plots did not compared with the control the major differ- differ from one another and that their values ence in specific leaf weight between control were greater than similar values found for the and sugar treatments occurred during abscis- control and sugar treatments in addition the sion of ephemeral leaves in late july and early development ofnew leaves on terminal shoots august fig 4 specific leafweight increased in both nitrate and ammonium treatments approximately 180 to 200 when ephemeral 199011990 EFFECTS OF NITROGEN availability 13

16 0 0 control 14 sugar A 0 M nitrate 12 A A ammonium CO 0 Q 10 E 33 z 8 4 0 6

2 t 0 april 15 april 20 april 25 april 30 may 5 date fig 2 ephemeral leaf numbers per terminal bud of artemisia tridentata sspasp wyomingensis during the early growing season standard errors are presented for each mean

T loo100 0 11 control

E sugar T M 9 nitrate Cc7ca 80 A A ammonium i con T 0Q 1 j 60 A E A u W 40

01 414.41afdf D cn 0 20

0 may 5 may 15 may 25 june 4 june 14 june 24 date fig 3 effects offouroffousoffourmour different nitrogen treatments on current year vegetative stem length of artemisiaofartemisia tridentata sspasp wyomingensiswyoiningensis standard errors are presented for each mean 14 P S doescheretalDOESCHER ETETALAL volume 50

02500.250 control

CT sugar 0 nitrate 0 02000.200 0 CM czicz3 ammonia E 0

0 0 01500.150 4 4

0 bi 01000.100

UQ 1 0 Q 0 V 00500.050

I1 0 0 0 I1

0 H H H 00000.000 U 1 tj ir apriapr155 apr2laprylapr21 apr22 jun03 jun09 junjun1111 junjun1818 jul13jul 13 jul27ju127 aug05 augaug1818 sepo I1 date fig 4 specific leafweightsleafweights ofArtemisia tridentata sspasp wyomingensis during the course oftheofodthethe 1987 growing season

TABLE 2 specific leaf weights of Artemisia tridentata artemisia plants table 3 analysis of main sspasp wyomingensis growing under four nitrogen treat- effect means revealed that plants in the nitrate ments mean values are averaged over the 1987 growing and ammonium had greater con- season treatments centrationscentrations than did the control and sugar treatment specific leafweightleaf weight gmegm2 treatments control 145 b water and photosynthesis sugar 1321321 relations ammonium 141 soil water depletion rates were generally nitrate 14145 more rapid in the nitrogen treated plots at means followed by similar letters are not significantly different at p5Ppa 05 both soil depths than in the sugar and control plots figs 5 6 differences were greatest TABLE 3 leaf nitrogen concentration aggmgggg of during rapid growth in mid may in the lower artemisia dentatatritridentatatridentate sspasp wyomingensis growing under 20 cm in the upper 20 cm the sugar treated four nitrogen treatments means are averaged over the plots 1987 growing season maintained the highest soil water content from april through may treatment leafnitrogen concentration aggmgg predawn plant water potentials were simi- control 19619.6196sa lar among treatments in mid may and late sugar 19619.619 8 july table 4 during rapid growth in june ammonium 25 lb1 however predawn plant water potentials in nitrate 272701127.011C nitrogen treated plots were lower than both means followed by similar letters are not significantly different at P 05 control and sugar treatments midday water potentials were also lower in june in both leaves abscised in august for all treatments nitrogen treatments compared with sugar and by late august when most ephemeral leaves control both predawn and midday readings had abscised specific leaf weight between declined in all treatments as the season pro- treatments was similar gressed application of both forms of nitrogen re- photosynthesis was significantly different sulted in greater leaf nitrogen contents of between treatments on only the 15 june 1987 1990 EFFECTS OF NITROGEN availability 15

11 16 control T 0 sugar 0 0 nitrate 14 A A ammonium Q

414 1 020.2 mpa

T c T 0 0 0 0 0 10 j 070.7 mpa A 8 151.5 mpa 0 in 6

4 arrilapril may june july aug months fig 5 seasonal pattern ofsoil water content in the upper soil profile 2 20 cm for four different nitrogen treatments in 1987 vertical bars are 95 confidence limits field capacity 00030.0303 Mmpapa 18 soil water content

15 020.2 mpa

Q 1513 070.7 mpa 00

c X 0 0 151.5 mpa 9 T A 202.0 mpa 0 7 0 0 control en 0 sugar 0 nitrate A A ammonium

5 i i i april maymoy t june july aug months

fig 6 seasonal pattern of soil water content in the lower soil profile 20 40 cm for four different nitrogen treatments in 1987 vertical barsbars are 95 confidence levels 16 PPSS DOESCHERdoescheretalETALET AL volume 50

TABLE 4 predawn and midday plant water potentials mpa in artemisiaartemisio tridentata sspasp wyomingensis through the growing season in 1987

treaticreatitreatmentstientsmients date control sugar nitrate ammonium predawn aa may 19 1241.241 24 1 0505aa i1 o4a04oba igligi121121a w june 3 131a1 31 1 44a44 1 77a77 1 77 1aaa july 21 1621.621 62 1 62a62 1741.741 74ba 11771.7777ba midday ab april 15 185185aa 1 67670 1 65b65 1621.621 62ab ab1b june 3 2 ogba09 2206ba06 2242.242 24 232.32 31bbibb numbers followed by the same lowercase letters are not significantly different P 05 between dates for each treatment numbers followed by the same uppercase letters are not significantly different P 05 between treatments for each site

TABLE 5 photosynthesis ingmg COcog m 2 seesec 1 of artemisiaartemisio tntritrldentata sspasp wyomingensiswyomingensts under four nitrogen treatments during 1987

date control sugar nitrate ammonium april 17 030 000 0360.36036s 054 june 15 00251125 003333b 1 7373a 2182.189182 bab8 june 20 02011 016olg0 16 028 002525 july 30 04204210 42 1 022a0 22 022a 0200 20 august 1 0430431 0430 43a 0200.20 051 means followed by the same letter are not significantly different between treatments at the same date

date table 5 increased photosynthesis on years application of water andor nitrogen this date corresponded to the period of rapid resulted in enhanced growth while in other leafand stem growth in early june years no response was observed fischer et al 1987 concluded this variable response was discussion due to yearly changes in available soil nitrogen in the great basin biomass production nitrogen availability is probably second of forage species has not always been related only to water as the most limiting factor to to precipitation charley 1972 sneva and biomass production in great basin plant com- britton 1983 miller et al 1990 sneva and munitiesmunities james and jurinak 1978 at our britton 1983 and miller et al 1990 reported study site addition of nitrogen was also found reduced herbaceous production in the third of to significantly promote aboveground growth three consecutive wet years it has been spec- ofaof A dentatatritridentatatridentate sspasp wyomingensis fertiliza- ulated that nitrogen may be limiting following tion increased ephemeral leaf size number of successive wet years as prolonged plant ephemeral leaves and stem elongation rates growth depletes soil nutrients and poor qual- fertilization also increased nitrogen amounts ity organic matter is slow to decompose in the subsoil and the leaves these responses parker et al 1984 fischer et al 1988 on a lend support to our original hypothesis reclaimed mine spoil carpenter and west namely that A tridentata responds favorably 1987 indicated no response to nitrogen addi- to increases in soil nitrogen tions for the species artemisia dentatatritridentatatridentate sspasp although our results showed a positive vastyanavaseyanavaseyana available nitrogen was probably growth response in A tridentata to nitrogen not limiting due to stockpiling of topsoil and the role nitrogen plays in influencing growth the lack of site occupation by an establishing of andaridarld species is poorly understood in the plant community in carpenter and west s desert southwest researchers have report- 1987 study leaf nitrogen concentration of ed a variable response of larrea dentatatritridentatatridentate control plants was 323.2 in contrast our DC covgov dominated communities to water study had leaf nitrogen concentrations of 202.02 0 and nitrogen inputs gutierrez and whitford 272.7 and 252.5 for control nitrate and am- 1987 fischer et al 1987 during certain monium respectively site and subspecies 1990 EFFECTS OF NITROGEN availability 17 differences may have also contributed to nitrogen additions have been reported to in-in the different response crease shoot to root ratios in larrea tndentridenanden A positive growth response was shown for tata fischer et al 1988 laethalajtha and klein the ammonium plots in spite of enhanced am- 1988 monium levels being found only in the 20 40 specific leaf weights averaged across the cm soil depth the nonsignificant concentra- growing season were significantly influenced tions of ammonium in the surface soils may by the sugar treatment prior to leaf senes- have been caused by losses due to volatiliza- cence specific leaf weights were similar be- tion absorption ofammonium on soil colloidscolloids tween treatments on seven ofeight dates mea- high biological activity andor ammonium be- sured specific leaf weights inm the sugar ing hydrolyzed to nitrate in the lower soil treatment however were significantly less depth also the growth response may have than in the control during senescence and been related to the application of ammonium abscission of current year s ephemeral leaves in the sulfate form whether or not A triden once the majority of ephemeral leaves had tata responds to sulfur additions has yet to be abscised in mid august for all treatments determined despite the lack ofenhancement specific leaf weights were again similar the in the upper soil depth of the ammonium decrease in specific leaf weights in the sugar treated plots the extensive root system of A ti eatmentestmenttreatment was probably a function of delayed dentatatritridentatatridentate would allow this species to readily leaf senescence and abscission rather than utilize ammonium in the lower soil depths leaves being lighter per unit of surface area xylem potential and soil moisture readings although not indicated by the soil nitrogen found in this study would at first appear data some reduction inm available soil nitrogen contradictory to the observed growth re- may have occurred on the sugar plots sponsessponses the more negative soil moisture and marschner 1986 reported an increase in plant water potentials reported for A triden leaf area indices in plant populations with an tata growing in the nitrogen plots suggest increase in nutrient supply increased leafde- that these plants were more water stressed velopment during the early part of the grow- and thus should have reduced aboveground ing season and a larger leaf area index in the growth however the opposite was found years when mineralizable nitrogen levels are we feel that a possible explanation for these relatively high may increase artemisia s com- results centers on the greater growth and petitivepetitive advantage for nutrient resources over physiological activity of plants in the nitrogen associated species miller 1988 reported that plots since specific leaf weights were simi- artemisia maintained a relatively high leaf lar between treatments during active leaf area early inm the spring compared with associ- growth we can assume that differences in leaf ated species allowing it to capture soil water area between treatments are approximately resources early in the growing season early proportional to biomass leaf biomass of increased leaf area also enhances its ability to A dentatatritridentatatfidentatatridentate in the nitrate and ammonium maximize photosynthesis when environmen- plots was found to have increased 520 and tal conditions are favorable depuit and cald- 230 respectively over control wang well 1973 in warm desert shrubs rate of leaf 1989 greater growth and leaf area of these area development was the primary factor lim- plants may mean that more soil water was iting whole plant carbon gain during the early used by plants in the nitrogen treatments portion of the growing season comstock et al this observation is supported by the research 1988 of svejcar and browning 1988 they re- in conclusion application of both nitrate ported a greater leaf area higher physiologi- and ammonium increased growth response cal activity and a subsequently more rapid of A dentatatntritrlfritndentatatridentatatridentate over control and sugar treat- soil water depletion in burned versus un- ments apparently this species can oppor- burned stands of andropogon gerardgerardiigerardiaii vit- tunistically take advantage of increased man the greater photosynthetic rates of A soil nitrogen by increasing amount of leaf dentatatritridentatatridentate in the nitrogen plots during early area available for growth increases inm avail- june also further support this observation in able soil nitrogen might occur following addition an increase in shoot to root ratio events such as several consecutive years of could have influenced plant water relations below average precipitation or weakening of 18 PSP S doescheretalDOESCHER ETALET AL volume 50

perennial grasses through overgrazing A tri communities dominated by big sagebrush soil dentata may enhance its competitiveness bbyy science society ofamericaof america journal 4865948.65948 659 663 responding favorably levels eissenstat D M AND M M CALDWELL 1988 compet- to increased of itive ability is linked to rates of water extraction soil nitrogen A field study of two fridlandaridland tussock grasses oe colodiacologia 757511 7 acknowledgments FISCHERPISCHER F M L W PARKERJPARKER J P ANDERSONandersonandwAND W G WHITFORDWHITFOBD 1987 nitrogen mineralization in a we thank T J svejcarSvejear W C krueger desert soil interacting effects of soil moisture and N fertilizer soil science society ofamerica jour- R H waring and A R tiedemann for their nal 51 1033 1041 helpful reviews on earlier versions of this FISCHER F M J C ZAK G L cunningham AND W G manuscript this is technical paper no WHITFORD 1988 water and nitrogen effects on 9006 of the oregon agricultural experiment growth and allocation patterns of creosotecreosotebushbush in station the northern chihuahuan desert journal of range management 41 387 391 GUTIERREX J R AND W G WHITFORD 1987 responses literature CITED ofchihuahuanofchihuahuan desert herbaceous annuals to rainfall augmentation journal of arid environment BAATH E V LOHM B LUNDREM T ROSSVALL B soder- 12127 139 strom B SOHLEUIUS AND A WIREN 1978 the HORNECK D A J M HART AND K TOPPER 1989 meth- effect of nitrogen and carbon supply on the devel- ods ofsoilofsoil analysis used in the soil test laboratory opment of soil organism populations and pine at oregon state university corvallis SM 894 seedlings microcosm experiment oikosbikos 31 HIRONAKA M AND E W TISDALE 1963 secondary suc- 153163153 163 cession in annual vegetation in southern idaho BREMMER J M 1965 total nitrogen appp 1149 1178 ecology 44 810 812 in C A black ed methods of soil analysis part JAMES D W AND J J JURINAK 1978 nitrogen fertiliza- 2 american society of agronomy madison wis- tion of dominant plants in the northeastern great consin basin desert appp 219319219231231 in N E west and CALDWELL M M D M eissenstat J H RICHARDS AND J skujins eds nitrogen in desert ecosystems M F ALLEN 1985 competition for phosphorous dowden hutchingson and ross inc strouds differential uptake from dual isotope labeled soil burg pennsylvania interspace between shrub and grass science 229 LAJTHA K AND M KLEIN 1988 the effect of varying 384386384 38638& nitrogen and phosphorus availability on nutrient CARPENTER A T AND N E WEST 1987 indifference of use by larrea tridentata a desert evergreen mountain big sagebrush growth to supplemental shrub Oeoecologyoncologycology 7534875 348 353 water and nitrogen journal of range manage- LENTZ R D AND J H SIMONSON 1986 A detailed soils ment 4044840 448 451 inventory and associated vegetation of squaw CARPENTER L H 1972 middle park deer study range butte range experimental station oregon state fertilization game research report part II11 col- university agricultural experiment station spe- orado division ofwildlifeofwildlife denver cial report 760 CHARLEY J L 1972 the role of shrubs in nutrient cy- MARSCHNER H 1986 mineral nutrition in higher plants cling appp 182203182 203 in C M mckell J P blais- academic press new york 674 appp dell and J R goodin eds wildlandWildwindlandland shrubs mcdermittMCDEBMITT D K 1987 photosynthesis measurement their biology and utilization united states systems performance comparison of the LI 6200 department of agriculture forest service gen- to the LI 6000 LICORLI COR inc application note eral technical report INT 1 6200 1 CHARLEY J L AND N E WEST 1975 plant induced soil MCGARITYMCCARITY J W 1961 denitrification studies on some chemical patterns in some desert shrub ecosys- south australian soils plant soil 14 1 21 tems in utah journal ofecology 63 945 964 MILLER R F 1988 comparison of water use by arte- 1977 micro patterns of nitrogen mineralization misia tridentata sspasp wyomingensis and chryso- activity in soils of some shrub dominated ecosys- thamnus viscidiflorusviscidiflorus sspasp viscidiflorusviscidiflorus journal of tems in utah soil biological biochemistry 9 range management 41 58 62 357 366 MILLER R F M R HAFERKAMP AND R F ANGELL 1990 COMSTOCK J P T A COOPER AND J R ehleringer effects of season of defoliation on soil water and 1988 seasonal patterns of canopy development plant growth crested wheatwheatgrassgrass journal of and carbon gain in nineteen warm desert shrub range management in press species Oeoncologyoecologycology 7532775 327 335 MILLER R FFANDLAND L M SHULTZ 1987 development and DEPUIT E J AND M M CALDWELL 1973 seasonal pat- longevity of ephemeral and perennial leaves on tern offetofnetofnet photosynthesis of Artemisia tridentata artemisia tridentata nutt sspasp tridentata great american journal ofbotany 60 426 435 basin naturalist 47 227 230 1975 gas exchange ofthree cool semidesertsemi desert spe- PARKERPARKEB L W P F SANTOS J PHILLIPS AND W G WHIT- cies in relation to temperature and water stress FORD 1984 carbon and nitrogen dynamics during journal of ecology 63 835 858 the decomposition of litter and roots of a chi doescherDOESCHERPP S R F MILLERM ILLER AND A H WINWARD 1984 huahuanhuaxuan desert annual lepidium lasiocarpum soil chemical patterns under eastern oregon plant ecological monographs 54 339 360 1990 EFFECTS OF NITROGEN availability 19

SAS 1988 general linear models procedures appp tlsdaleeTISDALEE WMW M HIRONAKA AND M A FOSBERG 1969 549 640 in SAS user s guide statistics release the sagebrush region in idaho problem in range 6036.03 edition SAS institute carygary north carolina resource management idaho agricultural experi- scholander P F H T HAMMEL E D bradstreet ment station bulletin 512 15 appp AND E A hemmingsen 1965 sap pressure in WALLACE A E M ROMNEY G E KLEINKOPF vascular plants AND science 148 339 346 S M SOUFI 1978 uptake of mineral forms of SNEVA F A AND C M BRITTON 1983 adjusting and nitrogen by desert plants in N E west and forecasting herbage yields in the in intermountain J skujins eds nitrogen in desert ecosystems big sagebrush region oftheodtheofthe ore- steppe province dowden hutchingson and ross inc strouds gon state university agricultural experiment burg pennsylvania station bulletin 659 61 appp WESTWESTNN E ANDANDJJ SKUJINS eds 1978 nitrogen in STEEL R D G AND J H TOKRIE 1980 principles and desert torrie ecosystems dowden hutchinson and ross procedures of statistics A biometrical approach ad2d inc stroudsburgStroudsburg pennsylvania ed mcgraw hill inc hightown new jersey STURGES D L 1977 soil water withdrawal and root char- WANG J 1989 response of Arfeartemisiamisiamisla tridentata sspasp acteristicsacteristics of big sagebrush american midland wyomingensiswyoniingensis and stipa thurberianathurberiana to nitrogen naturalist 98 257 273 amendments unpublished thesis oregon state SVEJCARSVEJCAB T J AND J A BROWNING 1988 growth and university corvallis gas exchange of andropogon gerardiiti as influ- WARINGWARINC R H AND B D CLEARY 1967 plant moisture enced by burning journal of range management stress evaluation by pressure bomb science 155 41239 244 1252125412521253 1254 TISDALE D AND M HIRONAKA 1981 the sagebrush grass region a review of the ecological literature university of idaho forestry wildlife and received 10 october 1989 range experiment station bulletin 209 accepted I1 november 1989 great basin naturalist soisol501soo 1990 appp 21 31

FORM AND dispersion OF MIMA MOUNDS IN RELATION TO SLOPE STEEPNESS AND ASPECT ON THE COLUMBIA PLATEAU

george W coxcox1coxa

ABSTRACT patterned ground consisting of mima type earth mounds and associated sorted stone circles and nets is widespread on the columbia plateau ofwestern north america studies ofthe geometric relationships of mounds and stone nets to slope aspect and steepness were earnedcarried out at the lawrence memorial grassland preserve north central oregon in ilinejune 1987 mound and roundfieldmoundfieldmoundfield characteristics were sampled on randomly chosen iha1 ha plots on slopes of different aspect and steepness mounds were largest most circular and symmetrical in form and most fully encircled by beds of size sorted stones on level sites on slopes of increasing steepness mounds decreased in size showed increasing asymmetry and downslope elongation and became connected into lines oriented up and downslope encircling stone beds became more weakly developed or disappeared on upslope and downslope sides ofthe mounds and the lateral beds developed downslope extensions that eventually merged with those of adjacent upslope and downslope mounds these patterns are interpreted as ireflecting changes in the manner of soil translocation by northern pocket gophers thomomys talptalpoidesoides due to their responses to tunneling on slopes and to the modification of the flow ofwater across the slope because ofthe presence ofmounds

mima type earth mounds are a characteris- soil translocation by pocket gophers cox and tic feature of grasslands with shallow soils or alienallenailen 1987a poor drainage in western north america cox our previous studies of mima mounds and 1984 these mounds containing stones up to sorted stone beds have been conducted on about 50 mm in diameter commonly range up level areas where mounds are circular in form to 2 m in height 20 m in diameter and 50 ha 1 and regular in spacing observations by previ- in density recent investigations at several ous workers cited above and patterns evi- locations have supported the hypothesis that dent on aerial photographs indicate that mima type mounds are formed over long pe- mound form and roundfieldmoundfieldmoundfield geometry are riods of time by the centripetal translocation modified considerably on slopes the objec- of soil toward centers of activity of geomyid tive of this study was to define variation in pocket gophers these centers located ini- mound form and roundfieldmoundmoundfieldfield geometry with tially in the deepest best drained sites avail- slope aspect and steepness and to determine able are gradually transformed into mounds if the activities of pocket gophers can account by soil translocation cox 1984 cox and alienallenailen for the variation 1987b cox and gakahu 1987 cox et al 1987 METHODS mima mounds are an extensive and prominent feature of the shrub steppe ofthe colum- studies were conducted at the lawrence bia plateau in eastern washington northern memorial grassland preserve LMGP and on oregon and southwestern idaho USA here adjacent ranch land of the fridaypriday brothers the mounds are frequently encircled by beds corporation southern wasco county ore- of sorted stones and ermoundintintermoundinterwound flats often gon 4457n 12048w 1 11 june 1987 exhibit polygonal networks of sorted stone this was the site of previous studies of the beds waters and flagler 1929 kaatz 1959 structure of mounds and associated beds of maldemaide 1961 1964 fosberg 1965 these fea- sorted stones cox and alienallenailen 1987a cox et al tures formerly interpreted as periperiglacialglacial fea- 1987 the LMGP a registered national land- tures have also been interpreted as a result of mark owned by the nature conservancy lies

department of biology san diego state university san diego california 92182005792182 0057

21 22 G W cox volume 50

at an elevation of 1036 1060 in on the sha- field the overall aspect of each plot was de- niko plateau formed of columbia river terterminedmined with a compass and the slope steep- babasaltsbasalessalts and includes several ravines that fall ness measured with an inclinometer A count steeply northward into the valley of ward of the mounds in each plot was obtained and creek 122 in below the preserve has a cold the mound nearest the plot center was desig- semidesert climate with an average annual nated for detailed measurements maximum precipitation of 280 mm the surface of the mound height was measured with a meter plateau is mounded biscuit scabland with stick and line level the orientation ofthe long mima mounds that range up to about 2 in in axis of the mound downslope direction was height and 20 in in diameter the mound soils measured with a compass maximum and are classed as condon eolian silt loamscoams and minimum diameters were measured with a the intermound soils as bakeoven residual meter tape and the components of these di- very cobbly loamscoams the vegetation of mounds ameters relative to the highest point on the and deeper upland soils is dominated by mound were also recorded mound top to up- idaho fescue festuca idahoensis and blue slope edge top to downslope edge etc dis- bunch wheatgrasswheatgrass agropyronagropyronspicatumspicatumspicatum the tances to the first and second nearest neigh- shallow ermoundintintermoundinterwound soils are dominated by bors between mound high points were also scabland sagebrush artemisia rigida sand- measured with a meter tape as was the mini- berg bluegrass poa scabrella several spe- mum distance between the highest points of cies of biscuitroot lomatium sppapp and bit- the two mounds that were furthest apart in terroot lewisia rediredivivoredivivaviva the northern this three mound set the fraction of the pocket gopher thomomys talptalpoidesoides is abun- mound encircled by beds of bare size sorted dant throughout the preserve A comprehen- stones cox and alienallenailen 1987a was recorded for sive physical and biotic inventory of the the upslope and downslope halves of the LMGP is given by copeland 1980 mound the length of stone beds diverging relationships of mound and roundfieldmoundmoundfieldfield from that surrounding the mound and extend- characteristics to slope and aspect were ex- ing downslope tails was recorded the plored by sampling mounds throughout the maximum length of this measurement was the LMGP and on a small area of adjacent ranch point at which this tail reached another land for this sampling an aerial photo with a mound finally the number of discrete superimposed 100 X 100loom m grid was used pocket gopher activity areas was recorded as some grid units that included very steep an estimate of the number of animals occupy- slopes or canyon bottoms were largely or ing the mound areas with surface heaps or entirely unmounded since the objective of plugged tunnel openings were considered the study was to examine mound characteris- separate activity areas when they were sepa- tics in relation to slope only mounded grid rated by more than 5 in units 50 of the surface showing mound from these measurements a number of ermoundintintermoundinterwound topography were considered for descriptive characteristics were calculated sampling this criterion also assured that the mound area was calculated assuming that the grid units selected were internally uniform in base was a circle or ellipse and volume was their slope these grid units with the aid of a computed on the assumption that the mound topographic map were also grouped tenta- was a segment ofa sphere or prolate spheroid tively into five aspect classes level with an elongation el was computed from the fol- overall slope less than 252.5 or north east lowing relationship south or west facing all grid units within the 153 ha LMGP werewere considered in addi- EelI1 v2va2va bb2abaa tion to allow adequate representation of where a and b are the major and minor radii south facing slopes a loha10 ha area of land north A second measure of elongation eae2 was also of ward creek was also included sets of 7 calculated as the ratio of the major to minor grid units were chosen by random coordinates diameters of the mound asymmetry AS was in each of the five aspect classes a total of 35 calculated grid units the aerial photo was then used to locate ASasi 1 salsylsyi 1 I ssf05ss2105 these grid units hereafter termed plots in the where slasland Ss aresflthe asymmetries on the 1990 MIMA MOUNDS 23

TABLE 1 characteristics of mima mounds and moundfieldsmoundfields on plots differing in slope steepness at the lawrence memorial grassland preserve north central oregon values are means t standard errors

slope steepnessste spnessaspness overaoverall 11 0 252.52 1 25 50 50 75 75 10010.0loo10 characteristic nan3n 355 n 6 n 15 n 8 n 63 mound features height cm 687286872.868768 7 28 817688176.881781 7 68 679 t 44 629 t 51 657 t 65 area mam2 1459 146 1880188.0188 0 2512525.11 1749 t 247 1182 262 6808968.089680 j 89 volume mamm3 528 62 786 137 606 10110.110loi 1 406 112 235532355.3235 53 elongation 1420111420.11142 oiioli0110 11 123 0180.180 18 146 t 020 173 020 107 oilolioii0110.11 asymmetry 0590080590.08059 008 014 004 059 t 008 078 023 077 j 027 sorted stone beds upslope side 366 56 592 17117.117 1 353 t 78 344 13113.113 1 200 j 82 downslope side 258492584.925825 8 49 458 166 282809800 t 75 175 70 11711 7 t 79 overall 315473154.7315 47 525 156 317 L 63 259 t 95 175 jt 77 downslope tails m 2172.172 17 064 0 165 t 073 4892134899134892.13489 213291313 202 L 1 11 roundfieldMoundmoundfieldfield features 1 density ha 216 121 2 188231882.3188 23 215 t 191 9 244 313.13 1 212 j 29 connection olioii0110 11 003 005 005 018 006 004 004 009 j 006 linearity 0840020840.02084 002 0740060740.06074 006 088 003 0770060.77006077 006 0091ogi91 j 0030 03 mounded surface 31531.531 5 353 37637.637 6 28828.828 8 1441414.44 long and short axes respectively calculated RESULTS

slS I1 aaaagaaaa7a Ss wb measurements ofthe overall slope aspect in the field revealed that several of the plots where a and b are the longer and a and fc tentatively placed in particular aspect classes the shorter distances from mound edge to actually fell in other classes As a result the peak along the major and minor axes connec- final set of samples comprised 10 N facing tion C of the sampled mound to the two 5 E facing 5 S facing and 8 W facing plots nearest mounds on opposite sides was cal- table 1 culated as the mean of ratios of heights of between mound divides to maximum mound mound and roundfieldmoundfieldmoundfield characteristics height linearity QL of alignment of the sam- showed a complex relationship to slope aspect pled mound and its two nearest neighbors was and steepness with respect to slope steep- determined as the ratio ofthe sum ofdistances ness mounds at the LMGP were confined to measured from center to center for a series of slopes less than about 10 in steepness mounds to the single straight line distance mounds exhibited greatest height basal area between the first and last in the sequence and volume on level areas or very gentle data on mound and roundfieldmoundfieldmoundfield character- 252.5 slopes table 1 basal area and vol- istics were analyzed by BMDP statistical soft- ume declined progressively with increased ware procedures dixon 1983 logarithmic slope steepness ANOVA fyf331 3623.62 and arcsinearearc sine transformations were employed P 05os05.05 for area F 3293.29 P 05os05.05 for vol- to achieve normality for some variables ume the mean height of mounds on level to means and standard errors were computed for very gently sloping areas was significantly plot data grouped by slope aspect and slope greater than on steep 50so505.0 757.5 slopes steepness classes and these classes were then t 2262.26 DF 12 P 05os05.05 volume the compared by t tests and ANOVA BMDP ID best overall indicator of mound size showed a and ad7d correlations among all combinations strong negative correlation r 496496.496 of variables and stepwise linear regressions P oi0101.01 with slope steepness fig 1 using selected variables as the dependent mound asymmetry and elongation were variable were also performed BMDP 2rar both greatest on steep 50so505.0 757.5 slopes plant names follow hitchcock and cron- table 1 asymmetry was significantly lower quist 1973 for mounds on level to very gently sloping 24 G W cox volume 50

overall

down upslope slopesiope

rock ring development

volume connectionD rlineaalinealinearity

nearest neighbor slopealope distance steepness

elongation density mound axis direction

P 005 Ppooiooi001 P 0001 asymmetryYJ

fig 1 correlationcoicol relation relationships between major variables of mound and roundfieldmoundfieldmoundfield geometry for plots sampled on slopes of differing steepness and aspect at the lawrence memorial grassland preserve north central oregon june 1987

0 252.5 areas than on gentle 25252.5 5505.0so 00 slopes between the development of sorted stone t 3303.30 DF 19 P oiol01.01 steep slopes beds and slope steepness was stronger on os t 2352.35935 DF 12 P 0505.05 or very steep the downslope side of mounds r 367367.367 75 10.0loo slopes 10 757.5 100 t 2332.33 DF P 05os05.05 than on the upslope side r 304304.304 P 05os.05 05 asymmetry was also strongly corre- P 05os05.05 lated r 449449.449 P oloi01.01 with slope steepness roundfieldmoundfieldMoundfield characteristics showed weaker fig 1 elongation defined as the ratio of patterns of showed long to short mound axis was significantly density mounds little greater on steep slopes than on level to very variation with slope steepness table 1 con- gently sloping areas t 3073.07 DF 12 P nection and linearity which were positively os oi0101.01 elongation was significantly greater on correlated r 343343.343 P 0505.05 exhibited steep than on very steep slopes t 2562.56 highest values on steeper slopes linearity DF 12 P 05os05.05 was significantly lower on level to very gently the development of encircling beds of sloping 0 252.5 plots than on plots with gentle bare size sorted stones was strongest on level 25252.5 505.05so 00 slopes t z 2292992.29 DF 19 P 05os05.05 to very gently sloping plots table 1 the de- or very steep 75757.5 10010.010loo 0 slopes t 2282.28 DF of overall 10 P 05os05.05 combination of mound gree development beingI1 negatively the correlated r 343343.343 P 05 with slope size basal area and density resulted in a steepness fig 1 the negative correlation greater surface coverage by mounds on very 1990 MIMA MOUNDS 25

TABLE 2 characteristics of mima mounds and moundmoundfieldsfields on plots differing in aspect at the lawrence memorial grassland preserve north central oregon values are means L standard errors aspect overall level north east south west characteristic n 35 n 7 n 10 n 5 n 5 n 8 mounds height cm 68768.768728282.8 80780.780758585.8 756 444.4 60 0 8 0 54254.2 545.4 63963.9 525.2 m2 75644 60080 54254 63952 area ma 1459145.9145 9 14614.614 6 1723172 3 26426.426 4 1466146.6146 6 24524.524 5 1171117 1 42342.342433 3 1210121.0121 0 4244242.44 iss1552155 2 39239 2 volume mamm3 582 2 62 5 571 1 1 183 307 58258.258 626.26 2 71571 13613 6 57157.157 ililii11111 40740 7 18318.318 3 30730.730 7 t 848.48 4 52152.152 1 15615.615 6 elongation 1421.421 42 olioii0110.110 11 1 10 t 0 20 1321 32 0200.200 20 1 72 0 32 1 60 t 0 18 1511.511 51 0280.280 28 asymmetry 0590 59 0080 08 0150 15 0030 03 0510 51 0080 08 loo1001 00 0340 34 0660 66 0300 30 0750 75 0140.140 14 rock circles upslope side 5 6 7 167 7 335 186 380 107 366563665.636636 6 56 50750 16716.716 33533.533 5 11511.5ils11 5 41041 0 18618.618 6 38038.038 0 10710.710 7 24424.424 4 626.26 2 downslope side 25825.825 8 494.94 9 39339.339 3 15415.415 4 20520.520 5 10510.510ioslos 5 20020 0 10510.510ioslos 5 25025.025 0 12212.212 2 25025.025 0 11311.311 3 overall 31531 5 474.74 7 45045 0 15215.215 2 27027 0 919.19gi 1 30530 5 13513 5 31531.531 5 10310.310 3 25925 9 616 1 rock tails m 21722.1717 0640.640 64 0 1951 95 106log1.061 06 4644.644 64 3253 25 3283.283 28 t 1631.631 63 2112.112 11 1031.031 03 roundfieldmoundfieldMoundfield features 1 density ha 21621.621612121.2 17817822222.2 200-2001313 2364123623.6 41 26826.826850t 5 0 2252622522.5 262.6 connection olioii0110 11 0030 03 0090.090 09 0060 06 0200.200 20 00900.0909 0060.060 06 0060.060 06 0080.080 08 0080 08 0060.060 06 0040.040 04 linearity 0840 84 0020.020 02 0780.780 78 0060 06 091ogi0.910 91 0030.030 03 0750 75 00900.0909 0820.820 82 0040.040 04 0860.860 86 0040.040 04 mounded surface 31531.531 5 307 293 276 324 349 gentle to gentle slopes than on steep to very mound density was positively correlated steep slopes with slope aspect expressed as deviation in data for mounds grouped by slope aspect degrees from north r 405 P 05 den- table 2 indicate that mound size height sity ranged from 17817.817 8 mounds ha on level basal area volume was greatest on level plots plots to 26826.826968 8 mounds ha 1 on south facing and next greatest in height and volume on slopes table 2 linearity of a sample mound north facing slopes mounds were smallest in and its two nearest neighbors was greatest for both height and volume on south facing north facing and least for east facing slopes slopes and next smallest on east facing slopes linearity was significantly greater for north the variation among heights on plots differing facing slopes than for level plots t 2262.262 26 DF in aspect was significant ANOVA ff41140 15 P 05 3483.48 P 05os05.05 mean volume on south facing several other important relationships were slopes was significantly less than on level sites not directly linked to either slope steepness or t 2312.31 DF 10 P 05os05.05 slope aspect fig 1 A number of these cen- mounds were least elongate or asymmetric tered on nearest neighbor distance and stone on level and north facing plots and most elon- circle development nearest neighbor dis- gate and asymmetric on east facing slopes tance was positively correlated with mound table 2 elongation expressed as the ratio volume r 349 P 05 in addition of longer to shorter axis was significantly nearest neighbor distance showed a strong greater for east and south facing plots than direct correlation with mound elongation for level plots t 3143.14 and 2762.76 respectively r 449 P 01 and stone circle develop- for east and south facing plots DF 10 P ment r 468 P 01 furthermore the 05os05.05 variation in asymmetry was significant correlation of nearest neighbor distance to among aspect groups ANOVA fo 3293.29 development of the stone circle on the P 0505.05 elongation of mounds was closely downslope side of mounds was very strong parallel to slope the long axis of the mound r 641 P 001 mound volume also being very highly correlated with the slope showed a direct relationship to stone circle direction fig 3 r 822822.822 P ooiool001.001 the development both overall r 346 P 05 development of sorted stone circles differed and on the downslope side r 415 P 05 little for slopes of differing aspect table 2 the more elongate a mound the greater was downslope tails of sorted stones were noted the development of the stone circle on its on slopes cfallofallofallaliail aspects downslope side r 379 P 05 the 26 GgecoxgwcoxW cox volume 50

greater the density of mounds however the maximum steepness of mounded slopes may poorer was the development of the stone cir- be real and may relate to soil texture and other cle on the downslope side of the mound r factors affecting vulnerability to erosion 423423.423 P 0505.05 linearity of mound arrange- mounds may occur only on slopes of200of20 steep- ment on the other hand was negatively re- ness or greater when soils are rich in clays as lated to development of the stone circle both they are in many locations in southern califor- overall r 339339.339 P 05os05.05 and on the up- nia cox 1984 slope side r 335335.335 P 05os05.05 finally dens- researchers also offer diverse statements ity showed a positive correlation with asym- on how mound shape varies with slope steep- metry r 364364.364 P 05os05.05 and mound axis ness scheffer 1958 states that mima mounds direction was negatively correlated with are generally circular in shape as seen from linearity of mound arrangement r 335335.335 above regardless of slope vitek 1973 in p05Ppos05os05.05 southern colorado found that mounds tend to be nearly circular even on slopes up to 10410.410 4 discussion in steepness on the columbia plateau however mounds are usually described as being the major patterns of variation of mound more elongate on slopes in southern wash- and roundfieldmoundfieldmoundfield characteristics with slope ington and northern oregon waters and steepness and aspect are listed below flagler 1929 reported that the mean ratio of 1 mounds show maximum size circular major to minor axes increases with increasing and symmetrical form low connection and slope to a value of 1431.43 on slopes of 6 steep- linearity and well developed stone circles on ness the long axis of these mounds is said to level sites be parallel to the slope malde 1964 noted 2 on slopes mounds become smaller and that mounds in southwestern idaho are typi- more elongate and asymmetrical with the cally circular but that on hillsideshillsides they are long axis parallel to the slope and show noticeably elliptical with the long axis di- greater connection and linearity of arrange- rected downslope kaatz 1959 in central ment washington found that the typical mound is elliptical in shape with a ratio of major to 3 on slopes stone circles become weaker minor axis of about 1.41 the long axis being especially on the downslope side of mounds minor 141 parallel to the slope 1963 found and beds diverge to form olmsted stone downslope that mounds tails in eastern washington are often elongate the ratio of major to minor axis being 4 slope effects general are in more inin- 11iili1.11 1 151.5lsis he also stated that the long axis is facing tense on south- and east slopes than on aligned with prevailing winds and is some- north and west facing slopes except for con- times across slopes rather than parallel to nectionnection which tends to peak on gentle north them fosberg 1965 stated that in twin falls facing slopes county idaho mounds elongate into down- much variation exists in the literature con- slope stripes cerning the steepness of slopes on which A degree of connection or confluence of mounds occur waters and flagler s 1929 mounds and their alignment into rows parallel data on the columbia plateau and nearby to the slope has been noted by several work- areas record mounds on slopes up to only 606 ers waters and flagler 1929 malde 1964 in steepness and kaatz 1959 stated that and fosberg 1965 describe mounds on the mounds occur on slopes up to about 770 in columbia plateau as forming beadlike rows steepness brown 1951 however reported along small drainage divides or between stone that mounds in this region occur on slopes up stripes on steeper slopes perhaps the best to 35 45450 vitek 1973 reported mounds in overall description of this pattern together southern colorado on mountain slopes up to with that of mound form is given by brown 20 in steepness and in southern california 1951 for sites near maupin oregon cox 1984 found mounds on slopes up to 30300 on in steepness price 1949 stated that mounds the steeper slopes they are oriented in more or less parallel lines along the rill divides tend to be elongate occur in the western states in mountain mead- and coalesce and are not as high nor as perfectly kept up ows with slopes up to 20 30 this variation in as on the level looking at these slopes from a distance 1990 MIMA MOUNDS 27

or studying aerial photos one gains the impression of a for a similar small mound on a slope how- continuous mound down the slope as though it consti- tuted the entire rill divide A close inspection however differences in translocation would result ever reveals that the crest ofthe strip is not even that even if the amount of tunneling activity re- it is divided by well formed mounds rising above the mained the same in terms of distance and level ofthe surrounding soil direction from the mound center on the fewer authors describe changes in the sides of the mound lying on the slope contour arrangement of sorted stone beds as slope moundwardmoun dward and upward translocation will be steepness increases malde 1961 1964 states similar to soil movements on a mound on level that in southwestern idaho the stone pave- ground on the downslope side of the mound ments surrounding mounds change progres- however an animal must move soil a greater sively to parallel stone stripes running up and vertical distance to achieve the same horizon- downslope implying the disappearance of tal displacement since this requires greater pavement sections on the up and downslope energy expenditure horizontal displacement edges of mounds kaatz 1959 stating that will probably often be less than expected on stone circles and networks change to sorted the upslope side of the mound in contrast a stone stripes on steep slopes also notes that given horizontal displacement will occur with sorted stripes may occur without any upslope less vertical displacement in some cases of connection to the former features brunn course much of the actual horizontal dis- schweiler 1962 describes a similar pattern placement will be downslope thus expendi- and diagrams a configuration in which stone ture of the same energy will lead to a greater 11 tails arise from stone rings encircling than expected horizontal displacement mounds or from intermound polygonal net- the consequence of differences in mean works to extend downslope pyrch 1973 displacement distance is that more soil will be noted that sorted stone stripes occur on slopes translocatedtranslocated onto the upslope side of the up to a maximum steepness of 15 33 at the mound than onto the downslope or lateral LMGP cox and alienallenailen 1987a found that on sides the mound should thus grow most in a level areas the development of stone circles is lateral and upslope fashion however this directly correlated with mound size and that growth could permit a circular form to be on slopes the initial pattern of modification is retained as long as the average ofupslope and the weakening of the bordering bed on the downslope addition rates equals the addition downslope side of the mound and the diver- rates to the lateral edges ofthe mound such a gence of downslope tails pattern will prevail whenever the mean hori- are these mound features compatible with zontal translocation distance of soil at a given the basic hypothesis of origin of both mounds distance from the mound center is linearly and associated stone circles polygonal nets related to the mean slope of the translocation and stripes by the soil translocation activities path fig 2 of pocket gophers and if so how does this if the relationship of mean displacement mechanism interact with site characteristics distance to slope is curvilinear and convex related to steepness and aspect to yield the however then additions to the lateral edges observed patterns will be greater than expected and to the up- let us first consider the implications of the slope and downslope edges less than ex- relationship of the intensity of mounmoundwarddward pected thus the mound will expand in width soil translocation by pocket gophers to dis- across the slope if the relationship is curvi- tance from the center of a small mound and linear and concave additions to the upslope elevation below its top as observed by cox and downslope edges will be greater than ex- and alienallenailen 1987b for a mound on a level site pected and those to the lateral edges less than average mounmoundwarddward translocation increased expected the mound will elongate up and with distance from the mound center and downslope in the latter case the total average upward translocation increased with amount of soil translocatedtranslocated onto the down- elevation below the mound top on a level slope side of the mound will still be much less site these tendencies would be distributed than that moved onto the upslope surface As symmetrically other factors being equal and the mound grows in height addition to the the mound would tend to enlarge symmetri- downslope side of the mound will also de- cally maintaining a circular shape cline at the same time slope conditions on 28 G W cox volume 50

the slope this arrangement will give a strong 1401 tu measure of linearity if the two nearest neigh- i bors of a mound are upslope and 120 given E downslope in the connected line strong lin- 0 of slope would 1 earity arrangement on a thus 15 100 a i imply that the distance E mean between D1 mounds in the same line is less than that be- 8 80180 i1 & tween mounds in different lines this should a i1 not be the case if uniformly spaced mounds 601goigol60 1 2 develop on a slope and those directly up and j i downslope from each other become con- 40640 1 A our 0 lo- x nected data show only a weak relation- 2 ship between slope steepness and linearity 20 suggesting that little more than the connec-

i i tion of mounds lying and downslope from n 5 L L i r i i up 00 1 0 20 40 50 60 70 each other has occurred of ri angle slope the presence of a mound on a slope would fig 2 horizontal displacement distance mound modify the flow of water across the slope ward for soil mined by thomomys bottaibottae min southern concentrating it along the upper and lateral california in relation to slope for data from cox and alienallenailen sides of the mound and producing a dry 1987b shadow downslope cox and alienallenailen 1987a the concentrated flow of water along the the upslope side ofthe mound will still permit sides of a mound would tend to continue di- heavy translocation onto the mound surface rectly downslope these areas of maximum thus slope relationships should cause ero- wetness favor the formation of sorted stone sional loss of soil to balance moundwardmoundward beds extending downslope several possible translocation sooner on the downslope side of mechanisms may contribute to the transfor- the mound than on the upslope side upslope mation of beds encircling the mounds into growth should therefore continue after down- elongate stripes paralleling them growth of slope growth has stopped fleshy rooted plants may lead to extensive data from soil translocation studies of tunneling by pocket gophers in these areas thomomys bottaibottae in southern california cox the collapse of deep tunnels and the down- and alienallenailen 1987b suggest that soil transloca- ward settling of soil and small stones during tion by pocket gophers of this genus varies the wet season may thus sort and expose with steepness in a curvilinear concave fash- stones near the surface cox and alienallenailen 1987a also play an ion over a range from less than 5 to more than erosion may important role par- ticularlyticularly as slope steepness 50 fig 3 these data werewe re obtained in stud- increases ies of soil translocation on level sites where figure 4 diagrams a hypothesis of how cir- the only slope was that of the mounds them- cular isolated mounds surrounded by sorted selves stone nets become transformed into elongate interconnected mounds bordered by linear elongation of mounds on the columbia beds of sorted stones hypothesis pre- plateau should be coupled with an upslope this dicts that lines of mounds on slopes should be movement of the mound high point at maxi- separated by two stone stripes one represent- size highest of elongated mum the point an ing the fusion of downslope extensions of the mound should thus be nearest its upslope encircling beds of each line of mounds exam- end ination of aerial photos of sloping areas on the elongation of mounds should lead to con- LMGP shows that this is generally true nection with adjacent mounds up and these observations strongly support the over- downslope when the soil mantle is deep all hypothesis that pocket gophers interact enough to permit the development of large with physical conditions and processes to pro- mounds such connection may create a linear duce the distinctive patterns of mounds and beaded arrangement of mounds parallel to sorted stone beds on the columbia plateau 1990 MIMA MOUNDS 29

1 0I D s ea e0 7 C 0 ay 5 c ca 404 0 0 C CO PclocluCIO Q g 0 caCQ 3 t3ta C 03 1 2 0 A downslope steep upslope slope steepness h

ac 3 03 2

00 404c 0 D 11E A 110 C IIcw 12aaC

0 CCO

B 0 downslope lateral upslope edge edges edge

fig 3 A possible relationships between mean horizontal translocation distance mounmoundwarddward and slope steepness B volume ofsoil moved onto mound surface at various points around mound perimeter by pocket gopher translocation in relation to slope steepness based on the relationships outlined in A for a mound of a given slope 30 G W cox volume 50

A 0 0 0 D B

LEVEL 0

VERY GENTLE SLOPE

D j f

GENTLE SLOPE 0 1

STEEP SLOPE

fig 4 diagrammatic model of the transition of unconnected circular mounds with encircling stone rings on level sites to linearly connected elongate mounds bordered by stone stripes on steep slopes 1990 MIMA MOUNDS 31

acknowledgments DALQUEST W W AND V B SCHEFFERSCHEFFEB 1942 the origin oftheodtheofthe mima mounds ofwesternofwestern washington jour- I1 thank elizabeth lucas for assistance min nal ofgeology 50 688468 84 field studies and in laboratory analyses of DIXON WJW J 1983 BM DP statistical software university mound and roundfieldmoundfieldmoundfield geometry and annan ofcalifornia press berkeley fridaypriday for permission to conduct much of the fosbergFOSBERCFOSBEBC M A 1965 characteristics and genesis of pat- work on areas of the fridaypriday brothers terned ground in wisconsin time in a chestnut soil ranch inm southern idaho soil science 99 30 37 adjacent to the LMGP B donald lawrence HITCHCOCK C L AND A CRONQUISTCKONQUIST 1973 flora of gave advice and encouragement throughout the pacific northwest university of washington the study J hunt J R mackay and V B press seattle scheffer provided valuable comments on an KAATZ M R 1959 patterned ground in central earlier draft of the manuscript this study was washington a preliminary report northwest carried science 33 145 156 out under a contract with the oregon MALDE H 1961 patterned conservancy E ground ofpossibleofpossible solifluc- chapter of the nature and was tion origin at low altitude in the western snake supported in part by grant INT 8420336 from river plain idaho united states geological sur- the USU S national science foundation vey professional paper 424 B 170170173173 1964 patterned ground on the western snake plain CITED river idaho and its possible cold climate literature origin geological society of america bulletin 75 BROWNBBOWN H C 1951 mound nncrorehefmicromieromicroreliefrelief of the columbia 191 207 plateau and adjacent areas unpublished manu- OLMSTED R K 1963 silt mounds of missoula flood sur- script 36 appp faces geological society of america bulletin 74 brunnschweilerBRUNN SCHWEILER D 1962 the periperlpenpeopenglacialperiglacialglacial realm in 47 54 north america during the wisconsin glaciation PRICE W A 1949 pocket gophers as architects of mima biuletyn peryglacjalny 11 15 27 pimple mounds of the western united states COPELAND W N 1980 the lawrence memorial grassgi ass texas journal of science 1 1 17 land preserve a biophysical inventory with man- PYRCH J B 1973 the characteristics and genesis ofstoneofstone agement recommendations unpublished report stripes in north central oregon unpublished the- revised 1983 oregon chapter nature conser- sis portland state university portland oregon vancy portland SCHEFFER V B 1947 mystery oftheofodthethe mima mounds cox G W 1984 the the distribution and origin of mima scientific monthly 65 283 294 mound grasslands in san diego county califor- 1958 do fossorialfossmessonalonai rodents type nia ecology 65 1397 1405 originate mima microreliefmicromieromicrorelieprelief american midland naturalist 59 cox G W AND D W ALLEN 1987a sorted stone nets 505 510 and circles oftheodtheof the columbia plateau a hypothesis VITEK J D 1973 mounds of northwest science 61 179 185 the ofsouthsouth central colorado an investigation of geographic and 1987b soil translocation by pocket gophers a geomorphic in characteristics unpublished dissertation mima roundfieldmoundfieldmoundfield oecologiaoecologia7272 207 2102 10 okla- homa state university stillwater coxcoxgcogg G W ANDANDCC G GAKAHU 1986 A latitudinal test of the fossorialfossboss onalonai rodent hypothesis of mima mound WATERS A C ANDANDCC W flaglerFLACLERFLAGLEK 1929 origin of small mounds plateau origin in western north america zeltzeitschnftzeitschriftZeitschrift farfur on the columbia river american geomorphologie 30 485485501solsoi501 journal of science 18 209 224 cox G W CG G GAKAHU AND D W ALLEN 1987 the ZAR J H 1974 Biostatistical analysis prentice hall small stone content of mima mounds in the co- englewood cliffs new jersey lumbia plateau and rocky mountain regions implications for mound origin great basin natural- received 10 june 1989 ist 47 609 619 accepted 30 november 1989 great basin naturalist soisol501 1990 appp 33 39

ESOX LUCIUS ESOCIDAE AND stizostedion VITREUM PERCIDAE IN THE GREEN RIVER BASIN COLORADO AND UTAH

harold M tyus and james M beardbeard1212

ABSTRACT northern pike esox luciusluctus stocked in the yampa river in 1977 invaded the mainstream green river by 1981 and subsequently increased in range and abundance the speed of this invasion is indicated by two recaptured pike that moved 78 and 110 km respectively downstream in about one year pike stomachs n 123 were usually empty 54554554.554 5 but some contained fish 43 and nonfish items 24242.42 4 red shiner notropis lutlutrensisrensis and fathead minnow pimephales kromelaspropromelaspronielaspronimelaselaseias predominated among the 12 fish species eaten walleye stizostedion green vitreum presumably introduced to the river drainage in the 1960s was widely distributed but low in abundance most of 61 adult walleye stomachs contained food 60760760.760 7 of 6 fish species eaten channel catfish ictaluruslctalurus punctatuspunctatus and fathead minnow were most frequently consumed northern pike and walleye were captured in habitats occupied by endangered colorado river fishes particularly colorado squawfish ptychocheilus lucius predation on endangered fishes was not detected but northern pike and walleye consumed at least threethithl ee otherothel native fishes the northernorthernn pike may pose a threat to endangered fishes due to its population expansion piscivorypiscivoiypiselpiscivory and resource sharing diets of northern pike and walleye species should be further evaluated if their abundance increases

northern pike were introduced into elk- carlson 1982 US fish and wildlife service head reservoir an impoundment on the 1987 however over 20 normative fishes yampa river drainage in 1977 P J mar- have been introduced into the basin for sport tinez personal communication and collected forage food or by accident tyus et al 1982 in the mainstream yampa river as early as fishes of upper colorado impacts of these 1979 E J wick personal communication introduced fishes on the native fauna are not their numbers increased in the upper yampa well understood but the presence of two river in the early 1980s wick et al 1985 and large piscivorespiscivores northern pike esox lucius a downstream movement into the green and walleye stizostedion vitreum in areas river was subsequently documented in 1981 presently occupied by endangered fishes is tyus et al 1982 green river fishery investi- cause for concern control ofnonnativeof nonnative fishes gationsgations northern pike reproduction has has been identified as a recovery measure been reported in the upper yampa river under provisions of an interagency recovery drainage where it has access to the main- program for endangered fish species in the stream river T P nesler personal commu- upper colorado river basin USU S fish and ninication wildlife service 1987 fish introductions in walleye presumably accessed the main- other locations have eliminated or partially stream green river by moving downstream extirpated native fish faunalfaunas and the instabil- from various tributatributariesries the fish was first ity of resultant communities has caused man- reported in utah in 1951 sigler and miller agement problems moyle et al 1986 1963 and reproducing populations of wall- the purpose of this study was to determine eye were established by fish stockings in diets of northern pike and walleye in the green duchesne river reservoirs fig 1 in the river and to evaluate the degree predationofpredationof 1960s and 1970s G M davis personal com- on native and endangered fishes we also doc- municationmunication ument the recent invasion of northern pike the green river basin of colorado and into the green river basin and the abun- utah is an important recovery area for four dance and distribution of northern pike and rare and endangered colorado river fishes walleye in the mainstream green river the reviewed by joseph et al 1977 carlson and results of this study are interpreted relative

1 USU S fishpish and wildlife service 1680 W highway 40 vernal utah 84078 present address 1361 vernon st emekaeureka california 95501 34 H M TYUSANDJTYUS AND J A BEARD volume 50 Y WYOMING

flamingFI gorge N reservoir

yampa riverriper aaa& 014 4140 dinosaur national monument

UTAH COLORADO

40 0 80km

fig 1 map ofoftheodthethe study area 1990 GREEN RIVER FISH ECOLOGY 35

to potential interactions of these species with TABLE 1 catches ofadult northern pike esox lucius sympatric endangered colorado river fishes and walleye stizostedion vitreum per hour of elec trofishing ch in the green river utah april june 1984 1988 n number offishof fishhishbish upper green river METHODS km 3378337.8 552 lower km 35 3377337.7 pike and walleye northern were primarily northern pike walleye collected by electroelectrofisbingelectrofishingfishing sampling was river hours location fished n n conducted from april to november 1979 ch ch 1981 and from april to june 1984 1988 in 517 1984 km ofthe mainstream green river the study upper 1012101.2 20 2020.20 15 lsis15.15 area included the mainstream green river lower 323 0 0 from confluence yampa its with the river in 1985 dinosaur national monument to a point 35 upper 8869 4 005005.005 8 oi0101.01 km above the confluence of the green and lower 289 0 0 colorado rivers fig 1 the lower 73 km of 1986 the yampa river was also sampled 1984 1989 upper 01.01 in 1989 spring sampling was conducted in the 7532753.2 5 oi01 17 0202.02 lower 359 0 0 lower 73 km of the yampa river and in the green river in a 208 km reach below its con- 1987 fluence with the yampa river fig 1 upper 7601 16 0202.02 8 oi0101.01 in 1979 1981 fishes were sampled with a lower 372 1 0303.03 1 0303.03 variety of gear including electroelectrofishingfishing 1988 seines trammel nets and wire traps depend- upper 4415441.5 9 02 0 ing on gear suitability sampling was con- lower 43043.0 3 0707.07 1 0202.02 ducted during preprerunoffrunoff runoff and post runoff conditions in 1984 1988 sampling included only alongshore electroelectrofishingfishing in the from 1979 to 1989 including 33 females in prerunoffpreprerunoflfrunoff and early runoff period and in- breeding condition mature ovaries with ripe volved continuous downstream coverage with eggs ripe females were captured in april a pulsed DC unit Electroelectrofishingfishing collections june in the mainstream green river at water in which all shoreline habitats were sampled temperatures of 10 19 C all pike were con- were considered representative collections sidered adults or large juveniles based on size and catches of fishes per hour ch sampled average 619gig mm TL range 321 10451045mmmm some 619mmtl were recorded opportunistic spring carlander 1969 average catch of north- electroelectrofishingfishing was also conducted in sus- ern pike increased 00050.0505 0140.14 fish per hour pected northern pike and walleye habitats from 1984 to 1988 table 1 seventy eight however no data were reported for these ch percent n 59 of the pike were collected in samples the upper green in 1984 1988 but all northern pike and walleye collected river many of these 43 were taken in shallow were measured for total length TL location low velocity shoreline habitats at the mouth of capture was also noted 1983 all fish after of ashley creek northern pike were spotty were sacrificed and stomach identi- contents in distribution but fied to lowest possible sometimes abundant in the taxon with the aid of semisemiimpounded habitats a dissecting impounded their captures 25x binocular scope the date were often associated with location prominent aquatic and water conditions at the point of and bank vegetation capture of all females with ripe eggs and fully we developed ovaries were recorded we also captured two tagged northern pike in this obtained 49 northern pike and 11 walleye study these adult fish 594 and 820 mm stomachs from other workers and identified TL were tagged by colorado division of their contents wildlife personnel in the yampa river in 1982 and 1988 E J wick and T P nesler per- RESULTS sonal communication one fish had moved about 110 km between 15 april 1982 and abundance and distribution 10 may 1983 when we recaptured it in the eighty four northern pike were collected yampa river at km 18418.4 the other pike had 36 H M TYUSANDJTYUS AND J M BEARD volume 50

TABLE 2 contents of 123 northern pike esox lucius stomachs taken in the green river basin utah and colorado 1984 1989

number frequency species status offreyofpreyofprey FISHES unidentified fish 40 106 notropis lutlutrensisrensis I1 24 73 pimephales kromelaspropromelasmelas I1 15 41 catostomus latiiatilatipinnislatipmmspinnis N 6 414.14 1 rhinichthysrhimchthys esculusosculus N 7 33 unidentified notropis sppapp I1 6 24 gila atrariaatranaadrana I1 3 24 catostomus discobolus N 2 16 ictalurus pmctatuspunctatuspunctatus I1 2 16 cyprinus carpio I1 3 08 notropis strastramineusstramineousstramstrammeusmineusmeus I1 2 08 unidentified cyprinidae I1 1 bu 08 unidentified cilacliagila1 sppapp 1 08 oncorhynchus mykiss I1 1 08 oncorhynchus clarki I1 1 08 Richardrtchardsonwsrichardsoniussonius balbalteatusteatus I1 1 08 OTHER empty 545 rana pipienspipkens N 1 08 lampropeltislampropeltts sppapp N 1 08 detritus 08 N native species I1 introduced species isuspecteduipected cdacliagilaacda lobustarobusta traveled 78 km from 16 june 1988 to 23 may mouth sucker catostomus latilatipinnispinnis blue 1989 when it was recaptured at km 484.8 head sucker C discobolus and speckled growth of these fish averaged only 10 mm TL dace rhinichthys osculus were the native walleye were also captured in the upper fishes consumed other prey items included a green river 90 n 50 and averaged 511 leopard frog rana pienspipipienspipkens a king snake mm TL range 395 686 mm these fish were lampropeltis sppapp and detritus thirteen presumed juveniles and adults based on size stomachs log10610.6 contained fish remains that carlander 1969 more widely dispersed than could not be identified pike walleye were usually captured in a vari- walleye in the green river primarily con- ety of slow shoreline runs usually associated sumed fishes including 5 nonnative and I1 with emergent or bank vegetation one ripe native species table 3 of61 stomachs exam- female walleye 577 mm TQTL was captured in ined 24 39339339.3 were empty and 10 16416416.4 the upper green river on 15 may 1984 at a contained unidentifiable fish remains chan- water temperature of 13 C we captured one nel catfish ictalurus punctpunctatusatus and fathead tagged walleye at the mouth of the duchesne minnow were the most frequently consumed river on 21 may 1984 this fish was tagged by nonnative fishes and flannelmouth sucker was BIOWEST incorporated on 13 april 1979 at a the only native fish consumed vascular plant point about 37 km upstream in the green material was found in one walleye stomach river L cristgrist personal communication this fish grew about 62 mm TL in five years discussion foods northern pike introduced in the yampa northern pike stomachs n 123 were river drainage in 1977 was presumed absent usually empty 54554554.5 but of the remainder in green river until first reported in 1981 97697.6 contained fishes table 2 red shiner tyus et al 1982 fishes of upper colorado notropis lutrensislutrensis and fathead minnow we captured the fish only in the upper green pimephales kromelaspropromelasmelas were most frequently river km 3378337.8 552 from 1981 to 1986 pike consumed of nine nonnative fishes flannel invaded the midsection km 192 3377337.7 by 1990 GREEN RIVER FISH ECOLOGY 37

TABLE 3 contents of 61 walleye stizostedion vitreum stomachs taken in the green river basin utah and colorado 1984 1989

number frequency species statusa offreyofpreyof prey FISHES unidentified fish 38 164 ictalurus punctatuspunctatus I1 37 164 pimephales kromelaspropromelasmelas I1 35 98 cyprinus carpio I1 4 66 lepomis cyancyanellusellus I1 4 33 catostomus latipinnislatipinmslatiiatipinnis N 1 161 6 unidentified cyprinidae I1 2 16 ictalurus melas I1 1 161.6iglg1 6 OTHER empty 393 fish eggs 16 vascular plant material 161.61lgig 6 anN native species I1 introduced species

1987 and it was first captured below green tyus et al 1982 fishes of upper colorado river utah kmkni 192 in 1988 this invasion and this may have resulted in the relative and downstream movement is supported by absence of roundtail chub as prey in pike both the absence oftheodtheof the fish in the green river stomachs we examined in the early 1970s holden and stalnaker northern pike may spawn in the main- 1975 and a first report of pike in the lower stream green river but if so recruitment is green in 1988 M moretti personal commu- low we did not capture small northern pike nicationni although movements of northern 321 minmm TL in this study and to our pike in large rivers remain poorly docu- knowledge pike reproduction has not been mented some studies in lakes and small noted by others however one 115 mm TL streams have shown that the fish can display specimen was seinedheined by HMT and others high mobility miller 1948 ross and winter from a shoreline area of the green river in 1981 but may move only short distances at a dinosaur national monument on 8 july 1988 time cook and bergersen 1988 our recap- it is not known whether this fish hatched in ture of two northern pike indicated that the the green river or was transported there fish can move long distances 75 kmyearkm year from another location also we captured sev- in the yampa river long distance upstream eral ripe female pike and it is possible that and downstream movement of radiotaggedradiotagged some of these fish spawned in the green northern pike has also been reported by river most ripe female pike 76 had empty T P nesler personal communication stomachs suggesting a reduction in feeding the majority of fishes consumed by north- activity with increasing water temperatures ern pike in this study were soft rayed forms and ripening ovaries frost 1954 lawler table 2 as previously noted by others 1965 beyerle and williams 1968 weithman and walleye were rare in the green river and anderson 1977 wolfert and miller 1978 their long period of residency suggests that channel catfish the only spiny rayed fish their numbers will probably not increase consumed was found in two stomachs we walleye were easily captured by electrofish could not positively identify roundtail chub ing and very few fish that we sighted escaped gila robusta in northern pike stomachs taken capture however it was difficult to capture from the yampa river but presumably one northern pike with electroelectrofishingfishing and many cilacliagilaglia sppapp was a roundtail chub T P nesler fish escaped A direct comparison of the rela- personal communication reported that tive abundance of walleye with that of north- roundtail chub were present in northern pike ern pike could be somewhat misleading and stomachs he examined from the yampa river it is noted that walleye were more rare and most of the pike we examined were from the northern pike more abundant than indicated green river where roundtail chub are rare by electrofishingelectrofishing catch rates we captured 38 H M TYUS ANDJAND J M BEARD volume 50 only one female walleye with developed acknowledgments ovaries and that was in may at a water tem- peratureperature of 13 C walleye in other locations this work was funded by the bureau of usually spawn at cooler water temperatures reclamation and the fish and wildlife ser- numerous 33333.3 727.2 C sigler and miller 1963 565.6sg lillii11111.1 vice fish and wildlife service C scott and crossman 1973 no small wall- employees assisted with data collection we eye 395 mm TL were captured in ththiss thank S R cranny and M moretti of the study utah division ofwildlife resources and R A valdez of young of the endangered humpback chub BIOWESTRIOWEST incorporated for pro- viding northern pike and walleye gila cypha bonetailbonybonytailtail chub G elelegansdelegansegans razor- stomachs G B haines and D moses back sucker xyrauchen tetanustexanustexanus and colo- aided in data summarization and word C rado squawfish ptychocheilus lucius may be processing A karp P pister and potential prey for northern pike and walleye an anonymous reviewer improved an earlier draft none of these fishes were identified in stom- manuscript achs of northern pike or walleye but our ability to detect such predation was constrained literature CITED by a small sample size of stomachs that con- BEYERLE G B ANDANDJ WILLIAMS some tained food of endangered fishes and J E 1968 observa- rarity tions of food selectivity by northern pike in inability to identify all of the fishes eaten aquaria transactions of the american fisheries sympatry of adults of northern pike wall- society 97 283128 31 CARLANDERCARLANDEB K D 1969 handbook of eye and endangered fishes is a cause for con- freshwater fishery biology vol I1 iowa state university press cern particularly if resource sharing occurs ames 752 appp during periods of limited availability we col- CARLSON C A AND E M CARLSON 1982 review of lected northern pike walleye and colorado selected literature on the upper colorado river squawsquawfishfisli in similar shoreline habitats in the system and its fishes pages 181 8 in W H miller green H M tyus andandcanacC A carlson eds fishes ofthe mainstream river in addition radio upper colorado river system present and future tagged northern pike and colorado squawfish american fisheries society bethesda maryland were syntopicsyncopicsyntopic in the green and yampa rivers 131 appp valdez and masslich 1989 wick and hawkins COOK M F AND E P BERGERSEN 1988 movements 1989 northern pike captured shal- habitat selection and activity periods of northern were in pike in eleven mile reservoir colorado transact- low flooded habitats also utilized by razor- ions of the american fisheries society 117 back sucker 495 502 stocking programs for northern pike and FROST W E 1954 the food of pike esox lucius L in windermere journal of animal ecology 23 walleye have been discontinued by state 339 360 agencies in colorado and utah G M davis HOLDEN P B AND C B STALNAKER 1975 distribution and P J martinez personal communication and abundance of mainstream fishes ofthe middle and the relative absence of small fish of both and upper colorado river basins 196719731967 1973 species suggests reproduction transactions of the american fisheries society that in the 104217 231 green is mainstream river low or nonexistent JOSEPH T W J A SINNING R J BEHNKE AND P B during most years however the continuing HOLDEN 1977 an evaluation of the status life invasion of northern pike and walleye into the history and habitat requirements of endangered green river from established reproducing and threatened fishes ofthe upper colorado river stocks should be system FWSOBS 77 62 US fish and wildlife monitored and their interac- service office of biological services fort tions with endangered fishes further evalu- collins colorado 194 appp ated until it can be more clearly demonstrated LAWLER G H 1965 the food of the pike esox lucius in that competition or predation on endangered hemming lake manitoba journal of the fisheries resource board of 22 fishes does not occur or pose a canada 2213579213571357 1377 serious threat MILLER R B 1948 A note on the movement ofthe pike the increasing abundance and spread of esox lucius copela 1948 62 northern pike the diversity of fishes con- MOYLE P B H W li AND B A BARTON 1986 the sumed and its syntopysyntomy with endangered frankenstein effect impact of introduced fishes fishes make this voracious piscivorepiscivore a poten- on native fishes in north america pages 415 426 tial in R H stroud ed fish culture in fisheries threat to endangered colorado river management american fisheries society be- fishes thesda maryland 481 appp 1990 GREEN RIVER FISH ECOLOGY 39

ROSS M J ANDJAND J D WINTER 1981 winter movements of USU S FISH AND WILDLIFE SERVICE 1987 recovery imple- four fish species near a thermal plume in northern mentation program for endangered fish species in minnesota transactions of the american fish- the upper colorado river basin USU S depart- eries society 101410 14 18 ment of the interior fish and wildlife service SCOTT W B AND E J CROSSMANCKOSSMAN 1973 freshwater division of endangered species denver colo- fishes of canada bulletin of the fisheries res- rado six sections various pagination I1 earch board ofcanada 18419661841184 19661 966 VALDEZ R A AND W J MASSLICH 1989 winter habitat SIGLER W F AND R R MILLERMILLEB 1963 fishes of utah study of endangered fish green river report utah state department of fish and game salt 1362136 2 BIOWESTBIGWEST incorporated logan utah lake city 203 appp 184 appp TYUS H M B D BURDICKBUBDICK R A VALDEZ C M HAYNES WEITHMAN A S AND R 0 ANDERSON 1977 survival T A LYTLE AND C R BERRY 1982 fishes of the growth and prey ofEsocidae in experimental sys- upper colorado river basin distribution abun- tems transactions of the american fisheries so- dance and status pages 127012 70 in W H miller ciety 106424106log 424 430 H M tyus andcandanacC A carlson eds fishes oftheodtheofthe WICK E JANDJJ AND J A HAWKINS 1989 colorado squawfish upper colorado river system present and future winter habitat study yampa river colorado american fisheries society bethesda maryland 1986 1988 final report contribution 43 larval 131 appp fish laboratory colorado state university I1 fort TYUS H M C W mcadMCADAA AND B D BURDICK 1982 collins colorado 96 appp green acad river fishery investigations 197919811979 1981 WOLFERT D R ANDANDTT J MILLER 1978 age growth and pages 1 99 in W H miller J J valentine D L food of northern pike in eastern lake ontario archer H M tyus BR A valdez and L kaed- transactions of the american fisheries society ing colorado river fishery project final report 107696 702 field investigations part 2 UUSS fish and wildlife service and US bureau of reclamationofreclamation salt received 25 july 1989 lake city utah 365 appp accepted 15 november 1989 great basin naturalist 501 1990 appp 41 46 INFLUENCE OF SOIL FROST ON infiltration OF SHRUB COPPICE DUNE AND DUNE interspace SOILS IN southeastern NEVADA wilbertwilberthH blackburnandblaekBlackburnBlackburnand1 and M karlkailkallkalikari woodwoodawood2

ABSTRACT the influence of soil frost on the infiltration rate of shrub coppice dune and dune interspace soils was evaluated near crystal springs nevada using simulated rainfall the infiltration rate of the coppice dune soil was greater than the dune interspace soil under frozen or unfrozen conditions because of different vegetation cover and surface soil characteristics coppice dune and dune interspace soils responded differently to freezing thus imposing a spatial and temporal responselesiesponse to infiltration laterate infiltration rate of soils with porous concrete frost increased as the soils thawed duringduidul ing simulated rainfalli but soils with nonporous concrete frost allowed very little infiltration to occur both coppice dune and dune interspace soils that were classified in january as having granular frost had a higher infiltration laterate than the same unfrozen soils in march

soil frost influences water infiltration rates tion during winter the study objective was to and is often a major influence on runoff gray determine the spatial variation in infiltration and granger 1987 harris 1972 haupt 1967 rates of frozen and unfrozen shrub coppice kane and stein 1983 klock 1972 kuzick and dune and dune interspace soils bezmenovBezmenov 1963 wilcox et al 1989 and zuzel and pikul 1987 areas where soil frost STUDY AREA strongly influences infiltration are characterized by cold winters transient snow cover the study area is located in southeastern and soils that may freeze and thaw several nevada about 7 kinkm west of crystal springs times each winter in addition to a diurnal stu3720STW N latitude 115201150201 W longitude at freeze thaw cycle pikul and allmaras 1986 1200 in elevation and 9 kinkm east of the zuzel et al 1986 infiltration rate of frozen 1850 2100 in ridgelineridridgelinggeline of the pahranagat soil is strongly influenced by the structure range the normal annual precipitation of of the soil frost which is determined in part 330 mm occurs mostly during winter as snow by the soil water content at the time of freez- or during july and august as thunderthundershowersshowers ing concrete frost has been identified as hav- winters are characterized by periodically cold ing the greatest impact on infiltration rates temperatures with frequent diurnal freeze haupt 1967 lee and molnau 1982 story thaw cycles 1955 blackbrushBlackbrush coleogyne ramosissimumramosissiinum is the spatial influence ofshrub coppice dune the dominant shrub with 22 crown cover and dune interspace soils on infiltration of associated species are joint fir ephedra unfrozen soil was originally established by nevadensis and box thorn lycium ander blackburn 1975 and verified by numerous soniijonii each with 2 crown cover herba- other investigators johnson and gordon ceous cover is sparse the study site is 1988 swanson and buckhouse 1984 thurow located on the tops of long narrow alluvial et al 1986 wood and blackburn 1981 wood fans with 5 slope to the east and the soils et al 1987 because shrub coppice dune and are loamy skeletal mixed thermic shallow dune interspace soils have different vegeta- typic durorthids tion cover and surface soil characteristics we two major types of surface soils are found hypothesized that they will respond differ- on the study site the coppice dune soil under ently to soil freezing and thawing thus impos- shrubs and the barren interspace soil between ing a spatial and temporal response to infiltra shrubs the coppice dune soil covers 35 of

SDAUSDA agricultural researchReseaichsearch service northwest watershed research center 270 south orchard boise idaho 83705 department of animal and range science new mexico state university las cruces new mexico 88003

41 42 W HhblackburnBLACKBURN AND MMKK WOOD volume 50

the surface whereas interspace soil covers cochran 1971 were used to test for differ- 65 the A horizon of the coppice soil is ences between infiltration rates of the coppice characterized by a weakly subangular blocky dune and dune interspace soils for the january structure and a gravelly sandy loam texture and march sample dates interspace soils have gravel pavement several pebbles thick over the mineral soil the RESULTS AND discussion 50 mm thick loamy crusted A horizon is massive has vesicular pores and is broken all plots during january were classified as into 80 150 mm diameter polygons the having soil frost 100 minmiumm thick located about crusted interspace soil slakes and disperses 50 minmm below the surface three of the cop- readily when wetted pice dune and five of the interspace plots were classified as granular frost the remaining METHODS plots of both soils were characterized as porous or nonporous concrete frost of which infiltration rates were determined using a 12 coppice dune and 6 interspace plots were drip type rainfall simulator blackburn et al classified as porous concrete frost 1974 with simulated raindrops 252.5 minmm in infiltration rate after 25 min was signifi- diameter drops falling 212.1 in reach 5255.25 in cantly greater for coppice dunes than for dune 1 sec or 71 of the terminal velocity achieved interinterspacesspaces under both frozen and unfrozen by raindrops in an unlimited fall laws 1941 soil conditions fig 1 table 1 similar rela- simulated rainfall was applied on frozen soil in tiontionshipsships between unfrozen shrub coppice january and on unfrozen soil in march 1974 at dune and dune interspace soils have been 1 a rate of 76 minmiumm hr for 30 min the rainfall reported by blackburn 1975 johnson and intensity was chosen to assure runoff from gordon 1988 thurow et al 1986 and each study plot runoff plots 280 X 500 minmm wood and blackburn 1981 the differences were randomly placed on 16 and 6 shrub cop- in infiltration of coppice dune and dune inter- pice dune soils and 13 and 5 interspace soils space soils have been attributed to differences during the january and march sample dates in vegetation and surface soil characteristics respectively this sample size is considered blackburn 1975 johnson and gordon 1988 adequate for rangeland conditions wood infiltration of unfrozen rangeland soil is usu- 1987 shrubs were cut at ground level and ally characterized by a high initial rate that removed from the coppice dunes to reduce decreases rapidly with time and stabilizes rainfall interception losses at some constant rate within 30 to 60 min fig volume of runoffwas measured every 5 min 1 however mean infiltration rates in janu- for 30 min infiltration rates were determined ary declined within the first 15 min for the as the difference between simulated rainfall coppice dune soils and within 20 min for inter- and runoff volumes soil frost was character- space soils in neither case did they stabilize at ized adjacent to each runoff plot prior to the a constant rate infiltration rates for both soils simulated rainfall event three structural increased during the latter part of the rainfall forms of soil frost were observed and subjec- due to thawing of the porous concrete soil tively classified according to criteria by hale frost layer of some plots figs 2 3 As a 1951 and haupt 1967 granular frost con- result there was no significant difference in sisted of scattered granules ofice binding min- 30 min infiltration rates between frozen and eral soil together nonporous concrete frost unfrozen dune interspace soils however 30 was characterized by dense thin ice lenses min infiltration rates tended to be lower in the and ice crystals porous concrete frost was less frozen coppice dune soils in january than in dense than concrete frost but frozen chunks unfrozen soils in march fig 1 of soil were harder to break porous concrete infiltration rates of coppice dune plots frost was further defined by resistance to re- classified as granular frost were similar to peated thrusts of a pick before being punc- the rainfall application rate and 10 minmm hr 1 tured water used for rainfall simulation aver- greater than when the soils were unfrozen aged 4 C in january and 9 C in march in march figs 1 2 coppice dune plots analysis of variance and least significant dif- classified as porous concrete frost and thawing ference mean separation tests snedecor and during the rainfall event reached a minimum 1990 INFLUENCE OF SOIL FROST 43

80so

Q X 60

iv I1 0 40 g COPPICE DUNE 0 JANUARY 0 I- MARCH jJ 20 DUNE interspace JANUARY A MARCH 0 T 0 5 10 15 20 25 30 35 TIME MIN

fig 1 mean infiltration rates for all coppice dune and dune interspace soils in january and march crystal springs nevada

TABLE 1 significant difference of five minute interval infiltration rates for coppice dune and dune interspace soils for the january and march sample dates crystal springs nevada

time minutes soilsampleSoil Sample date 5 10 15 20 25 30 coppice march vs coppice january nsnsfns22 ns ns ns ns ns coppice march vs interspace march ns ns ns coppice march vs interspace january ns ns coppice january vs interspace march ns ns ns ns coppice january vs interspace january ns ns ns interspace january vs interspace march ns ns ns ns ns ns sample size coppice january n 16 march n 6 interspace january n 13 march n 5 clevel2levellevellevei ofsignificance P 01 PsP 05os05.05 ns nonsignificant at P 0505.05 determined with a one way analysis ofofvariancevariance and a least significant difference bsdisd mean separation test infiltration rate after 20 min of44 minmm hr but interspace plots that were initially classified 30 min rates increased to a rate similar to that as porous concrete frost and thawing during of the granular frost plots fig 2 the one the rainfall event reached a minimum infiltra- porous concrete frost plot that remained tion rate of 12 minmm hr 1 after 20 min after frozen during the rainfall event reached a which rates increased and were similar to un- minimum rate after 15 min and then increased frozen soils in march infiltration rate of the slightly during the remainder of the event to interspace nonporous concrete frost plot de- 1 1 33 minmm hr creased to 2 minmm hr at 30 min 21 minmm hr 1 infiltration rate at 30 min of intersinterspaceenters aceaee soil lower than the 30 of soils pgreaterogreaterpg min rate unfrozen in with granular frost was 14 minmm hr greaterreater march other researchers have reported sim- than when the soil was unfrozen in march ilar infiltration response caused by soil frost 44 W H BLACKBURN AND A K WOOD volume 50 80

13 I 60

40 0z

0 GRANULAR 20 FROST faf4 U POROUS CONCRETE frostthawingFROST THAWING Z t f tf POROUS CONCRETE FROST

0 r I1 I1 1 1 r I I 0 5 10 15 20 25 30 35 TIME MIN fig 2 infiltration rates in january for coppice dune soils classified as having granular frost porous concrete frost that was thawing during the rainfall event and porous concrete frost crystal springs nevada 80 0 GRANULAR FROST xor POROUS CONCRETE Z 60 frostthawi NG NON POROUS CONCRETE FROST or 40 0z

01 20

U Z

0 0 5 10 15 20 25 30 35 TIME MIN fig 3 infiltration rates in january for dune interspace soils classified as having granular frost porous concrete frost that was thawing during the rainfall event and nonporous concrete frost crystal springs nevada 1990 INFLUENCE OF SOIL FROST 45

structure trimble et al 1958 in new JOHNSON C W AND N E GORDON 1988 runoff and hampshire reported infiltration rate of soils erosion fromhrom rainfall simulator plots on sagebrush with granular frost to be higher than that of rangeland transactions of the american society ofagricultural unfrozen soils haupt 1967 found for the ofagricultural engineers 31 421 427 KANE D L AND J STEIN 1983 water movement eastern slope of the sierra the infil- into nevada frozen soils water resources research 19 1547 tration rate of porous concrete frost to in- 1557 crease as the soil thawed trimble et al 1958 KLOCK G 0 1972 snow melt temperature influence on and stoeckeler and weitzman 1960 found infiltration and soil water retention journal ofsoil infiltration rates of nonporous concrete frost and water conservation 27 12 14 in northern minnesota to be very low KUZICK I1 A AND A I1 BEZMENOV 1963 infiltration of feltwatermeltwatermeltwater into frozen soil soviet soil science 6 665 670 conclusions LAWS J D 1941 measurements of the fall velocity of water drops and raindrops transactions of the the infiltration rate of shrub coppice dune american geophysical union 2270922 709tog 721 soils was greater than dune interspace soils LEFLEE R W AND M P MOLNAU 1982 infiltration into under frozen and unfrozen conditions con- frozen soils using simulated rainfall paper no crete frost located 50 minmm below the surface 82204882 2048 american society of agricultural engi- had a pronounced effect on infiltration rates neers st joseph michigan infiltration rates of soils with porous concrete PIKUIPIKUL J L JBJR AND R R ALLMARAS 1986 physical and frost increased as the soils thawed during the chemical properties ofofaa haploxeroll after 50 years of residue management soil simulated rainfall soils science society of but with nonporous america journal 50 214 219 concrete frost allowed infiltration very little to SNEDECOR G W ANDWAND W G COCHRAN 1971 statistical occur both coppice dune and dune inter- methods iowa state university press ames space soils classified as having granular frost stoeckeler J H AND S WEITZMAN 1960 infiltration had a higher infiltration rate than the same rates in frozen soils in northern minnesota soil unfrozen soils in march due to different veg- science society of america proceedings 24 etation cover and surface soil characteristics 137 139 shrub coppice dune and dune interspace soils STORYSTOBY H C 1955 frozen soil and spring and winter floods yearbook responded differently to soil freezing and thus in of agriculture 1955 USU S department ofagriculture washington DCD C imposed a spatial and temporal response to SWANSON S R AND J C BUCKHOUSE 1984 soil and infiltration rate nitrogen loss from oregon lands occupied by three subspecies of big sagebrush journal of range management literature CITED 37 298 302 THUROWTHUBOW T L W H BLACKBURN ANDANDCC A TAYLOR JR BLACKBURN W H 1975 factors influencing infiltra- 1986 hydrologic characteristics of vegetation tion and sediment production ofseniofsemiof semiaridsemi andaridarld range- types as affected by livestock grazing systems plateau lands in nevada watelwater resources research 11 edwards texas journal of range man- 929 937 agement 39 505 509 S SARTZ BLACKBURN W H R 0 MEEUWIG AND C M SKAU TRIMBLE G R R AND R S PIERCE 1958 how 1974 A mobile infiltrometer for use on rangelandi angeland types of soil frost affect infiltration journal of soil journal of range management 2732227 322323322 323 and water conservation 13 81 82 GRAY D M AND R J GRANGER 1987 frozen soil the WILCOX B P C L HANSON J R WIGHT AND W H problem of snow melt infiltration in Y S fok BLACKBURN 1989 sagebrush rangeland hydrol- ed proceedings international conference on ogy and evaluation oftheodtheofthe SPUR hydrology model infiltration development and application water water resources bulletin 25 653 666 resources research center university ofhawaii WOOD J C M K WOOD ANDJAND J M TROMBLE 1987 im- honolulu portant factors influencing water infiltration and HALE C E 1951 further observations on soil freezing in sediment production on and lands in new mexico the pacific northwest pacific northwest forest journal ofariaofaridof and environment 12 111 118 and range experiment station researchReseaichsealch note WOOD M K 1987 plot numbers required to determine no 74 portland oregon infiltration rates and sediment production on HARRIS A R 1972 infiltration rate as affected by soil rangelands in southcentralsoutheentralsouth central new mexico journal freezing under three cover types soil science of range management 40 259 263 society ofamericaofamerica proceedings 36 489 492 WOOD M K AND W H BLACKBURN 1981 grazing sys- HAUPT H F 1967 infiltration overland flow and soil tems their influence on infiltration rates in the movement on frozen and snow covered plot wa- rolling plains of texas journal of range manage- ter resources research 3 145 161 ment 34 331 335 46 W H BLACKBURN AND M K WOOD volume 50

ZUZEL J F ANDJAND J L PIKUL JR 1987 infiltration into a journal of climate and applied meteorology 25 seasonably frozen agricultural soil journal of soil 1681 1686 and water conservation 42 447450447 450 ZUZEL J F J L PIKUL JR AND R N GREENWALT 1986 received 12 october 1989 point probability distributions of frozen soil accepted 12 december 1989 great basin naturalist soisol501 1990 appp 47 56 SEED production AND SEEDLING establishment OF A SOUTHWEST RIPARIAN TREE ARIZONA WALNUT JUGLANS MAJOR

juliet C S trombergStromberg 1 and duncan T fattenpatten

ABSTRACT A four year study offive populations has revealed influences on seed production and seedling establishment of the southwest riparian tree juglans major germination is abundant after production of large seed crops masts but masts are produced infrequently within years germination is stimulated by summer rains enabling seedlings to establish on riparian terraces as well as streamstreambanksbanks traits such as capacity for dormancy during summer drought allow some seedlings to survive on terraces but abundant rainfall is essential for high rates ofseedling success ranges ofmoisture tolerance vary among seeds collected from different populations suggesting that ecotypesecotypes may exist between riparian sites with dissimilar moisture regimes population based differences are associated in part with differences in seed size

arizona walnut juglans major torr I1 hel- central arizona germination safe sites for ler is a member of the interior riparian de- walnut may be more restricted in drier ripar- ciduous forest an assemblage of trees ian sites such as along ephemeral streams for that grow along streams in the interior south- example seed burial may become a prerequi- west brown 1982 walnut sometimes domi- site of germination as soil moisture decreases nates this community szaro 1989 but more as is true for certain oaks barrett 1931 often it occurs at relatively low densities and germination and establishment of walnut frequencies this distribution pattern while or other riparian species may also vary as a indicating that establishment occurs infre- function of genetic differences between popu- quentquentlyly does not reveal the stage where relations the existence of riparian ecotypesecotypes is generation is limited sudworth 1934 sug- not a new idea eg hook and stubbs 1967 gested low rates of seed production and high and riparian populations may differ between rates of seed predation by tree squirrels as isolated southwest watersheds with different possible natural causes of infrequent estab- hydrologic characteristics other studies in lishmentlishment seedling mortality from cattle graz- this series on walnut reproduction have iden- ing also may play a role rucks 1984 tified differences between populations in flo- lack of suitable germination and establish- ral ratios and seed weight stromberg 198811988 ment sites may limit recruitment but little is indicating that J major shows either plastic known regarding the relationship between responses or genetically based responses to these requirements and JJ major certain environmental differences within and be- generalizations have been made about the dis- tween riparian sites the objectives of this tributiontribution ofwalnut trees but the habitat char- study were to 1 identify factors that limit acteristicsacteristics of trees may differ from those in seed production and seedling recruitment of which the seedlings established A study that J major between sites and between years specifically addressed riparian seedlings re- 2 determine how germination requirements vealed that seedlings ofaofj major a facultative vary between micrositesmicrosites and 3 determine riparian species establish in many micrositesmicrosites how seedling growth response to soil moisture throughout the riparian zone this is in con- differs between populations trast to seedlings of obligate riparian trees such as alder oblongifolia arizona alnus METHODS torr that occur only immediately adjacent to the stream larkin 1987 larkin conducted seed production germination and seed- her study however in a wet montane area of ling survival ofaofj major were studied through

center for environmental studies arizona state university tempe arizona 85287120185287 1201

47 48 J C stromberganddSTROMBERG AND D T PATTEN volume 50

field and greenhouse experiments and field buried 2 cm deep sites were planted with observations five sites were selected in cen- their own seeds except for workman creek tral arizona hitt wash an ephemeral stream and rock creek and seedling size and sur- near prescott national forest rock creek an vival were recorded through 1985 microlitemicrositeMicrosite intermittent stream in the Mazamazatzaltzal moun- soil moisture was recorded monthly with a tains and perennial streams south fork dew point psychrometer and canopy cover walnut creek in prescott national forest and was estimated visually stromberg 1988 two sites along workman creek in the sierra influences on germination and seedling sur- ancha mountains stromberg 1988 eleva- vival were analyzed through multiple regres- tions vary from 11001 loo100 in at rock creek to sion analysis of germination and survival per- 22100mataztecpeakloo100 in at aztec peak cencentagestages with average seasonal values for microlitemicmicrositerosite moisture level canopy cover and field observations herbaceous cover twenty trees were selected at each site greenhouse experiments and samples of 20 shoots per tree were selected for nuts were collected in 1983 from trees at monitoring of seed production from rock 1983 to 1986 average values per tree were creek hitt wash and walnut creek to test for influence of seed calculated for seed production per shoot and weight and seed source on and seedling for flower production flower abortion seed germination growth nuts were sown inm the greenhouse min loam soil weight and seed viability stromberg 1988 in polyethylene lined pots four A mast large seed crop is operationally de- in under water- ing one was watered fined in this study as a crop 33 larger than regimes regime every day to average for the study other maintain saturated conditions others were watered at less frequent demography of natural seedlings was stud- inter- vvalsalsais producing average soil ied to determine when where and how many moisture contents by dry weight of 80 40 and 20 seeds germinate and to compare survival seeds were sown on the surface and buried rates between micrositesmicrosites one walnut tree 2 cm deep seven nuts from each of 24 near the stream and one on the adjacent parent trees were sown per germination terrace with nearby seedlings were selected treatment percent and speed and seedling survival were at each site each marked the center of an monitored eight weeks after emergence 8 m radius circular plot all seedlings in the plants were harvested for of plots were tagged and scored monthly from measurement dry weight stem height and root length each may to october of 1983 1984 and 1985 for at soil level stem height leaf length and number and moisture regression analysis was used to test for influence of seed weight a mortality twenty seedlings in a plot at hitt continuous variable s multiple wash were excavated to measure depth of the duncan range test was used to test for effects of nut in the soil profile seed source on germination and seedling growth rates field experiments RESULTS seeds were sown in field exclosures in fall 1982 and 1983 to test for influence ofmicrolitemicrositeofmicrositemicmierosite seed production soil moisture and canopy cover and burial seed production differed substantially be- depth on germination and survival excio tween sites and years all sites had large an- sures protected seedlings from trampling and nual variationvanation in seed production with annual predation four micrositesmicrosites streamside open coefficients of variation ranging from 100 to canopy terrace open canopy streamside 140 among sites masts were produced in one closed canopy terrace closed canopy were or two of the four years among sites fig 1 selected per site with three replicate excio large crops of seeds irrespective of viability sures per microlitemicrositemicrosite closuresexclosuresenclosuresEx were con- were produced more frequently some excep- structedstructed from 1133 cm hardware cloth and tional trees produced four consecutive large were 060.6og X 060.6og X I11mamin high with 15 cm of seed crops number of large flower crops also mesh below ground seeds were planted varied substantially between sites ranging 36 per exclosure in 6 rows in 3 planting from one at walnut creek to three at hitt treatments surface sown partially buried or wash 1990 RIPARIAN SEED production 49

F 2- 0

161.6lgig W CL TT

V 1 121121.21 Wx 3 J 0.808 T U- 08 UJ 040.4 W LL 0 060.6og

05-05 0M V X 040.4 W CL 030.3 W Wn 020.2 W J M 010.1oiol

0 aj3j 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 A K W H R

fig 1 site means for female flowers and viable nuts produced per shoot for Juglans major from 1983 through 1986 at aztec peak A workman creek K walnut creek W hitt wash H and rock creek R 50 J C STROMBERG AND D T patienpatrenPATTEN volume 50

TABLL 1 average rates of seed production expressed as viable seeds produced per 100 shoots and abundance of juglans major seedlings per 200 m2ma at five sites in arizona

seeds seedlings seeds seedlings seeds seedlings 1983 1984 1984 1985 1985 1986 aztecaztecpeakpeak 11 0 0 0 2 0 workman creek 9 2 3 1 3 0 walnut creek 38 52 0 0 14 23 hitthittwashwash 46 207 10 48 6 24 rock creek 10 3 10 1 2 1

masts were produced at all sites in 1983 they were produced the mast year of 1983 with abundant viable nuts maturing on trees was followed by abundant seedlings in 1984 fig 1 seed production patterns in following whereas the largely inviable seeds produced years differed among sites seed crops were during dry 1984 resulted in low or no recruit- low in different years among sites and crops ment in 1985 rainfall in the year of germi- failed at different reproductive stages the nation may also have increased seedling most common causes of crop failure were re- numbers this is suggested by the greater ductductionsions in numbers of female flowers and abundance of seedlings in wet 1983 than in production of inviable low weight seeds dry 1984 eg 132 vs 52 at walnut creek high rates of abortion were a less frequent despite production of masts in 1982 and 1983 cause at walnut creek for example viable within years seeds germinated during wet nut production was lowest in 1984 a result of seasons table 2 most germinated during few flowers and inviable seeds 3 viable the late summer august september rains hitt wash trees produced a large crop of vi- seeds germinated infrequently in may and able seeds only in 1983 seed production was only during wet springs or in wet streamstreambankbank limited at sequentially earlier reproductive micrositesmicrosites none germinated during the july stages from 1984 to 1986 in 1984 many flow- dry period ers were produced and matured but seed germination increased with microlitemicrositemicrosite soil viabilities were low 28 viable in 1985 moisture although moisture did not exclu- 2 flowers were again abundant but many 96 sively regulate germination rarr2 2424.24 df 14 were aborted in 1986 hitt wash finally had a P 0404.04 values for closuresexclosuresenclosuresex few seeds sharp decline in female flowering resulting in germinated on open terraces particularly at a low crop size rock creek trees showed a the low elevation sites rock creek hitt third pattern following the mast of 1983 wash table 3 where soils were drier table moderate numbers offlowersofflowers and viable seeds 4 canopy cover was positively associated were produced in 1984 this was followed by with germination percentages within these large declines in flower number in 1985 mod- drier micrositesmicrosites and dense herbaceous cover erate numbers of viable seeds were produced was negatively associated with germination in 1986 rates together soil moisture canopy cover and herbaceous cover explained germination 45 of the field variation in germination percentages between natural seedling abundance differed sub- micrositesmicro sites stantialstantiallyly between sites and years these dif- few surface sown nuts germinated in any ferencesferences were related in part to seed produc- microlitemicrositemicrosite table 3 burial increased germination rates seedlings were abundant only at tion rates within all micrositesmicrosites at all sites sites that produced many seeds hitt wash except workman creek where burial in satu- and walnut creek table 1 few seed- rated soil decreased germination partially lings were present at rock creek despite buried seeds germinated in moderate num- moderate seed production because of seed bers in streamstreambanksbanks and under canopy com- predation by arizona gray squirrels sciurus plete burial was necessary for germination in arizonensis see stromberg 1988 annual open terraces lack ofburialofburial limited germina- seedling abundance was influenced by size of tion of naturalofnatural seed populations for example the prior year s crop of viable seeds since many seeds were ungerminated on the soil seeds generally germinated the year after surface at hitt wash terrace excavation of 199019901 RIPARIAN SEED production 51

TABLE 2 juglans major seedling recruitment by month may through october in streamside and terraceten aeeace plots at sites in arizona plots are 200 m2ma

1983IS 83 1984584 198519 85 site plots M J J A S 0 M J J A S 0 M J J A S 0 workman terrace 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 creek streamside 0 0 0 2 0 0 0 0 0 1 0 0 0 0 0 0 0 0 walnut terrace 0 0 0 53 44 0 0 0 0 18 12 0 0 0 0 0 0 0 creek streamside 0 0 0 22 13 0 0 0 0 14 8 0 0 0 0 0 0 0 hitt wash terrace 6 3 0 92 43 2 0 0 0 105 48 1 0 0 0 21 7 0 streamside 14 5 0 35 13 0 0 0 0 47 25 1 2 0 0 13 5 0 rock creek terrace 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 streamside 0 0 0 3 0 0 0 0 0 1 0 0 0 0 0 1 0 0

TABLE 3 germination percentages ofjuglansofjuglans major sown at three planting depths in four mieromicromicrositcsmicrositessitessltes average seed viability was 60 values are means and standard deviations for 3 groups of 12 nuts planting depths aiaree buried at 2 cm partially buried and surface sown sites are aztec peak A workman creek K walnut creek W hitt wash H and rock creek R NA not available

streambankStreambank canopy streambankStream bank open site buried partially buried surface buried partially buried surface A 0 0 0 0 0 0 0 0 0 0 0 0 K 4 661 25 001 8 0 NA NA NA W NA NA NA 29 616 13 661 8 0 H 45 30130 3 551 0 001 46 616 0 0 0 010 R 50 18118 5 551 0 0 50 15115 0 0 0 010

terrace canopy terrace open site buried partially buried surface buried partially buried surface

A 0 0 0 001 0 010 0 0 0 0 0 0 K 21 616 25 010 8 010 50 33133 0 010 0 0 W 35 igl19191 16 lilill11111 3 515 28 igl16116 0 0 0 001 H 53 5 25 14114 8 919 0 0 0 0 0 0 R 21 igl12112 0 lol1010 0 0 4 8 0 0 0 0

TABLE 4 soil water potential at 30 cm mpaM pa for terrace T and streamside S open canopy micimieromicromicrositessitessltes during may july and september of 1983 1984 and 1985 sites aieare aztec peak A workman creek K walnut creek W hitt wash H and rock cleekcreek R

1983 1984 1985 site microlitemicrositeMicrosite may jul sep may jul sep may jul sep mean SDS D A T 0020.020 02 0150 15 ooi0010 01 0680.680 68 0830.830 83 0060.060 06 olioll0110.110 11 0790.790 79 0060.060 06 0300.300 30 033103310.3310 33 S ooi0010.010 01 016olg0.160 16 0000.000ooo00 0540.540 54 0630.630 63 0030.030 03 0060.060 06 0540.540 54 0030.030 03 0220.220 22 025102510.2510 25 K T 0030 03 0190.190olg19 ooi0010.010 01 1031 03 1361 36 0060.060 06 0240 24 1161.16ilg1 16 0060.060 06 0460 46 052105210.5210 52 S ooiool0010 01 0170.170 17 000ooo0 00 0780.780 78 0960.96ogg0 96 0020 02 0120.120 12 0930 93 0040.040 04 0340 34 040104010.4010 40 W T 0030.030 03 0240.240 24 0020.020 02 1181 18 1261.261 26 0050.050 05 0420.420 42 1331.331 33 0040.040 04 0510.510 51 0540 54 S 0020.020 02 0180.180 18 0000.00ooo0 00 0840.840 84 0960.96ogg0 96 0020.020 02 0180 18 1021.021 02 0020.020 02 0360.360 36 04210 42 H T 0030.030 03 0320.32003332 ooi0010.010 01 1651 65 1841.841 84 0150.150 15 0480.480 48 1801.801 80 0120.120 12 0710 71 076107610.7610 76 S 0030.030 03 0220 22 0020 02 1451.451 45 1631 63 0050 05 0260.260 26 1581.581 58 0060.060 06 0590 59 069106910.6910 69 R T 0040.040 04 0430 43 0200.200 20 1861.861 86 2042 04 0130 13 0640.640 64 1831.831 83 0140.140 14 0810.810 81 08010 80 S 0030.030 03 0320.320 32 0020.020 02 1451.451 45 1631 63 0050.050 05 0260.260 26 1581.581 58 0060.060 06 0600ogo60 06810 68 mean 0030.030 03 0260960 26 0040.040 04 1271 27 1441 44 0070 07 0320.320330 32 1391 39 0070 07 SD ooi0010.010 01 0090 09 0060 06 0380.380 38 0390.390 39 0040.040 04 0160.16olg0 16 0350.350 35 0040.040 04 52 J C stromberganddSTROMBERG AND D T patienPAITENPATTEN volume 50

TABLI 5 germination percentages forjuglansfor juglans major buried in soil or sown on the surface at three soil water contents by weight in the greenhouse no seeds germinated at 20 soil moisture seeds are from streamstreambanksbanks S or terraces T from walnut cicreekeek W hitt wash H or rock creek R values are means and standard deviations

saturated 80 moisture 40 moisture seed source buried surface buried surface buried surface W S 21212121 50 717 50 7 0 0 7 7 0000 T 0 010 50 29 42 18118 0 0 21 21 0000 H S 565 t6ta 28 31 63 29 0 to0 24 33 0000 T 0000 18 12 61 16 0 0 19 27 0 010 R S 0 001 22 9 39 11 0 001 13 0 0 001 T 0 001 14 7 48 15 0 001 16 5 0 001

seedlings revealed that only 5 of the seed- example seeds weighing 3 g germinated in 16 lings were from surface nuts whereas 75 days compared with 38 days for those weigh- were from partially buried nuts and 20 were ing 7 g buried at depths of 9 to 15 cm in pocket gopher thomomys sppapp caches seedling survival in the field greenhouse germination seedling mortality had a major role in limiting regeneration of J major only I1 of 374 juglans major germinated abundantly in natural 1983 seedlings among the study sites soil with a moisture content of 80 100 by was alive as of fall 1985 mortality was high in weight table 5 few seeds germinated in the the year of germination and in the two years dry soil or saturated soil planting depth influ- following fig 2 most seedlings died in enced germination at all soil moisture con- june typically a dry month or in winter tents in the saturated soil surface sown precipitation was sparse in early 1984 table seeds germinated in substantially higher per- 6 and seedlings that germinated in 1983 had cencentagestages than buried seeds in contrast seeds high winter mortality only 7 of hitt wash required burial for germination at all other and 9 of walnut creek 1983 natural fall co- moisture contents horts survived until spring 1984 many that germination rates in saturated soil varied did survive remained dormant through the between populations and with seed weight spring and summer drought of 1984 with- seeds collected from trees along the banks of holding leaf out until the late summer rains perennial walnut creek germinated in higher in contrast more than 20 of 1984 cohorts at percentages than seeds from sites with both sites survived to spring of 1985 ephemeral hitt wash or intermittent rock seedling survivorship varied between creek stream flow table 5 many cohorts micromicrositessites table 7 as well as between years from these latter sites did not germinate in A large part of the variation between mi saturated soil germination percentage in sat- crositescrosites was attributable to soil moisture sur- urated soil also tended to increase as seed vivorship after two years increased with soil weight g per cohort decreased y 31831.8 moisture between exclosures rarr2 3131.31 df r2ra2 53x r 15 df 22 P 05os05.05 seed weight 14 P 0505.05 with seedlings on open stream was not related to germination in dry or moist banks having highest survival seedlings that soil did survive in dry micrositesmicro sites had high mor- speed of germination varied with soil mois- tality of buds and shoots during their first and ture level and with seed weight seeds in second winters and grew slowly on terraces moist soil 80 100 germinated rapidly 16 42 of year old survivors originated new 10 days whereas seeds in drier soil 40 spring growth from basal buds 49 from lat- by weight germinated 28 12 days after eral buds and only 9 from the terminal bud planting although variance in germination on streamstreambanksbanks in contrast 15 legrewregrew speed was high within a cohort median ger- from basal buds 58 from lateral and 31 minationmination speed increased significantly with from terminal buds second year seedlings on seed weight for a cohort y 050.50os 5 55x rr2ra the open terrace that regenerated from a basal 3434.34 df 22 P oi0101.01 in 80 moisture for bud had average stem height of 252.5 cm with 1990 RIPARIAN SEED production 53

a HITT WASH WALNUT CREEK 5

4 V Z UJ 3

Z 2 lda UJ V 1

1 0 I i i i i i i i i i i i 6 6 i i i i i i i i i i i i r asondjfmamji M A M J jasondjfmamjS 0 N D J F M A M J JdasoJASOA 0 19851983 1984 1985 fig 2 seedling survivorship forror juglans major that germinated in fall 1983 at hitt wash HW and walnut creek WC values are 109logloge of seedlings remaining each month

TABLE 6 precipitation cm from 1982 to 1986 at a butedtritributestributed to seedling mortality interestingly climatic station near the walnut creek study site aver- seedlings did not have greatest herbivoreherbherbivoryivory at age annual is 39 precipitation is cm sites where adult herbivoreherbherbivoryivory was high see 1982 1983 1984 1985 1986 renaud 1986 rather seedlings in specific micrositesmicrosites had greatest leaf loss from january june 21 22 3 14 16 cerbiherbi july december 19 31 37 31 30 vores seedlings on terraces under canopy total 40 53 40 44 46 had greatest herbivoreherbherbivoryivory damage leaf area consumed by october 1983 was 33 L 15 compared to 9 for all other micrositesmicrosites as few as two leaves I1 cm long these seed- effects of cattle grazing were included in lings did not markedly increase in size from the study out of necessity because of the al- 1983 to 1985 seedlings in moist open areas most ubiquitous presence of cattle in south- grew rapidly as evidenced by a two year west riparian areas two sites hitt wash and streamstreambankbank seedling in full sun that reached walnut creek had heavy to moderate cattle 44 cmcraeraern tall grazing seedlings in exclosures at both sites flooding killed some streamstreambankbank seedlings had substantially higher survival rates than but did not impact terrace seedlings fall those in similar unprotected areas however floods killed 10 of walnut creek and 5 of this was also true for the ungrazed sites table hitt wash 1983 streamstreambankbank seedlings mor- 7 adverse impacts of cattle on seedlings in- tality rates from winterspringwinter spring floods were not cluded trampling and grazing at hitt wash quantified physical impacts of floods in- 22 of 213 natural seedlings had broken or cluded stem breakage coverage with debris eaten stems as did 13 of 130 seedlings at and scouring of seedlings several seedlings walnut creek ability of seedlings to recover resprouted after stem topping from trampling and grazing varied between herbivoreherbivoryHerbivory damage from insects also con sites 40 of all stem damaged seedlings at 54 J C stromberganddSTROMBERG AND D T patienpatrenPATTEN volume 50

TABLETABLL 7 survival percentages one two and four years after germination forjuglansfor juglans major seedlings in exclosures and natural areas in four micromieromicrositesmicrositcssitessltes

Exclosures natural areas mieMicmicrolitemiciositemicrositeMici ositerosite lyrayr1 yr 22yrayryr 44yrayryr isrI1 yr 2yrayr2 yr 44yrayryr Strstieamstidamstreainstrearnearn open 51 28 16 0 0 0 stream canopy 22 15 0 7 0 0 tenterraceaeeace open 6 0 0 5 0 0 terrace canopy 14 0 0 3 0 0

tableTABLF 8 dry weight root length shoot height rootshootroot shoot weight ratio and mortality ofeightof eight weekjuglansweek juglans major seedlings in the greenhouse in soil at three moisture contents asterisk indicates significant difference between hitt wash H and walnut creek W values are means and standard deviations

moisture seed drydi weight root shoot RSR S mortality treatment source g cm cm ratio 40 H 0350.350 35 00510.0510 05 25 212 9 221 14 03030.30 3 0 0 W 0440 44 005100510.0510 05 29 3 11 212 LI1 1 0310 3 0 0 80 11 0830 83 0 14114 34 717 16 212 090.9og0 9 0210210.210 2 0 010 W 0760 76 0 igl12112 34 661 15 212 090 9 020 2 0 0 saturated H 0120.120 12 0 02021 4 2 7 212 0400.44 0110 1 80 22 W 0270 27 00410 04 13 331 10 313 07 02021 13 616 hitt wash regenerated a new stem during the ing abundance of seedlings between years and same year of breakage whereas none did so at sites the extent of fluctuation in annual seed walnut creek production by J major is similar to values for other mast cropping trees Silversilvertown 1980 greenhouse seedling survival town and frequency of mast production is similar to all seedling cohorts had greatest growth in other juglandaceae nixon et al 1980 sork intermediate soil moistures and poorest in sat- 1983 waller 1979 seedlings were abundant urated soil table 8 root growth in particular only after mast years substantiating the view was low in saturated soil and root to shoot that infrequent production ofviable crops lim- ratios were low compared to drier soils some its regeneration sudworth 1934 rainfall cohorts however grew better than others in which varies considerably between years in saturated soil similar to results for germina- the southwest appears to have an important tion rates seedling growth and survival in influence on mast production the evidence saturated soil were related to seed weight and for this although based on a limited number seed source size and weight of seedlings in of years of observation comes from associa- saturated soil increased significantly with de- tions detected between rainfall and the repro- creasing seed weight among cohorts eg ductive stages that are critical to production of seed weight in g 26526.5965 454.5 root length in successful masts flower production associ- 2 cm rr2ra 4949.49 df 11 P oi01010.011 with regard to ated with abundant prior and present year seed source cohorts from the perennially rainfall and seed weight which increases saturated streamstreambanksbanks of walnut creek had with abundant spring rainfall stromberg greater root development and survivorssurvivorshipsurvivorsbiphipbip 1988 seed number between sites is limited in saturated soil than did cohorts from drier variously by low soil moisture and high pre hitt wash dispersal seed predation eg rock creek and insect herbivoreherbherbivoryivory eg workman creek discussion stromberg 1988 low rates of germination also limit recruit- although a survey officeoffiveof five populations is not ment to some degree whereas moisture representative of a species as a whole this for germination and establishment of some study has highlighted factors influencing re- obligate riparian trees is provided by stream cruitmentcruitment ofaofjof J major low and fluctuating flow fenner et al 1985 these processes availability of seeds plays a large role in limit in JJ major and perhaps other facultative 1990 RIPARIAN SEED production 55 riparian trees are influenced in large part by and seedlings as a result of ecotypic differen- rainfall recent documentation of fairly high tiation in morphology or physiology hook regeneration rates for walnut in the south- and stubbs 1967 and differences in tolerance west larkin 1987 medina 1986 may be a of flooding and saturated soil are common result of longtermlong term moisture cycles arizona among trees that grow on sites with different being in an above average cycle at the time of flooding histories mcgee et al 1981 isola- these studies tion of southwest riparian populations within although moist areas provided optimum mountain islands with distinct moisture safe sites for germination and seedling estab- regimes may have led to development of lishmentlishment romejsomej major seedlings established physiological and morphological ecotypesecotypes on terraces and along ephemeral streams little 1950 thornber 1915 seedlings were somewhat drought tolerant whereas small seed size and tolerance of as indicated by high survival in greenhouse saturated soil are associated with wet riparian drought conditions high rootshootroot shoot ratios in sites the large seeds produced by walnuts on drier conditions and ability to survive sum- drier sites stromberg 1988 may increase sur- mer drought via dormancy nevertheless vival of seedlings stressed by factors such as seedling numbers in drier riparian sites were drought or grazing although purely specula- low recruitment may be abundant only after tive the greater ability ofoffittofhitthitt wash seedlings a sequence of several wet years two for pro- to recover from trampling and grazing com- duction of a large viable seed crop another for pared with walnut creek seedlings may have abundant germination and one or two more been a consequence of larger seed size large for high survivorship the low frequency of seeds have large cotyledons that remain at- such a sequence may explain the uniform age tached to seedlings for up to a year after ger- structure of adult walnut populations at some minationmination stromberg 1988 allowing young low elevation sites stromberg 1988 seedlings to regenerate stems wetzstein et an additional factor that may limit estab- al 1983 in any case differences in seedling lishmentlishment of seedlings in dry sites is lack of responses between walnut populations high- burial processes that bury seeds include light the need for study of many populations to deposition of flood debris rare on terraces thoroughly understand reproductive dynam- caching by pocket gophers rare except in ics of any riparian species sandy soils trampling by large animals and possibly caching by squirrels although tree acknowledgments squirrels commonly cache walnuts stapanian and smith 1978 there is conflicting evidence this study was supported in part by a coop- about the frequency at which they cache nuts erative agreement with the US forest ser- of J major in parts of their range where vice RM 80 133 CA winters are mild squirrels immediately con- sume gathered nuts brown 1984 be- this CITED havior may contribute to the decline in abun- literature dance of walnut at low elevations babbettBARRETTBARBETT L I1 1931 influence offoresthorestmorestafforestofforest litter on the germi- this study suggests that germination and nation and early survival of chestnut oak ouercusquercus establishment requirements oft major differ inonmontanatana willd ecology 12 476 484 between populations as well as between BROWN D E ED 1982 biotic communities oftheodtheofthe ameri- specifically can southwest united states and mexico des- micrositesmicrosites populations from pe- ert plants 4 11342342 rennial stream sites appear to be more toler- 1984 arizona s tree squirrels arizona game and ant of saturated soil at the seed and seedling fish department phoenix 114114ppappp stages this should be verified on a larger sam- FENNER P W W BRADY AND D R PATTON 1985 ple of populations effects ofregulated water flows on regeneration of the greater germination fremont cottonwood journal of range manage- and seedling survival in saturated soil for ment 38 135 138 seeds from such sites may be a consequence of HOOK D D AND R M CRAWFORD EDS 1978 plant life lower oxygen demands of their smaller seeds in anaerobic environments ann arbor science stromberg 1988 or of physiological adapta- publishers ann arbor michigan 564 appp HOOK D D AND J STUBBS 1967 physiographic seed tions hook and crawford 1978 of tolerance source variation in tupelo gums grown in various moisture level is known to vary among seeds water regimes pages 61 67 in proceedings of 56 J C STROMBERG AND D T PATTEN volume 50

the ninth southern conference on forest tree silvertownSILVER TOWN J W 1980 the evolutionary ecology ofofmasthofmastmast improvement knoxville tennessee committee seeding in trees linnean society biological jour- on southern forest tree improvement macon nal 14 235250235 250 georgia SORK V L 1983 mast fruiting in hickories and availa- LARKIN G J 1987 factors influencing distribution and bility of nuts american midland naturalist 109 legeneregenerationrahon of riparian species along mountain 81 88 in streams in central arizonaanzona unpublished thesis STAPANIAN M A ANDANDCC C SMITH 1978 A model for seed arizona state university tempe 84 appp scatterscatterhoardingscatterhoardmghoarding coevolution of fox squirrels and litileLITTLE E L JR 1950 Southwestsouthwestersouthwesternei n trees a guide to the black walnut ecology 5988459 884 896 new native species of mexico and arizona agri- STROMBERG J C 1988 reproduction and establishment culturaleuitcultuialuralurai handbook 9 united states department of arizona walnut juglans unpublished ofagricultureofagricultlire major washington DCD C dissertation arizona state university tempe mcgl I1 A B M R schmierbach AND F A BAZZAZ 183 appp 1981 photosynthesis and growth in in populations SUDWORTH G B 1934 poplars principal tree willows of populus deltoidesdeltoides from contrasting habitats and walnuts oftheodtheofthe rocky mountains region tech- american midland naturalist 105 305 311 nical bulletin 420 united states department of MEDINAMLDINA A L 1986 riparian plant the communities of Agnagricultureculture washington DCD C foifolfort t bayard watershed in southwest new mexico in SZARO R C 1989 riparian forest and scrubland commu- southwestern naturalist 31 345 359 nity types of arizona and new mexico desert NIXON C M M W MCCLAIN ANDANDLL P HANSEN 1980 plants 9 1 138 six years ofhickoryofbickory seed yields in southeastern ohio THORNBER J J 1915 walnut culture in journal ofwildlifeofwildlife management 44 534 539 in arizona university of arizona agricultural experiment station renaudRLNAUURLNAUD D 1986 the impact ofherbivoreofherbivoiyherbivory on the repro- herbivory bulletin 76 469 503 ductive and defense allocation oftheodtheofthe arizonaanzona wal- WALLER D M 1979 nut unpublished thesis arizona state models of mast fruiting in trees univer- journal of oftbeoretical sity tempetempo 54 appp theoretical biology 80 223 232 WETZSTEIN 11 Y D SPARKS AND G A LANG 1983 RUCKS M G 1984 composition and trend of riparian cotyledon detachment and growth of pecan vegetation on five perennial streams in southeast- seed- lings hortscienceHortScience 18 331 ern arizonaanzona pages 97 107 in R E warner and 333 K M hendrix eds california ripariani ilananipanan systems ecology conservation and productive manage- received 30 june 1989 ment university ofcaliforniaofcalifornia press berkeley accepted 1 january 1990 great basin naturalist soi501 1990 appp 57 62

FORAGE QUALITY OF RILLSCALE ATRIPLEX SUCKLEYI GROWN ON AMENDED BENTONITE MINE SPOIL

marguerite E voorhees

ABSTRACT at peak standing crop nllscalerillscalerillscaleseale atriplex suckleyisuckleyi foliage grown on amended bentonite mine spoil contained adequate digestible energy crude protein and all mineral elements except phosphorus necessary for cattle sheep antelope and deer amendments sawdust NPK gypsum geneigenerallygener allyaily did not affect forage quality iron manganese aluminum sodium and potassium concentrations were high and may have adversely affected forage quality forage utility would be limited to a few months during the growing season

atriplex suckleyisuckleyi torrey redbrydb com- lecterlected from plots on raw bentonite spoil and monly called rillrillscalescale is the dominant native on bentonite spoil that had been amended invader on bentonite mine spoil sieg et al with various combinations of gypsum fertil- 1983 rillscaleRillscale is a spreading annual plant izer NPK and sawdust during the year prior usually less than 30 cm in height that flowers to harvest and 2 to assess the effects of treat- from early june to mid august bearing ma- ments on growth of rillscalerillscale during the year ture seed before the end of july the plant is following treatment found only in southern saskatchewan southern alberta montana wyoming north da- METHODS kota south dakota and nebraska it has been observed that the plant grows in saline study area and treatments clayey and alkaline land where nothing else the study area is located just west of the seems to grow frankton and bassett 1970 central black hills near upton wyoming on little in known about the biology of this the mowry shale formation sagebrush arte- species misia dentatatritridentatatridentate is the predominant vegeta- forage quality is an important consider- tion on this grassland with scattered stands ation in the selection of species for use in of ponderosa pine pinus ponderosa annual revegetation of bentonite mine spoil since precipitation averages 350 mm national grazing is the major postminingpostmining land use in oceanic and atmospheric administration regions where bentonite is mined wildlife 1981 falling mostly during the growing sea- forage and habitat are also emphasized in son from may to september soils are gener- reclamation efforts twenty two species of ally shallow and poorly developed wildlife are known to use atriplex species for an area was selected on unreclaimed ben- food and cover robinette 1971 martin et al tonite mine spoil that was mined before 1968 1951 atriplex species are valued by range on the property of american colloid the ex- managers because of their high protein con- periperimentalmental design was that of a 223 factorial tent bidwell and wooton 1925 arrangement of treatments with each of three the peak forage value of rillscalerillscale as with spoil amendments at two levels voorhees most annual forbs is in all likelihood limited etct al 1987 one level was the absence of to spring and early summer months cook each amendment while the other level was 1972 stoddart et al 1975 when available it the presence of the amendment may make an important contribution to the the study site was rototilledrototilled to a depth of nutrition of livestock and wildlife the objec- approximately 5 cm and gypsum was applied tive of this study was twofold 1 to examine at a level of 31 metric tons per hectare intro- chemical properties of rillrillscalescale foliage col duction of ca in the form of cas04caso was

SDAUSDA forest service rockyrociy mountain foiestforest and range experiment station rapid city south dakota 57701

57 58 MMEE VOORHEES volume 50

intended to facilitate exchange with monova- estimated peak standing crop 17 august lent sodium which would encourage floccula- 1983 to determine the rate of decline in tion and water penetration brady 1974 and standing crop resulting from drying and shat- discourage surface crust formation fertilizer tering of foliage as the season progressed it was added at the rate of 114 kg nitrogen 23 kg was assumed that harvesting one half of each phosphorus and 50 kg potassium per hectare plot had an insignificant effect on plants on the nitrogen and phosphorus were added as remaining half all harvested biomass was ammonium nitrate NHNO and diammo- oven dried at 55 C weighed and ground nium phosphate nh4hponh4ahp04 potassium through a 20 mesh screen was added as potassium chloride KCI sawdust was added at the ratio ofone part sawdust plant tissue analyses to two parts spoil by volume inorganic nini- plant tissue analyses included total nitro- trogen nh4nonh4n03 corresponding to 060.60og 6 of gen by conventional micro kjeldahl in vitro sawdust by weight was added to the sawdust dry matter digestibility and percentage ash before mixing with spoil to prevent a large church and pond 1978 duplicate samples increase in the carbon to nitrogen ratio and of plant tissue were analyzed to determine subsequent tie up of soil nitrogen by micro- nitrogen ash and dry matter digestibility organisms allison 1965 this amount of dry matter digestibility was determined with nitrogen corresponded to 6 kg nitrogen per acid pepsin using two 48 hour digestions in a metric ton of sawdust the sawdust amend- rumen buffer solution taken from cattle eat- ment was intended to increase sti ucturalstructural sta- ing grass hay tilley and terry 1963 crude bility and tilth of spoil as well as air and water protein percentage was estimated from kjel- permeability voorhees et al 1983 1987 dahl nitrogen CP N X 6256.25623 digestible the effects of organic matter additions in the energy DE was estimated from dry matter form of sawdust might be expected to increase digestibility values rittenhouse et al 1971 the stability of the substrate where organic and converted to metabolizable energy ME less matter is than 2 marshall and holmes mcalkg dry matter using the following for- 1979 as in bentonite mine spoil uresk and mula swift 1957 yamamoto 1986 gypsum and sawdust amendments were demcalkg X 0790.79 memcalkg manually incorporated into tilled spoil elemental concentrations of nitric acid whereas the fertilizer amendment was ap- extractable aluminum arsenic barium bor- plied to the surface all eight combinations on cadmium calcium chromium copper of the threethiee amendments including control iron magnesium manganese molybdenum were replicated twice to give a total of 16 nickel phosphorus potassium selenium plots each 60 X 150 cm the plots were tilled sodium strontium titanium and zinc were amended and seeded on 8 may 1982 plots determined for the plant tissue samples were were self seeded in 1983 as no seed was analyzed in duplicate checks standards and planted that year blanks were included elemental con- rillscaleRillscale seed obtained from sites along the centrationscentrations of nitric acid extracts were mea- montana wyoming border during late sum- sured using inductively coupled plasma mer of 1980 was planted inm each plot so that atomic emission spectrometry ICPAESICP AES seed weights corresponded to approximately fassel and knisely 1974 jones 1977 on the 2 three live seeds per CMcm2 this weight of seed nitric acid digestion havlin and soltanpourSoltan pour was calculated from total peipercentageper centage germina- 1980 gestring and soltanpourSoltanpour 1981 tion and seed density determinations made within six weeks of planting seed was broad- statistical analysis cast on the surface and raked 1 I1 cm into spoil A three way factorial analysis of variance one half of each plot was harvested for was used to determine the effects of spoil chemical analysis by manually cutting off amendments gypsum sawdust and fertil- stems at ground level during estimated peak izer and associated interactions on each fo- standing crop 7 july 1983 the other half liage property significant differences were was harvested approximately six weeks after accepted at the 0505.05 probability level 19901 RILLSCALE FORAGE QUALITY 59

TABLE 1 chemical composition of the foliage ofrillsealeofnilscalerillsealescale grown on bentonite mine pollspoil averaged across treatment that did or did not include amendment with sawdust NPK fertilizerfertihertilizel or gypsum

sawdustsawelust n1NPKna K gygypsumhsumpsum property units without with without with without with standing crop aghakgha 9551955 1973 1297 1631 1534 1394 dry matter digestibility 73 72 73 72 73 73 digestible energy kcalkgdm2 2968 2923 297099702.970 2920 2954 2955 metabolizablemetabohzable energy mcalkgdmmealkgdm 235 231 235 231 232 233 kjeldahl nitrogen 1 70 187 1 74 183 1741.741 74 183 crude protein ll11 12 11 12 11 12 ash 42 35 40 37 38 39 ca 047 043 045 045 043 047 mg M 099 092 098 094 096 095 P 0190.190olg19 0 17 0180 18 0180 18 018 0180.180 18 cap 255 257 265 247147 247 264 na 855 8188.188 18 830 843 863 8118 11 K 1131 13 110llolio1.101 10 illlii1 11 1121.121 12 1121 12 liiiiiili1111.111 11 znan aggrggpgg 66 63 61 68 55 74 fe biggjiggmgg 10775 11128 9924 8188 3999 10775 mn elgglggpgg 496 297 450 343 332 461 nxggNnaggagglgg 8 6 8 7 7 8 crjiggcrcraiggaggrgg 5 5 5 5 5 5 cu aggrggugg 7 6 6 6 6 7 MO aggrggagg 16 17 21 13 17 16 cumo 05 05 05 04 04 05 Bbagbwgaggrgg 35 41 37 39 39 37 al aggrgggg 1296 1006 1176 1126 938 1365 bapggBapba aggrgggg 37 32 35 34 28 41 srSrpsrpggsraggaggpgggg 69 73 72 69 68 73 titilg11gg 10 8 9 9 8 10 cleansmeans foifor each property that are followed by an asterisk are significantly different p 05 2basedabasedbased on inin vitro dry matter digestibility

RESULTS centrationcentration of nitrogen in foliage from 1701.70 to 1871.87 and increased the crude protein rating peak standing crop averaged kg dry 1464 from 11I1 I1 to 12 relative to foliage on spoil not matter per hectare 1 and ranged from table amended with sawdust table 1 267 to kg dry peak 2913 matter per hectare calcium and magnesium levels in rillrillsealerillscalesealescale standing crop was 107 greater and ash aver- foliage averaged 0450.450 45 and 0960.960ogg96 respectively aged 17 lower on plots that had been table 1 the level of phosphorus was 00180.1818 amended with sawdust alone or in combina- and the ratio of calcium to phosphorus was tion with other amendments than on plots 2612.61961 levels of sodium and potassium in that had not been amended with sawdust rillscalerillscale foliage were 8378.37 and 11121.1212 respec- standing crop decreased without grazing by tively when sawdust and fertilizer amend- 21 from early july to mid august ments were used in combination with or digestible energy ofrillsealeofrillscalerillsealescale foliage at peak without the gypsum amendment the foliar of standing crop based on IVDMD was 2948 content of magnesium and potassium de- kcalkg dry matter table 1 digestibility of creased relative to concentrations in foliage dry matter and estimates of digestible and on spoils amended with either of these two metabolizable energy levels were all signifi- amendments alone cantly decreased when sawdust and fertilizer zinc levels in rillsealerillscalerillsealescale foliage averaged were used in combination with or without the about 65 uggu gg table 1 iron and manganese gypsum amendment relative to the use of levels were 9131 and 397 uggu gg respectively other combinations of amendments nickel levels averaged 7 uggu gg whereas crude protein of rillscalerillscale foliage ranged chromium levels were 5 uggu gg from 9 to 14 amendment of spoil with saw- the concentration of copper in foliage was dust alone or in combination with other 6 vtggvaggp gg while molybdenum concentration was amendments significantly increased the con 17 uggnggu gg table 1 the sawdust amendment 60 MmevoorheesMEE VOORHEES volume 50

alone or in combination with other amend- considered inadequate as forage after the ments decreased foliar copper from 7 to 6 fruiting stage cook 1972 stoddart et al 1975 puggolggjlggvlgg but did not significantly alter the ratio of the quantities of forage available from copper to molybdenum growth of rillrillsealerillscalesealescale on spoil would be inade- high levels of aluminum and iron in quate for most grazing uses except during a rillscalerillscale foliage caused severe spectral inter- few months of the year standing crop de- ferencesferences for arsenic and selenium thus it was clined by 21 without grazing from early july not possible to determine the concentrations to mid august this decline following matu- of these elements rity was attributed to drying and shattering of foliar aluminum levels ranged from 1000 foliage no evidence of grazing by insects was to 1300 agglggjlgg the gypsum amendment alone observed however rillrillscalescale could make an or in combination with other amendments important contribution to the nutrition of live- significantly increased foliar aluminum levels stock and wildlife during the short period of from an average of 938 to 1365 elggvlggpugggg table 1 time it is growing and available other plant relative to foliage on spoil that had not been species could be planted with rillrillscalescale uresk amended with gypsum amendment of spoil and yamamoto 1986 welch 1989 to help with sawdust with or without other amend- meet the nutritional requirements of cerbiherbi ments resulted in a decrease in the concen- vores tration of aluminum in foliage compared with the sawdust amendment increased stand- foliage from spoil not amended with sawdust ing crop and decreased ash by improving cadmium levels in rillsealerillscalerillsealescale foliage were plant water relations and increasing the below detection limits io10101.0 aggpgggg for the availability of nitrogen increased availability ICPAESICP AES procedure boron concentrations in of water reduces plant requirements for salts foliage averaged 38 olggpuggjlggvlgg and were signifi- conversely plants under water stress accu- cantly greater when sawdust was added to mulate other nutrients when nitrogen is limit- spoil with or without other amendments ing mengel and kirkby 1982 decreases in than they were in foliage grown on spoil not plant ash as a result of the sawdust amend- amended with sawdust table 1 ment would account for significant decreases barium levels in foliage averaged 35 aggrggxgg in plant uptake of copper and aluminum while strontium concentrations averaged 71 the foliage of rillscalerillscale at peak standing crop aggrggxgg when sawdust and fertilizer amend- contained adequate digestible energy based ments were used in combination with or on IVDMD crude protein percentage and without the gypsum amendment strontium concentrations of all mineral elements except levels were greater than when either of these phosphorus for cattle sheep and wild rumi- amendments was used alone nants national research council 1975 1984 titanium concentrations in foliage aver- dean 1980 stone et al 1983 the sawdust aged 9 lggelggpgg and increased by 25 when gyp- amendment either alone or in combination sum was added with or without other amend- with other amendments resulted in an in- ments relative to foliage from spoil not crease in the concentrations of nitrogen and treated with gypsum table 1 crude protein while addition of both sawdust the fertilizer amendment with or without and fertilizer with or without the gypsum other amendments had little effect on foliar amendment decreased dry matter digestibil- composition table 1 except through interac- ity and estimated digestible and metaboliz- tion with the sawdust amendment able energy phosphorus supplementation would be discussion advisable for animals foraging on bentonite mined lands revegetated with rillrillscalescale cal- the forage utility of rillscalerillscale as an annual cium levels exceeded most dietary require- forb is probably limited to a few months dur- ments oflivestock national research council ing the growing season culminating with 1975 1984 welch 1989 but were marginal peak of growth in late june and rapidly declin- for deer dean 1980 adequate fresh water at ing thereafter late in the growing season low salinity levels would also be necessary most species of forbs fail to meet the protein since rillrillsealerillscalesealescale contains high quantities of sodium and energy needs of gestating animals and are and potassium toxicities of electrolytes are 1990 RILLSCALE FORAGE QUALITY 61

considered unlikely unless water intake is re- ing native vegetation of the surrounding area strictstricteded or water is highly saline church and should be encouraged pond 1978 iron manganese and aluminum were also present in very high concentrations acknowledgments in the foliage of rillrillscalescale which may depress cellulose digestion national research coun- this study was conducted in cooperation cil 1984 martinez and church 1970 grace with the department of range science colo- 1973 amendment of spoil with gypsum in- rado state university fort collins thanks creased whereas addition of sawdust de- are extended to dan uresk and joe trlica for creased the concentration of foliar alu- their assistance and encouragement through- minum an important consideration because out the study aluminum can cause gastrointestinal irritation or produce rickets by interfering with phos- literature CITED phate absorption ifpresent in large quantities in the diets of some animals underwood ALLISON F E 1965 decomposition of wood and bark sawdusty 1977 finally the copper to molybdenum sawsawdustsdusts in soil nitrogen requirements and effects on plants USDA agricultural research ratio of the foliage of rillrillscalerillsealescaleseale was low at 070.7 service technical bulletin 1332 57 appp and could cause molybdenum induced cop- BIDWELL G L ANDANDEE 0 WOOTON 1925 saltbushesSaltbushes and per deficiencies in livestock and wildlife that their allies in the united states USDA bulletin do not have access to copper supplements or 1245 40 appp BRADY N C forages high in copper 1974 the nature and properties of soils ath8th concentration milt- ed macmillan publ co inc new york 639 appp more and mason 1971 stone et al 1983 CHURCH D C AND W G POND 1978 basic animal other micro and macro minerals were ade- nutrition and feeding 0 & B books corvallis quate to meet the requirements of most oregon 300 appp herbherbivoresivores COOK C W 1972 comparative nutritive values of forbs grasses and shrubs appp 303 310 in C M the fertilizer treatment with or without mckell J P blaisdell and J R goodmgoodin eds other amendments had little effect on foliage wildlandwindlandWildland shrubs their biology and utilization composition except through an interaction proceedings of a symposium july 1971 logan with the sawdust amendment fertilizer utah intermountain forest and range experi- the ment amendment may have been ineffective for station ogden utah inin- DEAN R E 1980 the nutrition of wild ruminants appp creasing the availability ofnitrogen phospho- 278 305 in D C church ed digestive physiol- rus and potassium in spoils or other condi- ogy and nutrition of ruminants ad2d ed 0 & B tions may have inhibited uptake of these ions books inc Corvalcorvallishs oregon added nutrients were probably not leachedbeached passelFASSELFASSFL V A AND R N KNISELY 1974 ICP optical below emission spectroscopy analytical chemistry 46 rooting depth since permeability of 1110aI1 I1 loaioa unamended spoil is extremely low loss of FRANKTONFBANKTON C AND I1 J BASSETT 1970 the genus atriplex fertilizer as runoffrunoffmaymay have been a factor and chenopodiaceae in canada II11 four native thus would explain the interaction between western annuals A argentea A truntruncatatruncatecata A nowellipowelhpowellipowelli and A dioicadionica canadian journal of botany fertilizer and sawdust amendments since 4898148 981 989 sawdust amendment has been shown to in- GESTRING W D AND P N soltanpourSOLTAN POUR 1981 boron crease infiltration and decrease runoff voor- analysis in soil extracts and plant tissue by plasma hees 1986 alternately these elements might emission spectroscopy communities in soil not have been limiting to plant growth science and plant analysis 12 733 742 the GRACE N D 1973 high mn latter hypothesis unlikely effect of dietary M n levels on the seems because the growth rate and the level of mineral elements in sawdust amendment was effective for increas- the plasma and soft tissues of sheep new zealand ing the level of foliar nitrogen journal ofagricultural research 16 177177184184 rillscaleRillscale would be a good choice to consider HAVLIN J L AND P N soltanpourSOLTAN POUR 1980 A nitric acid plant tissue digest method for use with inductively in revegetating bentonite mine sspoilsspolispoilsolis because coupled plasma spectrometry communities in it provides substantial quantities offorage and soil science and plant analysis 11 969 980 nutritional qualities generally adequate to JONES J B JR 1977 elemental analysis of soil extracts meet requirements of livestock and wildlife and plant tissue ash by plasma emission spec- troscopytros copy soil plant the few nutritional inadequacies boxi communities in science and for and toxi analysis 8 349 sealescale 365 cities of rillrillsealerillscale introduction of other plants MARSHALL T J AND J W HOLMES 1979 soil physics on bentonite spoils may be feasible also graz cambridge university press new york 345 appp 62 M E VOORHEES volume 50

MARTIN A C H S ZIM AND A L ARNOIDARNOLD 1951 ameri- STONE L R J A ERDMAN G L FEDERFEDEK AND H D can wildlife and plants a guide to wildlife food HOLLAND 1983 molybdenosis in an area under- habits mcgraw hill book co new york 500 appp lain by uranium bearing lignites in the northern MARTINEZ A ANDANDDD C CHURCH 1970 effect ofvariousof various great plains journal of range management 36 mineral elements on in vitro rumen cellulose di- 280 285 gestion journal ofanimalofanimal science 31 982 990 SWIFT R W 1957 the nutritive evaluation of forages MAYNARD L A ANDJAND J K LOOSLI 1969 animal nutrition pacific agricultural experiment station bulletin mcgraw hill new york 613 appp 615 34 appp mencelMENGEL K AND E A KIRKBY 1982 principles of plant TILLEY J M AND R A TERRY 1963 A two stage tech- nutrition ard3rd ed international potash institute nique for the in vitro digestion of forage crops worblaufen bern switzerland 655 appp journal of british grassland society 18 104 111 MILTMORE J E andaandj L MASON 1971 copper to molyb- UNDERWOOD E J 1977 trace elements in humanburnanhunnan and denum ratio and molybdenum and copper concen- animal nutrition fth4th4tb ed academic press new trationstrations in ruminant feeds canadian journal of york 545 appp animal science 51 193 200 URESK D W ANDANDTT YAMAMOTO 1986 growth of forbs NATIONAL RESEARCH COUNCIL 1975 nutrient require- shrubs and trees on bentomtebenberitoniteberiberlbenl tonitetomte mine spoil under ments of domestic animals no 5 sheep ath5th ed greenhouse conditions journal ofrangeoforrangerange manage- national academy of science washington DCD C ment 39 113 117 72 appp VOORHEES M E 1986 infiltration rate bentoniteofbentoniteofbentomte mine 1984 nutrient requirements ofdomesticofdomestic animals spoil as affected by amendments of gypsum saw- no 4 beef cattle ath6th ed national academy of dust and inorganic fertilizer reclamation and science washington DC 90 appp revegetation research 5 483 490 NAIIONALNATIONAL OCEANIC AND atmospheric administration VOORHEES M E D W URESK ANDANDRR M HANSEN 1983 1981 wyoming climatological data annual sum- atriplex suckleyisuckleyi torrey redbrydb a native annual mary environmental data and information cen- plant for revegetating bentonite mine spoils in ter national climatic center asheville north arthur R tiedemann E durant mcarthur carolina howard C stutz richard stevens and kendall rittenhouse L R C L STRETERSTREETER ANDDAND D C clanton L johnson compilers proceedings symposium 1971 estimating digestible energy from di- on the biology Atriplexofatriplexof and related chenochenopodspods gesgestibletible dry and organic matter in diets of grazing 2 6 may 1983 provo utah USDA forestfoiest ser- cattle journal ofrangeoforrangerange management 24 73 75 vice general technical report INT 172 USU S de- ROBINETTE W L 1971 browse and cover for wildlife partmentpart ment of agriculture forest service inter- appp 69 70 inm C M mckell J P blaisdell and mountain forest and range experiment station J R goodin eds wildlandWildwindlandland shrubs their biol- ogden utah 309 appp ogy and utilization proceedings of a symposium VOORHEESVOORHFES M E M J TRLICA AND D W URESK 1987 july 1971 logan utah intermountain forest glowthgrowth of nllscdlerillrillsealesealescale on bentonite mine spoil as and range experiment station ogden utah influenced by amendments journal of environ- SIEG C H D W URESK ANDANDRR M HANSEN 1983 plant mental quality 16 411 416 soil relationships on bentoniteben tomte mine spoils and WELCH B 1989 nutritive value of shrubs in C M sagebrush grassland in the northern high plains mckell ed the biology and utilization of shrubsofshrubs journal of range management 37 289 294 academic press inc new york STODDART L A A D SMITH ANDANDTT W box 1975 range management ard3rd ed mcgraw hill book co received 15 may 1989 new york 532 appp accepted 15 december 1989 great basinBBMII naturalist 501 1990 appp 63 65

SUMMER FOOD HABITS OF COYOTES IN IDAHOS RIVER OF NO RETURN wilderness AREA

charles L elliott and richard guetig

ABSTRACT summer food habits ofcoyotesofcoyotes caniscams latranslatranolatrans in the river of no returnn wilderness area idaho were determineddetel mined analysis of 51 seats fecal samples revealedleveievealed that columbian ground squirrels spermophilus colum bianasbianusbtanufibianus mule deer odocoileus hemionus and deer mleemice peromyscus maniculatusmamculatus exhibited the greatest frequency ofoccuirenceofoccurrence for identified food items being detected in 57 27 and 16 respectively of seats examined

one of the most ubiquitous and adaptable and diameter at the widest point was deter- predators of the american west is the coyote mined using criteria established in other caniscams latranslatranolatrans As man altered habitats in the western studies weaver and fritts 1979 western states the coyote adapted its behav- green and flinders 1981 danner and dodd ior and diet to take advantage of these new 1982 we classified all seats s 20 mm in di- environments being generally dietary oppor ameter as coyote seats were washed sepa- tunists johnson and hansen 1977 coyotes rated and prepared for analysis in a manner have found prey to their liking on man s range- similar to that described by johnson and land murie 1951 short 1979 green and hansen 1979 prepared seats were analyzed flinders 1981 and farms gipson 1974 and following the procedure of green and in his cities maccracken 1982 although flinders 1981 hair was identified by many aspects ofcoyote ecology in man altered medullary characters moore et al 1974 or man impacted areas of the west have been teeth were also used to verify the animal investigated less is known of the role of the species consumed each coyote scat was coyote in relatively undisturbed wilderness treated as an individual observation no at- the objective of this study was to determine tempt was made to determinedetel mine the density of the summer food habits of coyotes in idaho s potential prey items inm the big creek area river of no return wilderness area RNRWA hence it was not possible to determine pref- erence indices for the items identified in the STUDY AREA AND METHODS seats examined study was conducted in the big the creek RESULTS AND ranger district RNRWA formerly the idaho discussion primitive area A description of the RNRWA fifty one seats collected met the s 20 mm and big creek area has been provided by diameter criterion and were classified as coy- hornocker 1970 ote the average dry weight t SD of indi- canid seats were collected from trails vidual coyote seats was 15315.315 3 L 595.95sg 9 g soluble located in the big creek drainage of the endogenous material accounted for an aver- RNRWA trails were surveyed the beginning age 393.93 9 t 232.32 3 g 25 of dry weightscatweight scat of may 1977 and 1978 and all seats encoun- thirteen mammal species were identified as tered were removed after the initial clear- food items consumed by coyotes during the ing trails were surveyed at least once a month summer in the RNRWA table 1 percent for newly deposited seats scat collection con- occurrence of identified food categories was cluded at the end of august 1977 and 1978 as follows rodents 100 cervidae 41441.441 4 collected seats were air dried and weighed insects 39239.239 2 birds 27427.427 4 reptiles 393.93 9

department ofbotanyof botany and range sciences brighambi ighambigham young university provo utah 84602 present addressaddi ess department of biological sciences eastern kentucky university richmond kentucky 40475095040475 0950 department of biological sciences eastern kentucky university richmond kentucky 40475095040475 0950

63 64 C L ELLIOTELLIOTT AND R GUETIG volume 50

TABLE 1 percent occurrence of material identified in 51 coyote seats river of no return wilderness area idaho may august 1977 and 1978

monthManthonth species identified may 991 june 14 july 15 august 13 total 51 columbian ground squirrel 429 733 923 568 spermophilus columbianuscolumbianus mule deer 33333.333 3 35735 7 133 307 274 odocoileusOdo codeus hemionus deer mouse 22222.222 2 14214.214 2 200 76 156 peromyscus maniculatusmamculatus moose alces alces 33333 3 21421.421 4 76 137 northern pocket gopher 11111.1illili11 1 14214.214 2 200 11711.711 7 thomomys talptalpoidesoides montane vole 22222 2 21421.421 4 66 117 microtus montanus golden mantled ground squirrel 222 66 58 spermophilus lateralislaterabsrahsfahs audubon s cottontail 11111.1ililii11 1 76 39 Sylvisylmlagussylvilaguslagus audubonauduboniiaudubomtii horse equus caballus 11111.1liililiiiill11 1 71 39 longlongtailtail weasel 717.17 1 19 mustelafrenatamustela frfredatafrenataenata noinorthernthem water shrew 71 19 sorex palustnspalustrispalustris water vole 71 19 arvicola nchardsomrichardrichardsonirichardsonbsoni snowshoe hare 71 19 lepus americanosamericanusamericanus unknown mammals 71 133 58 reptiles 222 39 arthropods iiilii11111 1 21421.421 4 533 61561.561gis 5 392 birds iliill11111 1 71 333 538 274 plant matter liilil11111 1 71 400 230 215 number of scatsseatsscabs examined

domestic livestock 393.9 and other carnivorescarnivores early september elliott and flinders 1980 191.91ig 9 the results of this study correspond the seasonal importance of columbian favorably with those of ribic 1978 johnson ground squirrels as a prey species for other and hansen 1979 and short 1979 in that predators iei e mountain lions felis con- rodents were the most frequently identified color inm the RNRWA was noted by seiden food category in the summer diet of western sticker et al 1973 increased mountain lion coyotes activity during the day in summer was felt to the columbian ground squirrel sper- be related to the availability of columbian momophilusphilus columbianuscolumbianus was the most fre- ground squirrels as a food item seidenstickerSeiden sticker quently occurring food item identified in sum- et al 1973 the presence of lesser species in mer coyote seats being found in 29 56856856.8 of the summer diet of lions was thought to hold the 51 seats examined the percent occur- down any increases over the lion s winter kill rence of columbian ground squirrel remains rate of elk cermscervus elaphusclaphus and mule deer in summer seats reflects the seasonal availabil- odocoileus hemionus hornocker 1970 ity of the squirrel as a prey item sper- although coyotes and mountain lions uti- momophilusphilus columbianuscolumbianus within the RNRWA lize a common food resource it is doubtful emerge from hibernation in late may and re- that they are serious summer dietary competi- main active above ground until late august tors elk and mule deer are the major food 1990 RILLSCALE FORAGE QUALITY 65

items consumed by mountain lions during the HORNOCKER M G 1970 an analysis of mountain lion summer in the RNRWA hornocker 1970 predation upon mule deer in the idaho primitive area whereas rodents comprise the bulk of summer wildlife monograph 21 JOHNSON M K ANDANDRR M hanseenHANSEN 1977 foods of coy items consumed by 1 coyotes see table in otes in the lower grand canyon arizona ari- the hierarchy ofpredatorsofpredators in the RNRWA the zona academy ofscience 12 81 83 coyote appears to occupy a trophietrophicatrophic level below 1979 coyote food habits on the idaho national that of the mountain lion engineering laboratory journal ofwildlife management 43 951 956 maccracken J G 1982 coyote foods in a southern acknowledgments california suburb wildlife society bulletin 10 the senior author thanks dr berranjerran flin- 280 281 ders brigham young university for MOOREMOORF T D L E SPENCE ANDCAND C E DUCNOLLEDUGNOLLE 1974 use of identification of the dorsaldorsai guard hairs of some laboratory facilities in the preparation of mammals of wyoming wyoming game and fish scat samples and the university of idaho s department bulletin 14 177 appp wilderness research committee for permis- murleMURIEMUBIE A 1951 coyote food habits on a southwestern sion to use the facilities at the taylor ranch cattle range jouljournalnalnai of mammalogy 32 291 295 field station river of no return wilderness RIBIC C A 1978 summer foods of coyotes at rocky flats colorado southwestern naturalist 23 area 152 153 sfidenstickiseidenstickerSEIDEN STICKERR J C IV M G HORNOCKERHOBNOCKER W V WILES literature CITED ANDJAND P MESSICK 1973 mountain lion social orga- nizanizationtion in the idaho primitive area wildlife DANNERDANNEB D AANDNA AND N DODD 1982 comparison of coy- monograph 35 ote and gray fox scat diameters journal ofwildlife SHORT H L 1979 food habits ofcoyotes in a semidesert management 46 240 241 46240 grass shrub habitat USU S forest service rocky ELLIOTTELLIOTTCC LlandalandjAND J T FLINDERS 1980 seasonal activity mountain forest and range experiment of station patterns columbian ground squirrels in the research note RM 364 4 appp idaho primitive area great basin naturalist 40 WEAVER J L AND S H prittsFRITTSFBITTS 1979 175 177 comparison of coyote and wolf scat diameters journalnalnai ofwildlife GIPSON P &S 1974 food habits of coyotes jour in arkansas management 43 786 788 journal ofwildlife management 38 848 853 GBEENJcreenGREEN J S ANDJAND J T FLINDERS 1981 diets of sympatric foxes and coyotes in southeastern idaho great received I1 june 1989 basin naturalist 41412514135141.251251 254 accepted 10 october 1989 cleatcreatgreat basin naturalist 501 1990 appp 67 72

INFECTION OF YOUNG DOUGLAS FIRS BY DWARF MISTLETOE IN THE SOUTHWEST

1 robert L mathiasenmathiasen1 carleton B edminster and flankfrank G hawksworth 2

ABSTRACT studies in several dieasdreasareas inm arizona and new mexico show that dwarf mistletoe arceuthobium doug iasulasu is rare in young douglas firs growing under infected overstoriesoveroverstonesstoriesstones less than 5 ofthe douglas firs under 26 years old and less than 6 ofthose under 114m4 in tall were infected in 77 mistletoe infested stands both percent infection and mean dwarf mistletoe rating of young douglas firs increased as tree age height and stand dwarf mistletoe ratings increased

douglas fir dwarf mistletoe arceuthobium 1986 also related infection of douglas firs dougladouglasiisii engelm is the most prevalent and less than 26 years old to three factors affecting damaging disease agent in southwestern infection of young trees listed by wicker mixed conifer forests andrews and daniels 1967 these included exposure time over- 1960 hawksworth and wiens 1972 jones story inoculum levels and sapling density 1974 this parasitic flowering plant occurs during a study designed to collect growth throughout the range of its principal host data for the development of a regeneration douglas fir pseudotsuga menziemenziesiimenziesiasii birbmirb model for southwestern mixed conifer stands franco in the southwest andrews and additional data on the infection of young daniels 1960 estimated that approximately douglas firs were collected from 13 mistletoe 50 of the southwest s douglas fir type was infested stands in the white mountains ari- infested by dwarf mistletoe zona these data were combined with the douglas fir regeneration is a frequent com- original data collected by mathiasen 1986 ponent of the understory of southwestern and the results are reported here in addition mixed conifer stands moir and ludwig 1979 the entire data set was summarized using the gottfried and embry 1977 fitzhugh et al heights of sampled douglas firs because pre- 1987 when overstoriesoverstories are infested with vious investigators have suggested that height dwarf mistletoe spread to young and advance may be a critical factor influencing infection of regeneration perpetuates the infestation over young trees by dwarf mistletoesmistletoes graham time therefore management of mixed 1960 hawksworth 1961 childs 1963 wicker conifer forests should attempt to minimize the and shaw 1967 scharpfl969scharpf 1969 infection of new and established regeneration from already infested over 1974 overstoriesstories jones METHODS gottfried and embry 1977 mathiasen 1986 summarized previous re- during 1980 81 douglas fir regeneration search on this problem and the factors that was sampled in 64 mistletoe infested mixed influence dwarf mistletoe infection he also conifer stands in four national forests in ari- provided some preliminary information on in- zona and new mexico A total of364 douglas fection of young douglas firs and sprucedspruces in fir saplings were sampled for total age height the southwest he found that little infection and height to live crown in addition each of douglas fir occurs before saplings are 26 douglas fir was examined for dwarf mistletoe years old only 6 of the douglas firs he infection and assigned a dwarfmistletoe rating sampled that were less than 26 years old were DMR using the 6 class system hawksworth infected whereas infection of older douglas 1977 this rating system divides the live fir reproduction averaged 83 mathiasen crown of a tree into thirds and each third is

USS forest service foiestforest pest Managermanagementnent 324 25th st ogden utah 84401 2uau S forest service rocky MounmountainmountaidtaldtaintaiD forest and range experiment station 240 W prospect st foiforfort t collins colorado 80526

67 68 R L MATHIASEN ETAL volume 50

rated separately as 0 no mistletoe infection infected saplings represent less than 4 of 1 less than 50 of live branches infected 2 saplings under 21 years old sampled infec- more than 50 of live branches infected the tion of saplings less than 16 years old was only ratings for each third are totaled to obtain the 2 in stands with a stand DMR greater than DMR for a tree mean stand DMR and mean 303.0 also very little infection of saplings less sapling DMR are calculated by adding the than 26 years old was found table 1 only DMRs for all live overstory trees or saplings 10 of saplings 21 25 years old were infected and dividing by the total number of live trees and all were in moderately infested stand or saplings respectively infection intensity is DMR 161.6iglg 303.0 or severely infested stands defined here as the mean DMR of the over- stand DMR greater than 303.0 story or saplings in a stand infection of 26 30 year old saplings in- overstory data collected for the 1980 81 creased to 30 in lightly infested stands stands were from rectangular plots ranging stand DMR 010.10oi 1 151.5isls1 5 and to over 65 in both from 0 04 to 0 36 ha for each live tree over moderately and severely infested stands 14mm1 4 m in height the species diameter at breast table 1 generally infection continued to height dahdbh to the nearest 2542.542 54 cm DMR increase as sapling age increased table 1 and crown class dominant dominantcodominantco in- A total of 14 infected saplings under 26 tertermediatemediate or suppressed were recorded years old were sampled these saplings were these data provided information on overstory in severely infested stands were over 14m141.4 m dwarf mistletoe infection intensity species in height had high crown ratios greater than composition and stand structure 00700.7070 or were in stands with over 740 saplings in 1988 an additional 334 douglas fir sap- per ha many of these 14 saplings had more lings were sampled in 13 mistletoe infested than one of the above factors contributing to mixed conifer stands in the white mountains their infection potential arizona data were collected as in 1980 81 percent infection and mean DMR for overstory data collected were the same as in saplings demonstrated the same pattern for 1980 81 but 0 04 ha circular plots were used height classes as for age classes tables 1 2 stand dwarf mistletoe ratings were calcu- little infection 10 or less was found in lated using all live douglas firs greater than saplings less than 141.4 m in height except in 2542.542 54 cm dahdbh for 1980 81 plots and greater the most severely infested stands where we than 5085.085 08 cm dahdbh for 1988 plots sapling found 27 infection in saplings 1091.09log 141.4 m crown ratios were calculated by subtracting tall however saplings over 141.4 m in height height to live crown from total height and then had much higher infection levels percent dividing by total height percent infection and infection and intensities mean DMR than mean DMR for saplings were calculated by smaller saplings table 2 five year age classes and 3 m height classes for each of stand 1 1 three DMRDMR classes 0 5 discussion 161.61iglg 6 303.03 0 and greater than 303.03 0 sapling densities were determined for the number of wicker 1967 wicker and shaw 1967 douglas fir saplings inm 0040 04 ha circular sub and mathiasen 1986 discussed several of the plots nested in the center of larger plots in factors influencing the infection ofyoung trees each stand by dwarf mistletoesmistletoes including duration of exposure to inoculum amount of inoculum RESULTS target area density of regeneration and removal of seeds by wind snow and other envi- both the number of infected saplings per- ronmentalronmental factors infection of susceptible cent infection and infection intensity mean young trees is largely influenced by a complex DMR increased as total age total height and interaction of the above factors mathiasen stand DMR increased tables 1 and 2 no 1986 presented information on the influence mistletoe infection was found on saplings un- of exposure duration to inoculum as ex- der 21 years old in stands with a stand DMR pressed by tree age amount of inoculum as less than 303.03 0 and only five saplings under 21 expressed by stand DMR and regeneration years old were infected in stands with a stand density as expressed by number of saplings DMR greater than 303.03 0 table 1 the five per ha additional information is reported 1990 DWAJDWARF RF MISTLETOEMISTlemoelemmeLETOE INFECTIONI1NFECTIC N 69

TABLE 1 infectioninifecticin ofdouglasof dou elasglas fir saplingssaisaj hngsangs bby age classeselasclas es and standst md DMR classes

stand DMRD MR class 010.10oi 1 15 16 30 30 total age class mean mean mean mean years N infinf1info DMR N inf DMR N inf DMR N inf DMR 16 49 0 00 58 0 00 108 2 0 1 215 1 010 1 16 20 30 0 00 29 0 00 21 14 010.10oiol 1 80 4 0 1 21 25 15 0 00 43 9 010 1 35 14 01 93 10 0 1 26 30 27 30 03 38 79 08 35 69 07 100 62 06 31 35 8 25 03 39 80 11liii1.11 1 41 88 09 88 78 09 36 40 12 66 09 40 90 14 36 97 18 88 90 15 41 45 7 71 09 15 80 20 12 100 28 34 85 19 total 148 16 02 262 43 06 288 41 06 698 36 05 percent infection

TABLE 2 infection of douglasofdouglas firhirbir saplings by height classes and stand DMR classes

stand DMR class 010 1 15 16 30 303.03 0 total height class mean mean mean mean m N inf DMR N inf DMR N inf DMR N inf DMR 154515 45 23 0 00 20 0 00 45 0 00 88 0 00 46764676.467646 76 14 0 00 13 0 00 35 3 0 1 62 2 010 I1 77 1071.071 07 21 5 010 1 33 0 00 31 6 01 85 4 0 I1 1 10 1371.371 37 21 5 02 29 10 01 22 27 02 72 14 02 1 40 1681.681 68 15 27 04 44 61 121 2 32 66 131.31 3 91 57 101.0iolo1 0 1 71 1981.981 98 15 40 07 48 65 151 5 41 71 161 6 104 64 14 2012 01 22922.2929 9 44 06 23 70 141.41 4 24 75 20 56 68 13 229 30 23 06 52 70 121.2131 2 58 69 18 140 59 14 total 148 16 02 262 43 06 288 41 06 698 36 05 percentpercent infection here for these factors as well as for the in- agementargement plans that do not successfully pre- fluence of target area as expressed by the vent or significantly reduce the infection of total height and crown ratio ofyoung infected douglas fir regeneration only serve to perpet- douglas firs in most situations where in- uate mistletoe infestations although the total fected young douglas firs less than 26 years heights crown ratios and densities of re- old were found a severely infested overstory generation contribute to the potential for stand DMR greater than 303.0 was present reinfection of young underunderstoriesstories these fac- infected young douglas firs in stands with a tors cannot be managed on a practical basis stand DMR less than 303.0 either had high however management plans that remove the crown ratios greater than 00700.7070 were over most severely infected trees followed with 14m1 4 in tall or were in stands with regeneration intermediate sanitation removals can effec- densities over 740 saplings per ha all three tively reduce the level of dwarf mistletoe in- factors would increase the potential for infec- fection in stands hawksworth 1978 thereby tion of young trees in dwarf mistletoe reducing the potential for infection of new or infested stands because of greater available advance regeneration in addition because target area douglas fir regeneration is not frequently in- because severe infection by dwarf mistle- fected before it reaches ages over 25 years in toe significantly reduces the growth of mer the southwest cutting cycles of 20 years or chantable size douglas firs in the southwest less allow managers at least two management and increases mortality of all size classes an- entries for reducing the level of mistletoe in a drews and daniels 1960 mathiasen et al stand before new douglas fir regeneration 1990 its control is an important consider- will be affected because little or no infection ation for resource managers vegetation man will occur until douglas firs are over 20 years 70 R L MATHIASEN ETETALAL volume 50 old in lightly infested stands the removal of height in lightly and heavily mistletoe severely infected overstory trees will signifi- infested stands respectively and infected cantly reduce the potential for infection of pines averaged 232.32 3 and 202.02 0 in in the same new and advance douglas fir regeneration stands scharpf 1969 reported that only the age at which douglas fir regeneration 7 of true firs under 009m9 in tall were infected becomes infected by dwarf mistletoe in the in severely infested stands inm california but southwest contrasts sharply with results re- that infection intensified rapidly in taller ported for other tree species and regions regeneration the results of infection of weir 1918 found that the average age of 50 douglas firs by height classes reported here naturally infected douglas fir seedlings used indicate that little infection can be expected for assessing the effects of dwarf mistletoe on until the trees reach heights greater than seedling growth in the northwest was 18 141.41 4 in in the southwest years hawksworth and graham 1963 found because these findings have important very little infection in lodgepole pine pinus implications in managing dwarf mistletoe confortacontortacontorta dougl ex loud reproduction un- infested stands similar studies should be con- der 10 years old but infection increased ducted for other dwarf mistletoe host combi- markedly in older stands 9 at age 15 18 at nations in other regions of the western united age 20 and 32 at age 25 some infection of states the results show that the generally ponderosa pine pinus ponderosa laws by accepted recommendation that infected over- southwestern dwarf mistletoe arceuthobium story pines and true firs be removed before vaginatumvaginatum subsp cryptopodum engelm the young stand is 10 years old or 090.90og 9 in tall hawksw & wiens has been found in loyear10 year is more restrictive than need be for douglas old seedlings gill and hawksworth 1954 fir in the southwest where little infection hawksworth 1961 based on these findings occurred inm stands under 20 years old or less for pines johnson and hawksworth 1985 than 14m1 4 in tall recommended that mistletoe infected residual trees be removed before the young stand literaturelltfratube CITED is 10 years old however the results of this study indicate that for southwestern douglas ANDREWSANDRFWS S R AND J P DANIELSDANIFLS 1960 A survey of dwarf mistletoesmistle in new fir the infected overstory trees could be left toes in arizona and mexico USDA foiestforest service rocky mountain forest for up to 20 years because of the very slight and range experiment station paper 49 17 appp chance of infection CHI 1islisilsaisDS T1 W 1963 Dwarfdwarfmistletoedwaifmistletoemistletoe control portuniopportuniopopportune there is less published data for the relation- ties in ponderosa pine reproduction USDA ship of regeneration height and dwarf mistle- foiestforest service pacific northwest forest and range experiment station unnumbered report toe infection but the general recommenda- 20 appp tion is that mistletoe infected residual trees fitzhughpitzhugh E L W H moirmolltmolitMOIB J A LUDWIG AND should be removed before the young stand is F RONCOBONGOrongo 1987 foiestforest habitat types in the 090.9og in tall johnson and hawksworth 1985 apache gila and part of the cibola national new graham 1960 found that forests arizona and mexico USDA for- dwarf mistletoe est selvleeservice general technical report RM 145 infection in douglas fir increased as size class 116ppilg116 appp increased in northern idaho only 15 of the cillGILLcligil611 1 L S AND F G hawksworth 1954 dwarfmistleDwarfmistle saplings sampled by graham were infected toe control in southwestern ponderosa pine forests whereas 25 and 39 of small and under management journal of forestry 52 the large 347 poles respectively 3 353 were infected hawks- coigotrigoigodriri biedbledmedwedRILD G J AND R S EMBRY 1977 distribution of worth 1961 reported that 19 of the pon- douglas fufir and ponderosa pine dwarf mistletoesmistle toes derosa pines in the 254 cm diameter class in a virginvngm arizonaalizona mixed conifer stand USDA were infected in stands infested by southwest- forest seiselservicevleevice research paper RM 192 16 appp GRAHAM D P 1960 dwarfmistletoeDwarfmistletoe in nezpercenezperee ern dwarf mistletoe in suiveymsurvey Nezperce northern arizona but national foiestforest USDA forestfoiest service research infection increased to 57 in the 127 cm noteint75note INT 75 7ppapp7 appp diameter class childs 1963 working in the hawksworth F G 1961 dwaifmistletoedwarfmistletoeDwarfmistletoe of ponderosa pacific northwest found that uninfected pine in the southwest USDA forest service technical bulletin 1246 112 appp ponderosa pines averaged 151.5 and 111iili1.11 inm in 1977 the 6 class dwaldwarff mistletoe rating system USDA forest serviceselvlee general technical report Gragiahaingrahainhain did not specify the dumetdimetersdumetcisdiametersdiletersmeterseiscisels 0of the size classes lie used RM 48 7 appp 1990 DWARF MISTLETOE INFECTION 71

1978 intermediate cuttings in mistletoe infested and sprucedspruces by dwarfdwarfmistletoesmistlemistiemistletoestoes in the southwest lodgepole pine and southwestern ponderosa pine great basin naturalist 46 528 534 stands in R F scharpfandScharp fand J R parmeter tech MATHIASEN R L FK G hawksworth AND C B EDMIN coordinators proceedings of the symposium on STER 1990 effects ofdwarf mistletoe on douglas dwarf mistletoe control through forest manage- fir in the southwest in preparation ment USDA forest service general technical MOIRMOIB W H AND J A LUDWIG 1979 classification of report PSW 31 86 92 spruce fir and mixed conifer habitat types of ari- hawksworth F G AND D P GRAHAM 1963 spread zona and new mexico USDA forest service and intensification of dwarfmistletoedwarfmistletoe in lodgepole research papelpaper RM 207 47 appp pine leproductionreproduction journal of forestryForestly 61 SCHARPF R F 1969 dwarfdwarfmistletoemistletoe on red fir infection 587 591 and control in understory stands USDA forest hawksworth F G AND D WIENS 1972 biology and service research paper PSW 50 8 appp classification of dwarf mistletoesmistletoes arceuthobium WICKER E F 1967 seed destiny as a klendusic factor of USDA forest serviceselvleeSeivice agriculture handbook infection and its impact upon propagation ofarofafof ar 401 234 appp ceuthobium sppapp phytopathology 57 1164 1168 JOHNSON D W AND F G hawksworth 1985 dwarf WICKER E F AND C G SHAW 1967 target area as a mistletoesmistletoes candidates for control through cul- klendusic factor in dwarf mistletoe infection phy- tural management USDA forest serviceselvleeSeivice gen- totopathology 57 1161 1163 eral technical report WO 46 48 55 wibbwiebWIER J R 1918 effects of mistletoe on young conifers johesJONES fJ R 1974 silviculture of southwestern mixed journal ofagricultural research 12 715 718 conifers and aspen the status of our knowledge USDA forest service research paper RM 122 44 appp received 31 january 1989 MATHIASEN R L 1986 infection of young douglas firs accepted 16 january 1990 great basin naturalist 501 1990 appp 73 82

NEW MEXICO GRASS TYPES AND A SELECTED bibliography OF NEW MEXICO GRASS taxonomy1TAXONOMY1

kellawkellywkellykeily W allredailredalired

ABSTRACT collection data bibliographic citations and curatorial information on 52 names of new mexico grass types are compiled A bibliography oftaxonomicof taxonomic research pertinent to the study ofnew mexico grasses is cross refer encedencee with genera known to occur in the state

bibliographic and historical information collections of 9 new taxa from new mexico are an essential but often neglected resource grant and santa fe counties contain the most for the student of plant systematics the cor- localities of new grasses table 2 santa fe is rect application of plant names requires accu- one of the oldest towns in the united states rate information concerning nomenclatural and was visited by many collectors early in types and precise floristic and identification the 1800s william gambel passed through in work demands access to reliable monographic 1841 or 1842 on his way to california his or revisionaryrevisionary literature this is especially collections were described by thomas nut- so when changes are made in traditional tall wislizenus followed in 1846 august systematic alignments reference literature fendler made extensive collections there in allows others to understand and evaluate the spring of 1847 sending them to asa gray the revisions most of his collections came from the santa fe nearly every botanist engaged in the taxon- creek area and within 10 12 miles of santa omy of grasses gramineae during the past fe A A heller G R vasey S M tracy century described at least one novelty from and T D A cockerell were other botanists newnow mexico material the list of grass types who collected near santa fe in the late 1800s presented here includes 52 taxa known to or early 1900s the collections of vasey and have been described from specimens gath- tracy contributed new grasses many of the ered in new mexico As a point of compari- collections from grant county came from the son 19 grass taxa have been named from utah mining camp of santa rita 15 miles east of material welsh 1982 twenty one different silver city charles wright J M bigelow authors contributed new taxa but three and george thurber collected there in the george vasey 14 names john torrey 6 1850s mangas springs also in grant county names and ernst steudel 5 names ac- was visited by 0 B Metcametcalfemeteaifeioeloe in 1903 who counted for nearly 50 of the plant names collected several hundred sets of plants table 1 botanical publication of grass taxa C G pringle H H rusbybusby and J G smith from new mexico began in 1854 with species also collected new grasses from grant county of aristida muhlenbergia oryzopsis and standley 1910 poa steudel 1854 and has continued for well the ensuing list attempts to include all over a century the latest being in 1986 from grass names based on new mexico material the genus andropogon campbell 1986 the author of the name publication data of the authors only A S hitchcock paul collector and number locality of collection standley george thurber george vasey deposition of type material and current taxo- and E 0 wooton also participated as field nomic disposition of the name are given for collectors of new grasses from new mexico each type table 1 george vasey heads the list with following the list of types is a list of the

journal article 1474 new mexico agricultural experiment station las cruces department of animalofanimal and range sciences box 313 I1 new mexico state university las cruces new mexico 88003

73 74 K W ALLRED volume 50

TABITABLE 1 1 authorsAutholthoithols and collectorscollect oisors of new mexico glassgrass tabietableTABI 2 counties of collection of new mexico grass types types namenarne authored collected county number of types thomas antisell 0 1 Bernahbernalillolloiloiio I1 william J beal 1 0 colfax I1 john M bigelow 0 5 donadoniadofna ana 4 W S boyle 1 0 eddy 3 samuel B buckley 2 0 clantcrantgiantglantgrant 12 christopherChu stophel S campbercampbehcampbellcampbeli 1 0 hidalgo 1 kalelkaleikarelKaielkaieldomindomin 2 0 lincoln I1 william H emoiyemolyemory 0 1 otero 2 august fendler 0 7 rio arriba 2 william gambel 0 2 san miguel 3 eduardeduaidhackelhackel 1 0 santa fe 11 albeitalbert S hitchcock 2 2 socorro I1 edwin james 0 1 ivan M johnston 1 0 marcusmareus maicus E jones 1 0 andropogon glomeratusglomeratus walterwaltelwaiter BSPS P var 0 C louismarielowslouislomslouls mariemarlemane 1 0 scabriglumus campbell systcyst bot 11 291 1986 edgar A mearns 0 1 C wright 2100 in 1851 grant co probably near silver elmer D merrillmen illlii 1 0 city GH orrickornckbB metcalfe 0 3 andropogon neo mexicanus nash bull bot george torr nash 3 0 club 25 83 1898 E 0 wooton 583 26 aug 1897 otero thomas nuttall 2 0 co white sands NY schizachynumschizachyrium neomexicanumneonwxicanum cyuisgcyrusG pringle 0 1 nash nash H H rusbyrushybusby 0 1 andropogon wrighwrightiiwrightiiitii hackel flora 68 139 1885 L scribner 4 0 frankfianklfrankl C wright 2104 in 1851 grant co silver city US cornelius L shear 1 0 bothnochloabothriochloa wrighwnghtiiwrightiiwrightiiitiitil hack henrhenn jaredgjaredyjared G smith 1 2 aristida syn pipl glum 1 420 paulpaulcC standley 1 1 fendlerianafendleriana steudel clum 1854 fendler 973 in 1847 santa co santa einstgeinsigernst G steudel 5 0 in fe fe US jason swallen 3 0 aristida purpurpurpureapurpureanea nutt vaivalvar fendlerianafendleriana steudstaud vasey george thurber I1 I1 john torrey 6 0 aristida longisetalongiseta steudel syn pipl glum 1 420 1854 978 1847 samuel M tracy 0 1 fendler in pioploprobablybabisbablybabiy santa fe co santa fe geoigcvaseygeorgeceorge vasey 14 9 US aristida purpurpurpureapurpureanea nutt var longisetalongiseta staudsteud vasey wilkins 0 1 S W woodhouse 0 1 aristida pansa wooton & standley contr U S nati elmereimer 0 wooton 1 5 herb 16 112 1913 wooton s n 60ct6 oct 1904 donadonia ana chaileswnghtcharlescharies wright 0 6 co tortugasTortugas mt nealnear las cruces US aristida subuniflora nash in SIsmallsmailfiallflall fl southeast US 116 1903 vasey sn& n in 1881 new mexico grass genera of new mexico erosscross referenced probablypioprobablybabiy santa fe co near santa fe NY aristida purpurpurpureapurpureanea nutt var fendlerianafendlenanafendlependle nana staudsteud vasey to a selected bibliography the bibliography riana bouteloua ausillapusilla vasey bull torr bot club 11 6 is not intended to be exhaustive rather only 1884 G R vasey sn& n kugmankingmankmgman new mexico US significant revisionary or summary papers probably kingman arizonaanzona as there is no known king- pertinent to new mexico grass taxonomy are man new mexico bouteloua simplex lag listed further references may be obtained by bromus porteriponteri coulter nash var frondosus shear consulting the works listed here particularly USU S DAD A div agrostagnost bull 23 37 1900 J G smith sns n grant co mangas springs US gould and shaw 1983 and soderstrom et al calycodonCalycodon montanusmonmontanumtanum nuttall J acad nat sci 1987 phil n seiser 1 186 1848 W gambel sns n santa fe co near santa fe PH muhlenbergia montana nutt NEW MEXICO GRASS TYPES A S hitchehitchc chaetochloa grisebachii fourn scnbnscrilmscrimm var amalaampla the acronym in parentheses holmgren et scribner & merrillmen illlii USDAU S D A div agrostagnost bull 21 36 al 1981 referslefers to the locality of type material 1900 G R vasey sns n in 1881 dona ana co organ holotype isotype fragment or other dupli- mountains US one of two specimens cited setariasetona cate material gnsebachugrisebachii fourn chondrosium eriopodum torrey in emory notes mil agrostis minutissimammutissima cudelcudeieudelsteudelSt syn pipl glum 1 171 deconnreconn 154 1848 bigelow sns n in 1847 along the del 1854 fendler 986 in 1847 santa fe co nealnear santa fe norte rio grande river new mexico US US muhlenbergia ininutissimammutissima staudsteud swallen bouteloua erioerloenopodaeriopodapoda torr torrtorn 1990 NEW MEXICO GRASS TAXONOMY 75

chrondrosiumfoeneumchrondrosium foeneum torrey in emory notes mil bunoburro mountains NY digitariaDigitana californicacalifornica benth deconnreconn 154 1848 emoiyemory sns n in 1847 uplands bor- henr dering the valley of the norte grande del rio panicum plenum A S hitchcock & chase contr USU S bouteloua hirsutehirsuta lag nati herb 15 80 1910 metcalfe 739 sep 1903 grant epicampesEpicampes subpatenssubpatens A S hitchcock U S D A bullbuilbuli co mangas springs US Panipalcumpanlpanicuinpamcumcuin bulbuibulbosumbulhosumbulbhosumosum H B K 772 144 1920 A S hitchcock 13541 eddy co pleopogon setosum nuttall J acad phil n ser 1 189 guadalupe US mountains muhlenbergia emersleyi 1848 W gambel s vasey snn in 1841 or 1842 santa fe co mountains of santa fe PH lycurus phleoides eragrostisfendlerianaeragrostis fendlerianafendletianariana steudel syn pipl glum 1 278 H B K var glaucifoliusglaucifohus beal 1854 fendler 932 in 1847 mexico now new mexico pleuraphisPleuraphis jamesiijainesiijarnejainejamesiesii jtoiieytorrey ann lye NYN Y 1 148 probably santa co near santa US poafiendlepoa fendlependle fe fe 1824 james s n in 1852 sources of the canadian riananana steudstaud vasey in river texas 01or new mexico NY hilaria jamesiijamesnjameanjamesie torr eragrostis neomexicana vasey contr U S nati benth heibhelbherb 2 542 1894 vasey sns n in 1881 dona ana poa aridaarldaarvoarmoarma vasey contr 1 co organ mountains US conti US catlnatl heibhelbH erb 270 1893 eragrostis mexicansmexicanamexicana R vasey hornembornem link G s n in 1881 socorro co socorro US poa bigeloviibigelovii vasey & scribner U S fendleriaFendleria rhynchelytroides steudel syn pipl glumgium 1 conticontr US natnati 1 270 1893 931 1847 420 1854 fendler 979 in 1847 probably santa co heibherb fendler inm new mexico fe santa co probably along nealnear santa fe US oryzopsis hymenoideshymenoides roem & fe santa fe creek east of schult rickerrieker santa fe US poa koeleriaKoeleria macrura domin forma trifloratritrifloralflora domin bibi occidentalsoccidentaleoccidentoccioccidentalisdentalsalisails vasey conconn tr US nati heibhelbherb 1 274 vasey bot 65 238 1907 E 0 wooton 110 probablypioplobablybabiy 1897 1893 G R sns n min 1881 santa fe co near santa dona ana co organ mountains US koeleriaKoeleria fe US cranthamacranthama ledealedeb schult poa tracyl vasey conticontr U S nati herb 1 276 1893 koeleriaKoeleria nitidanatida nuttall var laxa domin bibbibi bot 65 S M tracy sns n in 1887 colfax co on mountain sides 235 1907 0 B metcalfemetcalfesMetcal feshess n perhapspelpei haps grant co new atratonstratonat raton US mexico another specimen from arizona also cited sitanionSitanion caespitosum JGG smith USDAU S D A div koeleriaKoeleria macrantha ledebledea schult agristagiostagrost bull 1816189918 16 1899 J G Smithsmithssnn min 1897 crantgiantgrant melica porteriponteri scribner vaivar laxa boyle Madromadronofioflo 8 25 co nealnear cliffuscliff US elymus longilongifoliuslongtfoliusfolius smith gould 1945 E 0 wooton 680 in 1897 lincoln co white sporobolus filiculmis beal grasses N amer 2 288 mountains US 1896 bigelow sns n sep 1853 rio arriba co plaza muhlenbergia abata I1 M johnston J arnold arbor larga muhlenbergiaMuhlenbergutbergui thurthurbenthuyberibefi redbrydb 24 387 1943 wright 1982 in 1851 valley of the rio sporobolus giganteusgiganteus nash bull torrtorn bot club 25 grande presl S GH muhlenbergiamtihlenbei gia rapensrepens presi A 88 1898 E 0 wooton 394 26 aug 1907 otero co hitchehitchc white sands US muhlenbergia acuminataacuminateacuminata vasey bot gazcaygay 11 337 sporobolus thurberithurthurbergberi scribner USU S DAD A div agrostagnost 1886 wright 1993 in 1851 probably grant co near bull 11 48 1898 bigelow sns n sep 1853 rio arriba santa rita US muhlenbergia dubiaduinadubla fourn co plaza larga US muhlenbergia thurbenthurberiberlbefi redbrydb muhlenbergia metcalfeimetealfeimetcalmetealfeifelfei M E jones conticontr west stipa cwvifoliacurvifolia swallen J wash acad sci 23 456 bot 14 12 1912 B metcalfe 1485 in 1904 grant 0 in 1933 wilkins 1660 eddy co guadalupe mountains co santa US rita mountains US muhlenbergia neo mexicansmexicana vasey bot gaz 11 337 stipa lobata swallen J wash acad sci 2319923 199 1933 1886 G R vasey sns n in 188p1881 i ocksockyoekyrocky hillsbills and moun- A S hitchcock amer gr tain sides new mexico US muhlenbergia nadnati herb 819 eddy co baucipauci queen guadalupe US flora buckl mountains pennata L muhlenbergia sinmsasinuosasinuose swallen contr U S nati stipa jennata var neomexicana thurber in coulter 29 mearns man rocky mount 408 1885 thurber 269 probably herbheibhelb 204 1947 E A 2457 in 1893 hidalgo grant co san luis mountains US co rio mimbresmembres stipa neomexicana thuibshuibtburbTburb scribn muhlenbergia sylvaticasylvatica torr tontorr var flexuoseflexuosa vasey in wheeler rptapt USU S survey W looth mendmerld 6 stipa scribneriscribnersscribneri vasey bull torr bot club 11 125 vasey 1 284 1878 wright 731 in 1851 peipelperhapshaps cianicrantgiantgranigrant co near 1884 s n in 1881881 santa fe co santa fe US silver city muhlenbergia monticola buckl tricuspistricuspid luticamutica toltoitorreyi ey USU S eiplexpl miss jacifpacif rptapt muhlenbergia sylvaticasylvatica torr torr var prinpringleipringleeglei 4 156 1856 bigelow s n 22 sep 1853 san miguel co scribner bull torrtorn bot club 9 89 1882 C pringle laguna colorado NY tndenstedenstridens euticusmuticuimuticusmutimuttcuicur torr nash 480 crantgiantglantgrant co santa rita mountains US muhlen- trisetum montanusmontanuinmonmontanummontafanumtanumnuin vasey bull torr bot club 13 bergia pauciflorapaucijlora buckl 118 1886 G R vasey sns n in 1881 san miguel co las muhlenbergia wrighwrightiiwrightiiitii vasey in coulter man rocky vegas US mount 409 1885 wright 1986 in 1851 colorado and trisetum montanimontantmontanusmontaniimmontanummontanumim vasey var pilosum louismarielouislouls mariemarlemane new mexico US rhodora 30 212 1928 P C standley 4536 aug 1908 panicum bulbbulbosumosum HBKH B K var minus vasey san miguel co cowles trisetum montanusmontanummontanum vasey USU S DAD A div bot bull 8 38 1889 rusbybusby s n in 1880 Urauralepislepis composita buckley proc acad nat sci grant mangas perhaps co near springs US phila 1862 94 1862 woodhouse sns n probably 1851 panicum lachnanthum torrey USU S eiplexpl miss jacifpacif new mexico PH leptochloaleptochroaLeptochloa fascicularfascicularisfasciciilansis lam rptapt 7 21 1858 T antisell s n in aug 1854 grant co gray 76 KWALLREDK W ALLRED volume 50

Urauralepislepis poaeoides buckley proc acad nat sci glyceriaglycerin 54 phila 1862 94 1862 fendler 932 in 1847 probably hackelochloa 118 santa fe co near santa fe PH poa fendlerianafendlenanafendlependle riananana helictotrichonhehctotnchon 118 staudsteud vasey heteropogonHetero pogon 102 118 vilfatricholepistorreysilfavilfa tricholepistricholepis torrey VUSS eiplexpl miss jacifpacif rptapt 4 hierochwhierochloe 230 155 1857 bigelow s n oct 1853 bernalillo co sandia hilaria 200 232 mountains hana NY blepharoneuron tricholepistncholepzstricholepis torr holusbolushofcus 118 nash hordeum 24 25 26 27 28 29 30 35 36 38 64 152 153 imperata 118 SELECTED bibliography OF koelenakoeleriajoelenaKoeleria 109 192 193 NEW MEXICO GRASS TAXONOMY leersia 172 leptochloaleptochroaLeptochloa 147 general literature 3 20 46 48 58 59 60 102 log 116 102106106 laymusleymusleyotms 111711 17 7815178 151 118 119 150 189 199 204 205 227 229 236 lomw83124218220lolium 83 124 218 220 237 238 239 237238239 lycurus 175 tnticeaetriticeae tribe 18 19 23 38 39 41 77 78 79 151 melica 40 agropyronslagropyronSl51 65 75 76 78 80 151 167 185 216 Muhlenbergmuhlenbergiamuhlenbergiamiamtawtam92 155 156156169169 ito170174197170174.197170 174 197 agrostis 32 165 214 233 munroa 8 alopecurus 194 oryzopsis 15 125 49 andropogon 47 panicumpanicum3737 44 52 117 142 143 144 145 225 240 anthoxanthum 107 242 pappophorum 53 182 aperaapera118118 paspalum 1 12 111 159 aristida 2 102 pennisetum 52 221 arrhenatherum 118 phalaris 5 arundoi18arundo118Arundo 118 PWUTOphleum 121 avenaavena2121 22 phragmites 57 91 BeckbechmanBeckmanbeckmanniabeckmanmamanniama 90 177 phyllostachys 60 blepharoneuron 118 piptochaetwmpiptochaetium 15 118 bouteloua 103 ilslis115115180180 poapoo 10 33 112 132 133 141 149 201 202 brachiariabrachtartaBrachiariataftalarlalafiatarta 229 240 polypogonPolypogon 118 bromus 34 84 113 146 157 166 191 195 198 207 psathyrostachys 78 81 208224208 224 Pucepuceinelliapuccinelhainellia 54 61 66 93 buchlbuchloebuchloh 171 Redfieldia 50 178 calamagrostis 108 131 206 rhynchelytrum 118 calamomlfacalamovilfa 222 saccharumSacchariim 60 catabrosaCatabrosa 118 schedonnardus118 cenchrus 67 schistusschismusSchismus 62 89 chionschloris 6 138 schizachneschtzachne 137 212 cinna 118 chizachyriumschizachyriumS 102 118 coixcow 118 sclerochloa 213 cortaderiaCortadena 63 scleropogon 181 botteacottea 118 Scafesecalescafe382113838211211 cynodon 74 setariasetonasemona 85 868886 88 129129168176184168 176 184 221 dactylis 209 243 sorghastrum 223 dactyloctenium 102 118 sorgwn697282226sorghum 69 72 82 226926 danthoniaDanthoma 56 94 spartina 154 deschampsia 130 135 sphenopholissphenophohs 87 dichantheliumdichanthehum 105 sporobolus 128 183 digitaria 228 stenotaphrum 45 187 241 Distichdistichlishs 31 stipa 13 14 15 16 125 188 echinochloa 104 torreyochloa 54 55 61 eleusine 102 118 trachypogonTrachypogon 102 118 xelyhordeum 18 19 110 tragus 7 xelymotrigiaxelymotrigta 18 19 tedenstndenstridens 215 elymus 75 76 78 98 100 151 158 196 234 trlTnTritnplasistriplasistriplasicplasis 102 118 elyonurusElyonurus 102 118 tripsacum 71 Elyelytngiaelytrigiatrigia 18 19 51 65 99 151 216 trisetum 120 135 140 173 Enneenneapogonapogon 53 118 itnticosecalextriticosecale 42 203 eragrostis 59 9713413616417923597 134 136 164 179 235 triticum 38 43 126 127 Eremoeremopyrumpyrum 18 19 78 urochloaUrochloa 229 erloeriochloaeroerwchloaErwErio chloa 190 vpiavulpiavulpic 139 erloerioneuronErwErioerwneuronneuron 186 zeazeo 122 123 festuca 9 95 96 124 148 160 161 162 163 217 219 zoysia 60 118 1990 NEW MEXICO GRASS TAXONOMY 77

literature CITED 21 1968 on some relationships between avena sattva and A fatuafadua gramineae as studied 1 ALLRED K W 1982 paspalum distichumdistichum L var from canadian material can J bot 46 indutum shinners poaceae great basin natural- 1013 1022 ist 42 101 104 2221 1977 oats wild and cultivated a mono- 2 1984 morphologic variation and classifica- graph of the genus avena L poaceae dept tion of the north american ariskrisanstidaaristidatida purpurpurpureapurpureanea agrie ottawa complex gramineae Britbnttonia36tonia 36 382 395 23 1978 taxonomy of the tribe tnticeaetriticeae brittoniabrittania poaceae 3 ALLRED K W AND J I1 COLUMBUS 1988 new using various numerical taxonomic tech- mexico grasses a synopsis of the classification and niques III111 synoptic key to genera and synopses a key to the genera new mexico J sci 28 21 43 can J bot 56 374385374 385 4 ALLRED K W ANDANDFF W GOULD 1983 systematics 24 1980 Multivariatemultivanatemultivariate morphometncmorphomorphometriemetriemetric relation- of the bothriochloabothnochloa saccharoidessaccharoides complex poa- ships between hordeum jubatum and hordeum ceae andropogoneae systcyst bot 8 168 184 brachyantherum in canada and alaska can J 5 ANDERSON D E 1961 taxonomy and distribution bot 58 604 623 25 1983 oftheodtheofthe genus PhaphalarislartS iowa state J sci 36 1 96 morphometncmorphometricMorphometriemetric relationships in hor- poaceae 6 1974 taxonomy of the genus chlorischlongchiong deum vulgare tnticeaetriticeae II11 hordeum gramineae brigham young univ sci bull agrioagnocnthonagriocrithoncrithon H distichumdistichum H lagunculiforme H biolabiol ser 1921192 1 133 spontaneum and H vulgare can J bot 61 2023 2031 7 ANTON A M 1981 the genus tragus gramineae 26 1 1986 kewkewbullbullbuli 36 55 61 taxonomy of the north and south american species of hordeum section hordeas 8 ANTON A M AND A T HUNZIKER 1978 eleigenelgeneioElgeneugeneiogenero hordes eioelo trum can J bot 64 174517591745 1759 munroa poaceae smopsissinopsissynopsisSinopsis morfologica y taxohaxo 27 BAUM B R AND L G BAILEY 1984a nomica bol acad nac ci 52 229 252 taxonomic studies in wall barley hordeum munnummurimurlnum and sea 9 ARGUS G W AND S G AIKEN 1987 noteworthy dunnummurinusmurinum barley hordeum marlmarimannumnum I1 character collections new mexico festuca mariaummarinum investi- minutifloraminutiflora gation assessment of new and traditional charac- redbrydb Madromadronofioflo 34 268 269 ters can J bot 62 753 762 10 ARNOW L 1981 poa secunda presl versus poa sand 28 1984b taxonomic studies in wall barley bergnbergiibergei vasey poaceae systcyst bot 6 412 421 hordeum hunnummurinummunnummurimurl num and sea barley hordeum 11 ATKINS R J M E BARKWORTHBARKWOBTH AND D R DEWEY mariaummarinummannummarinum 2 multivariate morphomorphometricsmorphometncsmetrics can 1984 A taxonomic study oflaymus ambiguusambiambiguous leymusofleyrnus guus and J bot 62 2754 2764 L sahnassalinussahnus poaceae systcyst 9 tnticeaetriticeae bot 29 1988 A taxonomic of 279 294 investigation ofhordehorde- um arizoarlzoarizonicumanzomcumnicum poaceae tnticeaetriticeae with reference 12 BANKS D J 1966 taxonomy of paspalum cetaceum setaceum to related species can J bot 66 1848 1855 gramineae 269 sidaksida2sida 2 284 30 1989 are hordeum brachyantherum and 13 BARKWORTH E M 1978 A taxonomic study of the H califorcalifornicumcahformcumnicum tnticeaetriticeae poaceae conspecmc large plumedglumed gramineae conspecific species ofsttpaof stipa occur- can J bot 67 465246- 52 in canada can J bot 606 625 ring 566 31 BEETLE A A 1943 the north american variations 14 1979 A proposal to reject stipa colum- of distichlishs poaceae Distichdistichhs spicatespicata bull torrey bot club 70 biana and nomenclatural changes affect- 638 650 ing three western north american ofsfapa species ofstipa 32 BJORKMAN S 0 1960 studies in agrostis and poaceae taxon 28 621 625 taxon28 related genera lymbsymb bot upsal 17 1 112 15 1982 embryological characters and the 33 BOIVIN B AND D LOVE 1960 poa agassizensis a taxonomy gramineae oftheofodthethe stipeae taxon 312 new prairie bluegrass le naturalisteNaturaliste Canadien 233 243 8717387 173180173 180 16 BARKWORTH M E MCNEILL AND MAZE J ANDJJ 1979 34 BOOKMAN P A 1980 variation in bromus tector A poaceae taxonomic study ofstipa nelsoniinelsominelsonianelsoniinil with um poaceae in eastern washington madronoMadroflo 27 a key distinguishing it from related taxa in western 36 42 north america can 2539 2553 J bot 5722 35 BOOTH T A AND A J RICHARDS 1976 studies in 17 BARKWORIHBARKWORTH M ATKINS E ANDANDRR J 1984 laymusleymus the hordeum munnum aggregate 1 morphol- gramineae hunnummurinum hochst tnticeaetriticeae in north america ogy bot J linn soc 72 149 159 taxonomy and distribution amer amei J bot 71 36 BOTHMER R N N JACOBSON R B JORGENSEN AND 609 625 E NICORA 1982 revision of the hordeum pusti 18 BARKWORTH M E AND D R DEWEY 1985 ge lum group nordic J bot 2 307 321 mically no based genera in the perennial tnticeaetriticeae 37 BOUGH M J C COLOSI AND P B CAVERS 1986 ofnorthofnorth america identification and membership the major weedy biotypesbiotypes of proso millet panlpani- amer 72 767 J bot 776 cum mthaceummiliaceum in canada can J bot 64 19 BARKWORTHBABKWORTH M E D R DEWEY ANDANDRR E ATKINS 1188 1198 1983 new generic concepts in the tnticeaetriticeae 38 BOWDEN W M 1959 the taxonomy and nomencla- gramineae of the intermountain region keys ture oftheodtheofthe cheatswheats barbarleysbaileysleys and ryes and their wild and comments great basin naturalist 43 561 relatives can J bot 37 657 684 hordeum 572 secale Tnttriticumicum 20 BAUM B R 1967 kalmkahnkaim s specimens ofnorthofnorth ameri- 39 1967 taxonomy of interintergenericmtergenencgenericgenerlegenerie hybrids of can grasses their evaluation for typification the tribe tnticeaetriticeae from north america can J canad J bot 45 1845 1852 bot 45 711 724 78 K W ALLRED volume 50

40 BoyboylBOYIjii W S 1945 A cytotaxonomic study of the 62 CONERT H J AND A M TURPE 1974 revision der north american species of melica Madromadionomadronomadrofiofioflo 8 gattung Schschistusschismusbchismusismus poaceae arundinoideae dan 1 26 thoniethonieaeae abaabh senckenbergSenckenberg naturfnatura ges 532 41 boylBOYIboyle i W S AND A flH holmgrenHOLMRYN 1955 A bytecyte 1 81 netie study of natural and controlled hybrids 63 CONNOR H E 1983 names and types in cortadecorrade lebetweentween agropyron trachycaulumtrachycaulum and hordeum riafiaflastapfgramineaenastapfgiammeaeStapf Gramineae II11 taxon 32 633 634 jubatum genetics 40 539 545 64 COVAS G 1949 taxonomic observations on the 42 BRIGGLEBRIGOLF L W 1969 TriticatuticaletriticaleTuticaiecaleealele a review cropscicrop sciseisel north american species of hordeum madronoMadro fioflo 91979 197 202 10110loi 1 21 43 BRIGCILBRIGGLE L W ANDANDLL P REITZ 1963 classification 65 daubenmire R 1960 an experiment study of and of triticum species of wheat varieties growngi own variation in the agropyron spicatumspicasptcatumtum A inermeinermae in the united states USDA tech bull 1278 complex bot gaz 122 104 108 135 appp 66 DAVIS J I1 1983 phenotypic plasticity and the 44 BRUNKINBRUNKEN ESJTLS J N ANDJAND J R ESTES 1975 cytological selection of taxonomic characters in Puccipuccinellianellia and morphological variation in panicum virgatwnvirgatumvirgatum poaceae systcyst bot 8 341 353 southw naturalist 19 379 385 67 DELISLE D G 1963 taxonomy and distribution busey BROSCHAT 45 busibusl P T K BBOSCHAI AND B J CENTER 1982 of the genus cenchrus iowa state J sci classification of st augustinegrassaugustmegiassAugustinegrass cropcropscisciseisel 22 3725937 259 351 469 473 stenotaphrum 68 dewiDEWET J M J 1968 biosystematics of the both 46 burzinBUTZIN F 19731971 die namen deider supiagenenschensupragenerischen nochrioriochloanochlotichloaiottloti barbinodisbarbinodis complex gramineae ameramnerammer einheitenEinheiten deider gramineae poaceae willdenowia J bot 65477 484 71137 113 168 69 1978 systematics and evolution of sor- campblCAMPBICAMPBELL S 47 LI C 1983 systematics of the andro- ghum sect sorghum gramineae amer J bot pogon virgimcusvirginicusvirginiousvirginicus complex gramineae J arnold 6547765 477 484 64 171 254 arbor 70 DEWET J M J D S borgaonkar AND W L 48 1985 the subfamiliessubfamilies and tribes of richardson 1963 chromosome number and poaceae grarnineaegramineae in the southeastern united mode of reproduction in the bothnochloimnaebothriochloininae states J arnold arbor 66 123 199 caryologiacaiyologial6Cary ologia 16 47 55 49 1986 phylogenetic constructionsreconstructionsie and 71 DFWLTDEWET J M J J R GRAY ANDahdeahdjJ R HARLAN 1976 two new varieties in the andropogon virginicusmrgimcusvirginiousvirginicus systematics of 1tripsacumripsawm gramineae phytolopbytolo complex poaceaepoaceae andropogoneaeandiopogoneae systcyst bot 11 giaglagia3333 203 227 280 292 72 dewiDEWET J M J J P HUCKABAY 1967 the origin of 50 CAMUS A 1954 etude du genre Redfieldredfieldiaredfwidwia vasey sorghum bicolor II11 distribution and domestica- amciicamam6ricain et malgacheMalgache not system 15 7 10 tion evolution 21 787787802802 51 CBAPMANCHAPMAN S R AND L J PERRYpi rbyRHY JR 1973 taxo- 73 DEWET J M J AND J R HARLAN 1970a both nomic and agronomic variation in agropyron spi rionochloariochloachloa interintermediaintermedialmedia a taxonomic dilemma taxon batumcatum and A inermeinermae J range manage 26 1933919 339 340 41 43 74 1970b biosystematics of cynodon L C 52 CHASICHASE A 1921 the linnaean concept of the peaipealpearlpeari1 rich gramineae taxon 1956519 565 569 millet amelamnelamerammel J bot 8418 41 49 75 dewlyDEWIYDEWEY D R 1963 natural hybrids Agropyronofagropyronofag spyronopyron 53 1946 Enneenneapogonapogon desvaux ii and pap trachifcaulumtrachycaulumtrachycaulum and agropyron scribbenscribnenscribneriscribnerinerlnen bull pophorum wrighwnghtnwrightiiwrightiiitiitil an agrostological detective torreytoireybotbot club 90 111 122 stolystory Madromadrofiomadroniomadiono8fiogio 8 187 189 76 1964 natural and synthetic hybrids of 54 CHURCH G L 1949 A cytotaxonomic study of agropyron spicaspicatumtum x Sitsitamonsitanionsidamonanionanlon hystrix bull tor- clycenaglyceria and Puccipuccinelliapuccinelhanellia amer J bot 36 reybotreynotrey bot club 91 396 405 155 165 77 1983 new nomenclatural combinations in 55 1952 the genus torreyochloa rhodora the noinorthth american perennial tnticeaetriticeae grami- 5419754 197 200 neae Britbrittoniabrittaniabnttonia35tonia 35 30 33 56 CLAY K 1983 variationVanation in the degree of cleis- 78 1983 historical and current taxonomic togamy within and among species of the grass perspectives of agropyron elymus and related danthoniaDanthoma amerarnerarmer J bot 7083570 835 843 geneiagenelagenera crop sci 23 637 642 57 CLAYTONCLAY lonION W D 1968 the correct name for the 79 1984 the genomic system of classifica- common leedreedfeed taxon 17 168 169 phragmites tion as a guide to interintergenericmtergenencgeneriegeneric hybridization within 58 1981 eailyearlyeally sources of tribal names in the perennial tnticeaetriticeae appp 209 280 in J P gramineae kew bull 36 483 485 gustafson ed gene manipulation in plant 59 clCLAYTONaytonAVION W D SMS M philllpsPHILLIPS anuanusANDANDSSAA RENVOIZERENVOIZF improvement plenum press new york 1974 floiafiolaflorafiora of tropical east africa gramineae 80 dewlyDEWEY D R AND K H ASAY 1982 cytogenetic part II11 crown agents foiforholbol overseas governments and taxonomic relationships among three diploid and administratorss london appp 177 450 crestedciested wheatwheatgrassesgrasses crop sci 22 645 650 60 claylonCLAYIONCLAYTON W D ANDANDSS A rlRENVOIZENVOIL 1986 genera agropyron J graminumgiammumgraminum gigrassesasses of the world kew bull additaddia 81 DEWEY D R AND C HSIAO 1983 A cytogenetic ser XIII 389 appp basis for transferringtransfer ing russian wildrye from elymus 61 clCLAUSENAUSI N R T 1952 suggestion foiformolpor the assign- to psathyrostachys crop sci 23 123 126 ment of torreyochloatorrcyochloa to Puccipuccinelliapuccinelhanellia rhodora 54 82 docdoggendoggbndoggerrDOGcerigeriGErr H 1970 sorghum longmans london 42 45 403 appp 199011990 NEW MEXICO GRASS TAXONOMY 79

8381 DOREDOBE W G 1950 persian darnel in canada sci 106 GOULD F W ANDANDRR B SHAW 1983 giasssystemgrass system agrieagne 3015730 157 164 loliumlonum I1 atiesatles ad2d ed texas a&maam university press college 84 ELLIOT F C 1949 bromus inermisinermia and B pumpelpumpejpumper station 397 appp lianusliamisdianus in north america evolution 3 142 149 107 GRANT M C ANDJAND J antonovicksANTO NOVICKS 1978 biology of 85 EMERY W H P 1957a A cytotaxonomic study of ecologically marginal populations of anthoxan- setaria macrostachya glamGiamgramineaemeae and its rela- thum odoratumodoratum I1 phoneticsphenetics and dynamics evo- tives in the southwestern united states and mex- lution 32 822 838 ico bull torrey bot club 849584 95 105 108 GREENE C W 1984 sexual and apomictic reproduction in calamagrostis grammeae 86 1957b A study of repireproductioneduction in setaria in Gramgraminealgramineacgramineaemeae from east-cast- ern macrostachya and its relatives in southwestern eineln north america amer J bot 71 285 293 united states and mexico bull torrey bot club 109 creuterGREUTEKGREUTER W 1968 notulae nomenclaturalnomenclaturaleses et bib 84 106log 121 easternliogiaphicaeliographicae 1 4 candollea 23 81 108 koele 84106 riaizanta 87 ERDMAN K S 1965 taxonomy of the genus ita 110 GROSS A T H 1960 distribution and ecology of sphenopholissphenophohs gramineae iowa state sci 39 J elymus macmunnii vasey can 289 336 macouniimacounmacounn J bot 38 63 67 111 GUZMAN R AND F J SANTANA M 1987 espe 88 D E 1959 morphological las fairbrothers variation ciescles mexicanasmexicanas del genero paspalum L grami- of setaria and S vindisviridisvindzsidis Brit 11 fabert ulfvir bnttoniabrittoniabrittaniatonia neae univ guadalajara folletocolleto tccmco no 1 48 tecnico 44 112 HALPERINHALPERIIS M 1933 the taxonomy and morphol- 89 FARUQI S A B& QURAISH AND H 1979 studies on ogy of bulbous bluegrass poa bullosabulbosabulhosa vtviparaviviparaviviparyuluivivipara libyan grasses V population variabilityvaival lability and dis- J ameiamelamer soc agron 2540825 408413408 413 tribtributionution of schistusschismusSchismus afaarabicsaraarabicusarabtcusbicus andan d S barbatisbarbarbatusbatus in 113 HARLANHABLAN J RK 1945 cleistogamy and chasmogamy pak libya J bot 112 167172167 172171 in bromus caricarinatusnatus hook & arn amer J bot 90 FERNALD M I1 1928 the american and eastern 32 66 72 asiatic BeckBeckmanbeckmanniabeckmanmamanniama rhodorarhodoia3030 24 27 114 HARVEY L 1975 eragrostis appp 177 201 in F W 91 1932 phragmites comcommuniscommoniscommumsmunis trin var gould grasses of texas texas aam university berlandienberberlandieriberlangieriberlanberianlandiendieridierl fournier comb nov rhodora 34 press college station 653 appp 211 212 115 HILL S R 1982 vegetative apomixis vivipary 92 1943 five common rhizomatous species of in boutelouaBoutelona hirotahirwtahirsuta lag poaceae sida 9 muhlenbergia rhodorarhodoia4545 222 239 355 357 116 HITCHCOCK A S 1920 93 FERNALD M I1 ANDANDCC A WEATHERBY 1916 the hitciicock AS the genera ofgrasses of the genus Puccipuccinelliapuccinelltanellia in eastern northnor th americaamenea united states with special reference to the eco- rhodora 18 1 23 nomic species USDA bull 772 117 HITCHCOCK A S AND M A CHASE 1910 94 FINDLAY J N AND B R BAUM 1974 the nomen- the north american of panicum U S claturalclatural implications oftheofodthethe taxonomy Danthoniaofdanthoniaof species conticontr US catlnatl heibherb 15115 1 396 in canada canad J bot 52157352137352 1573 1581 118 1951 manual of the grasses of the united 95 frederiksen S 1979 festuca minutifloraminutiflora rydb redb states ad2d ed UUSDAS DA discmisc publ 200 10511 051 appp a neglected species bot notisernoticer 132 315 318 iosi 119 HOLMGREN P K W KEUKEN AND E K SCHO 96 1982 festuca brarhyphyllabrachyphylla F saximonmaximon FIELD 1981 index herbanorumherbariorumHerbanorum part I1 tana and related north america nord the species in noid herbaria of the world bohn scheltema & hol- J bot 2 525 536 kema utrecht 97 FREEMAN D 1979 lehmann lovegrasslovegiasslonegrassloviovegrass range- 120 hulffnhuljcfcn E 1959 the trisetum spicapicapicatumspicatumtum complex lands 1 162 163 eragrostis sv bot tidskr 53 203 228 98 GABLE M L biosystematic 1984 a study of the 121 HUMPHRIES C J 1978 notes on the genus phleum genus elymus gramineae tnticeaetriticeae in iowa L bot J linn soc 76 337 339 proc iowalowalowaacadacad sci 9 140 146 122 iltis H H 1983 from teosinte to maizemalze the 99 GILLETT J M AND H A sewsegSENN 1960 cytotaxonomy catastrophic sexual transmutation theory science and intraspecificmfraspecificinfraspecificinfraspecific variation of agropyron smithiismithnsmithia 222886222 886 894 zea rydb canad J bot 38 747 760 redb 123 litislitlsILXIS H H annjANDJannAND J F DOEBIEYDOEBLEY 1980 taxonomy of 100 GOULD F W 1947 changes nomenclatural in zea Gramgramineaemeae II11 subspecific categories in the elymus with a key to the california species zea mays complex and a generic synopsis amer Madromadronofioflo 9 120 128 J bot 67 994 1004 101 loilol 1957 new northnortbnordb american andropogonsandiopogons 124 JAUKAR P P 1975 chromosomeChiomo sorne relationships be- of subgenus amphilophusamphtlophus and a key to those spe- tween lolium and festuca gramineaeGram meae chromo cies occurring in the united states madronomadrofioMadromadroniofio 14 soma52soma 52 103 121 182918 29 bothriochloabothnochloa J 125 JOHNSON B L 1945 natural hybrids between ory- grasses 102 1975 of texas texas a&maam uni- zopsis hymenoideshyme noides and several of sttpastapa press species stipa veiversityver sity college station 653 appp amer J bot 325993959932 599 608 103 1979 the genus bouteloua ann missouri 126 JOHNSON B L D BARNHARTBARNHARF AND 0 HALL 1967 gard 66 348 bot card 416 analysis of genome and species relationships in 104 COULDGOULD F W M A ALI AND D E fairbrothers the polyploid cheatswheats by protein electrophoresis 1972 A revision of echtnochloaechinochloa in the united amer J bot 54108954 1089 1098 7 fitrittriticumriticumicum states amer midi naturalist 87 36 59 127 JOHNSON B L AND H S DHALIWAL 1976 repro- 105 GOULD F W ANDANDCC A CLARK 1978 dichanthe ductive isolation of triticum boeoticumboeotic um and poaceae triti- liumhuinilum of the united states and canada cum urartu and the origin01 iamigm oftheodtheofthe tetraploid cheatswheats ann missouri bot gard 65 1088 1132 ameiamelamer J bot 63108863 1088 1094 80 K W ALLRED volume 50

128 JONES E K ANDANDNN C FASSFIT 1950 subspecific 150 MARTIN WWCC ANDANDCC R HUTCHINS 1980 A flora of variation in sporobolus cryptcryptandrusandrus rhodora 52 new mexico vol 1 vaduz west germany 125 126 J cramer l276pp1276 appp 129 KARLSSON T 1987 setaria adhaerans and S fabenpaben 151 MELDERIS A 1978 taxonomic notes on the tribe new to sweden svensk bot tidskr 81 305 311 tnticeaetriticeae gramineae with special reference to 130 KAWANO W 1963 cytogeography and evolution of the genera elymus L sensugensu lato and agropyron the deschampsia caespitosa complex can J bot gaertner sensugensu lato bot J linn soc 76 41 719 742 369 384 131 1965 calamagrostis purpurascens R br 152 MITCHELL W W 1967 on the hordeum juba and its identity acta phytotax geolotgeobot 21 tum H brachyantheruinbrachyantherum question Madromadronofio 19 73 89 108 110 132 KELLOGG E 1985a A biosystematic study of the 153 MITCHELL W W AND A C WILSON 1964 the poa secunda complex J arnold arbor 66 hordeumjubatumhordeum jubatum caespitosum brachyantherumbrachyantheruin 201 242 complex in alaska Madmadronoroilo 17 269 280 133 1985b variation and names in the poa 154 MOBBERLEY D G 1956 taxonomy and distribution secunda complex J range management 38 of the genus spartina iowa state coll J sci 30 516 521 471 574 134 KOCH S D 1974 the eragrostis pectinacea pilosa 155 MORDENMOKDEN C W AND S L HATCH 1986 vegetative complex in north and central america grami- apomixis in muhlenbergiamuhlenbergw rapensrepens poaceae era neae eragrostoideae illinois biolabiol monsgrmonogr 48 grostideaegrostideac sidallsida 11 282 285 1 74 156 1987 anatomical study of the muhlenber- 135 1979 the relationships of three mexican gia rapensrepens complex poaceae chlondoideaechloridoideae Aveneaveneaeavenearae and some new characters for distin- eragrostideaeeragroshdeae sida 12 347 359 guiguishingshing deschampsia and trisetum taxon 28 157 MORROW L A AND P W STAHLMAN 1984 the 225235225 235 history and distribution ofdownyofdowny brome bromus 136 KOCH S D AND I1 SANCHEZ V 1985 eragrostis tectoriumtectorumtectorum in north america weed sci 32 mexicansmexicanamexicana E neomexicana E orcuttianaorcuttiana and E suppi 2 6 virescentvirescensmrescensolfesvirescens the resolution of a taxonomic problem 158 NELSON EENN AND R J tyriTYBITYRL 1978 hybridization phytologiaphytologicphytologia58Phytol ogiaogla 58 377381377 381 and introgression between elymus canadensis and 137 KOYAMA T AND S KAWANO 1964 critical taxa of elymus virginicusvirginiousvirgzmcusvirginicus poaceae proc okla acad grasses with north american and eastern asiatic sci 583258 32 34 distribution can J bot 42 859 884 schiaschizschtz 159 nomenclature COMMITTEE laftIAPTJAPT 1983 report achneacene on proposal 528 rejection of paspalum distichumdistichum 138 LAZARIDES M 1972 A revision ofaustralianofaustralian chlo L gramineae taxon 32 281 name stands ndeaerideaerideau gramineae australian J bot suppi ser 160 PAVLICK L E 1983a notes on the taxonomy and suppi no 5 chlortsChchloreschlorisllorts nomenclature of festuca occidentoccidentalisoccidentalistalis and F ida 139 LONARD R I1 AND F W COULDGOULD 1974 the north hoensis can J bot 61 337337344344 american species of vulpicvulpia gramineae 161 1983b the taxonomy and distribution of Madromadronofionionno 22 217 230 festuca idahoensistdahoensis in british columbia and north- 140 louismarieloulsLOUIS maniemanikMARIE 0 C 1928 the genus trisetum in western washington can J bot 61 345 353 america rhodora 30 209 228228231231 245 162 1984 studies on the festuca ouinaovina com- 141 lush W M 1989 adaptation and differentiation of plex in the canadian cordillera can J bot 62 golf course populations of annual bluegrass poa 2448 2462 anduaannua weed sci 3754-37 545954 59 163 1985 A new taxonomic survey of the 142 mcgregor R L 1984a panicum capcamillarecapillarecapzllareillare L var festuca rubraaubra complex in northwestern north occidentale redbrydb poaceae an illegitimate name america with emphasis on british columbia contr univ kansas herb no 10 phytologiaphytologicphytologia57Phytologiaogla 57 1 17 143 1984b panicum camillarecapcapillareillare L var barbtpulbarbipul 164 PERRY G AND J MCNEILL 1986 the nomencla- umatum nash comb nov phytologiaphytologicPhytol ogia 554 256 ture of eragrostis cilianensis poaceae and the 144 1984c panicum dichotomiflorum poaceae contribution of bellardi to alliomallioniallianialbiomAllioni s flora pede variation in kansas including notes on the sterile montana taxon 35 696 701 palea contr univ kansas herb no 8 3 p 165 PHIIIPSONPHILIPSON W R 1937 A ierevisionvision of the british 145 1985 panicum billmannhillmannbiilmannhillmaniinil chase poaceae species of the genus agrostis J linn soc bot vailidity and distribution contr univ kansas 51 73 151 herb no 13 99ppappappp 166 PINTO ESCOBAR P 1976 nota sobrebobre el ejamplar tipocipo 146 MCNEILL J 1976 nomenclature of four perennial de bromus catharticuscatharticusvamvahlvahi caldasiacandasiacaldasiallcaiCaldasia 11 9 16 species of bromus in eastern north american 167 POHL R W 1962a agropyron hybrids and the with a proposal for the listing of B burganspurgans L as status of agropyron pseudorepens rhodora a rejected name under article 69 taxon 25 6414364 143 147 611 616 168 1962b notes on setariabetanasetana vindis and 147 1979 Dipdiplachnelachne and leptochloaleptochroaLeptochloa poaceae S fabenpaben gramineae Britbnttoniabrittoniabrittaniatonia 14 210 213 in north america Britbrittaniabrittoniabnttonia31tonia 31 399 404 169 1969 muhlenbergia subgenus muhlenber- 148 MALIK C P 1967 hybridization offestucaoffestuca species gia gramineae in north america amer eidlmidimidl can J bot 45 1025 1029 naturalist 82 512 542 149 MARSH V L 1952 A taxonomic revision of the 170 POHL R W ANDANDWW W MITCHEILMITCHELL 1965 cytogeog genus poa of united states and southern canada raphy of the rhizomatous american species of amer midi naturalist 47 202 250 muhlenbergia bnttoniabrittoniabrittaniaBrittonia 1710717 107 112 1990 NEW MEXICO grassfaxonomyGRASS TAXONOMY 81

171 POZARNSKY T 1983 buffalograssBuffalograss home on the 196 SNYDER L A 1950 morphological variability and range but also a turf grass rangelands 5 214 hybrid development in elymus glaucus amer J 21621qbuchloebuchlbuchabuchw bot 37 628 636 172 PYRAH G L 1969 taxonomic and distributional 197 soderstromsouersSODERS tromIROM T R 1967 taxonomic study of sub- studies gramineae in leersia iowa st J sci 44 genus PodospodosemumpodoseinumPodose einummum and section epicampesEpicampes ofmuhofmuh 215 270970 215270 lenlenbergiabergia gramineae contr USU S nati 173 RANDALL herb J L AND K W HILU 1986 biosystematic 34 75 189 studies of north american trisetum spicaspicatumtum 198 soderstrom T R AND J H BEAMAN 1968 the poaceae systcyst bot 11 567 578 genus broinusgroinus gramineae in mexico and central 174 REEDERREEDEK C CG 1949 muhlenbergia mznuttsstma Muhlenbergta minutissima america mich publ mus biolabiol ser 3 steudel swallen and allies st univ its J wash acad sci 465520465 520 3936339 363 367 199 soderstrom T R K W HILU C S CAMPBELL 175 1985 the genus lycurus gramineae in AND M E BABKWORTHBARKWORTH 1987 glassgrass north americaamenea Phytolphytologiaphytologicphytologia57ogiaogla 57 283 291 systematics and evolution smithsonian press 176 REEDERREEDEB J R 1951 setariabetanasetana lutescentluteslutescenscens an untenable washington D C 473 name rhodora5353 27 30 DC appp rhodora 200 SOHNS E R 1956 177 1953 affinities of the grass genus beck the genus hilanadilanahilaria gramineae wash nanniamannwrostnwnniamannwRosthost bull torreytoireybotbot club 80 187 196 J acad sci 46 311 321 201 SORENG R J 1985 poa new 178 1976 systematic position of Redfieldredfieldiaredfleldtaia L in mexico with gramineae madronoMadro fionionno 23 434 438 a key to middle and southernsoutheinthern rocky mountain poaceae 179 1986 another look at eragrostiseragrostzs tephrozephro species great basin naturalist 45 santhosgramineaesanthosmanthos gramineae phytologia60phytologia60153 153 154 395 422 180 REEDERRFEDER J R ANDCAND C G REEDER 1980 systematics 202 SORENG R J ANDSAND S L HATCH 1983 AAcomcomparisonpanson poa poa of bouteloua brevisetabreviseta and B famosa cramigiamigrami of traceitracyi and occidentoccidentalisoccidentalistalis sida 10 leaeneae systcyst bot 5 312 321 123 141 181 REEDER J R AND J TOOLIN 1987 scleropogon 203 STACI- C A 1987 Triticatnticaletriticalele a case ofnomenclatural gramineae a monotypic genus with disjunct dis- mistreatment taxontaxon3636 445 452 tnticosecaletriticosecaleTriticosecale 1 tribtributionution phytologiaphytologicphytologia62Phytologiaogla 62 267 275 204 STANDLEY P C 1910a the type localities of plants 182 REEDER J R AND L J TOOLIN 1989 notes on fustfirst described from new mexico contr USU S pappophorum gramineae pappophoieaepappophoreae systcyst nati herb 1314313 143 227 bot 14 349 358 205 1910b A bibliography of new mexico 183 RIGGINsBIGGINS R 1977 A biosystematic study of the botany contr USU S nati herb 1322913 229 246 sporobolus asper complex gramineae iowa 206 STEBBINS G L JR 1930 A revision of some north state J res 51 287 321 american species of calamagrostis rhodora 32 184 ROMINGER J A 1962 taxonomy of setaria 35 57 gramineae in north america illinois biolabiol 207 1947 the origin ofthe complex ofbromus monsgrmonogr 29 1 132 caricarlcarinatuscartcartnatusnatus and its phytogeographic implications 185 SAKAR P 1956 crested wheatgrasswheatgrass complex can contr gray herb 165 42 55 J bot 34 328 345 agropyron 1 208 1981 chromosomesChromosomes and evolution in the 186 SANCHEZ E 1983 dasyochloa willdenowWilldenow ex rydaydbyd genus bromus gramineae bot jahrbjahob systcyst 102 berg poaceae genero monotfpico de northamnorteam 359 379 enca lilloalillealilloa3636 131 138138j5nonewonerloerioneuronErioneuron 209 SIFBBINSSTEBBINS G L JBJR AND D ZOHARY 1959 cytoge- 187 SAUER J D revision of stenotaphrum gramineae netic and evolutionary studies in the genus Panicepaniceaeae with attention to its historical geogra- dactylis univ calif publ bot 31131 1401 40 phy Britbrittaniabrittoniabnttoma24tonia 24 202222202 222 210 STEUDEL J 1854 synopsis Planplantaiumplantarumplantariumpian tarum glumaceaglumaceae 188 SEARS P B 1977 sleepy grass rangeman s journal i HOTrum pars I1 gramineae Stuttstuttgartegarte 43 67 68 stipa robusta 1 211 stutzSIUIZS IU IZ H C 1972 on the origin of cultivated rye 189 SHAW E A 1987 charles wright on the boundary ameiamelamer J bot 59 59 70 secale 1849184918521852 or plantae wnghtianaewrightianae revisited 212 SWALLEN J R 1928 the grass genus schizachne mecklermeckiermockler publishing corp westport connecti- J wash acad sci 18 203 206 cut 44 appp 213 1931 Crascrassipessipes a new grass genus from 190 SHAW R B AND R D WEBSTER 1987 the genus utah amer J bot 18 684 685 sclerochloa Erioeriochloachloa poaceae Panicepaniceaeae in north and cen- 214 1948 agrostis varlavariawnabihsvariabilisbilis redbrydb a valid tral america Ssidallsidal2idalda 12 165 207 species lean W bot 5123531231251231255123125123 125 191 SHEAR C L 1900 A revision of the north american 215 TATEOKA T 1961 abiosystematicstudyoftndensA biosystematic study of tridensteidens species of bromus occurring north of mexico gramineae amerarner J bot 4856548 565 573 U S D A div agrostolagro&tol bull 23 1 66 216 TAYIORTAYLOR D R ANDanulANDLL W AARSSENAABSSFN 1988 an iteimteiinter- 192 SHINNERS L H 1954 notes on north texas grasses piepretationpre tation of phenotypic plasticity in agropyron rhodora 56 31 32 nomenclature ofkoeleriaofkoelenaKoeleria repensrapens gramineae amer J bot 7540175 401 413 193 1956 illegitimacy of persoon s species of 217 TERRELLTEBBELL E E 1967 meadow fescue festuca ela Kokoelenakoeleriajoelenaeleria gramineae rhodora 58 93 96 tiortwrbior L or F pratensispratensis hudson bnttoniabrittoniabrittaniaBrittonia 19 194 SIEBER V K AND B G MURRAY 1979 the cytology 129 132 of the genus alopecurus gramineae J linn 218 1966 taxonomic implications of genetics soc bot 79 343 355 in ryegiryogiryeryegrassesgrassesasses lolium bot rev 32 138 164 195 SMITH P 1970 taxonomy and nomenclature of the 219 1968a notes on festuca arundinaceaarundtnacea and brome grasses notes roy bot caidcardgaidgard edinburgh F pratensispratensis in the united states rhodora 70 30 361 376 bromus 564 568 82 K W ALLRED volume 50

220 1968b A taxonomic revision of genus lol- 233 WIDEN K 1971 the genus agrostis L in eastern ium U S D A A R S tech bull 1392 fennoscandiaFennoscandia taxonomy and distribution 221 fl 1976 the correct names for pearl millet fennicabennicafennica55 1 209 and yellow foxtail taxon 25 297 taxon25 304 setaria 234 WILSON F D 1963 revision ofsitanionsifsttanwnSitanionanlon tnticeaetriticeae 222 THIERET J W 1966 synopsis of the genus calam gramineae bnttoniabrittoniabrittaniaBrittonia 15 303 323 ovilfaovilda Gramgramineaemeae castaneacastanea3131 145 152 235 witherspoon J T 1975 A numerical 223 VELDKAMP J F 1984 the identity of Andropogon taxonomic study of the eragrostis intermediaintermedialmedia hutansnutans linnaeus gramineaeGrammeae taxon 339533 95 97 inter complex unpublished dissertation of montana 224 WAGNONWACNON H K 1952 A revision of the genus university missoula 483 pp bromus section bromopsisBromopsis of north america ap brittoniabrittaniabnttonia7Brittonia 7 415 480 236 WOOTON E 0 1903 range grasses ofnerofnewof new mexico 225 WALLERWALLEB F RK 1976 Aabiosystematicstudyofpawbiosystematic study of panipanl rptapt gov new mexico 1902 232 234 cum section diffusediffusa poaceae in north america 237 WOOWNWOOTON E 0 AND P C STANDLEY 1912 the unpublished dissertation texas a&maam univer- grasses and grasslike plants of new mexico new sity college station 123 appp mexico agrieagne eapexp sta bull 81 175 appp 226 WARWICK S I1 B K THOMPSON ANDLAND L D BLACK 238 1913 descriptions of new plants prelimi- 1984 population variationvaivar lation in sorghum halehalepensepense nary to a report upon the flora of new mexico johnson grass at the northern limits of its range contr USU S nati herb 1610916 109log 196 can J bot 62 1780 1790 239 1915 flora of newnow mexico contr USU S 227 WATSON L H T CLIFFORD AND M DALLWITZ J nati herb 19119igi 1 794 1985 the classification of poaceae subsubfamiliesfamilies 240 ZULOACOZULOAGO F AND T R soderstrom 1985 classifi- and supersupertnbessupertribestribes australian J bot 33 433 484 cation of the outlying species of new world 228 WEBSTERWFBSTER R D 1987 taxonomy Digitariaofdigitariaof section panicum poaceae paniceaeae smithsonian Didigitariagitana in north america poaceae paniceaeae Panice contr Panice 59 sida 12 209 222 bot 229 1988 genera of the north american pan 241 BUSEY P 1986 morphological identification of icealiceae poaceae panicoideaepanicoidede systcyst bot 13 st augustinegrassAugustinegrass cultcultivarsivars crop sci 26 28 32 576 609 stenoptaphrum 230 WEIMARKWEIMABK G 1971 variation and taxonomy of hie 242 FELBERFELBEB F 1988 phenology of flowering of diploid rochloerochlee Gramgramineaemeae in the northern hemi- and tetraploid populations of anthoxanthum sphere bot not 124 129 175 alpinumalptnum and anthoxanthum odoratumodoratum can J 231 WELSH S L 1982 utah plant types historical per- botboc 66 2258 2264 spectivespective 1840 to 19819811 annotated list and bibliog- 243 MIZIANTY M 1986 biosystematic studies on raphy great basin naturalist 42 129 195 dactylsdactyhsdactylis L 1 review of previous studies acta 232 WESTERN REGIONAL TECHNICAL COMMITTEE W 90 soc bot pol 55 467467479479 1972 galleta taxonomy ecology and management of hilanadilanahilaria jamesiijamesitjamesia on western rangelandsi angelands received 21 september 1989 utahutahagragr eapexp sta bull 487 38 appp accepted 25 january25january 1990 great basin naturalist 501 1990 appp 83 84

noteworthy MAMMAL distribution RECORDS FOR THE NEVADA TEST SITE

philip A medical

previous reports on the mammals of the 1934 and Arlemont esmeralda county ne- nevada test site nye county nevada jor- vada 1478 in fish lake valley hall 1951 gensen and hayward 1965 ofarrell and 290 which is 210 km northwest of rock emery 1976 indicate the presence of 46 spe- valley cies 42 terrestrial mammals and 4 bats the juvenile female specimen collected under a new project entitled basic envi- in rock valley 3641n 11611w ronronmentalmental compliance and monitoring pro- NSMLV M 9202 measured 3191103217319 110 32 17 gram at the nevada test site two previously and weighed 91 g it is important to note that uncollected species of mammals were ob- rock valley lies at the northern edge of the tained and a range extension for a third spe- specter range along the southern boundary cies was documented during the 1988 sam- of the nevada test site the vegetation con- pling season voucher specimens have been sists primarily of larrea tridentata lycium deposited at the nevada state museum in las andersoandersoniinii lycium pallipallidumdum and ambrosia vegas nevada dubosadumosa romney et al 1973 this region is mustela frenatafredata nevadensis hall small located in an area that contains no above- animal trapping was conducted on three con- ground permanent water source the nearest secutive nights in rock valley nye county source is a tinaja a small rock crevice with nevada elevation 1035 in 14 16 june an opening approximately 20 cm that stores 1988 on 15 june 1988 two M f nevadensis water collected from runoff it is approxi- an adult female and a juvenile female were mately 5 km west of the locality in which the captured in sherman traps this record ex- M f nevadensis were collected tends the known range on the nevada test urocyon cinereoargenteuseinereoargenteus scottiiscottie mearns site approximately 58 kinkm south of the previ- one specimen was collected dead on ous known locality prior records reported holmes road 131.3 kinkmkim northwest of the junc- M jf nevadensis from the vicinity just south of tion of holmes road and stockade wash whiterock spring in northwestern yucca flat road nevada test site nye county nevada in the crayla lycium community jorgensen 3710n 11613w116013w at 0700 hr on 2 au- and hayward 1965 the male M f nevadannevaden gust 1988 this immature female specimen sis 4298 BYU cited by jorgensen and hay- NSMLV M 9203 measured 72731012070727 310 120 70 ward 1965 was collected on 15 may 1961 and weighed 1733 g this location is at the dead on the road and measured 38613041386 130 41 southern end of rainier mesa at an elevation no ear measurement no skull jorgensen of 1993 in and at the lower edge of pinyonpinyon personal communication I1 observed an addi- juniper habitat hall 1946241 cited a trap- tional juvenile live specimen in early april per capturing gray fox at 2194 in 898.9 kinkm 1987 approximately 4 kinkm southeast of white- northwest of whiterock spring during the rock spring it was later released additional winter of 1930 31 in pinyon timber this published records for M f nevadensis exist locality was most likely on the top of rai- for the spring mountains clark county ne- nier mesa or aqueduct mesa in the belted vada 50km50 km southeast of rock valley where range on the nevada test site an addition- weasels were observed but not collected burt al sighting was recorded on kahutepahute mesa by

reynolds electrical & engineering co inc BECAMP box 495 mercury nevada 89023

83 84 NOTES volume 50

D badger on 29 september 1986 343.4 kmkni basic environmental compliance and moni- north of silent canyon 3719n IIGW toring program I1 thank D badger L bad- along dead horse flats road nevada test ger P hopkin D lopez and M saethresaethreforforborror site at an elevation of 2102 in assistance in the field C jorgensen and C this specimen NSMLV M 9203 verifies pritchett for data pertaining to the specimens the hall 1946 record and substantiates jor- at brigham young university R B hunter gensen and haywardHaywardss 1965 contention that and B maza for reviewing this manuscript this species would eventually be collected on and M J ofarrell for confirming the identi- the nevada test site which falls within the ficationfication of all specimens reported I1 also thank distribution illustrated by hall 1981 the nevada department of wildlife for the microdipodops megmegacephalusacephalus sabulonissabusabukonislonis necessary scientific collecting permit hall three specimens were captured on fahutepahrtepahute mesa 030.30 3 km north of the junction of kahutepahute mesa road and buckboard mesa literature CITED road 37 15 N 116 28 W at an elevation of BRADLEY W G AND K S MOOR 1975 ecological studies 1919 in one on 29 june 1988 and two on 19 of small vertebrates in pu contaminated study ar- august 1988 the two august specimens eas of NTS and TTR pages 151 185 in M G were preserved and provide the first voucher white and P B dunaway eds the radioecologyradioecology of plutonium and other transuranics in desert en- specimens for the nevada test site nye vironvironmentsments nevada applied ecology groupen county nevada these specimens measured ergy research & development administration NSMLV M 9200 male 147752510147 75 25 10 12 g report NVO 153 BUBI NSMLV M 9201 female 150 75 24 10 13 burtburlburi W H 1934 the mammals of southern nevada 150752410 san g transactions of the diego society of natural jorgensen and hayward 1965 reported history 7 375 428 collecting three specimens north of the HALL E R 1946 mammals of nevada university of nevada test site in kawich valley BYU california pipressess berkeley 710 appp 4485 4486 4487 two additional records 1951 american weasels university of kansas of publications M m sabusabulonissabukonislonis exist from the neihsnellis bomb- museum of natural history 4 and gunnery 14661 466 ing range northeast of the ne- 1981 the mammals ofnorth america johnwileyjohn wiley vada test site one specimen BYU 4097 was & sons new york 2 vols 11811 181 appp collected 7 km north of the northeast JORCENSENJORGENSEN C D ANDANDCC L HAYWARD 1965 mammals of boundary of the nevada test site and the the nevada test site brigham young university science bulletin 6 1 8811 second record was reported by bradley and OFARRELL T P ANDANDLL A EMERY 1976 ecology of the moor 1975 from 8 kinkm north of the northeast nevada test site a narrativenanative summary and anno- boundary of the nevada test site the two tated bibliography U S energy research devel- specimens of M m sabusabulonissabukonislonis reported here opment administration report NVO 167 249 NSMLV M 9200 NSMLV M 9201 PP extend ROMNEY E M V HALE known Q A WALLACE 0 R LUNT the distribution approximately 32232.2 J D CHILDRESS H KAAZ G V ALEXANDER kmkin southwest of kawich valley and on to J E KINNEAR AND T L ACKERMAN 1973 some kahutepahute mesa on the nevada test site characteristics of soil and perennial vegetation in northern mojave desert areas of the nevada test site university ofcalifornia los angeles report acknowledgments 1291612 916 340 appp

this study was facilitated by funding from received 10 august 1989 the US department of energy through the accepted 2 november 1989 great basin naturalist soisol501soo 1990 appp 85 87

FORMATION OF pisolithus tinctoriustinctoriousTINCTORIUS ectomycorrhizae ON california WHITE FIR IN AN EASTERN SIERRA NEVADA MINESOIL

R F walkerwaiker

the gasteromycete pisolithus tinctoriustincttinctoriousorius field and tueller 1980 vegetation is sparse pers coker & couch occurs in temperate on most of the spoils but in addition to the subtropical and tropical zones worldwide and two pine species mentioned previously cali- in ectomycorrhizal association with numerous fornia white fir abies concolor var lodianalowiana conifer and hardwood hosts marx 1977 fre- gord lemm singleleafsingleleaf pinyon pinus quent reports of the occurrence of its basidio monomonophyllaphylla torr afrem&frem& fremprem utah juniper ju- carps near various pine species on harsh sites niperus osteospennaosteosperma torr I1 little and quak- in the eastern united states lampky and ing aspen populus tremultremuloidesoides michamichx have peterson 1963 schramm 1966 hile and hen- become reestablished on the periphery of the nen 1969 lampky and lampky 1973 marx mine near adjoining undisturbed forest and 1975 medve et al 1977 have prompted woodland walker s 1989 report concerning extensive efforts to inoculate seedlings in examinations made in september 1988 of the forest nurseries with this mycobiont marx probable hosts of P tinctoriustincttinctoriousorius in leviathan etetalal 1976 1982 1984 1989a 1989b subse- mine noted that basidiocarpsbasidiocarps of this symbiont quentlyquently the improved survival and growth of were absent in the immediate vicinity of the pine seedlings with P tinctoriustincttinctoriousorius ectomycor latter four tree species rhizae after plantingoutoutplanting on surface mines has reexamination of leviathan mine spoils in been attributed to enhanced uptake of nutri- august and september 1989 however re- ents marx and artman 1979 and water vealed numerous P tincttinctoriustinctoriousorius basidiobasidiocarpscarps walker et al 1989 current research is near seedlings and saplings ofcalifornia white focused on development of more effective in fir typically one or two dark yellow to brown oculacocula and inoculation procedures discovery basidiocarpsbasidiocarps fig 1aaa matching the descrip- of locally adapted P tincttinctoriustinctoriousorius isolates and tion of coker and couch 1928 were ob- identification of new host species served around solitary white fir seedlings A recent report walker 1989 disclosed while as many as five encircled individual P tinctoriustincttinctoriousorius occurring in ectomycorrhizal white fir saplings stipitate substipitatesubstipitate and association with jeffrey pine pinus jeffrejeffreyyi sessile forms were observed grev varying in size & balf and sierra lodgepole pine from 9 to 17 cm in length and from 3 to 7 cm in pinus confortacontortacontorta var murmurrayanarayana grev & diameter approximately 100 basidiocarpsbasidiocarps balf I1 engelm on spoils of the leviathan were found associated with white fir and mine in alpine county california located these were rarely more than 2 in from the on the eastern slope of the sierra nevada host 384230n 1193915w at an elevation of strands of mycelia with gold yellow pig- 2200 in and consisting of approximately 100 mentation which compare favorably with ha this open pit sulfur mine has been inactive the P tinctoriustincttinctoriousorius rhizomorphs described by since 1962 the average annual precipitation schramm 1966 were traced through the of approximately 50 cm is primarily snowfall spoils from basidiocarpsbasidiocarps to the root systems and the mineminesoilsoil has a ph of 404.04 0 to 454.5 a of white fir seedlings and saplings these deficiency of plant available N and a poten- mycelia were connected to ectomycorrhizaeectomycorrbizae tially phytotoxic concentration of al butter of similar pigmentation that matched the

university of nevada reno department ofrangeoforrangebangerange wildlife and forestry 1000valie1000 valleyvaileyvallevaileValIe road reno nevada 89512

85 86 NOTES volume 50

figpig 1 pisolithus tincttinctoriustinctoriousorius associated with california white fir on an eastern sierra nevada mine spoils A basidiocarpsbasidio carps bar represents 5 cm B ectomycorrhizae on roots bar represents 1 em description of those formed by P tincttinctoriustinctoriousorius fig ab1b or the fungal mantle characteristic of reported by marx and bryan 1975 examina- these mycorrhizalmycorrhizae tion of the complete root system of an isolated an earlier attempt to inoculate white fir white fir seedling with a single associated ba seedlings at plantingoutplantingout with P tinctoriustincttinctoriousorius ba sidiocarpsidiocarp revealed numerous P tinctoriustincttinctoriousorius ec sidiosporessidiospores was largely unsuccessful alvarez tomycorrhizae with approximately 35 of the and trappe 1983 the evidence presented roots exhibiting the bifurcate or coralloid form here however indicates that this host and 1990 NOTES 87

symbiont association occurs naturally in the of pine seedlings on acid coal spoils in kentucky sierra nevada efforts to monitor the mycor- and virginia reclamation review 2 23 31 MARX D BRYAN rhizal development of sierra nevada and fl AND W C 1975 growth and ectoeato in- mycorrhizal development of loblolly pine seed- termountaintermountain woody flora will continue in or- lings in fumigated soil infested with the fungal der to further ascertain the host ranges ofthis symbiont pisolithuspisohthus tinctoriustinctonustinctinctorioustinctoriusorlustonus forest science 21 and other ectomycorrhizal fungi 245 254 MARX D H W C BRYAN AND C E CORDELL 1976 growth and ectomycorrhizal development ofpineorpineofpine acknowledgments seedlings inm nursery soils infested with the fungal symbiont pisolithuspisohthus tinctonustinctoriustinctinctorioustinctoriusorlustonus forest science 22 91 100 this paper contains results of the nevada MARX agricultural D H C E CORDELL D S KENNEY J G MEXAL experiment station research J D ARTMAN J W RIFFLE AND R J MOLINA project 612 funded by the mcintire S tennisstennis 1984 commercial vegetative inoculum of piso cooperative forestry research program the hthusfithusaithus tinctonustinctoriustinctinctorioustinctoriusorlustonus and inoculation techniques for author is indebted to P M murphy of the development ofectomycorrhizae on bare root tree seedlings division of forestry nevada of forest science monograph 25 101 appp department MARX D H C E CORDELL S B MAUL AND J L conservation and natural resources and to RUEHLE 1989a ectomycorrhizal development on D C prusso of the department of biology pine by pisolithuspisohthus tinctoriustincttinctoriousorius min bare root and con- university of nevada reno for their invalu- tainer seedling nurseries I1 efficacy of various able vegetative inoculum formulations new forests 3 assistance 45 56 1989b ectomycorrhizal development on pine by pisolithuspisohthus tinctonustinctoriustinctinctorioustinctoriusorlustonus in bare root and container CITED literature seedling nurseries II11 efficacy of various vegetative and spore meculamoculainocula new forests 3 57 66 ALVAREZ 1 I F AND J M TRAPPETBAPPE 1983 dusting roots of MARX D H J L RUEHLE D S KENNEY C E CORDELL abies concolor and other conifers with pisolithuspisohthus J W RIFFLE R J MOLINA W H PAWUK tinctoriustinctonustinctinctorioustincttonus spores at outoutplantmgoutplanting oriusorlusoflus planting time proves inef- S NAVRATIL R W TINUS AND 0 C GOODWIN fective canadian journal of forest research 13 1982 commercial vegetative inoculum of piso 1021 1023 lithushthusaithus tinctoriustincttinctoriousorius and inoculation techniques for butterfield R I1 AND P T TUELLEBTUELLER 1980 revegeta- development of ectomycorrhizae on container tion potential ofacidoxacidof acid mine wastes in northeastern grown treeti ee seedlings forest science 28 373 400 california reclamation review 3 21 31 MEDVE R J F M HOFFMAN ANDTAND T W GAITHER 1977 COKER W C AND J N COUCH 1928 the gastero- the effects of mycorrhizal forming amendments mycetes oftleoftbeofthe eastern united states and canada on the revegetation of bituminous stripstriplinestripminestnpmmemine university of north carolina press chapel hill spoils bulletin of the torrey botanical club 104 201 appp 218 225 HILE N AND J F HENNEN 1969 in vitro culture of SCHRAMM J R 1966 plant colonization studies on black pisolithuspisohthus tinctoriustinctonustinctinctorioustinctoriusorlusoflustonus mycelium Mycmycologicmycologiaologia 61 wastes from anthracite mining in pennsylvania 195 198 transactions of the american philosophical society 56 1 194 LAMPKY J R AND J H PETERSON 1963 pisolithuspisohthus tincto WALKER R F 1989 pisolithusptsohthus gastero- husriusnus associated with pines in missouri Mycmycologicmycologiaologia tinctonustinctoriustinctinctorioustinctoriusorlustonus a 5567555 675 678 mycete associated with jeffrey and sierra lodgepole pine on acid spoils in the sierra nevada LAMPKY S A AND J R LAMPKY 1973 pisolithuspisohthus in cen- mine in in great basin naturalist 49 111 112 tral florida Mycmycologiamycologicologia 65 1210 1212 WAIKERWALKER R F D C WEST S B mclaughlin AND MARX D H 1975 mycorrhizalmycorrhizae and establishment of C C AMUNDSEN 1989 growth xylem pressure trees on strip mined land ohio journal of science potential and nutrient absorption of loblolly pine 7528875 288 297 on a reclaimed surface mine as affected by an 1977 tree host range and world distribution of induced pisolithuspisohthus tinctoriustmctonustinctorioustinctoriusorlus infection forest the ectomycorrhizae fungus pisohthuspisolithus tinctonustinctoriustinctinctorioustinctoriusorlusoflustonus science 35 569 581 canadian journal of microbiology 23 217 223 MARX D H AND J D ARTMAN 1979 pisolithuspisohthus tincto received 14 october 1989 busriusnus ectomycorrhizae improve survival and growth accepted 15 november 1989 great basin naturalist solsoisot501 1990 p 89

BONE CHEWING BY ROCKY MOUNTAIN BIGHORN SHEEP

K A keating 1

bone chewing has not to my knowledge female ratios long suckling times male matu- been reported in wild north american ration rates and low concentrations of lung- bovidsbovins herein I1 describe an instance of bone worm protostrongylusProtostrongylus sppapp larvae in bighorn chewing by a rocky mountain bighorn sheep feces were all indicative of a high quality ovis canadensis an adult female consumed expanding population keating 1982 this two bones on mt evert s winter range in suggests that bone chewing was not a result of yellowstone national park during winter general nutritional deficiency in the popula- 1980 81 the bones apparently were from a tion though deficiencies in individual animals small ungulate probably bighorn sheep or in specific dietary components cannot be mule deer or prongpronghornhornborn antelope the first discounted bone was consumed in 5 10 minutes I1 inter- rupted consumption of the second sekulic acknowledgments and estes 1977 report that sable antelope frequently spend I1 hr chewing a bone and this research was supported by the rob described a case of a yearling male chewing on and bessie welder wildlife foundation the the same bone for 555.5 hr the rapid consump- national rifle association the montana de- tion observed here was likely related to the partmentpartment of fish wildlife and parks and weathered brittle condition of the bones yellowstone national park sekulic and estes 1977 also report at least two instances in which bone possession by literature CITED sable antelope led to aggressive displacement GEIST V 1971 mountain sheep a study in behavior and of younger animals by adult females during evolution university of chicago press chicago my observations several other bighorn sheep 383 appp continued to feed nearby but showed no inter- KEATING K A 1982 population ecology ofrocky moun- est in the bone chewing was tain bighorn sheep in the upper yellowstone river bone observed drainage montanawyomingMontana Wyoming during second unpublished the- the of two unusually mild win- sis montana state university bozeman 79 appp ters in which forage was generally free of SEKULIC R ANDANDRR D ESTES 1977 A note on bone chew- snow and range use by elk was minimal in- ing in the sable antelope in kenya mammalia 41 dices of population quality are believed to 537 539 reflect the nutritional status of a population received 1 september 1988 geist 1971 in this study high youngadultyoung adult accepted 17 october 1989

glacier national park science center west glacier montana 59936 REDUCED PRICES ON BACK ISSUES great basin naturalist and great basin naturalist memoirs

the prices listed below are min effect through december 1990

GREAT BASIN naturalist regularly 12 per back issue the great basin naturalist is published quarterly volume 1 1939 through volume 34 1974 4 per issue or 10 per volume volume 35 1975 through volume 48 1988 5 per issue or 15 per volume complete sets A few complete sets are available for information contact james R bamesbarnes editor 801 3785053378 5053

GREAT BASIN naturalist MEMOIRS selected memoirs 50 off the prices listed below no I1 the birds of utah 10 no 2 intermountain biogeography a symposium 15 no 3 the endangered species a symposium 6 no 4 soil plant animal relationships bearing on revegetation and land reclamation in nevada deserts 6 no 5 utah lake monograph 8 no 6 the bark and ambrosia beetles of north and central america coleoptera scolytidae a taxonomic monograph 60 no 7 biology of desert rodents 8 no 8 the black footed ferret 10 no 12 research inm the auchenorrhyncha homoptera a tribute to paul W oman 30

send orders for back issues to editorial office attnanttn carolyn backman great basin naturalist 290 MLBM brigham young university provoprovostprovoutProvoUTUT 84602 USA please include 2 handling and postage for each great basin naturalist volume or memoir great basin naturalist solsoisot501 1990 appp 91 92

distribution OF LIMBER PINE DWARF MISTLETOE IN NEVADA

robert L mathiasen 1 and frank G Hawksworth 2

limber pine dwarf mistletoe arceutho- note confirms hawksworth s suspicion that bium cyanocarpunicyanocarpum A nels ex redbrydb A limber pine dwarf mistletoe occurs in the nels viscaceae parasitizes several species toiyabe mountains of white pine subgenus Strobus in the west- in 1989 the senior author found arceutho- ern united states from montana and colo- bium cyanocarpum at several new localities in rado to oregon and california hawksworth nevada fig I1 on limber pine in the north and wiens 1972 mathiasen and hawksworth snake mountains white pine county near 1988 the principal hosts of this mistletoe are mount moriah in the bull run mountains limber pine pinus flexilisflexilis james whitebark elko county near porter peak in the santa pine P albialblalbicaulisengelmalbicauliscaulis engelm great basin bris- rosa mountains humboldt county south of tlecone pine P longaevalongaeva D K bailey and windy gap at three locations in the toiyabe rocky mountain bristlecone pine P ariaristataaristotastata mountains north toiyabe peak and bunker engelm it also occurs on western white hill lander county and near arearc dome pine P monticola dougl and foxtail pine nye county and on the southeast slopes of P balfourianabalfouriana grev & balf in northern boundary peak in the white mountains es- california hawksworth and wiens 1972 meralda county just east of the california mathiasen and hawksworth 1988 the lim- state line limber pine dwarf mistletoe was ber pine dwarf mistletoe causes local severe also collected on limber pine and great basin mortality in several areas hawksworth and bristlecone pine near mount washington in wiens 1972 mathiasen and hawksworth the south snake mountains probably near 1988 the same location where D K bailey col- the distribution of limber pine dwarf lected it on great basin bristlecone pine in mistletoe in nevada is poorly documented addition a population of limber pine dwarf hawksworth and wiens 1972 hawksworth mistletoe was found in the warner mountains 1988 kartez 1988 first collected in nevada of northeastern california modoc county on in 1881 on great basin bristlecone pine in whitebark pine specimens of dwarf mistle the spring creek charleston mountains in toes collected are deposited at the USDAUS DA clark county it was not collected elsewhere forest service pathology herbarium FPF in nevada until 1958 when it was found in the rocky mountain forest and range experi- ruby mountains near elko elko county ment station fort collins colorado four hawksworth and wiens 1972 it has since additional mountain ranges in nevada were been reported from the sheep mountains visited the toquicatoquima mountains mount jef- clark county copper and east humboldt ferson area and the monitor range monitor mountains elko county hawksworth and peak area in lander county the indepen- wiens 1984 and the south snake mountains dence mountains jack peak area in elko white pine county collection by D K bai- county and the pine forest mountains duf- ley hawksworth 1988 suggests that the fer peak area in humboldt county although mistletoe probably occurred in the toiyabe extensive populations of limber pine andor mountains of nye county because linsdale whitebark pine were observed in those areas et al 1952 illustrated a limber pine that ap- no dwarf mistletoe was found however it peared to be infected by this mistletoe this is probable that the parasite occurs in other

USU S forest service forest pest management 324 25th street ogden utah 84401 2uauU S foiestforest service rocky mountain foiestforest and range experiment station fort collins colorado 80526

91 92 NOTES volume 50

widely separated mountain ranges in nevada both limber and great basin bristlecone pines occurred some 800 to 1000 m lower 0 0 than at present during the late quaternary thompson and mead 1982 thus their 9 dwarf mistletoe was presumably much more widely distributed then also however as the climate warmed the pines and their associated dwarf mistletoe receded to the higher elevations of major ranges and the dwarf 0 mistletoe became restricted to scattered 0 0 relictualrelicrevictualtual populations

literature CITED critchfield W B AND G L allenbaugh 1969 the distribution of pinaceae in and near northern nevada Madromadronoflofio 201220 12 26 0 critchfield W B ANDEAND E L LITTLE 1971 geographic distribution of the pines of the world USDAUS DA forest service miscellaneous publication 991 97 appp hawksworthHAWKSWOMH F G 1988 mistletoesMistle toes of nevada northern nevada native plant society newsletter 14 3 hawksworthhawkswokth F G AND D WIENS 1972 biology and classification of dwarf mistletoesmistletoes arceuthobium fig 1 distribution ofarceuthobium cyanocarpwncyanocarpum in USDAUS DA forest service agricultural handbook 401 nevada and adjacent areas solid dots denote previously 234 appp reportedrepoidepoi ted locations hawksworth and wiens 1972 1984 1984 biology and classification ofarceuthobium and open circles indicate locations first portedreportedle in this an update appp 2 17 in biology of dwarf mistle study toes proceedings of the symposium USDA forest service general technical report RM 111 KARTESZKARTFSZ J T 1988 A flora of nevada unpublished dis- parts of these ranges as well as other isolated sertationsertation university of nevada reno 1729 appp mountain ranges in nevada that support size- dissertation abstracts international B 49 5119 able populations of limber pine great basin 1989 bristlecone pine whitebark LINSDALE M A J T HOWELL andaandjAND J M LINSDALE 1952 or pine critch- plants of the toiyabe mountains area nevada field and allenbaugh 1969 critchfield and wasmann journal of biology 10 129 200 little 1971 MATHIASEN R L AND F G hawksworth 1988 dwarf considering that limber pine dwarf mistle- mistletoesmistletoes on western white pine and whitebark toe is noted for its extremely disjunct distribu- pine in northern california and southern oregon tion in the western united states hawks- forest science 34 429440429 440 THOMPSON R S ANDJAND J I1 MEAD 1982 quaternary worth and 1972 1984 late wiens and its occur- environments and biogeography in the great rence in adjacent areas in western utah deep basin quaternaryQuateinary research 17 39 55 creek mountains and eastern california panamint mountains and warner moun- received 25 november 1989 tains it is not surprising to find it surviving in accepted 30 november 1989 great basin naturalist soisol501soo 1990 appp 93 95

SOREX PREBLEI IN THE NORTHERN GREAT BASIN

mark A ports and sarah B george2georgea

sorex prebleiprebles has been described as a rare tain ranges following a north to northeast pro- shrew of the columbia basin with a distribu- gressiongression and high valley floors are the prom- tion extending as far east as the northwestern inent physiographic features within the great plains junge and hoffmann 1981 its higher valleys and river drainages are mesic to presence in the great basin desert is delin- xeric shrublandsshrublands dominated by artemisia and eated by specimens that have been collected chrysothamnus the river bottoms are domi- primarily on the periphery of this region nated by more xeric halophytes such as sarco- zeveloff 1988 other specimens have been batus and chrysothamnus collected from the northern eastern and collection of S prebleiprebles specimens from western edges of the great basin these in- northern elko county suggests that this clude shrews from the northeastern corner of shrew is more common and widespread in california williams 1984 the northwestern the northern great basin than previously corner of nevada hoffmann and fisher supposed A total of 12 specimens ofsofjofsS prebleiprebles 1978 eastern oregon jackson 1922 1928 were collected near sheep creek 10 kinkm west hansen 1964 verts 1975 southeastern of haystack ranch 55 kinkm N of elko sheep washington armstrong 1957 west central creek drains the independence range idaho larrison and johnson 1981 montana seventy five sunken pitfall traps were placed hoffmann et al 1969 hoffmann and fisher in a grid and shrews were sampled from this 1978 western wyoming hoffmann et al area from june to october 1984 ports and 1969 and the south shore of the great salt mcadoo 1986 other sorex include 31 lake in utah tomasi and hoffmann 1984 S vagransvamagransgrans 13 S montimonticolusmonticuluscolus and 7 S merrimerrlmem there are no records from the snake river ami these specimens are housed at the natu- plain of southern idaho or from the bulk ofthe ral history museum of los angeles and great basin desert in nevada or utah south northern nevada community college of the 40th meridian S prebleiprebles was also collected approximately herein we report records of sorex prebleiprebles 49 km east of sheep creek in the perennial from elko county nevada in the northern riparian willow and wild rose of the mary s great basin these records fill in a major gap river at this locality approximately 87 kinkm in the distribution of the species and offer NE of elko a single specimen was taken in a additional information on their habitat and snap trap on 12 june 1986 also collected here sympatric associations with other species of were three specimens of S vagransmagrans and one sorex specimen ofsofjofsS montimonticolusmonticuluscolus the northern great basin is included in the these records fill in a distributional gap for great basin division of the intermountain S prebleiprebles in the great basin the only other floristic region Hoholmgrenlingren 1972 this area record for this species in nevada is from includes the closed river basins of the hum- washoe county hoffmann and fisher 1978 boldt river and the mary s river as well as the to the east S prebleiprebles has been collected from seasonally wet bonneville flats and the snake the southern shore of the great salt lake river plain the region has a continental tooelethoele co utah tomasi and hoffmann climate of fairly hot summers and cold snowy 1984 the specimen from the NW corner winters approximately 130 fault block moun of nevada washoe county is approximately

life science department northern nevada community college 901 elm st elko nevada 89801 2seetionsection of mammalogymam kalogymalogy natural history museum oflosoffosof los angeles county 900 exposition blvd los angeles california 90007

93 94 NOTES volume 50

345 km from the sheep creek locality mcadoo 1986 the most likely habitat associ- whereas the mary s river locality is approxi- ations for S prebleiprebles in this area seem to be mately 233 km from the great salt lake ephemeral and perennial streams dominated record to the north the nearest record of by shrubs primarily below 2500 m in eleva- S prebleiprebles is from the jordan valley malteurmalheur tion county oregon jackson 1922 approxi- although sympatry among species of long mately 223 km away from the north edge of tailed shrews is common in the western the great basin both the independence united states spencer and pettus 1966 range and the mary s river are on the border williams 1984 S prebleiprebles has rarely been ofthe floristic regions ofthe great basin to the captured in association with other shrews south and the columbia basin to the north junge and hoffmann 1981 found S prebleiprebles Hohnholmgrengren 1972 associated with its congener S cinereuscicinereousnereus some authors have suggested that S pre haydenihaydenshay deni in coniferous forest mountain shrub blei is found in marshy and riparian habitats habitats in yellowstone county montana in larrison and johnson 1981 whereas others california s warner mountains S prebleiprebles is have stated that this species prefers arid to sympatric with S vagransmagrans and S merriamimerriami in semiarid shrub grass associations or openings ecotonal habitats of forested riparian and dry in montane coniferous forest dominated by shrublandsshrublands williams 1984 sagebrush tomasi and hoffmann 1984 our herein we document the association of habitat observations in northeastern nevada S prebleiprebles with three other species ofsorexof sorex only partially agree with those of tomasi and it is difficult to explain why the plant commu- hoffmann 1984 nity at sheep creek which is a xeric ephem- at the sheep creek locality we collected eral stream habitat can support four species S prebleiprebles in a seasonally wet sagebrush dom of shrews whereas the more mesic complex inated community the stream exhibits a mountain habitat in the warner mountains snowbeltsnowmeltsnowmelt spring runoffrunofffromfrom early spring until and the perennial riparian habitat on the early summer and then it dries up big sage- mary s river support only three shrew spe- brush artemisia dentatatritridentatatridentate rubber rabbit cies certainly it is unusual to find S montico brush chrysothamnus nauseosus and ante- lus in a low elevation sagebrush community lope bitterbitterbrushbrush purshia tridentata provide this species is normally associated with more a dense overstory and a variety bunchgrassofbunchgrassof bunchgrass mesic habitats in nevada hall 1946 species and forbs provide a stable understory churchfield in press suggests that shrews by the end of summer the grasses and forbs are very flexible in their foraging habits and have seeded and the area is very dry the will decrease their dietary overlap when the elevation at this locality is 2150 m the number of shrew species in a community in- mary s river locality has a perennial source of creases dietary generalists usually are found water from the Jarjarbidgeharbidgebidge mountains to the in greater numbers than dietary specialists in north the soils here tend to be more fertile multi species communities this may be the with extensive meadows and the dominant case with shrew communities in the great vegetation includes willows salix sp wood s basin but these species assemblages will be rose rosa woodwoodsiiwoodsiesii greasewood sarcobatus explained only with detailed ecological stud- sp and great basin wildrye elymus cana ies examining micromicrohabitatshabitats diets and life dendensisbensissis other grasses and forbs provide an history patterns plus comprehensive trap- abundant cover in adjacent meadows that are ping to insure that all shrew species in a cut seasonally for hay particular area are well documented williams 1984 has collected S prebleiprebles in similar mixed sagebrush aspen and willow acknowledgments riparian habitats in the warner mountains of northeastern california we wish to thank marcus pete rawlins six years of shrew trapping by the senior and the nevada department of wildlife for author in wet meadows montane coniferous their contributions of shrew specimens and forest and high elevation mountain brush of habitat information from the mary s river northeastern and central nevada have failed we also thank loislotslols ports who assisted in the to turn up a specimen ofofsjofsS prebleiprebles ports and field 1990 NOTES 95

literature CITED JUNCE J A AND R S HOFFMANN 1981 an annotated key to the long tailed shrews genus sorex of the ARMSTRONG F H 1957 notes on sorex prebleiprebles in united states and canada with notes on middle washington state 38 6 murrelet american sorex occasional papers museum of churchfield S in press niche dynamics food re- natural history university ofkansas 94 1 48 sources and feeding strategies in multi species LARRISON E AND communities of shrews museum of J D R JOHNSON 1981 mammals of southwestern idaho press biology special publication university ofidaho moscow 166 appp HALL E R 1946 the mammals ofnevada university of PORTS M A AND J K mcadoo 1986 sorex merriamimerrimerrlami california press berkeley 710 appp insectivora Soncsoricidaesoneidae inm eastern nevada south- HANSEN A 1964 ectoparasites of mammals from ore- western naturalist 31 415415416416 gon great basin naturalist 24 75 81 SPENCER A W AND D petruspetousPETTUSPET iusrus 1966 habitat prefer- HOFFMANN R S AND R D FISHER 1978 additional ences in five sympatric species of long tailed distributional records of prebleprebless shrew sorex shrews ecology 47 677 683 preprebleipreblesbleibiel journal of mammalogy 59 883 884 TOMASI T E AND R S HOFFMANN 1984 sorex prebleiprebles HOFFMANN R S P L WRIGHT AND F E NEWBY 1969 in utah and wyoming journal of mammalogy the distribution ofsome mammals in montana I1 65 708 mammals other than bats journal of mammalogy VERTS B J 1975 new records for three uncommon mam- 505793057950 579 604 mals in oregon murrelet 56 22 23 HOLMGREN plant N H 1972 geography of the inter WILLIAMS D F 1984 habitat of appp associations some rare mountasmountainmountam region 77 161 in A cronquist shrews sorex from california journal of mam- A H holmgren N H Hoholmgrenholingrenlingren and J L malogy 65 325 328 reveal intermountain flora I1 pub- vol hafner ZEVELOFF S I1 1988 mammals of the lishing co new york intermountain west university of utah press salt lake city JACKSON H H T 1922 new species and subspecies of 365 appp sorex from western america journal of washington academy ofscience 12 262 264 1928 A taxonomic review of the american long received 10 february 1989 tailed shrews north american fauna 51 1 238 accepted 12 december 1989