Great Basin Naturalist
Volume 53 Number 2 Article 15
6-4-1993
Full Issue, Vol. 53 No. 2
Follow this and additional works at: https://scholarsarchive.byu.edu/gbn
Recommended Citation (1993) "Full Issue, Vol. 53 No. 2," Great Basin Naturalist: Vol. 53 : No. 2 , Article 15. Available at: https://scholarsarchive.byu.edu/gbn/vol53/iss2/15
This Full Issue is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. T H E
GREATR EAT BASINbaab7a S I1 N MTURALISTNA afufta FISTlyom
llemoe ak
VOLUME 53 n2na 2 JUNE 1993
BRIGHAM YOUNG university GREAT BASIN naturalist editor JAMES 11 BARNES 290 MLBM brigham young university provo utah 84602
associate editors MICHAEL A BOWERS BRIAN A MAURER blandy experimental farm university of department ofzoology brigham YoungyounguniversityyoungiuniversityUniversity virginia box 175 boyce virginia 22620 provo utah 84602
J R CALLAHAN JIMMIE R PARRISH museum of southwestern biology university of BIOWESTBIO WEST inc 1063 west 1400 north logan new mexico albuquerque new mexico utah 84321 mailing address box 3140 hemet california 92546 PAUL T TUELLER department of range wildlife and forestry JEANNE C CHAMBERS university of nevada reno 1000 valley road USDA forest service research university of ne reno nevada 89512 vada reno 920 valley road reno nevada 89512 ROBERT C WHITMORE JEFFREY R JOHANSEN division of forestry box 6125 west virginia uni department of biology john carroll university versityhersityversity Morganmorgantowntown west virginia 26506 6125 university heights ohio 44118 PAUL C MARSH center for environmental studies arizona state university tempe arizona 85287
editorial board richard W baumann chairman zoology H duane smith zoology clayton M white zoology lerran T flinders botany and range science william hess botany andan rangedRange science all are at brigham young university ex officio editorial board members include clayton S huber deandeanideann college of biological and agricultural sciences norman A darais director university publications james R babesbamesbarnes 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding ofthe great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1993 are 25 for individual subscribers 15 for student and emeritus subscriptions and 40 for institutionsin 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 natu- ralist through a continuing exchange ofscholarly publications should contact the exchange librarian harold B lee library brigham young university provo UT 84602 editorial production staff joanne abel technical editor jan spencer assistant to the editor natalie miles production assistant copyright 0 1993 by brigham young university ISSN 001736140017 3614 official publication date 4 june 1993 5935 93 750 4946 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university
ISSN 001736140017 3614
VOLUME 53 30 JUNE 1993 NO 2
great basin naturalist 532 appp 97 106
DEPLETION OF SOIL MOISTURE BY TWO COLD DESERT bunchgrassesBUNCH GRASSES AND EFFECTS ON photosynthetic performance
jay E andersonAnderandersoniandereonisoni and nancee L toftatoft2tofttoht
ABSIRACI this study compared the abilities of two cool season bunchgrassesbunchbuncherbunchgrgrassesasses to extract moisture from a drying soil and compared photosynthetic and stomatal responses of the two species as soil moisture supplies were depleted when grown in 49491 L pots in a greenhouse laymusleymus cinereus extracted more water from the sodsoilsoli and maintained higher gas exchange rates to lower absolute amounts otof soil water than did agropyron desertorumdesertorum the soil water content at the lower limit of extraction was 10310 3 for L cinereus and 13313 3 for A desertonondesertorumdesertonontorum when soil moisture was expressed as estrextractableextr actable soil water there was little difference between the species in pattern of water use both species maintained high stomatal conductances gw and photosynthetic rates A until extractable soil moisture was reduced to about 15 for field grown plants undertinderlinder severe watelwater stress A was higher in L cinereus than in A desertorumdesertortondesertorumtorton at comparable leaf watelwater potentials the relationship between A and gw was similar for the two species the higher A in L cinereus was a consequence of higher ggw thus higher A in L cinereus is achieved through some sacrifice of water use efficiency
keykegkeywordswords extractableeftievti actable soil water leafwaterleafleak water potential stomatal conductance water use efficiency leymuslaymus cinereuscinereouscinereus agropyron desertorumdesertorurndesertorum
plant species vary widely in their tolerance comstock and ehleringer 1984 ehleringer and of seasonal drought and in the mechanisms they cook 1984 delucia and heckathorn 1989 use to cope with declining supplies of soil mois- chaves 1991 as well as changes in the diurnal ture some species tolerate seasonal drought by patterns of gas exchange schulzeSchuize and hall maintaining high leaf water potentials through 1982 tenhunen et al 1987 it clearly would be stotstorstomatalnatal closure turner 1979 although they advantageous for such species to maintain pho- may maintain a high photosynthetic capacity totosynthetic rates as high as possible as soil and low stomatal conductance asliadliwill severely restrict plant water potentials decline carbon gain under prolonged drought in con- As plants extract water from a drying soil the trast other species allow their leaf water poten- amount ofplant available water decreases expo- tials to drop as soil water potentials decline nentially with decreasing water potential eg turner 1979 this enables the plant to con slatherslatyer 1967 fig 333.3 consequently the vol tinuedinue to extract water from a drying soil but ume of water gained by a plant in drying a given decreases in leaf water potential typically are volume of soil to 202.0 mpa over that gained in accompanied by decreases in photosynthetic ca- drying a soil to 151.5isls mpa for example is so pacity and stomatal conductance jones 1973 small that it would seem rather negligible in
I1 departnientdepirtrnentofof biologicbiological il sciences idahoidabo state university patelloP atello 1daho83209idahoidabo 83209 012011O 11 fifth street apt 2cac davis californiailifoi 95616
97 98 GREAT BASIN naturalist volume 53 terms of total carbon gain jordan and miller preliminary gas exchange data from field grown 1980 and jordan et al 1983 estimated that plants J anderson unpublished data sug- the additional water made available to a crop as gested that L cinereuscicinereousnereus plants maintained higher a consequence of lowering leaf water potential photosynthetic activity and had higher stomatal 11 a few bars would support transpiration only for conductance at low leaf water potentials than 3 or 4 days in the absence of additional root did A desertorumdesertorum plants to test those pospossibilisibili growth thus there would seem to be little ties we conducted a greenhouse experiment 1 advantage in making the necessary osmotic ad to compare photosynthetic and stomatal re- justment andor other leaf modifications to tol- sponses of these two species to drying soil and erate very low water potentials and we might 2 to compare the lower limit of extraction of expect little difference among drought toleranttoletoierant the two species in addition we compared pho- species in their lower limit of extraction of soil tosyntheticto capacity and conductance of the two moisture we use ritchieritchiesritchiesrRitchieschless 1981 definition of species under water stress imposed naturally in the lower limit of extraction the amount of the field both laboratory and field data support water remaining in the soil when plant growth the hypothesis that in comparison to A deser and activity completely stop I1 toriumcorium L cinereuscicinereousnereus plants deplete soil moisture on the other hand tolerance of very low reserves more completely and maintain higher plant water potentials may offer advantages photosynthetic rates as water supplies are di- other than gaining more water from a particular miniminishedshed but this is achieved through some volume of soil osmotic adjustment may enable sacrifice in water use efficiency by L cinereuscicinereousnereus a plant to maintain turgor in growing roots which in turn would enable the plant to explore METHODS the soil for additional water reserves sharpe and davies 1979 jordan et al 1983 westgate plant materials and boyer 1985 turner 1986 drought toler laymusleymus cinereuscicinereousnereus is a robust tussock grass ant sagebrush antemannemAftemartemisiaaftemisiaisia tritrldentatatmentata plants of native to cold deserts and lower mountain slopes the great basin of north america move water throughout the intermountainintel mountain west of north at night along a hydraulic gradient in the roots america it occurs on alkaline or saline lowland from deep in the soil to drier soil at shallow sites as well as nonsaline upland sites often on depths richards and caldwell 1987 caldwell deep soils young and evans 1981 walker and and richards 1989 the water deposited in brotherson 1982 plants used in this study were those shallow layers can be extracted the follow- transplanted from a near monocultural natural ing day to support carbon gain or other physi- stand at the idaho national engineering labo- ological activity finally caldwell 1985 ratory lneINEINELL additional information concern postulated that drying a soil to a very low water ing the physiologyecoecophysiology of L cinereuscicinereousnereus can be content may be a way of excluding competitors found in anderson et al 1993 these arguments and results suggest that sig- agropyron desertommdesertorumdesertommtorum is a tussock grass na- nificantnificant but possibly subtle differences could tive to the steppes of asia it is naturalized in exist in the lower limit of extraction of soil water western north america where it has been used among species indeed sinclair and ludlow extensively for rangeland rehabilitation stands 1986 found small differences in the lower limit established by seeding often persist as near of extraction among four tropical legumes monomonoculturescultures marlette and anderson 1986 grown in pots plants used in this study were transplanted from anderson et al 1987 compared the sea seeded stands at the INEL additional informa- sonai patterns of soil water extraction among tion about the ecophysiology of A desertorumdesertorum four drought toletoietolerantrant cold desert species they can be found in nowak and caldwellcalcaidwell 1984 found little difference in the lower limit of ex- 1986 and nowak et al 1988 the four when traction among species grown in greenhouse monomonoculturescultures on a common soil however there studies was some indication that the native buncligrassbunchgrassbunchgrass twenty four pots were constructed from leymuslaymus cinereuscicinereousnereus scribn & merr A love polyvinyl chloride pipes each pot was I1 in tall might be able to extract more water from a soil and 0250.25 rn in diameter the pots were filled than could the introduced species agropyron with a 1iliiii1111 1 mixture of bacciobaccto potting soil sand desettorumdesertorumdeserdesetttorumorum fisch ex link schult in addition and a clay loam soil used in experimental field 199311993 DEPLETION OF SOIL MOISTURE BY bunchgrassesBUNCH GRASSES 99 plots at the INEL see field studies the mean A and standard error dry mass of soil placed in 100 the pots was 44444.4 040.4 kg at field capacity that volume of soil 18.5185 0.1oi01 L of water held 185 01 or 80 39739.7 water by volume dormant A deser torum and L cinereuscicinereousnereus plants were collected from stands at the INEL in november and held 60 W at 5cac until 12 plants of each species were trans 0 40 planted into the 49491L pots in december the U plants a where they were placed in greenhouse U received natural sunlight by four 20 supplemented 0 A desertorumdeser torum 1500 W metal halide lamps the height of the L cicinereuscinereouscznereus lamps was set so that photosynthetic photon flux 0 density PFD at canopy height was 1500 2000 2 imollmolxmolalmol m s at midday the photoperiodphotoperiodwaswas 13 h B
1 1 1 1 plants were fertilized twice a week with full 0 strength ruakura nutrient solution smith et al c 1983 after the plants became well established six of each species were assignedb randomly to a well watered control and the other treatment 2 six were assigned to a water stress treatment the well watered plants received nutrient solution twice a week and distilled water once a week water stress was induced by withholdingbya4thholding water 0 from the plants for 50 days after bringing the 3 4 soil field because of the water content to capacity 0 large volume ofsoil and water in a pot water stress C n 5 was imposed gradually simulating soil drying that 0 5 10 15 20 25 30 35 occurs naturally under field conditions time doy the pots were weighed every 3 44 days dur- fig 1 A changes in total soilsodsoli water content with timetirrie ing the drying period to determine the amount after withholding water from agropyron desertorumdeserfommdeserdeserftorumomm and of held the soil of each soil water leymuslaymus cinereuscicinereousnereus plants dashed lines and for irrigated water in pot controls solid lines growing in 49491L pots in a greenhouse content was expressed in two ways percent of B soil water potential vs time for treatment means corre- total soil water TSW was defined as volume spondingsp to the closed symbols in A see table I1 for statis- of water in the soil volume of water at field tical analyses capacity X 100 extractable soil water ESW see ritchie 19819811 was expressed as a percentage from pools on the floor resulting from the wa- of the difference between the volumetric water tering of other plants after day 36 the amount content at field capacity and that when growth of water in the soil decreased to levels slightly of that species had stopped day 50 of the drying lower than those on day 29 because growth of period soil water potential was measured with individuals of both species had essentially singlesingie junction screen caged psychrometers stopped by day 33 and changes in soil water J R D merrill specialty equipment logan content after that date were negligible only data utah placed at soil depths of 150150350550350 550 and for 33 days are included in most analyses pre- 750 mm in three pots per species containing sented here water stressed plants psychrometricpsychro metriemetric output was rate of elongation of expanding leaves was monitored every 3 4 days using a model NT 3 used as an index of growth rate leafelongation nanonanovoltmetervoltmeter decagon devices inc pull- rate LER of the youngest leafwas determined man washington by measuring its length at two times and divid on day 33 there was a slight increase in the ing the difference in length by the time interval weight of pots in the water stress treatment length measurements were made on two vege- fig 1 but it was not until we noted a substan- tative tillers per pot on three consecutive days tial increase in weight of some pots ottortonorr day 36 each week leaf elongation rates reported here that we realized water had entered some pots were averaged over 48 h 100 GREAT BASIN naturalist volume 53
TABLF 1 general linear models analysis of soil water content expressed as percent of total soil water soil water content expressed as extractableestrextractable soil water elongation rate of the youngest leaf on a tiller and soil water potential forlenforleyforbeyforbeymuiforoor lebinusleyinusmuimul cinereus and agropyron deserdcsertorumdesertorumtorum plants growing in 49491 L pots in a glasshouse for each dependent variable mainmaln effects weiewelewere always included in the model but if interinteractionictionaction terms were not significant they were excluded from the model all independent variables in each model were treated as classification variables there were two levels of species SP A desertorumdesertorum and L nereuscincreuscinereuscincinereousci creus for all dependent variables in part a there were two levels of stress STR wellweilweli watered and water stressed and 10 days DAY afterifterwatelwater was withheld hornfromgorn the the water stressed plants 1 4 8 12 16 19 22 26 293329 33 for soil watelwater potential in partpaithb tlieieweiethere were foinfourfoun levels of sod depth DP 150350150 350 550 750750mmmin and 9 days DAY 4 8 12 16 19 22 26 29 33
a effect species stress day SP x STR SP x DAY STR x DAY SP x STR x DAY total soil water p0001P 0001 p0001P 0001 p0001P 0001 P 0002 P 0001 n 228 extractable soilsod water nsfns1ns p0001 p0001P 0001 P 0001 n 228 leaf elongation rate P 0001 p0001 p0001P 0001 p0001P 0001 P 0002 P 0001 P 0003 n 255 b effect species depth day SP x DP SP x DAY DP x DAY SP x DP x DAY soil water potential P 002 P 0001 P 0001 n 204
notigiflut
field studies of 2 kpa the concentration of coa inside the cuvette was 330 340 LLJLL L light was provided field studies were conducted at the the isow bulb INEL experimental field station where by a 150 W quartz halogen projector measurements were made periodically monomonoculturescultures of A desertorumdesertonimdeserdesertotorumnim andandlL cinereuscicinereousnereus were established by transplanting mature indi throughout the drying period between 0900 and visualsviduals from nearby stands to experimental plots 1600 hours TSW in the pot was determined having a homogeneous soil to a depth of 242.4 m immediately after gas exchange measurements see anderson et al 1987 for details the soil for gas exchange measurements in the field consisted of 26 sand 54 silt and 20 clay leaf temperature was 24c PFD was 1900 lmolalmol1 3 2 1 and had a bulk density of 128 g cm the mnrv1S or greater v was 232.3 kpa and 602coagoagog con- measurements reported here were made during centrationcentration inside the cuvette was 335 5 RLjulJLL the third growing season after the plants were Llil light was from sunlight or a 150 W projec- transplanted tor lamp measurements were made in june and early july between 0830 and 1500 hours gas and plant water exchange water potential was measured with potential measurements leaf 60 a pressure chamber PMS instruments co net photosynthesis A transpiration E corvallis oregon immediately after gas and leaf conductance to water vapor gvgw were exchange measurements on the same leaf measured on the youngest fully expanded leaves or on the penultimate leaf on a tiller after calculations and statistical analyses inflorescences developed with an open com- A E and gw leaf penpensatingsating gas exchange system which has been plus boundary layer were previously described nowak et al 1988 toft et calculated according to caemmerer and far- al 1989 gas exchange measurements of quhar 1981 ambient atmospheric pressure at greenhouse grown plants were made at a leaf the INEL is about 85 kpa field measurements temperature of25cof25c a PFD of 1900 2000 rmolamol1 and at pocatello idaho is about 86 kpa green- m 2 s 1 and a leaf to air water vapor gradient v house measurements statistical analyses were 199311993 DEPLETION OF SOIL MOISTURE BY bunchgrassesbunciigrassesBUNCH GRASSES 101
10 0 L cinereuscicinereousnereus 0 A desertorumdeser torumtotum 5030 08 25 H r 20 06 15 0 10 aa 1AI deserlorutndes ettogrid rum 0a 1 L 5 cinereus
02 0
00 0 10 20 30 050 5 doy daydoy 1 time 0 4 cn 04 0 fig 2 relative leaf elongation rate LER of the young- j est leaf on tillers as a of 0 vegetative expressed percentage C 030 3 controls vs time after withholding water from agropyron A deser cinereous 491L desertorumtorum and leginus cinereuscinereus plants growing in 49 05 020 2 pots in a greenhouse see table I1 for statistical analyses 010 S c performed with the general linear models pro- 00 cedure ofofsashofsasSAS SAS institute 1982 250 RESULTS
j 200 h following the withholding of water TSW j was consistently lower in pots containing L cin 1 A deser 150 areusereus than in those containing desetlonimdesertorumtorum U fig 1aaa the more rapid decline in TSW in L cinereuscicinereousnereus pots early in the experiment may re- 0 20 40 60 80 100 flect the larger size of that species but if plant iotototatotorotojota soi waierwaterwoter size or growth rate was responsible for observed fig 3 net photosynthesis A leaf conductance to water species differences in TSW differences would vapor ggw and intercelluintercellulartar concentration ofor carbon dioxide have diminished with time did not occur ci for agropyron deserdesendesertorumdesertcrumdeserttorumcrumorum and lebinusleymiisleyinus cinereuscicinereousnereus plants that vs total soil water in 49491 L pots the curves were fitted using thus the significant species differences and an inverse secant transformation of soil water content see species by treatment interaction table la table 2 for equations and values of constants measure- show that water stressed L cinereuscicinereousnereus plants were ments were made at a leaf temperature of25cof 25oc a leaf to air kpa capable of extracting more water from the soil vapor pressure gradient of 20 an ambient coz concen- tration of 330340330 340 JLRL L and a photosynthetic photon flux A desettorumdesertorumdeserdeseTt volumet- as than were stressedstressedastressedA torumorum the density of 1900 almollmoljimol photons inms2s ric soil water content at the lower limit of extrac- tion day 50 was 13313.3 and 10310.3 for A desertorumdesertorum and L cinereuscicinereousnereus respectively ereusareus 50550550.5 mm da 1 was nearly twice that for the time courses of soil water potential av- A desertorumdesetforumbeserdesertorum 26096026026.0 mm ifda 1 relative LER of eraged over all four positions in a pot and over both species decreased within 6 days after water all pots with water stressed plants are shown in was withheld from the stress treatment plants figure IB water potential depended on loca- fig 2 early in the period of soil drying rela tion within the pot table ib with lower water tive LER of stressed plants was lower for A potentials occurring at higher positions in the desertorumdesetforumdeserehsertorum than for L cicinereuscinereousnereus resulting in a soil profile soil water potentials in pots contain- significant species by stress interaction table ing L cinereuscicinereousnereus were significantly lower than 1 this suggests that leaf elongation may be those of A desertorumdesertorum after day 5 fig IB more sensitive to water stress in A desertorumdesettorumdeserdeseTttorumorum table ib than in L cicinereuscinereousnereus for well watered plants elongation rate of the youngest leaf on a tiller LER for L cin 102 GREAT BASIN naturalist volume 53
table 2 analyses of covariancecovanance of photosynthesis leaf conductance to water vapor and intercellular c02 concentration toifoifolfor lfcwlebinusleyinusLflellcw cinereusLicinereouscinereunereus and agropyronafoAropyron desertotitincle&ertorum plants grown in 49491 L pots in a glasshouse see fig 3 the classification vaivalvariablelablelabie in each model was species SP A desertorumdesertorum aadeagde and L cinereus leeileelalecileci the covariatecova nate NX was a transformation oftotalof total soil water TSW expressed as percent of the water content at field capacity the general form of the transformation was X seesec TSW QC where secsee is the inverse secant function and C is a constant determined by iteration and given heiehelehere toitolfolfor each species
n constant r2raR a effect species TSW SP X TSW aadeagde leeledlecl aadeagde ledleeileellem photosynthesis P 004 P 0001 P 004 260 78 86 83 n 16 conductanceC P 04 P 0001 265 139 83 83 it 16 intelintercellularcellulaicelceicellularlulai c02c021 P 002 P 005 P 004 271 195 67 86 n 16
the relationshiprelation ship between A or ggw and TSW water per unit of carbon gained than did A for both species is shown in figure 3 to facili- desertorumdesetforumdesertorum which reflects the additional cost of tate statistical analyses numerous models were water paid to achieve higher A fitted to the data by linear regression and trans- A and ggw ofplants sampled in the field in late formation ofthe dependent andor independent spring and early summer were positively corre variable the model that consistently provided edwithlatedwithlatedlat with ij measured concurrently fig 4 for the best fit considering all data sets was both species A was more closely correlated with f than was ggw and the correlation coefficients Y bo b TSW C seesec Q were higher for L cicinereuscinereousnereus than for A deser field sampling was initiated where Y is net photosynthesis or leaf conduc- torum at the time tance seclsedlsediseesec is the inverse secant function C is a L cicinereuscinereousnereus plants were considerably inorelnoremore water desett constant bo is the intercept and bib is the slope stressed than were A desettommdesertorumdesertorumomm plants be- the constants were determined iteratively by cause of differences in ESW in the plots at the substituting values for them until the highest beginning of the growing season data not coefficient of determination rar2R was obtained shown this difference is reflected by the high- values of C and r2Rra are given in table 2 for the est fl values recorded for the two species fig curves shown in figure 3 4 As aeonacona sequenceconsequence the highest rates ofA were analyses of covariancecovanance were performed for lower forlaorlfor L cinereuscicinereousnereus than for A desertorumdesertorum and each gas exchange vanablevariable using species as a were considerably lower than the maximum A classification vanablevariable and the inverse secant ofoflkoflL cicinereuscinereousnereus observed in the greenhouse fig transformation of TSW as the covariatecovanate table 3 or for well watered plants in the field ander- 2 L cinereus had higher A and gw than A son et al 1993 that most L cicinereuscinereousnereus plants desertorumdesettorumdeserdesettcesertorumorum at both high and low levels of TSW were stressed while some A desertorumdesettorumdeserdesetttorumorum were fig 3 table 2 when soil water availabilityavailabilitywas was not may account for the higher correlation coefboef expressed in relative terms iei e ESW the re- ficients forlaorl cinereuscicinereousnereus data in figure 4 indicate sponses of the two species were similar both that A and gw were generally lower in severely maintained high A and ggw until extractable water stressed A desertorumdesertorum than in L cinereuscicinereousnereus at content reached about 15 marked reductions comparable ajqji which is consistent with findings of A and ggw occurred at lower levels of ESW from the greenhouse study data not shown the relationship between A and gw is similar intercellular cog concentrations qc were for the two species when grown either in the higher in leaves of L cinereus than in A deser field or greenhouse fig 5 this again demon- torum over a wide range of soil water contents strates that the higher A in L cinereuscicinereousnereus was a fig 3 table 2 therefore under comparable consequence of higher ggw rather than a higher atmospheric conditions L cinereus lost more photosynthetic capacity field grown plants had 199319931 DEPLETION OF SOIL MOISTURE BY bunchgrassesBUNCH GRASSES 103
I1 I1 I1 I1 I1 I1 I1 I1 1 060 6 1 I1 0 A desertorumdesertorum r063ra 65 A desertorum field 5030 r0 desertorum 0 0 L cinereus ra 88 0 L nereusjetenere us fielde r0r088 05 cirete 0 1AI desertorumdesere s ertorumtottorU lalobb 25 0 L cz en cinereusneteustousrous lob 00 i 040 4 CN 0 0 E 20 0 E s 6 03 0 0 1 15 0 IE 0 vs 0 2 0 3 10 02 5 C 5 01
0 00 0 5 10 15 20 25 30 35 050 5 2 1 A desertorumdesertorum rarogo60 r0r060 A eolholamolpholmol nn s L cinereuscinereouscicznereus r0r078ra 78 040 4 h T fig 5 leaf conductance to water vapor ggw versus net U photosynthetic rate A for agropyron desertorumdesertorum and ley
1 in field closed symbols 030 5 h mus cinereus plants grown in the or E in a glasshouse open symbols gas exchange measure- ments were made over a wide range of sod water availabili- 0 ah022h N ties cuvette conditions were the same as in figure 4 trend lines for glasshouse grown plants solid line and field grown plants dotted line were fitted by linear regression 01 with A taken as the independent variable schulze and hall 1982
000 0 7 6 5 4 53 2 1 0 1987 that L cinereuscicinereousnereus plants could extract more water potentia mrampa leaf water from a soil profile than could A deser fig 4 net photosynthesis A and leaf conductance to torum in addition L cinereuscicinereousnereus maintained water vapor ggw for field grown agropyron desertorumdesertorum and higher A at low f compared to A desettorumdesertorumdeserdeseTttorumorum leymuslaymus cinereuscicinereousnereus plants as a function of leafwater potential measured concurrently gas exchange measurements were these data suggest that L cinereuscicinereousnereus might have made at leaf temperature of 24c a leaf to air vapor pres- a competitive advantage should the two species kpa sure gradient of 23 an ambient c02 concentration of co occur it is important to remember however 330 340 illliljil L and a photosynthetic photon flux density 1 A is 1900 jamoljimol photons ntm s trend lines were fitted by linear that desertorumdesertorum naturalized in western regression with leaf water potential taken as the inde- north america and so the species did not co pendent variable solid line L cinereuscicinereousnereus dotted line A evolve desertorumdesertorum L cinereuscicinereousnereus achieved higher A by maintain- ing a higher ci through higher stomatal conduc- higher A for a given gw compared to greenhouse tances thus there was an additional water cost grown plants fig 5 the intercept of the re- per unit carbon gained the large stature of L gressiongression of gw on A did not differ significantly cinereuscicinereousnereus plants suggests a high water require- from zero for field grown plants indicating that ment and indeed stands of this species can use WUE was quite constant over the range ofaofA and copious amounts of water anderson et al ggw measured schulze and hall 1982 however 1987 As a consequence L cinereuscicinereousnereus typically is the regression line did not pass through the found on deep soils or areas that receive runoff origin for plants grown in the greenhouse P water from adjacent sites miller et al 1982 0505.05 which suggests that WUE decreased with walker and brotherson 1982 roundy 1985 increasing A and gw on such sites its capacity to dry a soil to lower water contents may exclude potential competi- discussion tors indeed it often occurs in pure or nearly pure stands on such sites sirotnak 1990 results of the greenhouse study confirmed showed that there was intense intraspecific our suspicions based on data from monoculmonocur competition in a stand oflofkoflL cinereuscicinereousnereus and the L tures grown in the field see anderson et al cinereuscicinereousnereus plants reduced leaf water potentials of 104 GREAT BASIN naturalist volume 53 neighboring chtysothamnuschrysothamnus nauseosus shrubs first 3 weeks of the drying period fig 2 also the size and lower water use efficiency of L may reflect greater osmotic adjustment in L cinereuscicinereousnereus are likely disadvantageous however on cicinereuscinereousnereus sites where total water available over the grow previous studies have shown that leaf expan- ing season is more limited mortality of L cin sion frequently is more sensitive to water stress ereusareus plants was greater than that of A than is photosynthesis begg and turner 1976 leserdeserfonimdesertorumdesertorum on our experimental field plots dur- As reported here for two perennial tussock ing a severe drought in 1987 88 unpublished grasses kuang et al 1990 found that LER in observations wheat and lupin was reduced almost immedi in contrast to L cinereuscicinereousnereus A desertorumdesertorum atelyabely after withholding water they demon- establishes well and thrives on shallow soils and strated that LER decreased in response to very arid sites rogler and lorenz 1983 its drying soil even when leaf turgor was main- success on such sites probably reflects a smaller tained and they suggested that leaf growth size at maturity even under well watered con stomatal conductance and osmotic adjustment ditionsditeditionsions the ability to withstand prolonged are all controlled by the balance of leaf phytopayto drought and prolific production of viable seeds hormones as influenced by hormones produced hull and klomp 1967 marlette and anderson in the roots given the frequent observation that 1986 pyke 1990 caldwell and his colleagues A and ggw are closely correlated eg fig 5 it at utah state university have shown that A would seem reasonable to add photosynthetic deserdesertorumdesertonimdesendesertotorumnim is a vigorous competitor for water capacity to the list and soil nutrients reviewed by dobrowolski et the relationship between ggw or A and TSW al 1990 its competitive ability reflects at least fig 3 indicates a close coupling between leaf in part the production of thin roots that enable gas exchange and soil water content as has been it to extract water rapidly from the soil eissen- reported for a number of herbaceous andandwoodywoody stat and caldwell 1988 species growing in a variety of soils gollan et al it seems probable that the ability oflofkoflL cin 1985 turner et al 1985 sinclair and ludlow ercus to reduce soil water content to lower levels 1986 henson et al 1989 turner et al 1985 than those of soil supporting A &desertorumdesersertommtorum re- and gollan et al 1985 demonstrated that al- flects lower osmotic potentials in leaves of L though ggw and ajqjf often were correlated the na- cinereuscicinereousnereus we attempted to estimate osmotic po ture of the relationship was dependent upon tentiatertialtential1 of greenhouse grown plants from pres environmental conditions and the rate of soil sure volume curves but leaves of L cinereuscicinereousnereus drying thus they found no unique relationship were so brittle that we were unable to obtain between gw and tpi and postulated that ggw and A reliable data concurrent measurements of leaf are controlled by the level of water in the soil relative water content RWQRWC and water poten rather than in the leaf subsequent studies have tiai offield grown plants showed that fora given confirmed that hypothesis showing that the RWC L cinereuscicinereousnereus had lower i than did A de roots sense water availability or some related seTtonseytonsefsettontinsertorumsertorumlintin data not shown P 0404.04 by analysis of parameter in the soil and transmit signals to the covariance this difference between species leaves that control their behavior gollan et al could arise from a greater degree of osmotic 1986 masle and Passipassiouraoura 1987 Passipassiouraoura adjustment lower osmotic potential at a given 1988 henson et al 1989 zhang and davies RWCRWQ by L cinereuscicinereousnereus compared to A deser 1989 1991 tardieu et al 1991 our data are torum consistent with this model for both species ggw kuang et al 1990 postulate that the fac- and A were closely related to soil water content tors that causes a reduction in LER as the soil fig 3 rather than showing a cause anand deffeeffect dries also induces osmotic adjustment in the relationship the correlations between ggw and leaves their results show that the proportional fig 4 likely reflect variationcovariationco in response to change in LER per unit osmotic adjustment was declining soil moisture supplies much greater in lupin than in wheat which in conclusion this study shows that there suggests that LER in a species with greater may be small but significant differences in the osmotic adjustment might be less sensitive to extent to which cold desert species can dry a soil the 11 stress imposed by drying soil thus the profile such differences may be important in observation that LER was reduced relatively competitive interactions caldwell 1985 com less inlinbinlL cinereuscicinereousnereus than inA desettommdesertorumdeserdeseTttorumomm over the pared with A desertorumdesetforumdesertorum L cinereuscicinereousnereus maintains 199311993 DEPLETION OF SOIL MOISTURE BY bunchgrassesBUNCH GRASSES 105
higher photosynthetic rates as soil moisture sup field study of two andlandaddlandaridland tussock grasses oecologia plies decline but it does so by maintaining a 75175 1 7 GOLLAN TJT J B PASSIOURA AND R MUNNS 1986 soil higher conductance not through a greater pho- water status affects the stomatal conductance of fully tosyntheticto capacity turgid wheat and sunflower leaves australian journal of plant physiology 13 459 464 GOLLAN T N C TURNFR AND EDE D scilulrsciiul7e 1985 acknowledgments the responses of stomata and leaf gas exchangeexeliaexcliaexella nge to japourvapour pressure deficits and soil water content III111 in we thank teresa ratzlaff bob rousseau the sclerophyllous woody species neritunnerininnerinon oleander and mark shumar for technical assistance and oecologiaoecologia6565 356362 HENSON I1 E C R lensilensenJFNSIJENSEN N AND N C turnerTURNI 1989 drs tony condon josette masle rana munns leaf gas exchange and water relations of lupinsdupins and and two anonymous reviewers for comments on wheat I1 shoot responses to soil water deficits austra- the manuscript this paper is a contribution lian journal of plant physiology 16 40113401 413 from the idaho national labora- HULL A C J AND G J KLOMP 1967 thickening and engineering spread of crested wheatgrasswheat grass stands on southern idaho tory radioecologyRadioecology and ecology program the langesranges journal of range management 20 222 227 research was supported by the office of health lonesJONESJONFS H G 1973 moderate teimterm water stress and associ- and environmental research US department ated changes in some photosynthetic parameters in of energy and environmental restoration and cotton new phytologist 72 1095 1104 en JORDAN W R W A J DUGAS AND P snouseSHOUSE 1983 waste management US department of strategies for crop improvement for drought pioneploneprone ergy idaho field office regions agricultural water management 7 281 299 JORDANJO udanRDAN WRW R ANANDD F R MIMILLERLLE 11 1980 genetic variability in sorgsorghumhurn root systems implications for drought tol- literature CITED erance pages 383399383 399 in N C turnerandturnerTurnerandand PE J kra- mer eds adaptation of plants to water and high ANDERSONANDFRSON J E R S NOWAK N L tohtoftton AND K RAS- temperature stress john wiley & sons new york MUSON 1993 photosynthesis in laymusleymus cicinenereusfeitsreitsfelts di- KUANG JBJ B N C tubnerTURNFRTURNER AND I1 E HENSONNSON 1990 urnal and seasonal patterns manuscript influence of xylem water potential on leaf elongation ANDERSONANDFRSON J E M L SHUSIIUMARmailmallMAii N L tonTOFT AND R S and osmotic adjustment of wheat and lupin journal of NOWAK 1987 control of the soil water balance by experimental botany 41 217 221 sagebrush and three perennial grasses in a cold desert MARLFITFMARLETTE G mandlmanajmandjMAND J E ANDERSON 1986 seedbanksseed banks environment andarid soil research and rehabilitation 1 and proppropaguleagule dispersaldispel salgalgai in crested wheatgrasswheatgrass stands 229 244 journal of applied ecology 23 161 175 brocBEGG J EANDNE annAND N C TURNERTURNFR 1976 crop water deficits MASLE J AND J B PASSIOURA 1987 the effect of soil advances in agronomy 28 161 217 strength on the growth of young wheat plants austra- CAFMMFRFR S VON AND G D FARQUIIAR 1981 some lian journal of plant physiology 14 643 656 relationships between the biochemistry ofphotosynof photosyn- MILLERMILLFR R F F A BRANSON I1 S MCQUEENN AND C T thesis and the gas exchange of leaves planta 153 SNYDFRSNYDER 1982 water relations in soils as related to 376 387 plant communities in ruby valley nevada journal of CALDWFLLCALDWELL M 1985 cold desert pages 198 212 in B F range management 35 46246268468 chabot and H A mooney eds physiological ecology NOWAK R SJS J E andlannANDANDIE RSON ANANDI1 N L toftTOFI 1988 gas of north american plant communities chapman and exchange of agropyron desertontindewrtomm diurnal patterns hall new york and responses to water vapor gradient and tempera- CALDWFLLCALDWELL M M AND J II11 RICHARDSricrio HARDS 1989 hydraulic ture oecologiaoecologia7777 289 295 lift water efflux fromfroin upper roots improves effective- NOWAK R S AND M M CALDWFLLCALDWELL 1984 A test of ness of water uptake by deep roots oecologia 79 1 5 compensatory photosynthesis in the field implications CIIAVFSCHAVES M M 1991 effects of water deficits on carbon for herbivoreherbivoryherbivory tolerance oecologiaoecologia6161 311 318 assimilation journal of experimental botany 42 1 16 1986 photosynthetic characteristics of crested COMSIOCKCOMSTOCK JJANDJAND J eiilfringfr 1984 photosynthetic wheatgrasswheatgrass and bluebluebunchbunch wheatgrasswheat grass journal of responses to slowly decreasing leaf watelwater potentials in range management 39 443 450 enceliafrutescensenceha frutescentfrutescens oecologia 61 241 248 pa&siouraPASS louIOU RA J B 1988 root signals conticontrolol01 leaf expansion deluciaDFLUCIA E HANDSH AND S A hfckaiiiornheckxriiorn 1989 the effect in wheat seedlings growing in drying soil australian of soil drought on water use efficiency in a contrasting journal of plant physiology 15 687 693 great basin deseideserdesert t and sierran montane species plant PYKE D A 1990 comparative demography ofcooccoof co occur cell and environment 12 935 940 ring introduced and native tussock grasses persistence dobrowolski J P M M CALDWELLCALDWFLL AND J H and potential expansion oecologia 82 537543537 543 RICHARDS 1990 basin hydrology and plant root sys- RICHARDSri HARDS J H AND M M CALDWFLLCALDWELL 1987 hydraulic tems pages 243 292 in C B osmond L F pitelka lift substantial nocturnal water transport between soil and G M hidy eds plant biology of the basin and layers byartemisiaby ArtemisiaartemisiatndentatatHdentata lootsroots oecologia 7348673 486 range springer verlag new york 489 ehleringer J RANDCR AND C S COOK 1984 photosynthesis RIKIIIFRITCHIE J T 1981 soil water availability plant and soil 58 in encehafarinosaenceha fannorafannosa gray in response to decreasing leaf 327 338 water potential plant physiology 75 688 693 rooleRoCLEROCLFRii G A AND R J lorenLORFN 1983 crested wheat eissenstat D M AND M M CALDWFLL 1988 com- grass early history in the united states journal of petipetitivetive ability is linked to rates of water extraction a range managementM anagement 36 91 93 106 GREAT BASIN naturalist volume 53
ROUROUNDYNDY B A 1985 emergence and establishment ofbasin function stanford university press stanford califor- wildrye and tall wheatgrasswheatgrass in i elationrelation to moisture and nia salinity journal of range management 38 126131126 131 TOFTon N L J E ANDFRSONANDERSON AND R S NOWAK 1989 SAS insmuirdinsmuirins11ljrf 1982 SAS useiuser s guide statistics SAS water use efficiency and carbon isotope composition institute inc carygary north carolina of plants inm a cold desert environment oecologia 80 scnuilscouil EDE D AND A E haliIHALLlalllali 1982 stomatal re- 11 18 sponsesspon ses water loss and coa assimilation rates of plants turnellTuRNEurnerURNFRii N C 1979 drought resistance and adaptation to in contrasting environments pages lsiISI181 230 in 0 L water deficits in crop plants pages 343 372 in H lange P S nobel C B osmond and H zieglerziegleiedseds musselmussell and R C staples eds stress physiology in physiological plant ecology II11 encyclopedia of plant crop plants john wiley and sons new york physiology new setlesserlesseriesseisel lesies volume 12b springer verlag 1986 adaptation to water deficits a changing per- newnewyoikyork spectivespec tive australian journal of plant physiology 13 SHARPE 11 E AND W J davilDAVII s 1979 solute regulation 175 190 and growth by roots and shoots ofwaterof water stressed maize tuilnriiurnerUKNFR N C EDE D SCHULZESC IIULE AND T GOLLAN 1985 plants planta 147 43 49 the responses of stomata and leaf gas exchange to SINCIAIKSINCLAM T R AND M M LUDLOW 1986 influence of japourvapour pressure deficits and soilsod water content II11 in soil water supply on the plant water balance of four the mesophytic herbaceous species helianthusHehenthusanthus an tiotropicalpical grain legumes australian journal of plant nuns oecologia 65 348355348 355 physiology 13 329 341 alkeralgerWALKEALKFRii G R AND J D brothersonBROHII RSON 1982 habitat SIKOINAKs1sa ROTNAK J 1990 intraspecific and interspecific coinpeticompeti- relationships of basin wildrye in the high mountain tion in lcfmmslaymusleymus cinereus and chrysothamnus nauseousnauseowsnauseosus valleys of central utah journal of range management in a cold desert community unpublished mastermasterss the- 3562835 628 633 sis idaho state university pocatello WESTGKITrsicalf M eandieandjE AND J S boyeboyerboyenBOYFRR 1985 osmotic adjust- sikryiSIAIYIHii R 0 1967 plant water relationships Acadacademicernieernicernle ment and the inhibition of leaf root stem and silk press london growth at low water potentials in maize planta 164 SMHII G SCS C M JOIINSIONJOHNSTON AND I1 S cornfortilcornroiun 540 549 1983 comparisonCompanson of nutrient solutions for growth of YOUNGOUNG J AAANDRAND R A EVANS 1981 germination of great plants in sand culture new phytologist 94 537 548 basin vjldryewildrye seeds collected from native stands ZIIANGZIIANC agronomy journal 73 917 920 TAimiTAHDIIiuu F N kankailkallKAII aiji1111iijiKI 0 BETHENODbr mi NOD J AND z11ang DAVIFSDAVIES WWJJ DAVIFSDAV ES 1991 maize stomatal conductance in the lanclanoIANC JJANDWJAND W J 1989 abscisic acid produced field its lelationshiprelationship with soil and plant water poten- in dehydrating roots may enable the plant to measure tials mechanical constraints and ABA concentration the water status of the soil plant cell and environ- in the xylem sap plant cell and environment 14 ment 12 73 81 121 126 1991 antitranspirant activity in xylem sap of maiemalemaizemalze of 42 317 321 tlTENHUNENNHUNI N JV D R W PEARCYpi aiaala Y AND 0 L lancklangkLANGELANGP 1987 plants journal experimental botany 317321 diurnal variationsvanations in leaf conductance and gas ex- change in natural environments pages 323 351 in E received 1 december 1992 zeigelzeigerzelger G D1 farquhaiandlfarquhar and 1 R cowan eds stomatal accepted 13 january 1993 great basin naturalist 532 appp 107 117
ON THE RELATIVE importance OF FLORAL COLOR SHAPE AND NECTAR REWARDS IN attracting pollinatorsPOLLINATORS TO MIMULUS
steven D sutherland and robert K vickery jr 2
ABSIRACI pollinator preferences were observed for the six species ofot section erythrantheerythfanthe of the genus mimulus using greenhouse grown plants placed in a meadow in the red butte canyon natural area salt lake county utah thethepnncipalprincipal pollinapolhnatoispollinatorstors weiewelewere hummingbirds and bumble bees hummingbirds preferredpiefpreferred the species with the most reflexed tubular flowers regardless of color whereas bumblebees preferred pink lavender or yellow flowers to red flowers regardless of shape results for the six species were confirmed by observations of faf2 hybrid recombinant plants selected such that flower color could be held constant and flower shape varied and vice versa
kegkeykeywoidskeywordswords mimulus hummingbirds bumblebees flower color flower shape pollinatorspollinators speciation
the dependence of animal pollinated plants to have a aideaddewide spectrum of color vision kevan on biotic vectors for pollen transfer has in many 1983 cases led to the evolution of floral adaptations FLORAL morphology hummingbirds that benefit one class of pollinatorspollinators for example are generally considered to prefer tubular in this study hummingbirds or bumblebees but shaped flowers K grant and V grant 1968 discourage or exclude other potential visitors yet artificial experiments show that humming- medusemeeuse 1961 procter and yeo 1972 faegrifaegre birds freely visit radial relatively flat flowers and van der pijlbijl 1979 percival 1979 wyatt graham pyke personal communication bum- 1983 medusemeeuse and morris 1984 these adapta- blebees visit all shapes of flowers although they tions include 1 floral color which is important appear to prefer flowers with a landing platform for long distance recognition and attraction 2 andor a nectar guide percival 1979 floral morphology which may be an attractant FLORAL REWARDS nectar appears to be as well as a determinant of the effectiveness of the underlying attraction to hummingbirds and pollination by these visitors and 3 floral re- nectar andor pollen to bees free 1970 hein- wards the ultimate reason for pollinator visits rich and raven 1972 heinrich 1975 1976 FLORAL COLOR hummingbirds are com- stiles 1976 burquez and corbet 1991 monly said to have evolved a preference for red because these adaptations floral color flo- or orange red flowers K grant 1966 K grant ral shape and nectar rewards determine the and V grant 1968 raven 1972 faegrifaegre and van identity or kind of the pollinator changes in der pijlbijl 1979 medusemeeuse and morris 1984 how- floral traits can potentially cause a shift in polli ever in artificial hummingbird feeding prefer- natorsgators eg from bees to hummingbirds thus ence experiments no color preferences were leading to reproductive isolation and possibly detected bengbene 194119451941 1945 wagner 1946 col subsequently to speciation this well may have lias and collias 1968 miller and miller 1971 happened in Penpenstemonpenstenwnpentstemonstemon straw 1956 and stiles 1976 goldsmith and goldsmith 1979 Aquaquileciaaquilegiaacfuilegiailecia chase and raven 1975 and may be nor were color preferences detected in most happening in shapanusrhapanusRhapanus sativus color morphsmorphe george 1980 mcdade 1983 but not all vick stanton 1987 and in the monkey flower spe- ery 1992 natural experiments in contrast cies of section erythranthe bumblebees are commonly reported to eschew the six species of section erythranthe in- red flowers scogin 1983 although von frisch clude a single pink flowered bee pollinated heinrich 1979 some years ago had shown bees species M lewisii pursh and five red flowered
I1 nature conservancyconse y 21560 route 245 marywillemarysvillemarywildeMaryMarys villewille ohio 43040 2departmentdepartment ofot biology university ofutah0 utah salt lake city utah 84112 anthorauthor to wliointowhoin correspondencorrespondedconeon esponclence should be addressed
107 108 GREAT BASIN naturalist volume 53
1cmacm mimulus lewisii non reflexed
tat1
y 1cmacm mimulus verbenaceusverbenaverbenaceousceus partially reflexedreflexedflexed
1cmacm mimulus cardinalis fully reflexed
fig 1 the three primarypi imaryamary floral shapes in the enthrantheerythranthe section of the genus mimulus 199319931 pollinator preferences IN MIMULUS 109 hummingbird pollinated species M rupestrisrupestris mimulus cardinalis flowers are fully reflexed greene M eastwoodiae rydberg M nelsonnnelsoniinil that is the upper and lateral corolla lobes are grant M cardinalis douglas and M verben sharply turned back and the labellum is folded abeusaceus greene gieseyhieseyhieseyetHieseyetet al 1971vickery1971 vickery 1978 back on itself fig 1 the more reflexed the 1987 flowers are the more tubular they appear the the bee pollinatedpollinated species mimulus lewisii cocorollasrollas of all red flowered species have tongue has a wide distribution from southern california guides like the pink flowered species but lack northward to southern alaska and eastward to nectar guides under UV light northwestern colorado flowers vary in inten- all six species produce bisexual that is per- sity of color from light lavender pink sierra fect flowers and are self compatible the flow- nevada race to deep magenta pink rocky ers of all five red flowered species contain the mountain race the cocorollasrollas are non reflexed same set of six anthocyanin pigments chanicyani fig 1 and see color illustrations in vickery and din 3 glucoside cyanidin 3 rhamnoglucoside wullstein 1987 that is the corolla lobes are at cyanidin 3 caffeoyl glucoside pelargonidin 3 right angles to the axis of the corolla tube or are glucoside pelargonidin 3 rhamnoglucoside even thrust forward as in the sierra nevada race andpelargonidinandpelargonidin 3 caffeoyl glucoside and the the corolla throats have a pair of hairy ridges same carotene pigment pollock et al 1967 that serve as tongue guides for pollinatorspollinators also the flowers ofpink flowered M lewisii have the the throats display dark nectar guides under UV same three cyanidin pigments but they lack the light three pelargonidins and the carotene pigment is ofthe five hummingbird pollinated species restricted to red dots in the corolla throat vari three have restricted distributions mimulus ru ationsactions in flower color of the different species the pestrispastris is a narrow endemic that grows on shady populations and hybrids reflect variations in cliff faces in the sierra de tepoztlan 70 km presence amounts and floral locations of the south of mexico city mexico mimulus ru anthocyanin and carotene pigments in addi- have identified three different of pastrispestris flowers vary from pinkish red to cardinal tion we pairs M cardinalis M verbenaceusverbenaverbenaceousceus and M red in different populations the flowers fig genes in lewisii that off anthocyanin production 1 vary from non reflexed actually thrust for- turn when recessive leading to yellow ward in the pinkish red flowered population homozygous flowers the 9102 to partially reflexed in otherpopulationsother populations due to remaining carotene pigment M cardinalis and M verbenaceusverbenaverbenaceousceus and to in partially reflexed flowers the upper pair of in white flowers in M lewisii which lacks caro- corolla lobes is strongly reflexed but the laterallateralrai tene further information on the morphol- pair and the labellum are not mimulus east for ogy distribution physiology genetics and woodinewoodiae is endemic to moist areas hanging taxonomy of the group see A grant 1924 gardens in protected overhangs in sandstone pennell 1951 hieseygieseyhieseyetalet al 1971 and vickery shelter caves in southeastern utah and north- 19781987 eastern arizona it has cardinal red partially sisixx species of section erythranthe form reflexed flowers mimulus nelsoniinelsonianelsonii is a broad the a promising group for investigating the basic endemic found alongalonaion streams in limited areas 9 question of this study what floral traits are high in the sierra madre occidental in the states responsible for attracting particular pollinatorspollinators of and sinaloa mexico too has durango it specifically hummingbirds and bumblebees to reflexed flowers cardinal red partially the flowers other two hummingbird pollinated species of section erythranthe have wider distributions MATERIALS AND METHODS mimulus verbenaceusverbenaverbenaceousceus is distributed in northern mexico sonora through arizona into southern utah mimulus verbenaceusverbenaverbenaceousceus flowers are cardinal experimental design red and partially reflexed mimulus cardinalis is the experimental design for this study of distributed along the pacific coast from central pollinator attraction for the species and hybrids baja california to southern oregon and inland of section erythranthe has two parts first in into nevada and arizona mimulus cardinalis experiment 1 we plan to observe the pollinator flowers vary from yellow to orange to orange red preferences for the six species comprising the to cardinal red in different populations A few section in relation to their flower colors flower populations are mixed eg yellow and red shapes and nectar rewards second in experiexpert 110 GREAT BASIN naturalist volume 53
taniTABITABLE 1 1 culture numberss and origins of the species and hybrids of section erythranthe used in the experimental field studies
M cardinalis douglas 13106 yellow flowered morph growing in a wash with palms and running water elev ca 150 m on north side of cedros island baabeiabajabeja california del norte mexico used for experimental hybridization only 13249 red flowered morph from the same population M verbenaceusverbenaverbenaceufsverbenaceousceus greene 5924 growing along bright angel creek nealnear phantom ranch elev ca 612 m grand canyon national park coccinoboccino co arizonaanona M nelsoniinelsonianelsonii grant 6271 growing in a small brook in a pine forest elev ca 2555 m near the sierra madre divide on rt 40 sinaloa mexico M eastwoodiae rydberg 6079 glowinggrowing inm seeps in caves on high sandstone cliffs elev 1415 m bluff san juan co utah M rupestrisruperupefstrisstris greene 9102 growing on a conglomerate rock cliffoneliffcliff on tepozteco trailtrallfrail elev ca 2300 m sierra de tepoztlan morelos mexico M lewisii pursh 5875 growing by a small stream on albion basin road elev 2680 m alta salt lake co utah 6103 giowmggrowing by a small sandy stream leaving ice lake sierra nevada elev ca 2000 m placer co california used toitolfolforoor the experimental hybridization only FI hybrid 13258 M cardinaliscardinally 13106 cedros island yellow morph corolla lobes fully reflexed x M lewisii 6103 ice lake sienaslenasierra nevada lavender corolla lobes thrust forward fa hybrid population 13300 fiF 1 hybrid 13258 X self
ment 2 we plan to distinguish the relative effects lavender pink race ofofmM lewisii with thrust for- of color shape and nectar rewards this adliwill be ward corolla lobes 6103 the fiF hybrid had done by testing plants ofan fa hybrid population medium pink flowers with partially reflexed up- that recombine these traits such that we can 1 per corolla lobes the F hybrid was self polli hold shape constant and observe the effect of natedbated and from it a large fa hybrid population different colors 2 hold color constant and ob- ofclose to 1000 plants was grown the fa hybrid serve the effect of different shapes and then 3 plants recombined the parental flower shapes relate these results to the nectar characteristics and nectar characteristics and exhibited a wideavide of the plants array of transgressive variation in flower color red yellow light lavender pink light pink me plants diumdaum pink and deep magenta pink As would be anticipated from the genetic control of an- populations representative of each species thocyanintho production see above there were table 1 were selected in the university ofutah cyanin few red flowered fg hybrid plants greenhouse these populations included the relatively few with corolla lobes two color morphsmorphe red and yellow of M cardi- also there were very thrust forward nalis and the two races of M lewisii the sierra nevada light lavender pink race and the rocky mountain deep magenta pink race the various METHODS populations were grown from seeds collected either from the wild or from populations of experiment I1 greenhouse grown transplants of the different floral visitor preferences for the species species and color forms F and fg hybrid populations were grown to determine floral preferences of the po from an experimental cross made between the tertialtential pollinatorspollinators for the six species of section extreme floral color and shape forms of section erythranthe plants of each species except M enfihrantheeiyhranthe specifically between the yellow nelsoniinelsonianelsonii plus the F hybrid ofofmM cardinalis X color form ofofmM cardinalis population 13106 M lewisii were placed for observation in a moist with fully reflexed corolla lobes and the light meadow in red butte canyon natural area in 199311993 pollinator preferences IN MIMULUS iliillIII111 the wasatch mountains elevation 1980 in salt three major flower shapes non reflexed par- lake city utah tially reflexed and fully reflexed in each of the due to problems in initiating flowering in several colors such a set was obtained by self mimulus nelsoniinelsonianelsonii in 1985 this species was not pollinapollinatingting the Ffi hybrid of M cardinalis available for study until the next year fortuit- 13106 X M lewisii 6103 see table 1 we ously the 1986 results for M nelsoniinelsonianelsonii could be selected a subset of plants from the resulting fg included in the analysis because the findings for recombinants with fully reflexed partially re- the three other red flowered species common flexed and non reflexed flowers in red yellow to both experiments de facto controls were and four tints and shades of pink too few insignificantly different from 1985 to 1986 even thrust forward flowered fa plants were ob- though year to year changes in climate and pol- tained to analyze the fourth shape category in linator guilds usually might lead to significant this experiment insufficient red flowered fa differences hybrids were obtained and so three native but nine potted plants of each species and of the corresponding red flowered species non re FFI hybrid were placed on top of inverted one flexed M rupesttisrupestrisrupestris partially reflexreflexededMM verbenverren gallon black plastic pots to raise the flowers abeusaceus and fully reflexed M cardinalis were above the surrounding meadow vegetation and substituted mimulus nelsoniinelsonianelsonii was added to the increase their visibility to potential floral visivlsi experiment to fill the data gap in the previous tors pots were placed at Ilin1 in intervals in a 5 X year s experiment the fa hybrids also recom- 9 m grid within the first row the five species bined the nectar characteristics of volume con- and FIF hybrids were randomly positioned this centcentrationration and sugar content but these were relative sequence was maintained in the sub- not selected thus the second experiment in- sequent rows but the position was shifted three cluded in addition to M nelsoniinelsonianelsonii plants of the places in adjacent rows this arrangement re- three major shape classes in each of six color sulted in each species being surrounded by the categories combined with various but anseunse other four species or the Ffi hybrid lecterlected nectar traits the populations were observed for 2 3 plants of the fa hybrid population including hours at different times of day from early dawn the red flowered species to be tested were to dusk for seven days in late june and early july raised in the greenhouse in spring 1986 five for a total of 20 hours the weather was consis- plants ofmofM nelsoniinelsonianelsonii and five of each color shape tently clear warm and sunny numbers and category were placed in the study area in red identities of potential pollinatorspollinators making close butte canyon in late june the pots were ran- approaches to the flowers and numbers of open domly arranged at imI1 m intervals on inverted pots flowers for each species and the fiF hybrid were in an I111I1 X 9 m grid close approaches by major recorded daily an approach for this study of pollinatorspollinators hummingbirds and bumblebees floral traits that attract pollinatorspollinators was consid- were recorded daily for six days approaches or ered to be a close approach to a flower generally actual visits by other pollinatorspollinators eg flies were followed by an actual visit that is by a hum- rare for each major type of potential pollinator mingmingbirdbird thrusting its beak into the flower or a the total number of approaches for a given day bumblebee landing on and entering the flower was divided by the number of open flowers to at the end of the experiment day 8 flowers determine a daily average close approach rate were destructively sampled at dawn before pol- for each color shape category this value was linator visitation to detenaetendeterminenine the amount and standardized by dividing by the overall mean quality of available nectar the data were exam approach rate for that day to compensate for ined using chi square and spearman rank cor- daily variations in number of pollinator ap- relation methods sokal and rohlf 1969 for proproachesaches andor length of observation periods patterns of floral visitor preference in floral sutherland and vickery 1988 the daily stan- color shape andor nectar rewards dardizeddardi zed rates were then averaged to produce mean pollinator approach rates for each color experiment 2 shape test whether significant dif- pollinator preferences for flower colorshapeColor Shape category to ferencesferen ces in pollinator approach rates to each to discriminate between the relative attracattract color shape category occurred a two way analy- tivenessrivenesstiveness of flower shape and flower color it was sis of variance was performed sokal and rohlf necessary to have a population of plants with all 1969 if significant differences were detected 112 GREAT BASIN naturalist volume 53 a student newmannewm an keuls test was then con RESULTS AND discussion ducted to identify where the significant differ- ences occurred sokal and rohlf 1969 to test floral visitors for differences pollinator significant temporal in floral visitors observed in both experiments approach rates a kendall coefficient of concor- were principally hummingbirds and bumble- gibbons dance test was performed 1976 to bees with only negligible visits by flies they test whether hummingbirds were learning dur- were active all day the majority of humming- ing the experiment to associate flower shape bird visits were by broad tailed hummingbirds andor flower color with nectar rewards daily selasphorus platycercus a highly migratory approach rates to the 18 color shape categories species found throughout the range of the spe- 6 color classes and 3 shape categories were cies ofsection erythranthe in addition visits by ranked low to high and compared using the calliope Stellula calliope black chinned ar kendall coefficient of concordance to test the chilochuschilochus alexandalexandrilri and perhaps other species hypothesis that bumblebees were learning dur- of hummingbirds were observed in other areas ing the experiment ie that they were altering of the range other species would surely be ob their foraging preferences with time and expe- served as well johnsgard 1983 the majority of riencerience daily visitation rates to the 18 color observed bee visits were made by bombus ap shape categories 6 color classes and 3 shape polituspositus and B huntiihuntie although B bifarius B centraliscentralestralis insularisinsularismlaris P suckle categories were ranked low to high and com- cen psithyrus insu and suckleyiyi were observed also pared using the kendall coefficient of concor- dance experiment 1 nectar characteristics were ascertained at pollinator preferences for the six species the termination of each experiment day 8 for results table 2 of the overview of the six experiment I1 and day 7 for experiment 2 flow- erythranthe species indicate that humming ers were destructively sampled and nectar vol- birds did not approach flowers of the various umes and concentrations measured nectar species in proportion to their frequency in the p1pa1 volume was measured with a calibrated 5 experimental population xax2 22889288228.8 P ooi001001.001 micropipette nectar concentration for the hummingbirds significantly preferred cardinal same set of samples was measured as sucrose red fully reflexed M cardinalis flowers they equivalents weight per total weight basis with neither preferred nor avoided partially reflexed a pocket refractometer average sugar produc- cardinal red species but significantly avoided tion was calculated by converting nectar con- pinkish red non reflexed M rupestrisrupe stris and pink centrationcentration sucrose equivalents to weight per non reflexed M lewisii flowers table 2 these volume using values given in the CRC hand- results could be interpreted as showing an book of chemistry and physics 1978 and mul- avoidance of non pure red however amongst titiplying this value by nectar volume see bolten the cardinal red flowered species the strong et al 1979 for details to test whether signifi- preference for strongly reflexed M cardinalis cant interspecific experiment 1 or color shape compared to the equally red but partially re flexed species suggests to us that hummingbirds category experiment 2 differences existed in JL 00 0 nectar volume nectar concentration andor may be showing a preference for red but prob ably also an attraction to flower shape indeed sugar production a series of single classification ranking the species by increasing degree of re analyses of variance sokal and rohlf 1969 flexion M rupestrisrupestris M lewisii M verben were performed significant interspecific dif- if achus M nelsoniinelsonianelsonii M eastwoodiae M feren newman ferencesces were found a student keuls cardinalis nicely corresponds to the ranking of sokal test and rohlf 1969 was then conducted the species by increasing hummingbird ap- to identify where iiee between which species or proproachesaches table 2 floral shape appears to be groups of species or which color andor shape the main proximate cue for hummingbird ap- categories the significant differences occurred proproachesaches under these conditions then visitation rates were compared with nec bumblebees also were not attracted to all the tar volume nectar concentration and sugar pro species in proportion to their frequency in the duction using spearman rank correlation experimental population xax2 2972971297.11 P ooiool001.001 methods sokal and rohlf 1969 while individual bees would show particular 199311993 pollinator preferences IN MIMULUS 113
TABLF 2 results of experiment 1 flower number pollinator approaches and quantity and quality ofnectarof nectar rewards for the 6 species of section erythranthe and the fiF hybrid of M cardinaliscardinahsnabs x M lewisii means of 7 days of observationsofobservations within a column appiapplapproachoach rates with the same subscript do not differ significantly P 05
hummingbird visits bumblebee visits nectar
number number approaches number number approaches concentration volume amount of of per 100 of of per 100 W joljoi0jil of flowers approaches flowers flowers approaches flowers sugar 6pgg cardinaliscardinahs 260 442 1701 44 0 0oa 134 49 711 13249 verbenaverbenaceusverbenaceousceus 533 604 1131 95 1 la1 128 262 6 361 5924 nelsonnnelsonii 351 379 iosi1081 59 1 2i2aaa 119 282 8 34634 6 6271 eastwoodiae 170 193 114b 23 1 4i4aaa 110 31 330 6079 rupestrisruperupestnsstris 95 27 2828c 7 0 oi0 104 212 1 22822 8 9102 lewisii 156 53 3434c 155 208 1341 156 13 126 5875 fi hybrid 51 60 lisilisb1181 42 56 133b133h 136 41 565 13258 results from 1986 seisonselsonseasonserson controlsConti olsois showed themthein to be compiicompiacomparableubleubic preferences heinrich 1979 overall the bees mingmingbirdsbirds appear to be relying on floral shape significantly preferred pink flowered non cues specifically petal reflexion as an indicator reflexed M lewisii to the red flowered species of the hidden nectar rewards of the various table 2 whether non reflexed partially re species flexed or fully reflexed thus these results sug- when bee approach rates to the six species gest to us that bumblebees are attracted are compared with average nectar concentra- primarily by flower color and not shape under tion average nectar volume and average sugar these conditions production there are no significant correlations the ultimate reason for pollinator visits to r 4646.46 P iolo10.10 rir 3535.35 P 2020.20 and rrs 3838.38 the flowers is to collect floral rewards ie nec- P 2020.20 respectively using spearman rank cor- tar andor pollen in monkey flowers the nectar- relations perhaps this is because bumblebees ies are hidden from view therefore pollinapollinatorstors often collect pollen as well as or instead of cannot ascertain the quantity or quality of nectar nectar heinrich 1979 therefore their forag- rewards from a distance medusemeeuse and morris ing process is more complex than that of hum- 1984 so apparently pollinapollinatorstors have to associ- mingmingbirdsbirds which forage for nectar only ate visible floral clues such as shape or colbraithcoloraithcolocolor withrAith medium pink partially reflexed F hybrid nectar rewards waser 1983 flowers attracted both hummingbirds and bum when hummingbird approach rates to the blebees table 2 F hybrids showed a lower six species are compared with average nectar hummingbirdositationhummingbird visitation rate than their M cardi- volume and average sugar production of those nalis parent but a much higher rate than their species there are significant correlations r M lewisii parent the hummingbird visitation 96gg96.96 P ooi001001.001 rrs 9393.93 P oiol01.01 respectively rate to F hybrids was insignificantly different there is no correlation with average nectar con- from the rates to the partially reflexed species centcentrationration r 5757.57 P loio10.10 using spearman FIF hybrids have the same bumblebee visitation rank correlations in each case hummingbirds rate as M lewisii the partially reflexed flower seem to be making foraging decisions on the shape of the Ffi hybrids would seem to be at- basis of either nectar volume or sugar produc- tractingtr hummingbirds whereas their medium tion rather than nectar concentration inasmuch pink flower color would seem to be attracting as most of the species are red flowered hum bees 114 GREAT BASIN naturalist volume 53
rabirablFABI i 3 results of experiment 2 mean standardized hummingbird approaches per flower for six color and three shape categorieslesies within a row or column averages with the same subscripts do not differ significantly P 05
flower shape flowerplower non partly fully color reflexed reflexed reflexed mean red 0182 1056056 1347 0862 yellow 0730 0990 1213 0978 lavendellavenderLaven deidel pink 0584 1047 11081 ioslos108 0913 light pink 0658 1000 1414 1024 medium pink 0652 11261.1261 126 11451 145 0974 magenta pink 1086 1073 ligo11901 190igo illgilig11161 ilg116 x 0649 049b1049b 1236c
experiment 2 then their shape categories should have the pollinator preferences for flower colorshapeColor Shape highest ranking ifhummingbirds are perfect at distinguishing fully reflexed flowers then the six hummingbirds results table 3 of the colors analysis of approach rates in relation to floral offully reflexed flowers should have rank- 13 14 15 16 17 18 and the color and floral shape by means of a 2 way ings and sum of the would be 93 an ANOVA indicate that hummingbirds 1 did not rankings examination ofthe discriminate between flowers according to color sum of the rankings for fully reflexed flowers indicates significant 1 F 0960.96 P 2525.25 but 2 selectively visited a increase from 69 at day I flowers according to shape F 221122.11 P to 91 at day 5 spearman rank correlation coef- go oolooi001.001 hummingbirds preferred fully reflexed ficient rrs 9090.90 P 0505.05 apparently the birds flowers to partially reflexed flowers and partially are leaminglearning relatively quickly to select more reflexed flowers to non reflexed flowers as ad- reflexed flowers with their higher nectar vol- umbratedumbrated by the comparison of species results umes and sugar contents above with the minor exception of dark pink bumblebees if color is the primary fac- partially reflexed and non reflexed flowers tor attracting bumblebee floral visitors as ex- where visitation rates were reversed this pref- perimentperiment I1 suggestssuglyests to us and bees prefer erence was consistent for all color classes and non red flowers then bees should avoid the shape catecategoriesgonesi red flowered species M rupestrisrupe stris M verben the nectar characteristics of each shape achus and M cardinalis used in this experi- category of fa hybrid plants table 4 were not mental population as surrogates for the random recombinations as might have been ex- underrepresentedununderder represented red fg plants and preferen- pected but showed a significant correlation xax2 tially visit the yellow and pink flowered fa hy- 364203.6420 P 0505.05 between increasing degree brid plants results for the 18 color shape of corolla lobe flexionrereflexionpreflexion and increasing sugar categories of experiment 2 table 5 indicate content of nectar there appears to be genetic that bees selectively visit flowers according to linkage between floral shape and nectar reward color F 5065.06 P 00001.001ooi 1 but do not discrimi- of sugar content hummingbirds seem to be nate between flowers according to shape F taking advantage of this linkage to maximize 1031.03 P 2525.25 although bees discriminated their nectar intake against red flowers there were no significant hummingbirds appear to be leaminglearning to as- differences between visitation rates to non red sociate nectar rewards with flower shape re- yellow lavender pink light pink medium pink sults indicate a significant temporal association and magenta pink flowers in hummingbird floral preferences for the 18 results indicate a significant temporal asso- color shape categories W 0470.47 P oi0101.01 and ciation in bee foraging preference for the 18 the 3 shape categories W 0840.84 P 0505.05 but color shape categories W 0400.40 P oi0101.01 and no temporal association in hummingbird floral the 6 color classes W 0530.53 P 05os05.05 but no preference for the 6 color classes W 015ois0.15 P temporal association in bee floral preference for 5050.50 if fully reflexed flowers are preferred the 3 shape categories W 0280880980.28 P 20go20.20 this 199311993 pollinator preferences IN MIMULUS 115
TABLF 4 mean nectar concentration nectar volume and nectar sugaisugar production for plants in the faf2 hybrid experiment
corolla shape number of nectar concentration nectar volume sugar content flower color flowers SSDD J1 SD ramjiamjigmrgm SD
non reflexed corolla lobes red 15 10410442t 42 21-212020 22822 8 24324 3 yellow 45 1271276767 121312- 13 179220179 220 lavender pink 50 148-1488787 101510- 15 165165271271 light pink 50 17260 15151313 265249 medium pink 37 11258112 58 1436 19919 9 64664 6 magenta pink 40 10760107- 60 101310-loio 13 127-127217217
X 395 128 14 194
partially reflexed corolla lobes red 48 1281284949 2632 361361444444 yellow 45 iiilii1111116161 2332 364364677677 lavender pink 50 1791799090 1009loog10- 09 227227284284 light pink 50 12848128- 48 20-20201818 282282292292 medium pink 61 123-1231234646 1923 275328 magenta pink 50 107-1071073434 121212 12 147191147 igiigl191 x 506 12912 9 18 27627 6
fully reflexed corolla lobes red 41 134 31 494049 40 tii71171 1 621 yellow 51 11945119 45 334033 40 477 623 lavender pink 29 12448124 48 13 18 195603195 603 light pink 50 124 42 21 17 286 261 medium pink 48 105 53 09 09 120 179 magenta pink 50 126 46 14 16 207 289
x 448 12212 2 23 33333.333 3
TABLF 5 mean standardized bee approaches per flower for six color and three shape categories within a row or column means with the same subscripts do not differ significantly P 05
Reflexion color non part full mean
red 00 0065 0480 0182 yellow 1312 0798 1046 105211 052i lavender 1043 0905 0666 087110 871i light pink 1252 1644 1578 149111 4911 medium pink 1403 0613 1078 103111 0311 dark pink 1345 0779 1232 111911 1191 x 1059i1059a 0801 1013
2zwiyanovaay ANOVA chloicoloiol01 F 5065 06 PI1 ooi001 shapeslisllsil ipe F 1030I1 030 P 25 intelint ictionactiontion F 0630 63 P 75 implies that the bees were learning and that significant correlations between bee ap- their floral preferences for color shape catego- proproachesaches and nectar concentration r 08os08.08 P ries and color classes did in fact increase during iolo10.10 nectar volume r 2929.29 P iolo10.10 ororsugarsugar the experiment production rrs 2323.23 P iolo10.10 when bee approach rates were compared to thus under the conditions of red butte nectar characteristics table 3 there were no canyon in 1985 86 the guild of pollinapollinatorstors 116 GREAT BASIN naturalist volume 53 present and the community of meadow flowers bird archilochusArchilochus alexandrialexandn journal of comparative 130 209 220 present that is in this specific pollination mi physiology GRANGRANT 1 A L 1924 A monograph of the genus mimulus lieu handel 1983 hummingbirds showed annals of the missouri botanical garden 11 99 389 no color preferences among red yellow laven gnantGRANIGRANT K A 1966 A hypothesis concerning the prevalence der pink light pink medium pink or magenta of red coloration in california hummingbird flowers pink but foraged according to floral shape they american naturalist loo10000 85 98 reflexed GRAN K A AND V GRANIGRANT 1968 hummingbirds and preferred fully reflexed to partially to their flowers columbia new york non reflexed flowers on the other hand bum HANDELHANDFL S N 1983 Polhpollinationnation ecology plant population blebees showed no floral shape preferences but structure and gene flow in pollination biology aca- foraged according to floral color they avoided demic press orlando florida red flowers but visited yellow lavender pink hrnrkhrnakheniiicii II11 B 1975 the energetics of pollinationpolhnation annual review of ecology and systematics 6 139 170 light pink medium pink and magenta pink 1976 the foraging specialization of individual equally bumble bees ecological monographs 46 105 128 1979 bumblebeebumble bee economics harvard university press cambridge massachusetts acknowledgments HINRICII B AND P RAVENRAVFN 1972 energetics and pollina- tion ecology science 176 597 602 hirsfyhieseygiesey W M M A noimnonsNOBS AND BJORKMAN 1971 we piest and elaine 0 thank bob henry lin experimental studies on the nature of species V bio- sutherland for help with data collection lissy systematics genetics and physiological ecology of the coley vince eckhart monica geber ken erythranthebrythranthe section of mimulus carnegie institution paige and doug samson offered helpful criti- of washingtonofwashington publication 628 washington DCD P A 1983 hummingbirds of northofnorth amer- cism on the research was sup- joiinsgard PA the manuscript icaiealea smithsonian institution press washington DCD ported by an NSF grant BSR 8306997 to RKV kevanKVANKFVAN P G 1983 floral colors through the insect eye we thank terry L griswold research ento- what they are and what they mean pages 3303 30 in C E mologistmologist USDA bee biology and systematics jones and R J little eds handbook of experimental state pollination biology scientific and academic editions laboratory utah university logan utah new york for identification of the bees observed we mcdademe DAD F L A 1983 long tailed hermit hummingbird thank jeanette stubbe for much typing visits to inflorescence color morphsmorphe of Heliconia irrasairreasa condor 85 360 364 MFFUSFMEFUSE B J B 1961 the story of pollinationpolhnation ronald literature CITED new york MFFUSF B J B AND S MORRIS 1984 the sex life of plants on file new york BENEBENP F 1941 experiments on the color preference of facts 11 S MILLERMILLFR 1971 black chmnedhummingbiidschinned hummingbirds condor 43 237 242 MILLERMILLFR AND R E feeding activity and color preference of ruby throated humming- 1945 the role of learninglealleai angmng in the feeding behavior ofosblackotblackblack chinned hummingbirds condor 47 3 22 birds condor 73 309 313 PFNNFLLPENNELL F W 1951 mimulus L vol 3 pages 688 731 BOLTEN A B P FEIFFINSINCNSING ERR II11 G bakelBAKriBAKERandiANDANDl1 BAKERBAKFR in BP fel bakriandl flora odtheof the pacific states 1979 on the calculation of sugar concentration in L abrams illustrated ofthe stan- in press flower nectar oecologia beihnberlin 41 301 304 ford university stanford california PERAVALPFRper IVAL M S 1979 floral biology pergamon press ox- borquezBURQUEZ AANDAANDSS A CORBET 1991 do flowers reabsorb nectar functional ecology 5 369 379 ford england POLLOCK H G R K VKKFRYVICKERY AND K G WILSON CHASE V C AND P II11 RAVIRAVENN 1975 evolutionary and JR ani ecological relationships between aquilegia jortjorljontfornosafonnosajortnosanosa 1967 flavonoidFlavonoid pigments in mimulus cardlcardicardinalisnahsnabs and and A pubescenspuhescempubescentpubescens ranunculaceae two perennial its related species 1 Anthocyananthocyaninsins american journal plants evolution 29 474 486 of botany 54 694 701 PROPROCTER M YEO 1972 flowersofflowers COLUASCOLLIAS N E AND E C COLLIAS 1968 anna s hum- ierlerIFR manbMANDandpyroP the pollination of new york mingmingbirdsbirds trained to select differentdiffelbiffel ent colorss in feeding taplinger condor 70 273 275 RAVFNRAVEN P H 1972 why are bird visited flowers predomi- 26 674 CRC HANDBOOK oiOF CHEMISTRYgill mlsMIS ibyIKY AND PIIYSKPHYSICS s 1978 the nately red evolution chemical rubbelrubber co cleveland ohio SCOGIN R 1983 visible floral pigments and pollinatorspollinators in eds FAFGRIFAEGIU K AND L VAN debnebDERDF R pillPIIpilpli 1979 pollination ecol- C E jones and R J little handbook of experi- ogy pergamon pipressess oxford england mental pollination biology scientific and academic york preliFREEpree J B 1970 insect pollination of clopscrops academic editions new press new york SOKAL 11 R AND F J ROIILF 1969 biometry W H free- GEORGE M W 1980 hummingbird foragingfoifol aging behavior at man san francisco california malvaviscusMalvaviscus ararbarhboreusborcusoreus vaivalvar druinntondiidrnmnwncjii auk 97 790 SrSIANIONANTON M L 1987 reproductive biology of petal color 794 variants in wild populations of raphanus sativus I1 GIBBONS J D 1976 nonparametricnonparametncNonparametric methods for quantita- pollinator response to color morphsmorphe american journal tive analysis holt rinehartRinehaithalt and winston new york of botany 74 178 187 GOLDSMGOLDSMIgoldsmiiiigoldsmitiitilTiiIIII T II11 aniAND K M GOLDSMHH 1979 dis- sillesSIILFSSTILES F G 1976 taste preferences color preferences crimination of colors by the blaekblack chinned humming and flower choice in hummingbirds condor 78 10 26 199311993 pollinator preferences IN MIMULUS 117 strawSIRAW R M 1956 floral isolation in Penpenstenumstenum american vickeimvlckfry R K JR AND B M WULLSIMNWULLSTEIN 1987 com- naturalistnaturahst9090 47 53 parison of six approaches to the classification ofmimuof mimu- sutherland SSANDRAND R K VICKERYvicvie keinkeln JRJK 1988 tradeoffstiadetradefiade offs lus sect erythrantheerythranthc scrophulariaceae systematic between sexual and asexual reproduction in the genus botany 12 339 364 mimulus oecologia berlin 76 330335330 335 WAGNEwagnellWAGNFKwagnerii H 0 1946 food and feeding habits of mexican vickeimvl keryKFRY R K jilJR 1978 case studies in the evolution of hummingbirds wilson bulletin 58 69 132 species complexes in mimulus pages 405 507 in M K WASER N M 1983 the1 he adaptive nature of floral traits hecht W C steere and B wallace eds evolution- pages 242 286 in L real ed pollination biology ary biology vol 11 plenum new york academic press new york 1987 comparison of six approaches to the classifi- WYATIWYATT R 1983 plantpiant pollinator interactions and the evo cation of mimulus sect erythranthe scrophu- lution of breeding sysystemssteinssterns pages 51 95 in L A real lariaceaelanaceaelamiaceaeae systematic botany 12 339 364 ed pollination biology academic press new york 1992 pollinator preferences for yellow orange and cardinally red flowers of mimulus verbenaverbenaceusverbenaceousceus and M cardinalis received 2 october 1992 great basin naturalist 52 145 148 accepted 7 december 1992 great basin naturalist 532 appp 118 130
RESPONSE OF A SONORAN RIPARIAN FOREST TO A loyear10 YEAR RETURN FLOOD
1 2 1 J C stromberg B D richter D T patten and L G woldenwolden1
3 1 ABSIKACI in march 1991 a loyear10 year return flood 368 m s occurred in the hassayampaHassa yampa river a perennial stream 3 1 0 1 in s base flow within the sonoran desert depth of the floodwater ranged from 2649642 64 0200 20 in mean SD near 1 the stream to 0470.470 47 0310 31 in in the highest floodplainfloodplain zone prosopis forest flow velocity was 171 7 060.6og0 6 inm s and 090.9og0 9 I1 t 0404msin s in these same oneszones an average of 8 cm of sediment was deposited on the floodplainfloodplain with maximum deposition to 050 5 inni on densely vegetated surfaces 1 2 in above the water table native riparian vegetation showed resistance and resilience to the flood disturbance plants on high floodplainsfloodplains eg prosopisPioplosopis velutina trees and saplings and populus frearcgrefremonthfreirumtnfreemonthmonthmontu and sahsabsobsalix g0oddingiiooddinii trees had low mortality populusfremontiipopulus fremonfremontefremontntn and S gooddinggooddingngooddingiiii pole trees and saplings weiewelewere on less aggraded floodplainsfloodplains and sustained varying mortality depending on floodplainfloodplain elevation and depth of flood waters foifor example P fremonfremontiifrernontnaremonfremontiatiitil pole trees on 1 2 m high floodplainsfloodplains averaged 6 mortality compared to 40 for those on low floodplainsfloodplains 1 I1 in above the water table where standing water was 2 in seedlings of populus fremontpremont ti and salix gooddingii established abundantly after the flood along overflow channels and main channel sediment bars contributing to age class diversity foiforoor these episodically recruiting species the exotic species tatwirixtamanatamanx penpentandrapentandriatandra had greater mortality of pole trees 62 and lowmostlowpostlow post flood recruitment compared to pfremontiipfremontn andandsS gooddingii survivorship of shrub species also corresponded to floodplainfloodplain elevation zizyphus obtusifoliaobtu&ifolia grew on high elevation floodplainsfloodplains and had no mortality shrubsihsin abub species afloweroflowerof lowel elevation floodplainsfloodplains underwent mortality but revegetated aftelafterahter the flood via asexual reproduction foifolfor example stein density of the dominant shrub baobaccharisBacchans salicisalicifoliasalicifohafolia declined by half but irecoveredecovered to preprefloodflood levels by late summer primarily via stem sprouting dominant herbaceous plants on stream banks and low floodplainsfloodplains iei e the ihiomatousrhizornatous perennial grasses Paspapaspalumluin distichumdistichum and cynodon daktylondactylondactylon similarly compensated for a 50 decline in cover by vegetative spread the post flood herbaceous understory vegetation in high elevation floodplainfloodplain zones iei e prosopis velutinavehgehgeb itina forestsfoifor ests remained sparse throughout the summer and shifted in composition from nearly monotypic stands of exotic annual species to more diverse mixtures of native and exotic annual grasses and forbs
netikeifneifkegkey word parianparlanriparianii vegetationvegetationaodflowflood flowjiow disturbance populus fremonfremontnfremontiifremontiatiitn salix gooddingugooddigooddy nafingfi floodfloodplainplainlainiain aggradation resilience
flood flows have been said to be the prin decreased flooding is often a greater perturba- cipal driving force responsible for the existence tion to riparian floodplainsfloodplains than is flooding productivity and interactions of the major biota sparks et al 1990 as indicated by the substan- in riveriverniver r floodplain systems junk et al 1989 tial vegetational changes that occur when rivers with respect to floodplainfloodplain vegetation flood are dammed and flooding is suppressed reily flows play an integral role in the dynamics of and johnson 1982 pastoupautou et al 1991 dam seed dispersal plant establishment and species construction may result in increased riparian replacement patterns maintenance of species acreage in sediment deltas at upstream ends of and patch diversity and nutrient cycling and reservoirs debano and schmidt 1990 but al- productivity bell 1974 johnson et al 1976 tered flow and sedimentation patterns down smith 1980 long 1982 reichenbacherReichenbacher 1984 stream can result in decreases in plant kalliolaKalpalliolaliola and puhakka 1988 lisle 1989 skog- establishment rates and loss of riparian forests lund 1989 stromberg et al 1991 riparian rood and mahoney 1990 howe and knopf forests in andaridarld regions are particularly depenbepen 1991 dent on flooding because forest regeneration the size differential between base flows and often depends on periodic inundation of other- flood flows of a given recurrence interval is wise dry floodplainfloodplain surface soils hughes 1990 greater in andarid regions than in humid regions
I1 centocentercentorcwrccntccar i loiiolfor lnviionniditil studies arizonaanona stitestate umveisityuiiiversity tempetempo allailalialinaanonana 85287 3211 alifrlif nitiitnahuknahuv t coonselonskionsci vmcyrvanyevany boiikkB id i 0101nudocoltiatlonudo 80302832
118 199319931 RESPONSE TO loyearIOYEAR10IO YEAR RETURN FLOOD 119
graf 1988 large desert floods can cause sub- return flood 193 m3srrsms 1 in augustan ust 1988 and and strate erosion and plant removal in systems in a 2 year return flood 68 ms in july 1990 which stabilizing vegetative cover has been re- cattle duced by grazing base flow reduction or STUDY AREA other factors platts et al 1985 gordon ish shalom and gutterman 1989 stromberg and the hassayampaHassayampa river lies within the gila patten 1992 this occurred in large scale in late watershed of central arizona s basin and range nineteenth century when floods arizona large province and drains portions of the bradshaw on denuded floodplainsfloodplains and watersheds con- date creek and weaver mountains it arises at tritributed to regional erosion and downcuttingdown of cutting about 2350 in and flows freely and intermit- and reeves streams cooke 1976 floods also tently through bedrock canyons interspersed can cause local extirpation of aquatic species in with deep alluvial basins to its confluence with areas has where habitat fragmentation reduced the gila river at about 240 in south olwickofwickof wickwiek their ability to recolonize disturbed areas col- enburg in northwest maricolamaricopaMaricopa county arizona lins et al 1981 in general however because a shallow bedrock layer causes perennial surface desert stream ecosystems evolved with flooding flow for about 8 km the bedrock confined per- they are able to resist or rapidly recover after ennial reach is supplied with alluvial and basin flood events fisher and minckley 1978 fisher fill groundwater stored in a deep basin located et al 1982 reichenbacherReichenbacher 1984 around wickenburg jenkins 1989a 1989b few desert streams in the southwest have the watershed above this point is about 1800 not been modified to some degree by human kmkm2 approximately one third of which is com- activities the opportunity arises infrequently to posed of mountains vegetated by pinus ponder- study large floods in relatively unimpacted sys- osa forests the remainder is rolling hills and tems in february and early march 1991 rain valleys vegetated by interior chaparral and son- storms caused extensive flooding in arizona oran desertdesertscrubscrub species three day rainfall totals within the watershed of the study was conducted along a gaining the hassayampaHassayampa river were titl717.1 cm wicken- section of the perennial river reach base flows burg station to 10110loi10.11 cm prescott station com- increase from 0 to 0oiioli0110.1111 mns3sas at an elevation of prising about 25 ofthe annual average rainfall 600 in within the nature conservancy s has 1 preserve this resulted in peak stream flows of 368 msm3smv1 sayampa river jenkins 1989a the 1 3000 times base flow level at the nature riserriverniver has a gradient of 6 in km the primary has 1 conservancy s hassayampaHassayampa river preserve a channel sandy bed sediments a width of 3 relatively unmodified riparian system for which in and depth of about 030.3 in the floodplainfloodplain there are prefloodpre flood baseline data stromberg et which ranges from about 150 to 200 in in width this is al 1991 A continuing series of storms and in paper defined morphologicallygeomorphologicallygeo as that surface spring snowbeltsnowsnowmeltmelt produced several smaller flood adjacent to the channel and built of materials deposited in the ofthe peaks through the middle of april this event present regime riverniver graf 1988 this encompasses surfaces provided the opportunity to study the response vegetated by prosopis velutina that are up to 3 of the riparian ecosystem to a loyear10 year return in above the water table 1 based on flood our primary objectives were to quantify fig evidence that substrate in such areas was flood 1 changes in floodplainfloodplain topography resulting deposited burkham 1972 minckleyminckleyandand clark from sediment deposition and scour 2 purvi survi 1984 the adjacent uplands slope down to the vorshipworship of dominant trees populusfre riparian floodplainfloodplain with varying gradients the climate is montiinwntiimontai salix gooddinggooddingiiii prosopis velutina and arid with average annual rainfall of29 cm at the the exotic tamarix penpentandrapentandriatandra and shrubs bac- wickenburg station charis salicifoliasalicifolia Hymenhymenocleahynwnocleaocleaociea monygyramonygyra tes the hassayampaHassayampa river preserve was histori- saria sericea and zizyphus obtusifolia 3 cally grazed and used recreationallyrecreation ally but both post flood seedling recruitment and vegetative impacts were eliminated in 1987 when the area reproduction of trees and shrubs and 4 was acquired by the nature conservancy changes in cover and composition ofherbaceous richter 1992 the system may still be recov- species secondary objectives were to compare ering from these prior impacts however there the effects of the loyear10 year return flood to those are no streams in the area that have been un- of smaller prior year floods including a 5 year grazed for long time periods with which the 120 GREAT BASIN naturalist volume 53
popultisfiemontiipopiiiitsfiemotitii I1 salix gooddinngooddinggoodgoodelingiidinn foiestforest willi bioPiobloprosopispiosopissopis saplings
A
prosopis velainavehaina woodland young pdpidus salix forest
baccharis salicifoliasalicifolia shrubland willi Popopullispullis salix saplings
watertableWater tabieTable scale 5 in
abandoned chchannelmnelamnel llymenoclea11yinenoclea nionogyramoiiog la river channel shriblandshrublandsluublShRi blandincl
fig 1 representativere pi chentcsent itiveetive cross section of ai portion of the hassayarnpahdssayampd river floodfloodplfloodplainplainun tree heights and depth to the water table aiealeare to scale
hassayampaHassayampa river ecosystem could be com rod per plot throughout the floodplainfloodplain depth pared the herbaceous understory contains was measured in august 1990 pre flood and many exotic plant species but the overstory late march 1991 and may 1991 post flood species are predominantly native eg P velu- fifteen of the rods could not be relocated after tina P fremonfrenwntiifremontiifremontiatiitil and S gooddingooddingiiii except for the flood relationships of sediment deposited a small component of T penpentandrapentandriatandra portions of or scoured by the 1991 flood with floodplainfloodplain the watershed are grazed by cattle and urban elevation ie height above the water table izediced processes that may result in creasedinincreased sedi- distance from the primary channel and woody ment yield or increased peak flow velocities plant stem density were determined with uni- von guerard 1989 Kondolkondolfkondolfandfandand keller 1991 variate nonlinear regression analysis multivari- leopold 1991 ate analysis was not utilized because variables were not independent eg stem density and METHODS floodplainfloodplain elevation the relationship between average sedimentation within the floodplainfloodplain data were collected on stream discharge and flow discharge was quantified with univari- floodplainfloodplain aggradation and degradation woody ate regression using data for the 1991 flood and plant survivorship and recruitment and herba- for five smaller floods in prior years stromberg ceous plant cover species richness and shan etetalal 19919911 non weiner species diversity these data were tree and shrub Survivorsurvivorshisurvivorshipshishl obtained during the 1991 flood year and during p and recruitment three prior years with smaller floods plant no mencmenclaturelature follows lehr 1978 stem density of woody plants was sampled within 100 permanent nested plots distributed flow data and floodplainFloodplain sedimentation throughout the floodplainfloodplain large trees 10 cm daily stream flow rate msmv1 during the stem diameter at a height of I1 in were sampled study was measured automatically at a stream in 1989 and after the flood in 1991 in 10 X 40 in gage located ca 060.6og linkinkm downstream of the per- plots density ofshrubs tree saplings plants 1 ennial flow emergence depth of sediment de- cm stem diameter at a height of I1 m and 1 yr posited or scoured by the 1991 flood was old and pole trees 1 10 cineinelncm stem diameter at measured by sedimentation rods ie steel re a height of I1 in was sampled in late march or inforcement rods at 100 permanent plots one early april 1988 1989198919901990 19919911 and again in 199311993 RESPONSE TO loyearIOYEAR10IO YEAR RETURN FLOOD 121 july or august 1991 in 2 X 2 m plots saplings j50 shrubs and trees were mapped in all years allowing for more precise calculation of post loo100 flood revegetation and annual survivorship in years with different flood magnitudes tree and L shrub seedling densities were measured j 0 monthly in 1991 in 4 X 4 dm plots to document t 10 post flood seedling recruitment survivorship of shrubs saplings and pole trees from 1990 to 1991 was analyzed in relation to several environmental variables stem den- sity floodplainfloodplain elevation and distance from the 0 N D J 1 1 A M i i A channel and water depth velocity tractive MONIH 01 iiiili111 WA I1 I1 k Y al shear stress stream power and sediment depos- fig 2 daily hydrograph of the hassayampahassayarnpaHassa yampa river during ited during the 1991 flood with nonlinear re- the 1990 91 water year data for 12 26 marchmarelimarell were not gressiongression analysis the flood flow parameters available were calculated from a calibrated HEC 2 flood plain model and floodplainfloodplain elevation and dis- RESULTS tance from the channel were determined from cross sectional surveys of the floodplainfloodplain strom berg et al 1991 survivorship was analyzed for flow data and floodplainFloodplain topography a composite data set of all shrub sapling and the 1991 flood had peak discharge of 368 pole tree stems and separately for three individ- 1 my 1 2 value S inasin3s on I march fig a about 3000 ual species P fremontiifrenwntiifremontiatiitil gooddinggooddingiiii and B 1 fremon times greater than the base flow rate oloi010.1 msm3s salicifoliasalicifolia survivorship was not analyzed for ns about 12 times greater than the 15 yr bankfull species with low mortality P velutina and 1 eg discharge 30 my and with a recurrence in- Z or those in fewer than 20 study obtusifolia terval of slightly less than 10 years jenkins monog plots eg H nwnogyraufaura and T sericea 1989a discharge remained above base flow herbaceous cover values through mid april the flood inundated nearly all of the floodplainfloodplain in contrast to a 5 year cover of herbaceous vegetation was esti- return flow that did not inundate high flood mated visually within 100 penpermanentnanentcanent plots 1 I1 X plains vegetated by P velutina table 1 peak I1 in distributed among several different over- flowflowvelocityflowvelocityinvelocity in the riparian zone in march 1991 story vegetation types herbaceous cover in four as calculated from the HEC 2 floodplainfloodplain model og 1 overstory types B salicifoliasalicifolia stands H ranged from 171.7 060.6 in s in the near stream og monogyramonogyra stands populus salix forests and P herbaceous vegetation type to 090.9 040.4 in s in velutina forests was sampled in march and the high floodplainfloodplain P velutina forests for these same areas 26 02 september 1988 1989 1990 and 1991 cover peak water depth was 262.6 020.2 in and 050.5os 030.3 in table 1 and tractive shear in the streamside herbaceous type was sampled 2 stress was 12712.7 737.3 kg in 2 and 343.4 343.4 kg monthly during these years to document rates in surface topography was altered during the of post flood recovery to compare within year flood as a result of deposition of sediment and effects of flooding herbaceous cover by species woody debris on floodplainsfloodplains scouring of sedi- was sampled I1 X 1 m plots in flooded and in ten ment from channel banks and creation of scour P forests unflooded velutina in march april pools along the main channel and in overflow and 1991 sediment jjuneune july depth of was channels the loyear10 year return flood deposited used as the indicator of flooding prosopis velu- more than twice as much sediment 8 cm as a tina forests were chosen for this analysis be- prior 5 year return flood 3 cm fig 3 depo- cause they occupied the highest floodplainsfloodplains and sition peaked maximum of47 cm on floodplainfloodplain encompassed areas with and without flood im- sites that were 1 2 in above the water table and pact mean cover and species richness per plot declined in bell curve fashion on higher and were statistically compared between flooded lower surfaces fig 4 this pattern differed and unflooded forests with studentstudentss t test from that for smaller prior year floods in which 122 GREAT BASIN naturalist volume 53
TABITABLE 1 1 water depth and flow velocity on the Hassahassayampahdssdyampdyampa river floodplainfloodpldinfloodplain during floods ofvarying recurrencei echiecui i enceenee intervals by riparian vegetation type values dreare means standardstdnddrd deviationdeviadevladevidtion
1 water depth in flow velocity in s
2 year 5 year loyear10 year 2 year 5 year loyear10 year vegetation type flood flood flood flood flood flood streamside herbaceous 160216 02 240224- 02 260226- 02 522052- 20 60-goit602727 551955 19 populus sahsalix saplings 110311- 03 190319- 03 210321 030.3 382338- 23 523452- 34 542254- 22 bartmasbaccharis saliciwhcifoliasalicifoliafolia 08080404 1505 17-170505 190619 06 27 07 331033- 10 populapopulwpopubts salix pole trees 08080404 16-160505 18-180606og 22-221616 34-342424 36-36361717 hipnenodeahymenocleaHymen ocleaociea monogyramono gyra 030.3030202 07-070404 090409- 04 23-231212 260326- 03 3212391232- 12 pps salixsailysnfoiestforest 04-040404 08080505 090609 06 24-241313 341334- 13 351235- 12 prosopis velutina forest 00 00OM 0000oooo00 00 050305 03 000000oooo00 000000-00oo 00 30-301414
2tat
20
10 Z
0 z 1 0
0 7 loo100 200 300noo 400 050 5 45 1 1 15 15 2 2 255 255 3 3553 j55S DISCI ARGI infrnarn5rn 5 H OODPLAINFLOODPLAN ILEVATION m
fig 3 sediment deposition in relation to flood magni- fig 4 sediment deposition means and standard devia- tude of the iiassayampahassayampaHassayampa river values represent average tion bars on floodplainsfloodplains ofofvariousvarious heights above the water sedimentsedini ent deposition in the floodplainfloodplain during the six largest table during a loyear10 year return flood in 1991 in the has floods in the period 1987 91 the regression equation is y sayampa river 0008 0022x r2ra 97 df 5 P 01
tion vegetation types with abundant woody with sediment decreased curvilinearly increas- stem density eg B salicifoliasalicifolia and P fremon ing floodplainfloodplain elevation stromberg et al 1991 hitii S gooddinggooddingiiii pole stands accumulated many low floodplainfloodplain surfaces 1 in high adja- more sediment than did types with lower stem cent to the main channel had a net loss of sedi- density eg H ffwnogyramonogyramono gyra stands table 2 ment in 1991 although these areas regained some sediment during small flood surges in tree and shrub survivorship april the channel bed became wider and shal plants on high floodplainsfloodplains lower in many areas but had deepened and again woody growing where flood were least had highest sur- become entrenched in areas by midsummer of impacts vivorship of the 1991 flood the composite 1991 for sample of shrubs saplings and pole trees sur- the 1991 flood sediment deposited during vivorship increased significantly as functions of was related to density of woody vegetation and flood water depth fig 5 floodplainfloodplain elevation to topographic position in the floodplainfloodplain flood and distance from the primary channel table plain elevation and woody stem density were 4 mature trees of P wlvelutinautina P fremonfrenwntiifremontiifremontiatiitil and negatively correlated rar2 2525.25 P oi01oi01.01 df 84 S gooddinggooddingiiii grew on floodplainsfloodplains higher than 2 and deposition was related to both factors t2tar2 in above the water table and had 100 survivor- ilii11.11 P 0202.02 df 84 positive relationship for ship table 3 saplings of P velutina grew pri- ra woody stem density and r2T lsis15.15 P oi0101.01 df marily in the understory of P fremonfremontiifremontiatii S 84 negative relationship for floodplainfloodplain eleva gooddinggooddingiiii stands and also had high survivorship 199311993 RESPONSE TO loyear10 YEAR RETURN FLOOD 123
TABLETABLF 2 depth of sediment deposited or scoured on the hassayampaHassayampa river floodpldinfloodplainfloodplain during floods of varying recurrence intervals by vegetation type mean floodplainfloodplain height above the water table distance from the stream channel and density of woody stems are indicated for each vegetation type values are means standard deviation
height above distance woody sediment cm waterwdlcjl from steinstern1 l in table channel densitydensitynesity 2 year 5 year 10 year m Wm no in hoodflood flood flood streamside herbaceous 040104- 01 444 4 23 27 98982727 128-12 8 474 7 112 110 populus salix saplings 07070303030.3 989 8 99 125 434343 t 43 54-543030 100-10 0 626 2 besbecbaccharisBacchenschans salicisahcifohasalicifoliafolia 10-100505 22 16 48 49 27273030 585557585757 134 122 populus salix poles 13-130505 222122 21 89 46 424249t 49 51 42 14714 7 125 Hymenhymenocleaoclea monogyramono gyra 2005202.0 050.5 35 9 20 21 09 t 18L 8 09 11 585 8 545 4 populus salix forest 220722 07 48 t 23 03 08 03 t 13 17 26 788278- 82 prosopis velutina forest 27 06 72 20 04 02 000000oooot 00 06 16 23-234242
82 pole trees ofsofjofsS gooddingriiniinil Pfremontiifremontiapfremontiifremontiitil loo100yoo and T penpentandrapentandriatandra grew on mid height flood plains 1 2 in above the water table and had 80 respective survivorship of 93 73 and 38 W tamarix penpentandrapentandriatandra was the only one of these three species that had much lower survivorship cn ofpole trees in 1991 than in prioryearsprior years saplings of these three species grew on floodplainsfloodplains 1 I1 in 35 above the water table and each had about 20 survivorship of the 1991 flood of the 1991 flood by poles and survivorship 0 saplings of P fremonfrenwntiifremontiifremontiatii was significantly related 0 05 0 5 I1 1 15 I1 5 5 to floodplainfloodplain elevation distance from the I1 LOODFLOOD waltaWATIRWATTR drpiiiDFP iiilii111 m 4 salix stream and depth of floodwater table fig 5 survivorship of riparian shrubs saplings and gooddinggooddineii survivorship showed the same small trees poles in the hassayampahassayarnpaHassayampa river floodplainfloodplain trends but relationships were not significant 1990 91 in relation to flood water depth classes populus fremontremonfrenwntiifremontiifremontiatiitil poles on floodplainsfloodplains 1 2 in above the water table had 94 10 survival dingiidingis saplings had greater survivorship than P compared to 60 40 for those on floodplainsfloodplains frenwntiifremontiifremontiatiitil saplings 1 high values for saplings were 54 46 fremon in survivorship by shrub species in 1991 corre- for the higher floodplainsfloodplains and 30 38 for the sponded to topographic position in the flood lower to flood water depth P with respect plain stem survivorship averaged 100 for and S gooddingiiii poles and saplings fremonfremontiifrernontiifremontiatiifil goodding zizyphus obtusifolia a species that grew on high showed a threshold response in which sur- og type floodplainsfloodplains 32323.2 060.6 in above the water table vivorship declined sharply where water was vegetated by prosopis velutina forests 80 15151.5 in deep fig 6 sediment deposition for H monogyramonogyra a species that grew on flood shear stress stream power and velocityvelocitywerewewerere not plains averaging about 2 in above the water significantly related to survivorship for either table and 20 for T sericea a low floodplain species between years annual survivorship for species 131.3 020.2 in that sustained much stem P fremonfremontiifrenwntiifremontiatiifil and S gooddinggooddingiiii saplings de- breakage stem survivorship averaged 50 for creased significantly as annual maximum flood B salicifoliasalicifolia the most abundant shrub in the magnitude increased with for example 30 of floodplainfloodplain this species formed dense stands P fremonfremontiifremontiatii saplings surviving the 1991 flood primarily on low floodplainsfloodplains ca I1 in high but 43 surviving the 5 year return flood in 1988 also grew in lesser densities on higher flood and 58 surviving during the 1 year return plains stem survivorship ofboflofbB salicifoliasalicifolia was not flood year in 1989 fig 7 in all years S good significantly correlated with any flood parameter 124 GREAT BASIN naturalist volume 53
tabietableTABI 1 3 annual survivorship of dominant riparian trees and shrubs in the hassayampaHassayampa river floodplainfloodplamfloodplainplam data are for years with a 5 year return flood 1988 89 I1 year flood 1989 90 and loyear10 year return flood 1990 91
size class survivorship or growth species form 1988 89 1989 90 1990 91
1 a1 prosopis vcvelutinavclutinavehitinavehlutinalufinaitina mature tree NSnsfns1 NS 100 srtnsafiasafixsransaan gooddinggooddingiiwcldiniiii mature tree NS NS 100 popithisfiremontiipojxdu frcmontn mature tree NS NS 100 prosopis vebitinavelutma pole tree 100 91 100 sahsaksahit gooddinggooddingiigooddmgnii pole tree 87 80 93 populusfiremontiipopulapopulm frcmontu pole tree 86 89 73 nainatatnaTainafamcimtainafixfixniy pcntandrapentandrapentandriapentandra pole tree 95 87 38 prowpipnsopis ventinavelutina sapling 76 87 82 sahsaksalix gooddinngooddinggooddingiigood dinudinnii sapling 64 78 36 populapopulusfreniontiipopulu freinontn sapling 43 58 30 tamarix pentapenpentarulipentandrapentandriatandraruliyuliyull a sapling 84 75 37 zizyphusAjphu obtusifoliaohtumfolia shrub 100 100 100 htiiilnodeahyntenoclea vinionogyraonora shrub 96 100 83 baccharissalicifoliabacchant alicifoliaalicitoliatollafolia shrubshishl abub 100 100 51 tessariasericeaflariafkariafK aria icruefi shrub 100 100 17 matlirctresM itiikitzik ti s haveli ii stnisetnissto ins 10cni1 cin dianieterdiadladi nieter pollpoilpole tins hivehave stenisstems 1 locinkocin diadjacilell iiticttit saplingssipsup lings hI iveuve stemsestemsesteems 1 lincin cliimltcrdim terandberandind aivgreaterthaUL gieitei th in I1 yeuyeny rfiafiin ageuge hnsNS notsiiiipldlnot pldaldpid
takitableTABI I1 4 regression coefficients r values relating dessicationdessication by the end of summer there were 5 survivorship of a 10 return 2 threetin ee physical parametersparame teis to loyearyear seedlings m on floodplamsfloodplainsfloodplamsplains 1 m above the flood in the hassayampaiiassayampaHassa yampa river by saplings and pole trees of 5 to populusfirenwntiipoduinpopuinPopuiupuin frenumfrenumtntn and sahsalix gooddingoocldingngooddingiiii and by a composite water table table a value sufficiently high group of riparian shrubs saplings and pole trees eventually produce a mature forest with charac- 2 teristicte density of 030.30 3 stems rn table 2 salix water gooddinggooddingiipooddingizii seedlings also germinated abun- distance height depth flomfrom above during dantly in 1991 after the flood pulse in Mmayay 1991 channel streamstreambedbed hoodflood there were 618 S gooddingngoo&1ingii seedlings m 2 inm plots 1 m above the water table table 5 P fninoutupfiremonth survivorship 0200 20 0250 25 0450 45 seedlings by the end of summer were most S gooddinggooddingiigooddinguii survivorship 002 010 010 abundant on floodplainsfloodplains 040.40 4 060.60og 6 m above the composite riparian survivorshipsurvivorship011oiioli011 001313 001313 water table tanwrixtamanxtamana penpentandrapentandriatandra genninagerminatedted in september after P fremontiatiitiltn march 1 june fremontremonfremontiifremontn 11l M narnnann ooi00001 april and S gooddinggooddingiigooddingttiitt april may tarnarixtamanxtamaneTarnarix penpentandrapentandriatandra had maximum seedling density of 5 t 13 m 2 in june 1991 but none were alive by tree and shrub revegetation the end of summer baccharis salicisalicifoliasahcifohafolia stems recovered to pre- 2 in 1991 the germination period pfrenwnofpfremonof flood densities 484844.88 454.54 5 m measuredadthinmeasured within hitu march april coincided with the period of B salicisahcifohasalicifoliafolia vegetation zones by july 1991 peak floodplainfloodplain inundation As a result P fre primarily a result of stem sprouting and in part numtiirrumth seedlings had high density and broad a result ofseedling recruitment stem density of distribution throughout the floodplainfloodplain in april Hymenhymenocleaoclea monogyramonogyra increased by late sum- P fremontiifremontiatii seedlings were on floodplainsfloodplains up to mer 1991 to a value somewhat higher than pre fremon 2 35353.5 in above the water table fig 8 and 99 m flood levels 23232.32 3 252.52 5 stems m as a result of from the main channel as well as along stream vegetative reproduction tessafiatessanagessanaTessafiasahiasahna sericea also banks and overflow channels seedling density had post flood vegetative spread but stem den- declined steadily throughout the growing sea- sities had not attained pre flood levels by late son of 1991 with mortality due primarily to summer 199319931 RESPONSE TO loyear10IOYEARIO YEAR RETURN FLOOD 125
loo100
SALIX GOODDING 11 POPU LU S I1 remoRLMO nf11 80 75
CL 60 T
0 o
40
0 SALIXsalSAI IX SAPLINGSAFILING 25 20 SALIX pallpanlPOLIPOLL 0 POPUPOPUI1 US SAPHNGsael inoING POPUPOPUI US POPOIFF
0 0 0 a 0 0 L L 0 150 00oo 00.00 5 0 400 00 05 10 1 5 20 25 30 35 50 100 10 20000 vl50 FLOOD WATER DEPTH m DISCI aroIARGLARC msm5sm3s fig 6 survivorship of saplings and small trees poles of fig 7 annual survivorshipsumrivorship of saplings of populus fre populuspopulusfremontiifremonfremontiifremontiatiltii and salixsabby gooddinggooddingiigoodclingiiii 1990 91 in relation montu and salix gooddinggooddingiigooddingnii along the hassayampaHassayampa river to maximum water depth during a loyear10 year return flood floodplainfloodplain in relation to maximum annual flood flow latetaterate regression equations are y 59 008x0 08x rr2ra2 99 df 2 P 01 P fretnontnpfiremonffl and y 82 012x0 12x r2ra 97 df 2 400 P 05 S gooddinggooddingngooddingiiii rEMARCHJ wanchwancoMARCH MAY MJULY cover in these OCTOBER monspeliensisntonspeliensis areas increased nearly to preprefloodflood levels by september cover 300 within higher elevation vegetation types eg z populus salix forests remained low as of late summer within P velutina forests areas that loo100 z were flooded had lower cover but greater rich- ness and diversity of species throughout the ln summer compared to areas that were not flooded table 7 unflooded and flooded areas 0
1 1 1 5 0- 5 1 151 15 27 27 252 255 3 35 in the PF velutina forest were both initially domi- FLOODfloodplainfloodpiainrhainPLAIN IIEGHTHE shiGHT rn nated by two exotic winter germinating annuals hordeum leporinumleponleporinum and sisymmumsisymhriumsisymbrium irio fig 8 density of populusfretrwntiiPopulus fremontremonfremontiifremontiatiitil in relation to water these two species continued to dominate un- table depth by month during 1991 flooded areas throughout spring and early sum- mer flooded areas in contrast had about l6th16thlath the cover of unflooded areas and about 454 5 herbaceous cover 2 times as many species eg 929.2 191.91iglg 9 MrnvsravsVS 191.91lgig9 2 spring herbaceous cover in all vegetation 05os050.5 in april data these included severaseveral1 types except that of the highest floodplainsfloodplains P native annual forbs eg amaranthus palmeripalmere bowlesiaBowlesia amsinckiaarnsinckia interra gilia velutina forests was less abundant in 1991 than incaniincana interintermediainterraediaintermedialnedianedlamediaediaedla cilia sinuatasinuate humistratushumihumistratousstratus microserisMicro seris linearilineard in prior years table 6 herbaceous cover un- lotus jolia xanthium strum and P S foliajaliajaila strumahumstrumariumstruaHumanummarium verbesina der gooddinggooddingiiii forests I1 for exam- frenwntiifretnontii encelioidesencelioides and several exotic annual forbs and ple averaged 8 in 1991 compared to 25 43 grasses eg bromus rubens Herniherniariaatia cinerea on streamstream in prior years herbaceous cover solanum rostratum and tribulus terresttisterrestrisferreterrestris banks and in B salicifoliasalicifolia stands was 16 and I111 I1 respectively in late march 1991 compared to 38 and 34 in the prior year cover in these discussion two areas was composed primarily of rhizoma- tous grasses the native paspalum distichumdistichum and riparian systems are noted for their resil- the exotic cynodon dactylondaktylondactylon and also con- iency ie the ability to quickly return to predispre dis tained lesser amounts of other natives eg ty- turturbancebance conditions rapid growth rates high pha domingensisdomingensis and species of juncus and fecundity and capacity for asexual reproduction exoticsexotica eg melimeilMetimelilotuslotusiotus albus and polypogonPolypogon are among the factors that allow rapid recovery 126 GREAT BASIN naturalist volume 53
2 taini11 5 average seedling densities no m during the period of maximum abundance and at the end oftheodtheof the growing season fortor three riparian tree species on hassayampaHassayampa river floodplainfloodplamfloodplainplampiam surfaces 1 m above the water table
pop wits1 fretkntiifrcimmtn salix gooddinggooddingiigooddingnii tanuinxtarnaristarnarixTarnarixarly penpentandrapentandriatandra apiilapril october may october june october
1988 illliilil111 0 385 0 11 0 1989 12 0 2 0 1 0 1990 0 0 1 0 0 0 1991 205 5 618 3 5 0
IABIITABLE 6 herbaceous cover W along the iiassayampahassayampaHassayampa river floodplunfloodplainfloodplain by vegetation type from 1988 to 1991 values areai means t standard deviation
1988 1989 1990 1991 streamside herbaceous march 202420 24 293629 36 38 31 16 31 april 38 26 662866 28 683068 30 29 37 may 3825138- 25a 682868 28 7229728972 29 35 41 july 662266 22 74 30 43 30301 41 38 sept 693469 34 773077 30 60 25 603560 35 baccharissalicifoliabacchant wlicifoha shrubland marchmaich 22 18 19 19 34 26 11 24241 sept 38 36 41 35 25 21 41 43
Hymenhymenocleaainuhinu ocieaocleanocten monogiframonogyramonoayragyra shrubland march 19 16 202420 24 24 21 4 441 sept 7 11 15 t 13 12 11 13 13
a popiilwpopubts salixsalsaa foiestforest march 43 27 25 21 34 31 8 12121 sept 19 29 21 28 21 23 565 6 prosopis vdutmavehitinanebvehitina forestfontonjon st marchmaichmalch 84 17 352135 21 58 26 52 45 sept 7 12 12 16 10 17 6 13 post foodhoodflood salplillgsiniplinlitdatess
taboTABH1 1 1 7 covelgovelcovergover richness and shannon weinerwelner diversity of herbaceous understory species min prosopis velutinavehitina forests that weiewelewere and were not inundated during a I1100 year return hoodflood in march 199iggi19911 values for cover and richness are means standardstan daiddald deviations
species covergoverover species richness diversity month flood no flood flood no flood flood no flood marchmaich 77 71 30130 23230505 19-190606 053 091 april 18 t 4 79 36136 929 2 19 19 t 051oso0 5 246 073 lunejuneune 12 6 84-84240241 loi10110 1 313 1 22-2 2 0810 8 253 045 july 16916 9 88 illlio11 797 9 30 262 6 2312 3 231 123 suliksnllksignificantsigniflcantlllilftikdifficrowcat 05 of riparian plants after disturbance stromberg the loyear10 year return flood in the hassayampaHassayampa and patten 1989 gecy and wilson 1990 river inundated most of the floodplainfloodplain and de- densely vegetated floodplainfloodplain ecosystems also posited a net average of 8 cm of sediment maxi can be resistant to floods in the sense that floods mum of 050.5os m low elevation floodplainfloodplain pass over them without scouring vegetation or surfaces had greatest flow velocities to 7 m s 1 substrate hendrickson and minckley 1984 and water depths to 282.8 m the native riparian 199311993 RESPONSE TO loyearIOYEAR10IO YEAR RETURN FLOOD 127 vegetation showed a mixture of resistance and flood stromberg et al 1991 this present resilience to this flood disturbance species on study confirms the role oflarge floods in increas- high floodplainsfloodplains eg P velutina and Z obtusiobtuse ing age class diversity for these episodically re- folia had no mortality while those on lower ele cruitingcr species vation floodplainsfloodplains variously had mortality the exotic T penpentandrapentandriatandra co occurred with followed by seedling recruitment P fremontremonfrenwntiifremontiifremontiatiitil populus and salix but had greater mortality of and S gooddinggooddingiiii or by vegetative reproduction pole trees than did the native trees mortality of eg baccharis salicifoliasalicifolia T penpentandrapentandriatandra more likely resulted from intoler- prosopis velutina was the dominant tree on ance to physical flood effects than from physiphaysi high floodplainsfloodplains ca 3 m above the water table ological intolerance to inundation warren and and had high survivorship of trees and saplings turner 1975 irvine and west 1979 tamarix it did not show post flood seedling recruitment penpentandrapentandriatandra had low post flood seedling estabbestab consistent with prior studies indicating that P lishmentlishment due in part to a low density of mature velutina seeds germinate primarily after late seed producing trees in the hassayampaHassayampa flood summer floods stromberg et al 1991 populus plain and in part to the fact that the flood oc- fremonfremontiifremontiafiretrwntiitii and S gooddingii trees grew on flood curred several months prior to T penpentandrapentandriatandra plains 2 3 m high and also had high survivor seed germination and thus did not moisten po- ship young trees and saplings of these two tential gengerminationninationbination sites at an appropriate time species were on younger less gradedaggradedag flood june through october additionally much of plains and sustained some mortality salix good the available germination space during its ger- dingiidingis saplings and poles had lower mortality minationmi period was preempted by herbaceous than did populusfrenwntiipopulus fremonfremontiifremontiatiifil perhaps because of cover and by seedlings of P fremonfrenwntiifremontiifremontiatiitilfil and S greater stem pliability and tolerance to satura- gooddinggooddingiiii species that precede tamarix pen tion mcbride and strahan 1984 hunter et al tandra in the chronosequencechronosequence of tree species 1987 survivorship of both species was greater germination at the hassayampaHassayampa river on sites where flood waters were shallowistshallowestshallowest a vegetative reproduction is a common post factor reported to be an important determinant disturbance revegetation mechanism in flood of flood survivorship in other riparian systems plain systems gecy and wilson 1990 and was stevens and waring 1988 the relationship demonstrated by all shrub species in the has between water depth and survivorship may be sayampa river floodplainfloodplain that had flood mortal- an expression of effects of flood hydraulic force ity extent of flood mortality of shrub species at on plant removal or mortality via abrasion and the hassayampaHassayampa varied with their topographic stem breakage rather than of a causal relation- position in the floodplainfloodplain zizyphus obtusifolia ship between root saturation and mortality al- a species of high floodplainsfloodplains ca 3 m above the though correlations of mortality with flood water table had no mortality Bacbaccharischarischarls salici velocity and shear stress were not statistically folia undenunderwentventment a 50 decline in stem density significant this may have been due to chaotic during the flood but increased to pre flood den- movement ofwater and sediments on the flood sities by late summer primarily via stem sprout- plain which are not adequately represented by ing Hymenhynwnocleahymenocleaocieaoclea monogyranwnogyramonogyra and T sericea are flood simulation models such as HEC 2 both clonal shrubs that spread via root sprouts the 1991 flood created optimal seedling re- after mechanical injury gary 1963 and via cruitmentcruitment conditions for populusfrenwnffipopulus fremonfremontiifremontiapremontiitil and shoot sprouts after stem burial Hymenhynwnocleahymenocleaoclea salix gooddinggooddingiiii by scouring channel banks and monogyramonogyra compensated for flood mortality by depositing new sediment on stream banks re vegetative reproduction but this was not the dueing herbaceous and overstory competition case for tessariatessatlatessoriaTessatiasarlasariasafiasaTla sericea a low floodplainfloodplain spe- at least temporarily and moistening flood cies that had high flood mortality other studies plains at an appropriate time during seed dis- also have reported low flood survivorship for persal and place moderately high surfaces tessariatessoriaTessaria sericea stevens and waring 1988 above the zone offrequent summer flood scour vegetative reproduction also was the domi- stromberg et al 1991 tree ring studies have nant revegetation method for herbaceous plants shown that P fremonfremontiifremontiapfretwntiitiifil and S gooddinggooddingiiii estab- along stream banks and low elevation flood lish in large scale about once a decade within the plains cover in these areas declined by about hassayampaHassayampa river system during or after years half after the flood but recovered to preprefloodflood with large flows 250 msmv1 7 year return levels by late summer flood tolerant perennial 128 GREAT BASIN naturalist volume 53
rhizomatous grasses P distichumdist ichum and C dactydacay established in 1959 1952 and earlier strom- lon dominated these areas during and prior to berg et aal 1991 the loyear10 year return flood prob- the flood but species less tolerant of high flow ably reached a geomorphic threshold that velocities eg typhatypha domindominqensisdomingensisgensis may altiulti being the level at which substantial change in mately increase in abundance during flood floodplainfloodplain morphology and vegetation begins to interim periods fisher et al 1982 hen- occur based on studies of other desert rivers dricksondr and minckley 1984 understoriesUnderstories of that implicate the 5 year return flow as a thresh- high elevation floodplainsfloodplains showed changes in old discharge for channel and floodplainfloodplain instaansta cover and composition after the 1991 flood bilityability graf 1983 prior to the flood P velutina forests were domi- other potential effects of the flood on ripar- nated by dense nearly monotypic stands of ex- ian vegetation such as changes in plant produc- otic annual species eg hodeumhordeum murinusmurimurinumnum tivity as a result of nutrient or water pulses were that probably had become established during not addressed in this study nor was the role of past years of cattle grazing and other exogenous vegetation in moderating flood processes exilicexplic disturbances wolden et al 1991 after the itly addressed flood these areas had lower cover but greater data in this paper suggest that floodplainfloodplain richness of herbaceous species and greater rela- vegetation aided in stream bank stabilization tive abundance of native annuals we speculate and sediment trapping important functions of that compositional changes were due to reduced wetland and riparian vegetation fisher and competition with entrenched exoticsexotica an influx minckley 1978 cooper et al 1987 sullivan and of flood bomeborne seeds from upstream areas or stromberg 1992 the vegetation also may have other vegetation types within the floodplainfloodplain or enhanced groundwater recharge and reduced altered edaphic conditions resulting from depo the downstream impact offlood flows by redue sitionaition of sediment with different texture or nu ing flow velocities and increasing water reten- trient content stevens and waring 1988 tion time within the floodplainfloodplain burkham 1976 fluvial processes including floodplainfloodplain aggra- beschta and platts 1986 dation and formation of micromicroreliefrelief patterns eg backwater depressions contribute to the acknowledgments diversity and mosaicism of riparian plant com- munitiesmunities in many flood driven ecosystems kal we thank the nature conservancy for al- liola and puhakka 1988 within the lowing the use of the hassayampaHassayampa river pre- hassayampaHassayampa floodplainfloodplain as well variable sedi- serve as a research site review comments by ment deposition and scour patterns contributed W carter johnson and anonymous reviewers to 11 patchiness within the riparian floodplainfloodplain are greatly appreciated for example localized light gaps were formed in areas with major debris deposition and scour literature CITED pools ie backwater depressions were formed along main channels and in overflow channels BELL D T 1974 freetree stratum composition and distribu- floodplainFloodplain sedimentation accentuated the ex- tion in the streamside forest american naturalist 92 istence hydrologicalofhydrologicalof gradients eg gradients 35 46 BFSCIIIA R L annAND W S PLAIIS 1986 morphological of depth to groundwater which contribute to LANDW plalis significance ofsmallof smallsmail streams significance and function floristic diversity within riparian ecosystems of water resources bulletin 22 369 379 the hassayampaHassayampa river and elsewhere hupp BRAVARD JCJ C AMAROS AND G PAUIOU 1986 impact of and ostercamp 1985 bravard et al 1986 civil engineering works on the successions of commu- fluvialfluvijsystem stromberg et al 1991 nities htain a system oikos47oikosbikos 47 92 111 BuRKIBUKKIIAMIAM D E 1972 channel changes inm the gila rivelriver the 10 year return flood resulted in greater in safford valley arizona 1846 1970 united states floodplainfloodplain aggradation and greater woody plant geological survey professional paper 6556 1 24 mortality than smaller prior year floods effects 1976 hydraulic effects of changes in bottomlandbottomland on three floods gila river south- of floods than the 10 vegetation major in larger year event can only western arizona USU S geological survey professional be speculated upon A 100 year return flood in paper 655j 1 14 1970 in the hassayampaHassayampa river jenkins 1989a COLLINS J P C YOUNG jilJH J HOWELL AND W L apparently did not cause extensive loss of ripar- MINCKLEYMINCKLFY 1981 impact of flooding in a sonoran desert stream including elimination of an endangered ian vegetation as indicated by the presence of fish population peocihopsipeociliopsis 0 occidentaloccident alisallsails poecilii- extensive stands populusofpopulusof and salix trees that dae southwestern naturalist 26 415423415 423 199311993 RESPONSE TO loyear10IOYEARIO YEAR RETURN FLOOD 129
COOKICOOKE R U AND R W RFFVFSREEVES 1976 arroyos and envi- proceedings on headwaters hydrology american ronronmentalmental change in the american southwest claren- water resources association bethesda maryland press don oxford jonnsonJOIINSONJOHNSON WCW C R L BURGESSBURGFSS AND W R KFAMMIKEAMMERERRI R COOPERCOOPFR RRJJ W gilliam R B DANIFLSDANIELS AND W P 1976 forest overstory vegetation and environment on robargeROBARGFROBARCF 1987 riparian areas as filters for agricul- the missouri river floodplamfloodplainfloodplamplainpiam in north dakota ecol- tural sediments soil science society of america jour- ogical monographs 46 59 84 nal 51 416 420 JUNK W J P B BAYLEYBAYLFY AND R E SPARKS 1989 the DFBANO L F AND L J SCHMIDT 1990 potential for flood pulse concept in rivenver r floodplain systems cana- enhancing riparian habitats in the southwestern united dian special publications in fisheries and aquatic sci- states with watershed practices forest ecology and ences 106 110 127 management 3334 385 403 KALLIOLA R AND M PUPUIIAKKAIIAKKA 1988 river dynamics and FISIIFRFISHER S GGANDWLAND W L MINCKLEYMINCKLFY 1978 chemical char- vegetation mosaicism a case study of the river kainakamakalna acteristicsacteristics of a desert stream in flash flood journal of jockajohkaohka northernmost finland journal of biogeography andaridarld environments 1 253325 33 1570315 703 719 FISIIFRFISHER S G GRAY B GRIMM GLL J N AND D E buscilbusciibuscai KONDOLF G M AND E A KFLLFRKELLER 1991 management 1982 temporal succession in a desert stream ecosys of urbanizingurban izing watersheds california water resources temterntermtemm following flash flooding ecological monographs center report 75 27 40 52 93 110 LEIIII J H 1978 A catalogue of the floraotfloraofflora of arizona desert GARY H L 1963 root distribution of five stamen tamarisk botanical garden phoenix arizona seepwillowwillow and arrowweed science 9 311 seep forest leoboldLEOPOLDLEOPOLDL L 1991 hydrology and physical effects of urban- 314 ized watersheds california water resources center arcygrcy L AND M VVWILSON 1990 GFCYJJ WILSON initial establishment report 75 13 15 of after disturbance by debris flows riparian vegetation LISLFLISLE T E 1989 channel danaidynaidynamicnic control on the estab- in oregon american midland naturalist 123282123 282 291 lishmentlish ment of riparian trees after large floods in north- GORDON ISII SIIALOM N AND Y GUTIFRMANGUTTERMAN 1989 cordon western california USU S forest service general survival of the typical in a wadi bed canyon vegetation in technical report PSW 110 9 13 after a violent flood at en moor waterfall area in the in LONGLONC M C 1982 white alder alnus rhombifoliarhombifolia re- centralgentral negev desert pages 423 431 in E spanier Y growth following 1968 1969 floods crossosoma 8 1 steinberger and M eds environmental quality lunalurialurla 3 and ecosystem stability vol IVBIV B ISEEQS pub jeru- RIDE R AND SIRAIIANSTRAIIAN 1984 and salem israel mcbmabmcbludf J randlRANDJann J establishment survival of woody riparian species on gravel bars of an GRAF W L 1983 flood related channel change in an aridarldai id intermittent stream american midland naturalist 112 river earth surface processes and landformsLandforms region 235 245 8 125 139 MINCKLEYminoMINC kleyKLFY W L AND T 0 CLARK 1984 formation and 1988 definition of flood plains along arid region destruction of a gila bosque pages 231 242 mvin V R baker R C kochel and river mesquite commitcommu- rivers plants P C patton eds flood geomorphology john wiley nitymty desert 6 23 30 and sons inc PAUIOUPAUTOU GGJJ cirelgirel J L borelBORFL 0 manneivilleMANNIMANNE VILLIIVILLE AND J 1991 changes in floodplainfloodplamfloodplamplainpiam hendrickson D A AND W L MINCKLEYminoMINC KLFY 1984 cllalfmonrciialemont in vegetation caused by basis for cienegasCienegas vanishing climax communities of the damming a predictive diagnosis pages american southwest desert plants 6 131 175 126 134 in 0 ravera ed TerrterrestriateirestnalterrestrialestriA and and howe W H AND F L KNOPF 1991 on the imminent aquatic ecosystems perturbation recovery ellis horwood west sussex decline of rio grande cottoncottonwoodswoods in central new ltd england mexico southwestern naturalist 36 218 224 plaitsFLAIISPLATTS WSKW S K A GFBIIARDI annANDANDWLW L JACKSON 1985 effects oflargeof large storm events on basin range HUGHES F M R 1990 the influence of floodingoffloodingoffloading regimes the ripar- ianlan general on forest distribution and composition in the tana ian stream habitats USDA forest service river floodplainfloodplain kenya journal of applied ecology technical report RM ibo120 30 34 2747527 475491475 491 reiciienbaciierreioreloRFIC iifnbaciifr F W 1984 ecology and evolution of hunlenHUNIFRHUNTER W C B W ANDERSONANDFRSON AND R D olimarlOIIMART southwestern riparian plant communities desert plants 6 15 1987 avian community structure changes in a mature 22 floodplainfloodplain forest after extensive flooding journal of rellyRFILYREILY P W AND W C JOIINSONJOHNSON 1982 the effects of wildlife management 51 495 502 altered hydrologic regime on tree growth along the HUPP C R AND W R OSIFRCAMPOSTERCAMP 1985 bottomlandBott omland missouri river in north dakota canadian journal of vegetation distribution along passage creek virginia botany 60 2410 2423 in relation to fluvial landformslandforms ecology 66 670 681 RICIIIIRicIRICIIIIRlterITERR H 1992 development ofaof a conceptual model for IRVINEIRVINF J RRANDNAND N E weslWFSIWEST 1979 riparian tree species floodplainfloodplamfloodplamplainpiam restoration andaridarld lands newsletter 32 13 distribution and succession along the lower escalante 17 river utah southwestern naturalist 24 331 346 ROOD S bandjbandlbabdjB AND J M MAHONEYMAIIONFY 1990 collapse of ripar- JFNKINSJENKINS M E 1989a ground and surface water assess- ianlan poplar forests downstream from dams in westeinwestern ments supporting instream flow protection at the has prairies probable causes and prospects for mitigation sayampa river preserve wickenburg arizona environmental management 14 451 464 unpublished mastermasterss thesis university of arizona SKOGLUND S J 1990 seed dispersing agents in two regu- tucson larly flooded river sites canadian journal of botany 68 1989b surface and groundwater assessments sup- 754 760 porting instream flow protection at the hassayampapiassayampaHassayampa SMIIIISMITII R L 1980 alluvial scrub vegetation of the san river preserve wickenburg arizona pages 307 316 gabriel river floodplainfloodplain california madrono 27 126 inainwin W W woessner and D F potts eds symposium 138 130 GREAT BASIN naturalist volume 53
SPARKS R E PR B BAYLIBAYLEY Y S L kolllikollaikohlerK AND L L Os the society for wetland scientists 13th annual meeting 13 0 H N E 1990 distudistilbistu rbanceroance and recovery otof large flood in press plain rivers Environmentenvironmentaldl management 14 699 709 VON guebardGUFKARDGUERARD P 1989 effects oflandeflandof land use on sediment STEWENs L ELANDGAND G L WARINGWAKING 1988 effects of post yield southeastern colorado pages 223 241 in W W iainlaindaindarn flooding on riparian substrates vegetation and woessner and D F potts eds symposium proceed- invertebrate populations in the colorado river corri- ings on headwaters hydrology american water res- dor in clandgiandgrandgland canyon USDI bureau of reclamation ources association bethesda maryland glen canyon environmental studies report 19 wahrenWAHRFNWARREN D K AND R M TURNER 1975 salt cedar SIKOMBISTROMBERG J CANDCANDDD T palirnpkrtenpaliern 1989 earlyrecoveryearly recovery tamarixtamanytamony chinenchmensischinensissis seed production seedling estab- of an eastern sienaslenasierra riparian system following forty lishmentlishment and response to inundation journal of the years of stream diversion USDS forest service gen- arizona academy of science 10 135 144 eral technical report PSW iloiio110 39939904404 WOLDENWOLDFN LJ sirombfrgsriiomberg D PATTENANDpatienpatlenPAIIFN AND H RicRKIIIFRilterlITER 1992 mortality aldageandageand age of black cottonwood stands 1991 understory restoration in three riparian forest along diverted and undiverted streams in the eastern types arizona restoration and management notes sienslensierra a nevada california madrono 39 205 223 8 116 117 STROMBERG J C D T pailen AND B D RKIIIFRRICHTER 1991 flood flows and dynamics ofot sonoran riparian received I1 november 1991 forests rivers 2 221 235 accepted 1 december 1992 suisulSULLIVANi IVAN M E AND J C sirombnustrombf11g 1992 wetland functions in southwest riparian forests proceedings of great basin naturalist 532 appp 131 136
HABITAT SELECTION OF MERRIAMS TURKEY MELEACRISMELEAGRIS CALLOPAVOGALLOPAVO MERRIAMI HENS WITH POULTS IN THE BLACK HILLS SOUTH DAKOTA
mark A rumble 1 and stanley H anderson 2
ABSTRAGEai nacinaclRACI we studied hibithubithabitat it selection patterns of mehmerriammen urn s turkey meleagrisMele agns gallopavogallopavo mernaminwrriconimernami hens with boultspoults in a ponderosa pine pinus ponderosa ecosystem thirty six radioradlo mariedmarked hens produced 19 broods and we obtainedabtobt lined 230 locations oftensofbensof hensbens with boultspoults we described vegetation of habitats using entencriteriai from the rocky mountain region USU S forest service for determining effects of forest management and monitoring of wildlife populations most habitat units were 443232 ha and corresponded to third orderordel habitats as described by johnson 1980 hensliensilens with boultspoults selected large meadows and rarely selected dense ponderosa pine habitats younger boultspoults used meadows more frequently than did older boultspoults implementation of the black hills national forest land and resource management plan in pondeiosapincponderosa pine habitats will not negatively impact hens with boultspoults grazing by livestock reduces herbaceous biomass necessary for invertebrate food items of boultspoults and cover for boultspoults habitat selection patterns ofhens with boultspoults should be evaluated by age categories of boultspoults
worelsworcls selection keykegkeywords merriammernamss turkeys meleagrisMele agns gallopavogallopavo meimelmerrimerrlmerriaminamiami boultspoults habitat I1 forestfirrestfibrest inanweinentmanagement inverteinverne brates ineameadowsdows radioracho telemetry grazing
merriamMerriamslamss turkeys meleagris gallopavogallopavo phasis on old growth timber values and im- merriaromerfimerrimerriarmnwrriamiarm historically ranged as far north as provementsprove ments in technology of harvesting timber southern colorado when settlers moved into the have renewed concern for the habitat require- southwestern united states macdonald and ments of merriam s turkeys shaw 1986 in jantzen 1967 in 1948 1950 and 1951 wild addition to loggers other users also are increas- turkeys were transplanted to the black hills of ing their awareness and use of national forest south dakota in three separate releases of881515 lands and 6 birds respectively petersen and richard the objective of this study was to describe son 1973 by 1952 estimated turkey popula- habitat requirements of turkey boultspoults in a pon- tions in the black hills of south dakota and derosa pine ecosystem at the same resolution wyoming were 1000 birds and by 1960 popula- level as that used by the US forest service in tion estimates were 5000 7000 birds petersen making management decisions and monitoring and richardson 1973 suggesting excellent re- the effects of those decisions on wildlife productive potential in habitats of this region current pressures from society and statutory METHODS mandates eg national forest management act renewable resources planning act na study area tionaldional environmental policy act require that the effects of management activities such as this study was conducted in the central grazing and timber harvest on the various wild- black hills of south dakota approximately 16 life species be considered in management deci- km west of rapid city most of the land is under sions recently the value of ponderosaponderosapinepine has management ofthe black hills national forest increased placing greater demands for logging pactolapacoola ranger district although private hold- in the black hills G gire silviculturist black ings associated with ranch operations and sev- hills national forest personal communica- eral private homes and cabins exist in the study tion increased value of timber resources em area
IUSDAUSDA foiestforest simceservice 501 E st joseph st southS th dakota school ofofmineschiesmhies rapidityrapid0tyrapid ityliy south dakota 57701 rusdi2usdiSDI coopelchopelcooperativcooperativeitiveetive fishedesandfisheriesFishe desandand wildlife researchRese aldiaidi unit univeisityuniversityulli sity of wyomingofwyoiniiig nailiranijnar imieimle wyomingnvyanvy ing 82071
131 132 GREAT BASIN naturalist volume 53
vegetation in the study area is primarily pure coordinates were recorded to the nearest 100 in ponderosa pine forest 84 meadows and as- for each location only one location was re- pen populus tremuloidesvcdtremuloidesybirch betula corded for each bird on any given day we con- papirpapyrpapiriferapapyriferaifera habitats occur in drainages but sidered observations of more than one somesorne monotypic aspen habitats also occur on radio marked bird at a location as one observa- north exposures bur oak quercus mcilromcicrotnacro tion to assure independence among observa- carpa and white spruce picea glaucaglanca consti- tions alldredge and ratti 1986 each location tute less than 1 of the study area climax of a hen with boultspoults was assigned to a corre- vegetation or potential natural communities in- spondingsp compartment and stand see habitat clude the following pinus ponderosalsymponclerosasym descriptions below we collected habitat use phoricarpos albus P ponderosaponderosaarctostaphylosponlerosalarctostaphylosArctostaphylos data of hens with boultspoults over a three year pe- umuva ursi P ponclerosajuniperusponderosaponderosaijuIjunip efuserusevus communiscommoniscommunis riod 1986 1989 during the months of june populus tremuloidescorulustrentuloideslcorylus cornuta quercus september macrocarpal0stryanwcrocarpaostrya virginiajavirginianavirginiana Q macro carcarpaspalSpays albus and picea glaucalinnaeaglaucallinnaea bore- habitat descriptions alis types as described by hoffmanHoffinan and alexander 1987 habitats were detendeterminedaetennineddined by US forest service criteria and assigned to numerically geologic material is predominantly precanprecam identified geographical units boundaries were brian and cambrian granite schistsschiets and meta defined by watershed topography ridges and sediments homhoffmanfinanminan and alexander 1987 Hof drainages or distinct changes in vegetation elevation ranges from approximately 1300 to type private lands in the study area were as- 1800 climate is continental with cold win in signed to habitats based on interpretation of ters and 1959 is warinwarm summers orr january aerial photographs boundaries of adjacent habi- typically the coldest month with mean tem tats were extended if the vegetation type was ture extremes of I111 1 ac and peraperaturepesature 2c july august continuous new boundaries were assigned if are the warmest with mean ex- temperature changes in vegetation were apparent typically tremes of 15 average annual 29c precipita- these habitats are 4 32 ha land units although tion is 50 55 cm which 70 80 falls between of smaller size habitats inclusion habitats were april and september south dakota clima delineated if distinct vegetation types could be tological summary no 20 39 6 20396 US weather identified on 124000 aerial photographs in- bureau snowfall averages 84 cm but may clusion habitats were riparian areas meadows range from 25 to 200 cm aspenbirchaspen birch draws and monotypic aspen com- ties hundred habitat trapping and locations munitiesmuni five thirteen units encompassing 4380 ha were delineated on the we trapped wild Merrmerriamsiams turkeys during study area late february or early march ofof19861986198619871987 and vegetative descriptions of each habitat unit 1988 with rocket nets and drop nets over corn were obtained from five plots uniformly as- bait piles thirty six females were fitted with signed to each habitat unit and marked on backpack radio transmitters weighing approxi- 124000 contour maps in the laboratory plots mately 108 g radio transmitters were attached in each habitat unit were then located using to hensliens with 15215.2 cm bungee cords looped un these maps diameter breast height dahdbh of der the wings trees and tree basal area were estimated at each hen turkeys were monitored until behavior plot using a 10 factorprismfactorbactor prism sharpe et al 1976 or a constant signal pulse suggested the hen had when habitat units were too small to effectively initiated a nest after eggs hatched radio place five plots fewer plots were used marked hensliensilens were located three times each habitat assignments were made based on week we attempted to obtain one location in dominant species of vegetation DSV and over- each of three time periods sunrise to 0959 story canopy cover OCC buttery and gillam 1000 to 1359135914001400 to sunset each week turkey 1984 criteria for describing habitats allowed locations were determined by plotting bearings for further stratification by dahdbh but the most from known locations on USGS 124000 con useful resolution level in determining habitat tour maps in the field bearings were usually selection patterns of turkeys included DSV and taken from positions within 300 in of the esti- OCC rumble and anderson 1992 DSV cate- mated location universal transverse mercator gories were ponderosa pine aspenaspenbirchbirch oak 199311993 HABITATS OF MERRIAMS TURKEY POULTS 133
table 1 use and selection of habitats by merriam s turkey hens with boultspoults in the central black hills south dakota habitats selected less P 10 than expected are indicated by and those more than expected by
ageAge habitat percent proportional 030 3 weeks 4 7 weeks 8 12 weeks 0 12 weeks canopy cover area N 106a106logloga d N 65 N 59 N 230 aspenbirchAspen birch 0 40 00148 0 1 1 2 aspenbirchAspen birch 40 70 00191 2 1 0 3 aspenbirchAspen birch 70 100 00177 4 0 2 6 oak 0 100 00044 3 0 0 3 spruce 0 100 0 005600056 1 0 0 1 meadows 01016 36 17 9 62 ponderosa pine 0 40 01199 16 7 5 28 pondeiosapmeponderosa pine 40 70 03760 31 32 29 92 ponderosa pine 70 100 03412 13 7 13 33 sapisamplesampie siesslesizcazc reportedcp td are thetiietile numberber ofradioofrasioof radioradlo locations ofliensof ilensliens bothwithvoth boultspoults by age category spruce and meadows OCC categories were we assured correct classification of the habi- 0 40 41 70 and 71 100 and were esti- tat at turkey locations in several ways habitat mated based on the following equation boundaries were marked on field maps when OCC 05105pbasalBASAL areaft2ac the location of a hen with boultspoults was near the 1941.94 bennett 1984 nine habitat categories habitat boundaries we verified field location by were determined these habitats correspond to walking around the radio marked bird while third order habitats as described by johnson continuously monitoring changing directions of visual 1980 the signal andor by obtaining locations without disturbing the bird more than half of analyses the 230 data points used for analyses were visual locations chi square tests for independence were the tennterm habitat use will be used when used to test the hypotheses that 1 habitat use habitats in which turkeys were observed were patterns by hens with boultspoults were similar among not compared to availability habitat selection age categories of boultspoults and 2 habitat use pat- will be used when habitats in which turkeys terns by hens with boultspoults were similar among were observed were compared to availability time periods of the day an evaluation of chi thomas and taylor 1990 square residuals with a G standardization to a critical Z statistic mosteller and parunak 1985 was used to determine significant differences of RESULTS habitat use among age categories of boultspoults chi square goodness of fit tests corrected thirty six radio marked hens in this study for 1963 used to test continuity cochran were produced 19 broods from which habitat use and the hypotheses that habitat selection patterns of habitat selection patterns were evaluated A to- hens with boultspoults were similar to random ex- tal of 230 independent observations of habitat pected use for 1 all hens with boultspoults and 2 use were made smaller sample sizes of older three age categories ofboultspoults confidence inter- age categories of boultspoults table 1 were due to vals around proportional use were used to de- mortality and movements out of the defined tenterminenine habitats selected more or less than study area expected neu et al 1974 byers et al 1984 habitat use differed among age classes of bonferroni corrected Z statistics were com- boultspoults P ooi001001.001 ponderosa pine habitats with pared to the standardized chi square residual 71 100 overstory canopy cover were used mosteller and parunak 1985198 to determine more by hens with 8 to 12 week old boultspoults than if habitat selection deviated from expected ran byyounger boultspoults Ponponderosaderos a pine habitats with dom use when 5 observations occurred in a 0 40 overstory canopy cover were used rela- habitat statistical significance for all tests was tively more by boultspoults younger than 8 weeks than aat iolo10.10 by older boultspoults 134 GREAT BASIN naturalist volume 53
habitat selection patterns of hens with acre hengel 1990 reported hens with boultspoults boultspoults differed significantly P 00ooooiool00001.00 1 from ran in wyoming using riparian areas and meadows domdorn use patterns summed across all age cate- in this study several hens with boultspoults 2 A4 days gories hens with boultspoults selected meadow old moved long distances to large meadows habitats more than expected but rarely were one hen took her brood 565.6 kinkm 35353.5 miles in observed in ponderosa pine habitats with 70 less than 4 days to a large meadow and another overstory cover habitat selection patterns of moved approximately 18lsis1.8 km in less than 2 days hens with boultspoults younger than 8 weeks old were to the same meadow the longest movement by identical to those for all boultspoults no significant a hen with boultspoults was more than 23423.4 km over a patterns of habitat selection were apparent for 6 week period they went to the same large hens with boultspoults 8 12 weeks old P 11 kentucky bluegrass meadow as the previous Hhabitat use did not vary P 5.5511 among the two day et al 1991 reported direct move- three daily time periods for all hens with boultspoults ments of up to 353.5 km from nests to centers of nor did habitat use patterns vary P 30 habitat for turkey broods among daily time periods when evaluated for hens with boultspoults select meadows or other individual age categories of boultspoults meadow habitats because herbaceous vegetation sup- habitats selected by hensa4thhens with boultspoults were more ports an abundance of invertebrates inverte- than two times larger x 36 4 ha P 02 brates are necessary in diets of young than the average x 16 7 ha within the gallinaceous birds for proper growth and sur- study area vival johnson and boyce 1990 invertebrate abundance in this study area was greater in discussion meadows than in other habitats rumble 1990 boultspoults have high protein requirements during edges of kentuckyofkentucky bluegrass meadows best the 4 weeks after hatching robbinsbobbins 1983 hurst poe characterized habitats selected by hens with and 1985 based on changes of habitat boultspoults open ponderosa pine habitats with ex selection patterns presented here use of tensive understory vegetation were selected to meadow habitats and subsequent feeding on a lesser degree hens with boultspoults selected dense invertebrates continued through 7 weeks of age ponderosa pine habitats infrequently except as grazing by livestock reduces the herbaceous escape or occasional loafing cover hens with cover and associated invertebrate abundance boultspoults in meadows were seldom observed moiemolemore habitat selection and survival of boultspoults are cor- than 10 in from the forest meadow edge related with abundance of herbaceous cover use of forest meadows or forest meadow metzler and speake 1985 therefore exces- ecotonesecotones by hens with broods is common for sive grazing by livestock would be detrimental neailynearlyneally all subspecies of turkeys jonas 1966 to wuitsmuitspults less than 7 weeks old hillestad and speake 1970 williams et al 1973 habitat selection patterns ofboultspoults changed scott and boeker 1975 speake et al 1975 as the boultspoults became older proportional use of mccabe and flake 1985 daydayetcayetet al 1991 hens meadows declined from 34 for boultspoults less than with boultspoults in this study selected the largest 4 weeks old to 26 for boultspoults 4 7 weeks old kentucky bluegrass meadows 2 188 ha and and 15 for boultspoults older than 8 weeks similar rarely selected small clearings inm the forest or trends in habitat selection patterns have been upland dry meadows small openings and up- noted for merriamMerriamslamss turkeys and other subspe- land dry meadows are common on the study cies of wild turkeys pack et al 1980 healy and area often occurring within 200 inm of kentucky nenno 1983 mccabe and flake 1985 campo bluegrass meadows selected by hens with et al 1989 increased selection of forested boultspoults these upland dry meadows in the black habitats may be related to lower requirements hills are less productive than kentucky blue- for protein following feather development grass meadows hamm 1973 healy and robbinsbobbins 1983 increased selection of forested nenno 1983 found that boultspoults prefer natural stands with 40 overstory canopy cover by clearings in the forest over artificial openings boultspoults older than 7 weeks is within the range of findings from this study contradicted results overstory canopy cover management of ponder- reported for merriam s turkeys in other regions osa pine directed in the black hills national Schemmschemnitztz et al 1985 and mackey 1986 both forest land and Beresourcesource management plan noted boultspoults using small openings less than 25 forest plan the forest plan calls for manage 199319931 HABITATS OF MERRIAMS TURKEY POULTS 135 meritment of pine stands to 60 or 80 growing stock BOLDT C EANDJE AND J L VAN DUESEN 1974 silviculture of in the black hills the status of our level GSQGSL GSL is the basal area of a stand ponderosa pine in knowledge USDA forest service research paper projected to 25 cm dahdbh trees boldt and van RM 124 rocky mountain forest and range experi- duesen 1974 ment station fort collins colorado 45 appp because of changing habitat selection pat- butleryBUIIERY R F AND B C GILLAM 1984 forest ecosys- pages terns of hens with age ofpoults habitat tems 34 71 in R L hoover and D L wills boults require- eds managing forested lands for wildlife colorado ments of wild turkey hens with boultspoults should be division of wildlife in cooperation with USDA forest assessed by age categories of boultspoults including service rocky mountain region denver colorado boultspoults older than 8 weeks in analyses ofyounger 459 appp poults can obscure some habitat relationships BYERSBYFRS C R R K steinhorst AND P R KRAUSMANKKAUSMAN boults 1984 clarification of a technique for analysis of utiliotili zationbation availability data journal of wildlife manage- management implications ment 48 1050 1053 CAMPO J J W G SWANK AND C R HOPKINS 1989 brood habitat use by eastern wild turkeys in eastern because hens with boultspoults selected meadows texas journal of wildlife management 53 47 482 during critical growth and development phases COCIIRANCOCHRAN W G 1963 sampling techniques john wiley sons new york of boultspoults 7 weeks and because older boultspoults and inc 413 appp DAY K S L D FLAKEFLAKF AND W L tuckerTUCKFR 1991 move- selected forest stands with 40 overstory can- ments and habitat use by wild turkey hens with broods opy cover managing ponderosa pine stands to in a grassland woodland mosaic in the northern plains 60 or 80 GSL would not negatively impact habi- prairie naturalist 23 738373 83 tats of hens with boultspoults however dense pon- HAMHAMMm D C 1973 evaluation of cattle use of a deer winter inm the black s thesis atac range hills unpublished master derosa pine stands 100 fiacftac basal area south dakota state university brookingsBrookmgs 69 appp should be left along meadow edges to provide HEALY W M AND E S NENNONFNNO 1983 minimum mainte- escape and loafing cover for hens with boultspoults nance versus intensive management of clearings fortor grazing by livestock reduces the standing wild turkeys wildlife society bulletin 11 113 120 HFNGFLHENGEL D A 1990 habitat use diet and reproduction of biomass herbaceous in meadows hencel of vegetation merriammernamss turkeys near laramie peak wyoming un- excessive grazing would also reduce the abun- published mastermasterss thesis university of wyoming dance of invertebrates and cover necessary for laramie 220 appp growth and development of boultspoults HILLESIADHILLESTAD H 0 AND D W SPEAKESFFAKF 1970 activities of wild turkey hens and boultspoults as influenced by habitat proceedings of the southeastern association of game acknowledgments and fish commissioners 24 244 251 HOFFMAN G R AND R R ALEXANDERALFXANDFR 1987 forest vegetation of the black hills national forest of south financial support for this research was pro- dakota and wyoming a habitat type classification vided by the US forest service rocky moun- USDA forest service research paper RM 276 rocky tain forest and range experiment station and mountain forest and range experiment station fort collins colorado 48 appp the national s grants wild turkey federationfederations HURST G A AND W E POFPOE 1985 aminoammo acid levels and in aid program R hodorff T mills C patterns in wild turkey boultspoults and their food items in oswald K thorstenson K jacobson and L mississippi proceedings of the national wild turkey harris assisted with collection of field data M symposium 4 133 143 green volunteered considerable on this JOJOIINSONHNSON D H 1980 the comparison of usage and avail- time ability measurements for evaluating resource prefer- project and R taylor graciously allowed us on ence ecology 61 65 71 his property for trapping and fieldwork dr L JOHNSON G D AND M S boyceboygeBOYCF 1990 feeding trials flake and H shaw and two anonymous review- with insects in the diet of sage grouse chicks journal ers commented on earlier drafts of this manu- of wildlife management 54 89 91 JONAS R 1966 merriammernamss turkeys in southeastern mon- script tana technical bulletin 3 montana game and fish department helena 36 appp MADONALDmaoMAC DONALD DANDRD AND R A JANTZEN 1967 management CITED JANFFN literature of the merriammernamss turkey pages 493 534 inm 0 H hewitt ed the wild turkey and its management the ALLDREDGEALLDRFDGE J R AND J T raitirattiRAIIIRATTL 1986 comparison of wildlife society washington DCD C 589 appp some statistical techniques for analysis of resource MACKEYMACKFY D L 1986 brood habitat of merriammernamss turkeys in selection journal of wildlife management 50 157 south central washington northwest science 60 108 165 112 BENNETT D L 1984 grazing potential of major soilssods mcgabeMCCABE K F AND L D FLAKEFLAKF 1985 brood rearing within the black hills of south dakota unpublished habitat use by wild turkey hens in southcentralsouthcentral south master s thesis south dakota state university brook- dakota proceedings ofthe national wild turkey sym- ings 199 appp posium 5 121 13113 1 136 grestGREAT BASIN nafnxfnaturalisturalistmuralist volume 53
METZLERml li i i R AND D W SPEAKEspi aklAKJ 1985 wild turkey poult rumble M AANDA AND S H ANDERSON 1992 stratification mortality latesrates and their relationship to brood habitat of habitats for identifying habitat selection by mer structure in northeast alabama proceedings of the riambamsariamnamss turkeys great basin naturalist 52 139 144 national wild turkey symposium 5 133 143 SCHEMNITZ S D D L GOPRNDIGOERNDT AND K H JONFS mosilMOSTELLER1 LI H F AND A PARUNAKPAKUNAK 1985 identifying ex 1985 habitat needs and management of merriam s tieme cells in a sizeablesieableabie contingency table prob turkey inm southcentralsouthcentral new mexico proceedings of abiablabihsticlistic and exploratory approaches pages 189 224 in the national wild turkey symposium 5 199 232 1 D1 C Iloaiioaghniloaglinglin F mostellerMostelierllerIIer and J W tukey eds SCOTT VVFI AND E L bofkerboekerboakerBOFKFR 1975 ecology of mer s lxplexploring01 ing data tables trends and shapes john wiley nam wild turkey on the fort apache indian reser and sons inc new yorkyolk 527 appp vation proceedings of the national wild turkey symposium 3 141 158 nlneuu C W C 11 BY i RSHS AND J M PFIKPEEKpeigpelg 1974 A tech- loilolfoifor of utilization SHARPESIIAHPF G W C W henderHENDEE AND S W ALLENALLE N 1976 nique analysis availability data jour- mcgrawmegraw nal of wildlife management 38 541545541 545 introduction to forestry hill book co 544 orrohnOROHRR II11 K 1959 pi ecipitationprecipitation and stream flow in the black PP SHAW H G 1986 impacts oftimberof timber harvest on merriam s hills USDA forest service station paper 44 rocky turkey populations problem analysis arizona mountain forestfoiest and range experiment station fort report game and fish department tucson 44 appp collins Coloiacoloradodo 25 appp SPIAKFSPEAKE D W T E LYNCH G A wrichtWRICIIIWRIGHT AND W J PACKPA K J C 11 P BUKKI IU W K IGOI1 0 AMAND D J PYBUS burkert HAMMKKIIAMMRICK K 1975 habitat use and seasonal move 1980 habitat utilized by wild turkeytintun key broods within ments of wild turkeys in the southeast proceedings of oak hickory forests of west virginia proceedings of the the national wild turkey symposium 3 122 130 national wild Furkeykey 4 213 224 wildfurkeytiutin symposium tj1omastjhomasIIIOMAS D LANDL annAND E J TAYLOR 1990 study designs and PETERSENPI A mer- IIKSIN L E AND H rlricilardsonIIAKDSON 1973 mer-mei- tests for comparing resource use and availability jour riami lamiam s wildrurkeywild turkey in the black hills of south dakota nal of wildlife management 54 322322330330 pages 3 9 G C sanderson and 11 C in II schultz eds WILLIAMS L E jlJR D 1H1 auslinaustinAUSIIN T E PEOPLESPFOPLFS AND wild tulkeyturkey management current problems and pro- R W PIIILLIPS 1973 observations on movement and grams university of missouriofmissouri press columbia 355 appp behavior and development of turkey broods pages ROBBINS C T 1983 wildlife feeding and nutrition aca- 79 99 ingin G C sanderson and H C schultz eds wild demic pipressess inc new york 343 appp turkey management current problems and programs RURUMBLIM 13 LE M A 1990 ecologybeoeco logy of merriamMernmernarnsarnsaross tuitulturkeyskeys me- university of missouri press columbia 355 appp leagris gallopavogallopavo inernaimtnerriandinernaim in the black hills south dakota unpublished doctoraldoctoiallallai dissertation university received 8 may 1992 of wyoming Larlaramiearniearnle 159 appp accepted 10 february 1993 great basin naturalist 532 appp 137 144
SQUIRRELS AS PREDATORS
1 J R callahan
abslranabsrbatr A literature review and field observations indicate that most sciundssciurids are facultative predators on small vertebrates this behavior isis documented fortor at least 30 sciurid species in 8 genera the frequency of predation apparently is influenced by various factors including climate season gender reproductive condition and availability ofplantof plantpiant sources for certain nutrients such as calcium and nitrogen although sciundssciurids assimilate as much energy from animal foods as do obligate carnivorescarnivores behavior associated with predation appears to be less efficient min sciundssciurids and may rely partly on prey habituation and other adaptive behaviors
key words squirrel sciuridae predator carnivore omnivore
predators utilize various strategies that maxi- sonai communication there is universal ac- mize the probability of successful prey capture cepceptancetance that squirrels eat meat the question is while minimizing the probability of injury sci- how they obtain it squirrels are often seen urid rodents many of them facultative and op- eating carrion on roads but are rarely seen at- portuniportunisticstic predators are not morphologically tacking live prey stomach contents analysis may specialized for this role and should therefore overlook vertebrate flesh and cannot distinguish possess a wide variety of adaptive attack behav- live prey from carrion thus each new observa- iors As discussed below one of these behaviors tion of a squirrel acting like a predator becomes observed in tree squirrels appears similar to a journal note see literature cited although insinuation curio 1976 a strategy more often most of these notes imply that such behavior is associated with invertebrates than with mam- aberrant collectively they describe a significant mals component of the sciurid repertoire the same predation as defined here means the killing feeding adaptations that enable squirrels to and eating of active vertebrates including con crack nuts are sufficient for opening skulls lan- specifics or other relatively large mobile prey dry 1970 by free living squirrels this definition excludes TREE SQUIRRELS the following field ob- the consumption of eggs nestling birds small servation which prompted this review adds a insects or any animal that is already dead prey species to the list of reported sciurid predators offer some resistance eating carrion or aphis is on 6 april 197911979 1 saw a lactating female west- similar to browsing predation also excludes kill- ern gray squirrel sciurus grigriseusgrineusseus stalk and at- ing that appears unrelated to feeding as in de tack an adult mountain quail oreortyx dictuspictus fensebense of the nest harris 1985 or as a in mixed conifer forest at black mountain riv- reproductive strategy balfour 1983 weissen erside county california elevation 1800 m bacher 1987 finally behavior of caged squir- the quail was standing on a 60 cm stump at the rels is often abnormal and is excluded here as edge of a clearing five or six other quail were evidence of predation although it can provide nearby none of the quail appeared to react as clues to dietary deficiencies the squirrel crossed the clearing in an odd not all biologists accept the idea of squirrels crouched posture rustling the pine needles and as frequent predators despite the 70 year lit- leaves loudly enough to attract my attention erature record summarized in table 1 when it was 20 cm from the stump it leapt up odonoghue s recent 1991 finding that squir- and pounced on the quail after a brief struggle rels are the chief predator ofjuvenile snowshoe the quail escaped and the covey moved off in hares elicited general shock C krebs per sixyearssix years in the same area on two other occasions
musenmuseumin of Ssouthwestern biology university of new mexico albuquerque new mexico 87131 mailing address box 3140 1hemet califoniia92546california 9254692545
137 138 GREAT BASIN naturalist volume 53
TABLE1 1 published reports ofot predation by sounds this table is limited to behavior offree living squirrels and includes only piepredationdation and predationpiedation attempts as defined in text
species prey source tiitittrlTRI 1 SQUIHKI is1 s eastern gray squirrel conspecificsConspecifics birds holm 1976 bailey 1923 schinusschirusSchirus carolicarolineaiscdrohnenwneaisnenisnemis westelwestern n gray squirrelsaunsqun relrei mountain quail unidentified small this papelpaper Scsaurushirus riseus mammal peromyscus Elelasternpastempastedastern foxsquirrelfox squirrel dove blue jay other birds borell 1961 seton 1929 shattershaffershafter sciurus niger and baierbaker1991baker 1991 european ledred squirrelsaunsqun relrei squirrels birds guinellgurnell 1987 schlogel 1985 scittrusScittscmrusscyrusrus vulgaris douglas squirrelsquiireiirel ground squirrels roest1951roest 1951951 1 tandaschfrus dougdouumdouglasbdouglasilasiiasi pine redlied squirrel chipmunks tree squirrels cotton- hatt 1929 seton 1929 hamil- tanuaciiiniitandascittrus huaonicushwlomcus tail snowshoe hare mourning dove ton 1934 0 donoghueodonoghue 1991 other birds nero 1987 taylor 1988 little greengi een squirrelsaunsqun i el frog live emmons 1980 aetliociurusaethoschirus poensispoentispoenwpoen sis ghoundsquikrrisGROUND SQUIRRELS whitetailWintetailtalltali antelope squisquirrelireiirel pocket mouse kangaroo rat lizards bladleybradley 1968 morgart 1985 anivwspemiophilusammosperniophilus leucurwleucurusleucufusurus nelson antelope squirrel lizards hawbeckerhawbecker19471947 aininopennophiluananosperniophilus nehomnedomnelneinelvonivoni yellow bellied marmot Conconspecificsspecifics armitageetalarmitage etetalal 1979 marntotaflaviventrismarniottmarniotaMarniota flaviventn califcalifornia01 ma ground squisquirrelirelirei ground squirrels rabbits pocket grinnell & storer 1924 fitch spernwphilusperuwphilus beeche yi gophers moles budsbirds lizards 1948 sumnersumner&sumners& dixon 1953 sandberg and banta 1973 trulio etetalal 1986 belding ground squirrel chipmunks other small mammals sharsmith 1936 howell 1938 sperirupfiiluisperirwphilus beldmibeldbeldingibelbeidingimi junco warbler sherman & morton 1979 michener 1982 suslik field mice birds snakes Calmescalinescucu 1934 herzig straschil speniiophiluspernwphihts citellusottelcitellu Conconspecificsspecifics 1976 columbian ground squirrelsqui ireiirel fish live 13 howell 1938 spehspernwphibisiriophilu colwiibianuicolunibianus franklinfrankilnfrankiin gioundground squirrel voles domestic fowl ducks other polder 1955 johnson 1922 spenrwphiussperirwphilus franklifrankliniifranklinicnii birds howell 1938 sowls 1948 choromanski & sargeant 1982 coldengoldengoiden mantled squirrel voles deer mleemice chipmunks junco cameron 1967 tevis 1953 spenruphilusperawphibts lateralislaterally lizards mexican ground squirrel cottontail packard 1958 spenruphilusperirwphihts mexicanusmexicanus round tailed ground squirrel sparrows bradley1968bradley 1968 spernuphilitspernwphilus tereticaudwtereticaudus townsend ground squirrel conspecificsConspecifics michener 1982 spernwphibisspernwphihis towntownsendsenduii 13 lined ground squirrelsquiireiirel rabbits birds bridgewater & penny 1966 sperniophiluspernwphilus triecemlineatuitrideceinfineatus bafley1923baileyballey 1923 arctic ground squirrel conspecificsCon specifics snowshoe hare odonoghue 1991 holmes spertrwphilusbperrmphilus undulatesundulatusundulcitui collared lernlemmingming 1977 boonstraboonstraetaletalet al 1990 michener1982michener 1982 rock squirrelsaumsqum el wild turkeys cook & henry 1940 spenfiophilusperuwphilus variegatevariegatuvariegatusvariedaniedanlegatus 199311993 SQUIRRELS AS PREDATORS 139
TABLE 1 continued
species prey source
washington ground squirrel conspecificsConspecifics aleornalcornalcornf9401940 spermophilus washingtoniwashingwashingtondtoni south african ground squirrel domestic fowl other birds turtles shortridge 1934 ryan 1987 abrusaerusxerus inauris other reptiles african ground squirrel cobra stiles 1987 xerus rutilus
CHIPMUNKS cliff chipmunk crabs jenkins 1989 tamias dorsalisdoidol sahssabs
merriammernamss chipmunk lizards sparrows larsonlarson19861986 tamias merriamimerrinwrriamimerrtemiami least chipmunk tree swallows lederlelederleetaletetalal 1985 tamias minimusininimus asian chipmunk voles birds lizards frogs ognevognev19661966 tamias sibisibiricusricus eastern chipmunk conspecificsCon specifics voles swallows star- krull 1969 seton 1929 ginevan tamiasTainias stristatusnatus ling snakes frogs salamandeisalamansalarnandersalarsalamanderinanderdeidel 1971 hesterberghesterbeig19401940 liarfiarflarharriotriot 1940 shacklefordshackleford19661966 torres 1937
I1 saw a western gray squirrel stalk a bird briefly killing a young blue jay avoided the skeletal but then retreat without completing a predation muscle and gnawed on the joints and bones near attempt on I1 april 1993 S B compton per- the surface of the skin this is in agreement with sonal communication saw a western gray squir- other reports of tree squirrels eating bone and rel with a small live mammal in its mouth the antlersanglers cross 1969 leach 1977 in other cases size of a young peromyscus beside a road in the however sciundssciurids have consumed specific prey san jacinto mountains 2100 in organs such as the brain hamilton 1934 elliott cross ingles 1947 1969 jaeger 1929 1978 or viscera hesterberg 1940 or the flesh and stienecker and browning 1970 reviewed of the head holm 1976 the food habits of the western squirrel but gray predation by tropical tree squirrels appears reported no predation although the latter to be rare emmons 1980 saw no predation by found feather fragments in one stomach at any of nine african although one least six other tree squirrel species take live prey species aethosciurus stomach contained a frog table table 1 but the frequency of such behavior is 1 glanz et al 1982 wrote that sciurus unknown meat constitutes 2 11 ofthe diet of rarely eats animal foods paraxerus the eastern gray squirrel sciurus carolinensis granatensisgranagranatenszstensis Paraxerus packard 1956 nixon et al 1968 however celapcepapicepap eats eggs nestlingsnestlings and insects but ap- stomach contents analysis does not reveal how parentparentlyly no prey as defined here shortridge meat was obtained moreover squirrels are er- 1934 male P cepapi sometimes kill juvenile ratic predators and not all studies are in agree- conspecificsconspecifics but weissenbacherWeissenbacher 1987 regards ment whereas borell 1961 and odonoghue this as a reproductive strategy rather than pre- 1991 reported predation by the fox squirrel S dation per se viljoen 1978 reported no preda- niger and red squirrel tamiasciurusTamiasciurus hussonihudsoni tion by funisciurusfumsczurusFunisciurus congicuscongiouscongicus small arthropods cus respectively reichard 1976 saw no pre- and annelidsannelidaannelids are the only known animal foods of dation by either species Sundasundasciurussundasczurussciurus loum LaTislanscuslatiscuslanscumcus obswrusobscuresobscurus callo it is not entirely clear whether a tree squirrel sciumssciurussciume melanogaster whitten 1981 callosci attacks live prey to obtain meat per se or calcium urus erythraeus setoguchi 1990 and tamiopstanuops andor phosphorus from the bones shattershaffer and mcclelmcclellandinwclellandilandi moore andtate 1965 borges 1990 baker 1991 noted that a fox squirrel after stated that ratufaraoufa is an obligate herbivore 140 creatgrestGREATGRFAT BASIN naturalist volume 53
the infrequency of sciurid predation in the chipmunk taznaztamiasTaTnias speciosusspeciosus is somewhat spe- tropics could be an artifact based on the geo- cializedcialized as an arboreal nest predator grinnell graphic distribution of observers certain other 1908 grinnell and storer 1924 but it has been marnmammalsmayn malsmais however are facultative predators in reported to eat eggs rather than adult birds only part of their geographic range eg the similar behavior is reported for the uinta chip chimpanzee curio 1976 for sciurids the most munk tamias umbrinusumbrinus smith and anderson likely explanations include the following 1 1982 certain tropical plants andlindtind tree barks are rich in significance there are really two ques- calcium and other nutrients see borges 1990 tions here 1 why eat meat 2 why catch it 2 carnivorecarnivorycarnivory in the tropics may be associated while it is still alive with increased numbers of stomach nematodes A frequent answer to the first question is that Ernemmonsmons 1980 3 colder climate necessitates squirrels especially reproductive females may a high fat diet or 4 facultative predation is need a concentrated source of protein andor partly a learned behavior that can spread certain minerals this view is supported by stud through a local population but need not occur ies of calcium self selection by male and female over the entire range of a species the first malabar giant squirrels ratufaraoufa indica borges hypothesis is supported by the fact that tropical 1990 smith 1968 and carlson 1940 re- tree squirrels turn to predation when caged and ported that only pregnant and lactating tree deprived of a normal diet eg keshavakushava bhat squirrels regularly eat animal food studies cited 1980 by gurnell 1987 showed that female tree GROUND SQUIRRELS meat live prey and squirrels cannot always obtain enough calcium carrion is a major food source for ground squir- phosphorus sodium or nitrogen from a diet of rels worldwide although fewer data are avail- seeds goodrum 1940 speculated that female able regarding old world species table 1 lists squirrels may need meat to reproduce success- reports of predation by 18 species of spermo- fully keymer and hime 1977 reported aabildwildavild philus ammospermophilusamnwspernwphilusAmmospermophilus marmota and european red squirrel sciurus vulgaris with xerus A possible exception is the mohave nutritional osteodystrophy suggesting that die ground squirrel spermophilus twhavensismohavensis tary calcium may be a limiting factor in the which has been studied intensively leitner et distribution of certain species al 1991 but is not known to take live vertebrate A second viewpoint is that the seasonal in- prey the cheek pouches and stomachs of four crease in meat consumption whether of live sciurotamias davidianusdavidianus specimens contained prey or carrion compensates for a seasonal de- only plant material callahan and davis 1982 cline in the quality of plant food especially but no field data on this endemic chinese genus protein content and is not specifically related are available to reproduction nutrient density and water FLYING SQUIRRELS the southern flying content of plants eaten by squirrels decline in squirrel giaugianglaucomysClaucomys volans eats eggs nest springspying and summer bintz 1984 in the mojave lings and carrion bailey 1923 landry 1970 desert plant foods evidently contain sufficient but not consistently harlow and doyle 1990 calcium but nitrogen is likely to be limiting for I1 have found no record of predation as defined desert ground squirrels that are active year here round karasov 1985 conversely tree squirrels CHIPMUNKS the eastern chipmunk tamias in more mesic environments have fungi avail- stristriatusstriatumatus and the asian chipmunk T sibiricussibiricus able as a source of nitrogen but are more likely take a variety of prey table 1 lederle et al to need calcium seasonally carlson 1940 1985 reported that least chipmunks tamias coventry 1940 keymer and hime 1977 phos- minimus prey upon adult tree swallows as well phorus also may be a factor in food selection as eggs and young jenkins 1989 observed cliff cano and colome 1986 attribute the con chipmunks tamias dorsalis in coastal sonora sumption of carrion by cattle in parts of south mexico eating crabs and other marine inverne africa to phosphorus deficient soils when kept brates in tide pools because the exact size of the on a herbivorous diet belding s ground squir- crabs was not documented this is a borderline rels select plant parts highest in protein and case of predation larson 1986 reported that water eshelman and jenkins 1989 gumellgurnell merriaromerriarnMerrmerrlmerriamsMerrimerniamsalnarn s chipmunk tamias merrimerrlmerfimerriamitrwrriamiami occa- 1987 wrote that tree squirrels use animal food siosionallynally eats lizards and birds the lodgepole mainly in the summer weeks and kirkpatrick 199311993 SQUIRRELS AS PREDATORS 141
1978 studied the salt drive phenomenon in to some extent but neither the apparent low fox squirrels and marmarmottmarmotsmots clark 1968 found success rate nor the situation specific response that the proportion of animal food in spermo- to potential prey is unique to squirrels both philus richardrichardsonirichardsonbsoni stomachs increased from 3 phenomena are reported for many obligate in april to 24 in august tevis 1953 reported predators as well curio 1976 moreover kara- a similar phenomenon for chipmunks and gold- sov 1982 found that antelope ground squirrels en mantled squirrels assimilate energy from animal foods just as effi- A third hypothesis is that predation by squir ciently as do obligate predators reis is often incidental to killing for some other there is a learned component in predator reason usually territorial defense or reproduc- recognition and avoidance by birds and mam- tive competition in other words once the other mals curio 1976 robinson 1980 it would be animal is dead it provides an energetic bonus a waste of energy for rabbits to avoid deer for that can be consumeconsumed withoutda4thout further risk this instance even though one deer ate one rabbit explanation applies mainly to certain ground but if squirrels undergo dietary stress every year squirrel species holmes 1977 michener 1982 and begin eating peculiar things one might balfour 1983 harris 1985 expect prey to catch on the limited evidence the other question concerns the advantages available suggests this is not the case birds of live prey carrion contains protein and other apparently respond to models of squirrels near nutrients and it does not run away or fight back their nests hobson et al 1988 a not unex- carrion also has disadvantages nutrient content pected result since nest predation is a frequent diminishes due to desiccation and removal of sciurid behavior smith 1970 reported that two organs by the original predator or by earlier cactus wrens campylorhynchus brunneicapillus scavengers carrion occurs in high risk situ- attacked and injured a harris antelope squirrel ations near ravens predator dens or cars and ammospermophilusamnwspernwphilusAmmo spermophilus harriharrisiiharrisiasii near an old nest it may contain harmful bacteria but the worst in other contexts however birds and other po- thing about carrion is that it may not be available tential prey often seem to ignore squirrels when needed most reports of scavenging by A clue to this blasebiase response may be found free living squirrels involve road kills an artifi- in the stalking behavior occasionally observed cially concentrated phenomenon in tree squirrels including the western gray quasi prey such as small frogs and most squirrel as described above this behavior is not arthropods are a fairly safe bet when available associated with the routine operation of nest nestling birds are somewhat more difficult be robbing but seems limited to the relatively in- cause of nest defense by the parents smith frequent attempts on larger prey klugh 1927 1970 shaffer and baker 1991 the hardest similarly wrote that red squirrels sometimes ap- prey to explain are adult birds rodents and pear to stalk grouse or partridges repeatedly rabbits since these have defenses sufficient to advancing on the bird and then retreating this inflict injury on a squirrel the prevalent view is is similar to my own observations ofwestern gray that squirrels turn to live prey only as an emer- squirrels reported above the squirrel engaged gency food source reichard 1976 when other in this near caricature of a stalking predator is resources are depleted this view implies that actually more conspicuous than usual at least to predation is a freakish event that has no real the human observer it is possible that the bearing on the squirrelsquirrelss role in the food web to effect of obvious repeated stalking is to habitu- paraphrase landry 1970 isolated events of ate potential prey cacarrmivoryavory do not a carnivore make to support most published reports of sciurid predation this statement landry cites an observation of a are brief notes since it is difficult to conduct a deer eating a rabbit quantitative study of any rare phenomenon again however a growing body of evidence however some tentative inferences can be suggests that predation is a normal component drawn the proximate significance of stalking of the feeding repertoire for most sciurids at may be that the squirrel is showing conflict least outside the tropics this does not imply behavior advancing and then retreating if the that squirrels are fundamental predators but bird or other prey appears in a position to de- simply that they are opportunistic one re fend itself in the ultimate sense the squirrel viewer of this paper commented that squirrels should benefit from this behavior if its effect is are lousy predators undoubtedly this is true to condition the local prey population to disregard 142 grestGREAT BASIN naturalist volume 53 skulking squirrels if most squirrels acting like CANO R J AND J S colomeCOLOMFCOLONIF 1986 page 665 in microbi- co paul predators do not follow through then prey ology west publishing st minnesota CARLSON A J 1940 eating of bone by the pregnant and should learnleam not to respond this behavior is lactating gray squirrel science 91 573 analogous to the hoarding of nuts in that the ciioromanskic11oromanski J AND A B SARGFANISARGEANT 1982 gray go- squirrel is hoarding prey confidence later phers and prairie ducks north dakota outdoors 45 when normal food items are in short supply the 6 9 CLARK T W 1968 food uses of the richardson squirrel can exploit this conditioning the ground in squirrel Spennspermophilusophilus richardorichardoniincharchoniinii eleganseldelegansegans in the longer term selection for such behavior poten- laramie basin of wyoming southwesternSouth westein naturalist tially represents an entry point to a new feeding 1324813 248 249 niche particularly in marginal habitats where COOK A HHANDWHAND W H HEhenryHFNRYN RY 1940 texas ground squir- rels catch and eat young wild turkeys journal of mam- tree squirrels may be more likely to resort to malogy 21 92 predation COCOVFNIRYVENTu A F 1940 the eating of bone by squirrels science 92 128 CROSS S PR 1969 behavioral aspects of western gray squir- acknowledgments rel ecology unpublished doctoral dissertation univer- sity of arizona tucson 168 appp this paper was begun with the assistance of curlocuinoCURIO E 1976 the ethology of predation springer ver an NSF grant BNS 781746978 17469 to the university lag new york ELLIOTTel LIO I1 r L 1978 social behavior and foraging ecology of of and was resumed under a 1990 91 llo georgia the eastern chipmunk tamias strisfnatusatus in the adiron- theodore roosevelt memorial grant to the dack mountains smithsonianSmith soman contributions to zool- author ogy no 265 107 appp EMMONS L H 1980 ecology and sourceleresource partitioning among nine species of african rain forest squirrels literature CITED ecological monographs 50 31 54 ESIIFIMANESHELMAN B D AND S H jenkingJENKINSJFNKINS 1989 food selec- aicoalcoal ornOKNR N J R 1940 life history notes on the chitephitepnite ground tion by beldingbeiBeldingss ground squirrels in relation to plant squirrelseninesqnine journalouloui nalnai of mammalogy 21 160igo16 170 nutritional features lounjoinjournaljoun nalnai of mammalogyMarnma logy 70 846 AKMIIACI K BBDD JOHNS AND D1 C ANDERSEN 1979 852 cannibalism among yellow bellied juarrnotsjournalofmarmottmarmots journal of fnciifl 1 c II11 II11 S 1948 ecology of the california ground squir- mammalogymainmaln inalogyanalogy 60 205 207 rel american midland naturalist 39 513513596596 ilalilaiBAIIIYBLEY 13 1923 meat eating propensities of some rodents cinglnginevanGINGINFVANEVAN M 1971 chipmunk predation on bank swallows olOf01 minnesotaofminnesota journal of mammalogy 4 129 wilson bulletin 83 102 BAIIOUHBALFOUR 1D 1983 infanticide in the columbian ground glanCLANGLANZ WWEE R WwtliorinclontiioringtontilTiI ORINGTON J MADDFNMADDEN AND L R squirrelsquill el Sperinspermophilusperinophilusspermophilusophilus cohanbianuscolumhianus animal behavior HEANEYHPANFY 1982 seasonal food use and demographic 3194931 949 950 trends in sciurus atanaaranagranatensisgrancitenwgranatensis pages 239 252 in E BINTZninnlnBIN I1 CG L 1984 watelwater balance waletwater stress and the leigh A S rand and D M windsor eds the ecol- evolution of seasonal torpor in ground dwelling seiscisel ogy of a tropical forest seasonal rhythms and longtermlong term mids pages 142 165 in J 0 muriemurlemune and G R mich- changes smithsonianSmith soman institution press washington ener eds the biology ofgroundof gi aundound dwelling squirrels DC university of nebraskaofnebraska press lincoln GOODRUM PR D 1940 A population study of the gray BOONSTRABOONS 1 HA RRCC J kiu13sKRI BS AND M KANIIRKANTE11 1990 arctic squirrel in eastern texas texas agricultural experi- ground squirrelsaimisqimi el predation on collared lemmings cana- ment station bulletin 591 1 34 dian journaljoin nalnai of zoologyofzoology 68 757 760 cliGRINNFLLI1 NNE L L J 1908 the biota oftheodtheof the san Bernardbernardinomo moun- bom llii11 A E 1961 fox squirrel attacks mourning dove tains universityuniver sitygity of california publications in zoology journal of mammalogyofmammalogy 42 101 5 1 170 bolBoiBOHCI s R M 1990 sexualandsexual and site differences in calcium clunngrinnellGiuNNGRINNFLLellYLL J AND T I1 slorerSIORFRSTORER 1924 animal life in the isconsumption by the malabar giant squirrel ratufaraoufa yosemite university of california press berkeley indica oecologia 85 8680 86 gurnellGURNFLL J 1987 the natural history of squirrels facts on bradleybranleyBRADLFYbladlbludl 11 W G 1968 food habits of the antelope ground file publications new york squirrel in southern nevada journal of mammalogy HAMILIONHAMILFON WJW J 1934 red squirrel killing young cottontail 49 14 21 and young gray squirrel journal of mammalogy 15 BRIDGI waliwallWAII K D D AND D F piPENNYnm 1966 predation 322 by Citelcitclhicitellitslilslitss tndccemlineatutrideceinlineatus on other vertebratesvertesivertebi ates joinjour HARLOW R F AND A T DOYLEDOYLF 1990 food habits of nal of mammalogy 47 345 346 southern flying squirrels Clauglaucomysgiaucomys volans collected calCACAIll11 NnlFISCUS u R11 J 1934 taxonomischebiologischeundbioraxonomisclic biologischebiologische und bioblo from red cockaded woodpecker ficoidesPicpicoidesoides boreaborhaborealishs geographische forschungenForschungen ubeluber die gattung citellus colonies in south carolina amerleanamerican midland natu- oken inm rnmanicnhllbliriimamenRiimamen zeltZeitzeitschriftzeitschnftschrift fuiful saugetierk 9 87 ralist 124 187 191 141 HARRIOIIHARRIOTT S C 1940 chipmunk eating a red bellied callananGAIIAHANCALLAHAN J R AND R DAVIS 1982 reproductive tract snake journal of mammalogy 21 92 and evolutionaryevolutional y relationships of the chinese rock HARRIS M A 1985 possible occurrence of interinterspecificspecific samnsqmnsquirrelel sciurotaimaseffirotarnias dadidavidavidianusclavichanwdianus journal of mam- kilhigkilhngkilling by a columbian ground squirrel Spennspermophilusophilus malogy 63 424742 47 columbcolumbianuscolumhianusianus canadian field naturalist 99 250 252 camea10nCAM nonKON 1D M 1967 carnivorous behavior of the gold ilarrilxrrhalHAI 1 R T 1929 the red squirrel its life history and habits en mantled squirrel murrelet 48 13 roosevelt wildlife annals 2 1 46 199311993 SQUIRRELS AS PREDATORS 143 hawbeckerHAWBFCKFR A C 1947 food and moisture requirements morgaiamobmorgadamorgarrMOR galaGAia J R 1985 carnivorous behavior by a white of the nelson antelope ground squirrel journal of tailed antelope ground squirrel aininosperinophibisamino&pennophilus mammalogyM amm alogy 28 115 125 leucunisleucunis southwestern naturalist 30 304 305 henigHFRIGHFIIZIG sirasciiilstiiasciiil B 1976 nahrung und nahrung nelloneroNFRO R W 1987 house sparrow killed by red squirrel seiserterhserserwerhwerb des zieselsdiesels actathenologica217aetaacta theriologica 217 131 139 blue jay 453 180 181 hfstfrbfrchesterbeii G A 1940 chipmunk eats frog journal of NIXON C M D M WARLEYWARLFY AND M W MCCLAINM CLAIN 1968 mammalogy 31 350 351 food habits of squirrels in southeast ohio journal of HOBSON K A M L BOUCIIARBOUCIIARII AND S G SEALYSFALY 1988 wildlife management 32 294 305 responses of naive yellow warblerswarblers to a novel nest odonogiiufodonogiiue M 1991 reproduction juvenile survival predator animal behaviour 36 1823 1830 and movements of snowshoe hares at a cyclic popula- HOLM R F 1976 observations on a canmbihsticcannibilisticcannibalistic grey tion peak unpublished inastersmasters thesis university of squirrel natural history miscellany chicago acad- british columbia vancouver OGNFV S I1 1966 mammals of the USSR emy of sciences 197 1 2 and adjacent countries vol IV rodents israel program for HOLMFSHOLMES W G 1977 cannibalism in the arctic ground scien- tific translations jerusalem squirrel spermophilus parryiparrys journal of mammalogyofmammalogy PAPACKARDKARD R L 1956 tree squirrels 5843758 437138437 438138 gard the of kansas ecology and economic miscellaneous publica- HOWELL A H 1938 revision of the north american economic importance tions museum of natural history university ofkansasof kansas ground squirrels north american 56 1 256 fauna 11 1 67 INGLES L J 1947 ecology and life history oftheodtheof the californiaCahforma 1958 carnivorous behavior in the mexican ground gray squirrel california and game 33 139 158 fish squirrel journal of mammalogy 39 154 jaegeiiJAEGFR E C 1929 denizens of the mountains charles C jaegerlaeger polderpoldfiiPOLDFR E 1965 vertebrate coactions with the franklin s springfield illinois thomas ground squirrel proceedings of the iowa academyAcaderny of lenkinsJFNKINSJENKINS P 1989 unpublished memowrittenmemo written to didr russell sciences 72 202 206 dated 18 march 1989 davis REICIIARDRFICKARD TAT A 1976 spispringing food habits and feeding be- JOIINSONJOHNSON A M 1922 an observation on the carnivorous havior of fox squirrels and red squirrels american propensities of the gray gopher spermophilus fran midland naturalist 96 443150443 150450 klim journal of mammalogy 3 187 ROBINSON S R 1980 antipredatorAntipredator behavior and predator KARASOV W H 1982 energy assimilation nitrogen re- recognition in belding s ground squirrelssqui nelsneis spermophi- quiquirementrement and diet in free living antelope ground lus bekbeklingilingi animal behaviourbehavioui 28 84840 852 squirrels ammospermophilusammospennophilusAmmospermophilus leucurusleucurus physiological roestROFSI A I1 1951 mammals of the oregon caves area zoology 55 378378392392 josephinejoseplrine county journal of mammalogy 3234532 345351345 351 1985 nutrient constraints in the feeding ecology of RYAN B 1987 the gabar and the squirrel witwatersrand an omnivore in a seasonal environment oecologia 66 bird club news 136 7 280 290 sandlsandbergSANDBFRGSANDI lenciERG SSANDBAND B H BANIABANTA 1973 instances of south- keslKESiKFSIIAVAlavaiAvA bhatBIIAT S 1980 cannibalistic behaviour in captive ern californiaCahforma ground squirrels sperrnophihispetmophihis western chats squirrel funambulus tristtiatustristnatustristnatus water- beechebeecheyiheecheyiyi nudnudipesnudipedipes eating iguanid lizards hernetonheipetonherpetonHeiHer peton house comparative physiology and ecology 5 44 45 7l71al 7 8 keymerKFYMFRKFYMER I1 F AND J M HIME 1977 nutritional SCIILOGELSCIILOGFL N 1985 Eichhorneichhorncheneichhoinchenchenehen fnsstfrisstfaisst amsel falke osteodystrophy in a free living red squirrel sciurus 329 321 vulgarisvuivulgans veterinary record 1002 31 32 sfioguciiisetoguciii M 1990 food habits afredofredof led bellied tree squir- KLUGII A B 1927 ecology of the red squirrel journal of rels on a small island in japan journal of mammalogy mammalogy 818 1 32 7157071 570578570 578 SETON of kiikilkliKHULLULL J N 1969 observations of tamias striatusstristriatumstnatsanatatusus feeding silonsitonsl lonton E T 1929 lives game animals doubleday upon condylura cristatacnstatacricristanastata transactions of the illinois dordoranan and co garden city new yoriyorkyolk academy of sciences 62 221 shacklefordsiiacklfford N 1966 eastern chipmunk feeding on a starling journal of mammalogy 47 588 LANDRY S 0 JR 1970 the rodentia as omniomnivoresvores quar- terly review of biology 45 351 372 SIIAFIER B S AND B W BAKERBAKFK 1991 observations of predation on ajuvenileajuvenile bluebl cyanocitta cricrlcrestatastata by LARSON E A 1986 merriam s chipmunk on palo escritoescntoescoto tietle jay criftatacristatacriftata a fox squirrel scum niger texas journal of science part II11 the individual in relation to its environment 43 105 106 dacobawacoba press big pine california 43105 SIIARSMHHSHARSMITH C 1936 carnivorous habits of the belding leaoLFAC II11 D 1977 osteophagyosteophage in the red squirrel blue jay leacil ground squirrel yosemite nature notes 15 12 14 35 102 shenmanSHERMANSIIFRMAN PPWANDMW AND M L MORIONMORTON 1979 four months LEIDERLE P E B C pijanowskipljanovvski AND D L BEAVER braver of the ground squirrel natural history 886 50 57 1985 predation ot tree swallows the least chipmunk oftree by shortridge G C 1934 the mammals of southwest afr- jack pine warbler 632 135 icarica 2 volumes william heinemann london LFHNFRLFITNER P B LEIINERLEITNER AND HARRIS 1991 lehnerlennerleitner andlANDJJ harmsharnis third year SMITISMIIII C C 1968 the adaptive nature of social organiza- coso exclosure baseline report grazing monitoring tion in the genus of tree squirrels tamiasliurntamiaschinisTamias chinis ecol- study coso known geothermal resource area inyo ogical monographs 38 31 63 county californiaCahgahgab forma unpublished report dated 24 may SMIIIISMITIL E L 1970 cactus wrens attack ground squirrel 1991 condolcondor 72 363 364 MICIIFNFRMICIIENER G R 1982 infanticide in ground squirrels sminiaminiSMITII K G AND D C ANDERSONANDFRSON 1982 food preda- animal behaviour 30 936 938 tion and reproductive ecology of the dark eyed junco MOORFMOORE J C AND G H H taieTAIFTATE 1965 A study of the in northern utah auk 99 650 661 diurnal squirrels sciunnaesciuridaesciurinae of the indian and indochi- SOWLS L K 1948 the franklinfrankiinfrankiln ground squirrel Citelcitelhiscitellnfbishis nese subregions fieldianaFiel diana zoology vol 48 field mu- frankfrankhnifrankonifrankliniliniilni sabine and its relationship to nesting ducks seum of natural history chicago illinois 351 appp journal of mammalogy 29 113 137 144 GREAT BASIN naturalist volume 53 shishlSniSTIENECKNiCKiF HWH WANDBAN j B M BHOWNINC 1970 foodfoodhdbitshabits VILJOENVILJOFN S 1978 notes on the western striped squirrel of the western gray squirrelsainisqini lel california fish and game funisciurusfumsciurusFuniFumsciurussciurus congiouscongicuscongicus kuhl 1820 madoqua 112 563656 36 48 119 128 srS i inslosimsi i i s 1D 1987 thetherikkitikkitaviofkorarikki tikkibikki tavi amoraomora swara 102 werksWEEKS II11 PPANDCAND C M kirkpatrick 1978 salt prefer- 29 ences and sodium drive phenology in fox squirrels and SUMNI R L AND J DIXON 1953 budsbirds and niammammalsmalsmais of woodchucks journal of mammalogy 59 531 542 thetiietile sierra nevada university of california press wfissfnbacweissenbacimi lierllerIIFR B K H 1987 infanticide in tree squir berkeleyberBei keley relsaels a male reproductive strategy south african 1 taywiiI AY 1 OK P 1988 predatorypiedatoryiedsquinelsred squirrels bluejay46097blue jay 462 97 journal otof zoology 222 115 118 ti11 vis L jiljl 1953 stomach contents of chipmunks and WIIIIIFNWHITTEN J E J 1981 ecological separation of three diur- mantled squirrels in northeastern california journal of nal squirrels in tropical rainrainforestforest on siberut island mammalogy 3431634 316 324 indonesia journal of zoology 193 405 420 tomostomusTOHHI s JJ K 1937 A chipmunk captures a mouse journal of Maminmammalogyalogy 18 100 received I1 december 1992 7 1992 tiwiioTitnuianuiwiioK L A W J louimyLoulouolouamy IKYimy D F HENNESSY D H accepted december OWINGS 1986 infanticide in california ground squir- rels animal behavior 34 291 294 great basin naturalist 532 appp 145 161
LATE quaternary vegetation AND CLIMATE IN THE ESCALANTE RIVER BASIN ON THE CENTRAL COLORADO PLATEAU
1 2 kim withers and jim I1 mead
ABSTRACT five alcoves rock shelters in the forty mile canyon willow gulch area of the escalante river basin in southeastern utah yielded rich deposits of late quaternary macromacrobotanicalbotanical remains the deposits were sampled and the contents identified in order to construct a chronology of vegetational change fourteen radiocarbon dates indicate that the fossils were deposited between 12690 and 7510 yr BPB P years before present ninety one plant taxa were identified 62 to species six species were common to all alcoves gambel oak quercus gambelngambgambeliigambeanelnhineliihiueill box elder acer negundo prickly pear opuntia subgenus Platyplatyopuntiaopuntia skunkbush rhuirhuschui arotnaticaaromcttica var tntrilobata serviceberry amelanchieramelanchiertitahensisutahensis and indian ricegrassricriencegrass oryzopsis hymehymenoideshymemndesnoides late pleistocene samples 11000 yr BPB P contain extraextralocalextrafocallocal elevationallyelevationally depressed species such as douglas fir pseudotsuga menziemenziesnnwnziesiisn spruce picea sp and mountain mahogany Cercocercocarpuscarpus ledifoliusledifohusledifolius and mesophytic species such as rose rosa woodwoodssiizi and water birch betula occidentoccidentalisoccidentalistalis early holocene samples 11000 8000 yr BPB P contain no elevationallyelevationally depressed conifers and the remaining mesophytic species decreasedecldecideel ease in relative abundance reticulated hackberry celtisreticulataceltis fetiretireticulatereticulataculata becomes common the terminalearlyterminaterminal earlylEarly holocene sample 8000 7000yr7000 yr BPB P contains abundant gambel oak and prickly pear but little else paleoclimatic interpretations for the late pleistocene correspond well to those of most other workers on the colorado plateau climates that weiewelewere wetter and at least seasonally cooler than they are today are inferred from the macromacrobotanicalbotanical assemblage however the increased moisture is attributed to higher stream base levels and increased groundwater rather than directly to increased precipitation early holocene climates are interpreted as warmer and drier than those of the late pleistocene but still wetter than the present climate groundwater levels appear to be decreasing due to stream entrenchment terminal early holocene climates were much warmer and at least seasonally drier by the end of the period groundwater levels had decreased so much that the alcoves were unable to sustain plant communities stream base level was probably near the present level
key words quaternary colorado plateau plants oak quercus pleistocene
the colorado plateau has been the focus of the macromicrofossilsmacrofossilsfossils in this study withers 1989 late quaternary paleopaleoecologicecologic work in the last were excavated from sandstone alcove rock decade however the late quaternary plant shelter alluvium and thus can represent only communities of the central plateau southeast- the community of the micromicrohabitathabitat where the ern utah are still inadequately known because deposit was found the objectives of this re- of the few case studies conducted late pleisto- search were to describe and to explore the pa cene early holocene plant communities of the leoenvironmental implications of changes in central plateau have been described using plant communities through time macromicrofossilsmacrofossilsfossils found in the alluvial deposits from cowboy cave spaulding and peterson STUDY AREA 1980 spaulding and van devender 1980 from scattered packratpankrat neotoma sppapp mid from 1986 to 1988 field crews from quater- dens from the paradox basin betancourt nary studies program northern arizona uni- 1984 1990 and from the megaherbivoremegaherbivore versityversity explored several alcoves in the forty dung blankets from bechan cave davis et al mile canyon and willow gulch areas of the 1984 mead et al 1986 cowboy cave han- escalante river basin glen canyon national sen 1980 and various alcoves mead and recreation area fig 1 which is the eastern aeenbroadagenbroad 1989 1992 boundary of the kaiparomtskaiparowits basin because
I1 wildlife and fisheries sciences texas aam uniuniversitysity college station texas 77843 2quatemaryqiiaternaiy studies Prpilgrampiogramprograinograin and department of geology northern arizonaanona university box 5644 flagstaff arizonaanona 86011 address for allaliailtilltuiltuii coicot respondenlerespondencerespondcorrespondenceenLeence
145 146 GREAT BASIN naturalist volume 53
12c 115 lioiio110 105
40 40
C M B M F W
35 3eae
7
c10doclodociodo ritepitePl te
30 30 k r N 0 200
0 115 lloilo110 105
fig 1 map of the southwest locating the colorado plateau delineated by stipples F forty mile canyon W willow gulch B bechan cave C cowboy cave M midden studies ofoi betancourt 1984 1990 our study is on lands administered by the na- agenbroad and mead manuscript stratified tional park service we were requested not to fluvial sediments are usually found on either locate the alcoves on a map or in a description side of steep talus slopes that are composed of with any precision somewhat more detailed sandstone blocks and boulders overlain by eo descriptions of the alcoves are in mead and lian sand these talus slopes often contain agenbroad 1992 anyone needing further de- prominent layers of macromacrobotanicalbotanical fossils and scriptionsscriptions of locations may contact the regional other remains such as bones hair and dung scientist at the rocky mountain regional off- five alcoves were chosen for study based on the ice denver colorado at least 851 plant taxa abundance of plant remains evident in the de- presently live in the kaiparowitskaiparona4ts basin resulting posits in many diverse local plant associations welsh etct al 1978 taxonomic terminology follows welsh etetalal 1987 MODERN environments the canyons cut through the navajo sand- stone and the upper units of the more resistant climate of the escalante basin is warm and kayettakayenta formation the alcoves appear to have semiarid with highly variable amounts of annual been formed in the cross bedded sandstone by precipitation strong orographic gradients are spalling wall exfoliation facilitated by the ac exhibited by both temperature and precipita- tion of moisture infiltration through joint con- tion mean annual temperature oflow elevation trolled surface drainage all alcoves in this sites along the river is 12c with 250 mm or less drainage area appear unique because they con- mean annual precipitation at high elevation tain perched terraces which are remnants of sites in the nearby aquarius plateau mean an- ancient fluvial deposits it is estimated that the nual temperature is 2c20cac with 575 mm of mean age of the fluvial deposits exceeds 30000 years annual precipitation webb 1985 199311993 quaternary vegetation ON COLORADO PLATEAU 147
vegetation in the basin ranges from great widespread but uncommon usually occurring basin desertdesertscrubscrub to spruce fir picea sppapp in sheltered shady areas abies sppapp forest low areas between 1400 in gambel oak quercus gambehigambeliigambgambelinehiehleliihill is common and 1700 in elevation are a mixture of in wet shady areas and hanging gardens in wil- shrublandsshrublands and grassland tanner 1940 pin low gulch but is restricted to a relictualrelicrevictualtual popula- yon juniper pinus sppapp juniperus sppapp wood- tion in a valleylike area in the upper part of land occurs between 1800 in and 2300 m forty mile canyon there it is found only on followed by a yellow pine oak manzanita pinus the shady north facing side of the canyon ponderosa quercus sppapp arctostaphylos pun against the rocks and on the sides and bottoms gens association between 2300 in and 2700 in of deep sandy washes shrub live oak Q tur- above 2700 in the southern face and top of the bibinellanella is common in the upper reaches ofboth aquarius plateau support a spruce fir forest canyons but becomes rare as the alluvium be- webb 1985 comes wetter shinnery oak Q havardhavardiiii is re- topography in the immediate area of forty strictstricteded to the driest uppermost portions of the canyons mile canyon and willow gulch is diverse ele- the wettest the vation ranges from 2300 in at the top of the in parts of canyons near seeps and at the edge of the creek important kaiparoaftskaiparowits plateau where the peperennialrenni I1 perenniperanni species include horsetail streams that carve the canyons begin to about equisetum sppapp common reed phragmites australis sedges 1100 in at the confluence of the two canyons at carex the escalante river sppapp scirpus sppapp birchleafbirch leaf buckthorn rhamnusrhamnusbetulaefoliabetulaefoliabetulaefoliaenolia andandsaltgrasssaltgrasssaltsaitgrass distichlis pinyon juniper woodland dominates be- spicataspicatespicata box elder is found scattered along the tween 2300 and 1500 below the woodland in in canyon bottom and occasionally in hanging gar- on benches and in slick rock areas is desertscrubdesert scrub dens common hanging garden species include dominated by blackbrushblackbrush coleogyne ramosisramonis ranwsis various ferns and mosses as well as poison ivy simaslom within the canyons habitats range from toxicodendron rydbergrydbergiiii cardinal flower nearly vertical rock faces to valleylike areas lobelia cardinalis watercress nasturtium of from dry rocky slopes and creekbedscreekbeds with ac ficificinalenale cardinal monkeyflower mimulus tive steep sand dunes to riparian woodlands and cardinalis and columbine aquilegiaaquilegiamicranthamicrantha hanging gardens valley areas are dissected by sandy washes many plant such as skunk- species FOSSIL bush rhus afoaromaticaaroffwticaaromatica var trilobata single leaf localities ash fraxinus anomala barberry mahoniafre s grobot grotto nwntiimontiimonmontaitii and serviceberryserviceberry amelanchier utahenutahan hooperhoopers hollow drobot and BF alcove are in forty mile canyon shrub ox sis are found throughout the creek system in a al- cove and oak haven are in willow gulch variety of habitats while others such as willow fig 1 all are located between 1100 and salix 1300 in in sppapp tamarisk tamarixTamatix ranwsisshnaramosissima in elevation troducedtrodproduceduced box elder acer negundo and seep willow baccarisbaccalisBaccaris emoryl are found only near forty mile canyon permanent running water cottonwood popu- the alcoves in forty mile canyon are diffi- lus fremontiifrenwnfflfremontiatiifil is common along permanent fremon cult to reach approximately 10 in of vertical or watercourseswatercourses as well as areas where ground nearly vertical sandstone rises from the deeply is the surface water near entrenched streambedstreambed to the base of the al- on slopes above the streambed dry streambed impor- coves streamsidestreams de vegetation below BF alcove tant species include skunkbush sagebrush and drobotgrobot grotto is dominated by willow and ATte gu- artemisiamisia sppapp blackbrushblackbrush snakeweed tamarisk but there is little streamside vegeta- tiertierreziarezia sppapp prickly pear brickellbushbrickell bush tion below hooperhooperss hollow brickelliaBrickellia B califomicacalifomica grangrandifloradiflora and in- hooperhooperss hollow and drobotgrobot grotto are dian ricegrassricegrass oryzopsis himebimehymehymenoideshynwnoidesnoides in val- large 100 200 in aideahdewide southwest facing al- leys many of these same species are common coves fig 2 tops of the deposits in both are along with winter fat ceratoidesCeratoides lanata and at approximately 1200 inm elevation in both al- scattered individuals of utah juniper singlesingie leaf coves modem vegetation is characterized by ash netleaf hackberry celtis retireticulatereticulataculata and desert grassland species with a few scattered cliff rose purshia mexicansmexicananwxicanamexicana hackberry is shrubs active and grass stabilized sand dunes 148 GREAT BASIN naturalist volume 53
1020.10cm
W
m W
661
cem ILI 1 4
V
fig 2 above and on facing page photographs of alcoves mentioned inm text A hooperhooperss hollow B drobotgrobot grotto C BF alcove and D oak haven note people for scale in center of B units containing macromaeromacrobotamcalmacrobotanicalbotanical remains occur about the rockfall smile in the centelcenter of the deposit dry preserved botanical remains occur stratigraphically above the layered fluvialil and lacustrine units that occupy the basal units at all alcoves sediments are of sangarnonianSangasangamomanmoman age 19931 quaternary vegetation ON COLORADO PLATEAU 149
u
14
77
7111 150 GREAT BASIN naturalist volume 53
as well as large areas of slick rock sandstone are 30000 yr BPBY they were deposited when the also found indian ricegrassricegrass is the dominant canyon stream was dammed downstream pos- species other species noted in the area include sibly by either a sand dune or canyon wall col- indigobushindigobush psorothamnusfremontupsorothamnusfrenwntii monnonmormon lapse at any point during deposition behind the tea ephedra virviridisidis snakeweed porcupine dam the canyon would have contained fluvial prickly pear opuntia erinaceaerinacea buckwheat and possibly eolian units in the stream channel eriogonum inflatum and dogweedhogweed dyssodia area but there would have been lateral fadesfacies of pentachaetapentachaetcipentachaeta colluvial deposition in the alcoves once the BF alcove is relatively narrow and faces sediment depth had reached the threshold of northeast fig 2 it is the highest alcove in this the dam height fluvial deposition would have study at about 1250 in elevation the area just ceased however lateral colluvial deposition outside the alcove is a steep grass covered sand would have continued we are recovering fossils dune species present include indian ricegrassricegrass in the lateral colluvial fadesfaciesfages dropseed sporobolus sp and occasionally when the dam was breached sediments mormon tea and snakeweed were probably downcut rapidly in at least the main streamstreambedbed remaining sediments were willow gulch eroded from behind and beneath by groundwa- ter flowing between the sediments and sand- the alcoves in willow gulch are presently stone seeps and springs they were also fairly easy to enter usually requiring only a short eroded by channeling and water flowing over climb up a steep embankment or sand dune the surface especially at the alcove triplinedriplinedripline shrub ox alcove is located in a wideaaideaaice valleylike remnants were left behind as predominantly area at 1190 elevation and faces northwest in lateral fadesfacies the alcoves oldest dated gambel oak and shrub live oak grow near the the in the layer from our study ca 23000 BY stream below the alcove skunkbush barberry yr BP soabsoa4 as well as mammoth Mammammuthusmuthus dung from and brickellbushbrickellbush are common cottonwood is below the profile in drobotgrobot grotto which dates abundant near the creekbedcreekbed and indian rice ca 26000 yr BPBY mead and agenbroad 1989 grass is widespread juniper trees are scattered is evidence that the valley sediments had already across the valley porcupine prickly pear and is been extensively eroded by the last full glacial hedge hogbog cactus echinocereus sp are found ca 21000 yr BPBY growing in cracks in the sandstone oak haven is the lowest alcove in this study depositional environments at 1140 in elevation and faces nearly due west and taphonomy fig 2 it is narrow and shelflike with a fairly large rubble filled shelter on the north end it the taphonomy what happens to organic is a wet active alcove although not so wet as remains after death of the localities is impor- to destroy the fossils about 50 in above the tant to understand organic remains are pre- present streamstreambedbed served in the alcoves behind the triplinedriplinedripline that this is the only alcove in which species found is inside the shelter and away from direct effects in fossil samples still occur alive today gambel of precipitation fossil remains were incorpo- oak is abundant throughout the alcove other rated into the alcove sediment layers conceiv- arboreal species found in or near the alcove are ably by various methods hackberry and box elder common shrubs in 1 A plant taxon could have washed into the clude seep willow sacred datura datura me alcove meaning that the plant actually grew at teloides serviceberry skunkbush roundroundleafleaf a different location possibly at a much higher buffaloberrybuffabuffailoberryloberry shepherdia rotundifoliarotundifolia and elevation and in a different community and was sagebrush A plunge pool is located to the south carried by stream action to a lower elevation to of the alcove and contains common hanging finally come to deposition in one of our alcoves garden species had this been the scenario the matrix immedi- the alcoves contain two major sedimentary ately around the plant remains would be fluvial units 1 lower fluvial layers without organic in nature the sediments would show some sort remains and 2 upper colluvial layers with fos- of stream action or deposition even the plant sils the multistratified fluvial sediments of remains would show some sort of transport which there are remnants remaining filled the damage which they do not show this is not alcoves and stream valley sometime prior to the case 199311993 quaternary vegetation ON COLORADO PLATEAU 151
2 A plant taxon could have been carried were dry screened through imm1 mm mesh at the into the alcove by wind action for this scenario site because the sand was so loose it was nearly to realistically happen the taxon in question impossible to avoid mixing the layers along the would have had to grow nearby enough to be edges of the pit when sterile sand was found at carried into the alcove by wind long distance 40 cm the excavation was terminated and the transport ofso many fossils is not likely given the pit backfilled A similar pit was excavated at the present physiographic situation certainly the top of the deposit holocene at drobotgrobot grotto taxon could have lived on the land above the however the same slumpageslumstumpagepage problems were alcove which is now mostly narrow slick rock encountered since recoveryrecoverywaswas minimal exca- having any of the fossil plants recovered in the vation was tenterminatedninatednineted at 30 cm alcove actually growing above the alcove in in the laboratory of quaternary paleontol- stead of in the canyon does not really alter our ogy northern arizona university bulk sam- overall conclusions ples were dry screened through a 20 mesh and finally 3 a plant taxon could have been 084 mm soil screen they were identified by growing in or immediately outside the alcove using the modemmodern collection housed in the mu- plants in this scenario would have pristine seum of northern arizona herbarium consult- macrofragmentsmacrofragments except for situations where ingingarthinga4thwith various members of the biology faculty spalling wall rock damaged the specimens the at northern arizona university and examining encasing sediments imply that only a colluvial regional literature gould 1951 martin and spalling depositional environment is present barkley 1961 morris et al 1962 delorit 1970 it is our opinion that given the three possible elmore 1976 albee 1979 welsh 1986 welsh depositional scenarios outlined above the al et al 1987 table 2 plants were identified from cove and the fossils discussed here are the result twigs leaves seeds flowers flower parts espe- of at least predominantly in situ deposition cially involucresinvolucred and fruits since the original that is species actually grew inside or in the mass and the number of unidentifiable frag- immediate vicinity of the alcove ments varied from sample to sample all identi- fied parts were counted and assigned a relative METHODS abundance according to the following scale I1 rare 1 2 fragments 2 uncommon 3 10 Macromacrobotanicalbotanical fossil collection fragments 3 common 11 50 fragments 4 and analysis very common 51 100 fragments 5 abun- dant 100 fragments following van deven- microfossilsMacromacrofossilsfossils were found in unconsolidated der 1973 layers leafleafmatsmats in the deposits or mixed with spall sands and blocks samples were difficult to radiocarbon dating obtain because the truncated deposits were very when possible radiocarbon dates were ob- steep and loose at angle of repose exposed tained on a single from a single layer surfaces were cleaned to remove loose con- species in cases where no single was abundant a taminatingtaminating slumpageslumstumpagepage the in situ leafleafmatsmats were species composite sample of a single species from two sampled by removing the mass of leaves twigs layers BF alcove or a plant and some and other plant materials by hand or with a species fecal material from a single layer trowel in between the leafleafmatsmats or where plant oak haven were used remains were found mixed with the sand bulk multiple dating procedures were performed in each alcove establish a chronol- samples of sediment were collected to radiocarbon done samples were taken from various locations ogy all dating was by beta analytic incorporated coral gables florida within each site so that an accurate description table 1 of the plant community and a chronology of the site could be constructed generally layers werewere sampled along a vertical line profile from the RESULTS top of the deposit at oak haven drobotgrobot grotto and shrub ox alcove sampleswerealsosamples were alsoaiso ninety one fossil taxa were identified 62 to taken to the side of the main profile to test as species withers 1989 of those only six were much of the variability of each layer as possible common to all alcoves gambel oak box elder at BF alcove a 025 m pit was excavated and serviceberry prickly pear skunkbush and in- bulk sampled at 10 cm intervals these samples dian ricricegrassegrass fourteen radiocarbon dates were 152 GREAT BASIN naturalist volume 53
TABLETABLF 1 radiocarbon dates all from beta lab in stratigraphic order for forty mile canyon and willow gulch samples GG drobotgrobot grotto BF birBFBIT alcove HH hooper s hollow SOA shrub ox alcove OH oak haven
radiocarbon date sample yobpyryrbpBP lab no matenalmaterialmanenal dated drobotgrobot grotto GGO none 661GGI none 662 7510 160 20999 quercus gamberigambehigambelngambean branch 663 9730 170 20998 quercus gamberigambehigambelngambean branch 664 none 665 9920 100 20999 quercus gamberigambelngambehigambean trunk BF alcove BFbidBIT 11790 190 14727 dung 12130 170 20995 pseudotsuga menziemenziesnmenziesiimenziesiasiislisllsn needles hoopers hollow hill 10630 110 25412 quercus gambellgambelngambehi twigs HIh11212 12010 110 25411 quercus gamberigambelngambehigambean twigs shrub ox alcove soalsoai 8520 80 25415 quercus gambellgambehigambeln twigs soabsoa2 8830 190 25656 quercus gambellgambelngambehi twigs soabsoa3 12690 180 25416 quercus gambellgambelngambehi twigs soabsoa4 23100 660 25413 quercus sp limb fragment oak haven olliOH0111 9180 100 28929 quercus gambellgambehigambeln branch 0112 11690 120 25418 rosa woodsnwoodenwoodsii twigs 0113 94701947009470 150 25657 picea sp twigs and dung 0114 none dattdtdat notalltpteclt ac ptapt& Ssee ttttextt obtained to determine the age of the deposits thirty five plant taxa were identified table materials dated included gambel oak wood 2 pncklypucklypfickly pear and gambel oak were the most rose rosa woodwoodsiiwoodsiesii branches douglas fir nee- abundant species in the oldest layer blackbrushblackbrush dles spruce twigs and dung table 1 and mormon tea were common more species of grass four were identified from this sample forty mile canyon than from any other indian ricegrassncegrassncricrieegrass was com- mon GROBOT GROTTO excavation into holo the remaining samples reveal no discernible cene sediments at the top of the deposit yielded change until all plant species disappeared from few macromicrofossilsmacrofossilsfossils and there was no obvious since the alcove sometime after 7500 yr BYB P gambel stratigraphy they were treated as one sample oak was the most abundant fossil species prickly GGO dated sample is identical to the not this pear was common in an undated sample from modem flora as observed elsewhere the in can- the bottom of the profile 664ggsgg4 and in the yon youngest sample GGI other species present four samples were collected from the 275 in small numbers include box elder hackberryhackhaekbeny cm thick gambel oak leaf roof spall layer that fishhook cactus sclerocactus sp solomonsolomonss began 157 cm downslope from the datum at the seal smilacina sp and indian ricegrassricncegrassncegrass indian top of the deposit two ofthese samples contain- ricriericegrassncncegrassegrass was the only species present in all ing gambel oak were submitted for radiocarbon samples dating twigs from the top and near the bottom BF ALCOVE the four samples from this of the oak leaf layer dated their deposition from alcove showed little nationvanationvenationvariationva except for a de- ca 9700 to 7500 yr BP table 1 A piece of oak crease in overall macromicrofossilmacrofossilfossil abundance as the stump located in situ approximately 17m17 m west sterile horizon was approached the lack of dis- of the profile dated ca 9900 yr BP cercerniblenible stratigraphy in the test pit and low 199319931 quaternary vegetation ON COLORADO PLATEAU 153
weights of any one fossil species resulted in the to the youngest sample hackberry was com- decision to pool the douglas fir needles from all mon in the sample dated 8830 yr BP but be- samples for radiocarbon dating the two dates came uncommon by 8520 yr BPBY prickly pear merge around ca 12000 yr BP table 1 was never abundant as in the other alcoves but thirty ninedine taxa were identified table 2 was only rare or uncommon in the youngest most abundant were douglas fir bigtoothbigtooth ma- samples an increased number of grass taxa ple acer grandidentatum box elder prickly five was observed in the youngest sample pear mountain mahogany Cercocercocarpuscarpus ledifoledino mountain mahogany was rare or uncommon in husuus and indigobushindigobush other common species all samples sagebrush and saltbush were rare included gambel oak currant ribes cf or uncommon in the youngest samples as were cereum skunkbush hackberry and single leaf servicebenservicemenserviceberryy rose and brickellbushbrickellbush ash spruce and fir were present represented several species were recorded only from this respectively by one needle and one cone scale alcove willow in the late pleistocene sample HOOPERS HOLLOW the oak layer at and utah fendlerellafendlerella fendlerellaFendlerella utahensis hooperhooperss hollow begins 190 cm from the da- two sedges carex bella and cladiuscladium californi tum is up to 100 cm thick and has been burned cum and cottonwood in the youngest sample extensively two samples record plant commu- all wewerere rare nities ofcaof ca 10600 and 12000 yr BPBY table 1 OAK HAVEN oak haven has both an oak only 13 taxa were identified from this alcove layer and a rose layer the stratum containing the low number reflecting the lack of preserva- the rose layer was chosen for profiling the tion because of burning table 2 gambel oak deposit began 20 cm from the datum and was was very common in the late pleistocene sam- about 155 cm thick the oak layer is located to ple uncommon or rare species identified from the north of the profile in a pile of wall fall this sample were prickly pear rose box elder overlain by eolian sand water birch betula occidentoccidentalisoccidentalistalis spruce and three samples were submitted for radiocar- indian ricegrassricegrass bon dating table 1 the gambel oak sample the early holocene sample records changes from the oak layer dated ca 9200 yr BP the in the vegetation during the pleistocene holo- two lower radiocarbon dates from the profile are cene transition rose water birch box elder reversed relative to the stratigraphy rose from spruce and mountain mahogany were no longer the top of the unit dates ca 11700 yr BP while at the site gambel oak and prickly pear were a sample of spruce twigs combined with ungu- common uncommon or rare species included late dung from below the rose layer dates ca poison ivy solomonsolomonss seal skunkbush and in- 9500 yr BP we feel that the spruce date beta dian ricegrassricegrass 25657 is equivocal as it is a combination of two entirely different species and should not be as- willow gulch signed to spruce or the stratigraphic position until confirmation SHRUBOXSHRUB OX ALCOVE the oak layer in shrub ox alcove began 120 cm from the datum forty taxa were identified from all samples and was about 350 cm thick gambel oak was table 2 the youngest sample contained abun- dated from four samples the oldest date dant fossils of gambel oak water birch and 23100 660 yr BP was on what appeared to bigtoothbigtooth maple were common rose is the most be oak wood from a highly degraded part of the abundant species from the dated rose layer gam- macromicrofossilmacrofossilfossil layer only three taxa oak hack- common species from the layer include berry and mountain mahogany were recovered bel oak bigtoothbigtooth maple and douglas fir rare from this sample but the community appears to or uncommon species include sagebrush box have been similar to those of the latest pleisto- elder water birch prickly pear and spruce the cene in this and other alcoves the remaining two lower undated layers contain abundant rose samples record the communities from the late and spruce water birch and box elder were pleistocene and the early holocene ca 12700 common to 8500 yr BPBY table 1 the forty mile canyon willow gulch thirty eight plant taxa were identified ta- sequence ble 2 gambel oak was abundant or very com- mon in all samples box elder and bigtoothbigtooth the fossil abundances of selected species maple decreased in abundance from the oldest from both canyons were grouped into a seriaseriatedsericatedted 154 GREAT BASIN naturalist volume 53
takiTABI I1 2 plants recovered from late pleistocene and holocene deposits in BF alcove drobotgrobot grotto hooperhooperss hollow oak haven and shrub ox alcove southeastern utah
660ccoGGOGCO 661GGI 662 663 664 BF hh1 hh2 soaisovSOM soabsoa2 soaSOM sokSOM ohlOHI ohaoh2 ohaoh3 ohaoh4 graze andropogonAndropoon gerardgefardgerardiigerardia01 arduii andropogon ctcf alomaiomdomeglomeratusglomlomeratulomeeratusratu I1 Boutboutelouaeloita barbatabenhen bata I1 Bouboutelouatelona erioerloenopodaeriopodapoda bouteloua tafidatnfidatrifuta I1 caletcalexcal ex sp 1 1 cladiuscladiuntcladiumCladi unt californiewncaliformcwn I1 deiclunnpiadeschairqsia caespitosacaeipitosa 1 distdiitclihiichlisnichlis pipicaiapicaliaspicatacaiacata I1 elbinuselyinusmug mus cf virginiansvirginicnsvirginicusvirginiousvirginicus 1 ehmuselbinuselyinus sp includes agropyronafroaffoAgropipon I1 1 hilaria nida 2 muldenbergiamiildenhergiaMulden beralabergia cf wrighwrightiiwrightnwrightetiifil 1 onopworyzopsisoqyzopsis jiuiwnoideshyntenoides stipa 3 2 2 1 2 1 1 2 1 1 2 1 Parlpanicumpunicumparlicumpari icum bulbhulhowmbulbosumosum 2 poa cf bigeloviibigelovii I1 setariasctaria sp I1 sporobolusporo hoiuholuboiu cf flexuousflexuowscfflexitosus 1 sporobolus sp I1 Trideris puuiellupulchelluspulpuichelluscbelius hnonewon 1 1 browse abiesahtesablesahles sp I1 acer randidentatuinrandidentafton 5 1 1 3 3 3 aceraiernemdonegundo 1 1 1 5 1 3 3 4 1 2 2 amaranthus sp 1 I1 Aniainbrowianibrosiabrosia sp I1 2 Aaadeaavegave sp amelanchier ulautaulahemiutaheitsisbemihemi 3 2 I1 1 1 1 I1 ai ctoilaphluarctostaphylusarctostaphylos pungentpuncnspungens I1 ArtenArtemiartemisiartemiwiartentisiatisiawi cf carnitcarmthiicarnithiihii 2 1 2 Artenaritimwiartentisiatisia cf dracunculudracuncubtsdracunculus 1 artedArtenartentisiaartciniwitisia tridentata 1 I1 aster cf spinospinosusspasmsspmsmssus 1 2 2 1 atriplex caneceizicanescentcanescenscanescens I1 1 1 1 2 atriplex cf conferticonfertifoliaconferttfoliafolia I1 baccharis sp I1 betula occidentoccidentalisocctdentnhsoccidentalistalisallsails 1 1 3 2 1 3 brickelliaBric kellia grangrandifloradiflora I1 2 1 1 1 Bricbrickelliakellia sp 2 2 1 ceanotliuceanothusfefenfendieripendieri 1 celtis retireticulatereticulataculata I1 I1 1 2 3 2 3 1 1 Ceratceratoideceratoidesoideoidesoldes lanata 1 1 ccrucarpwCercocercocarpuscarpus ledifohuledifiblius 3 2 2 1 1 1 1 2 2 chenopodiumclienopodiuiii atrovirens 1 chenopodiuntchenopodium cf muralemufale 1 chenopodium sp 2 chnwthamnuchiysothainnus sp I1 coleogynecoleogifne amowramosissimammow anawna 3 2 I1 comandraCoi nandra hallidapalhdapallida 2 CorycondaricondaryCondarydahydahidailsdatisdalls aurea I1 Grypcnptanthagryptanthatantha pterouirifaptemearya 1 1 criptanthnciyptantha etmsimasetosissima 1 cnptanthrtcqyptantha sp 2 I1 1 datura metehidesmethmeteloidesmetehidesnidesloides D wnghtiitvrightii I1 1 dithyreadtthijreaDithyrea fornicacalifornicacalifomicncali dyssodiaDiwdim dia sp ephedraEphedia sp 3 1 1 3 1 eriogeronEriogeron sp I1 I1 escltscholtlFM tc hottholtllaziakiaia ininutiflora I1 fendlerellaFendlerella utahenwutahensis fraxinus anoinalaanoiruila 3 i 199311993 quaternary vegetation ON COLORADO PLATEAU 155
tabletailleTABLF 2 continued
GGO oci661GGIGGJ 662 663 664 BF HH hh2 soaiSOM soabsoa2 SOM SOM OH ohaoh2 ohaoh3 0144
Gutiergutierreziagutierrezbareziafuzfaveziaeziauzia sp I1 haplopappus heterheterpogonpogon 1 haplopappus cf drummondrummonddrummondsdrummondiidrummondndnii haplopappushapkpcippus sp I1 2 1 2 1 1 heterothecaHetero theca sp I1 hymenoxys sp 1 juniperusJumperuspurus scopulorum 1 juniperusJumperus sp 1 lycium pallidumpallidum I1 mentzelia sp opuntiaopuntiaspsp 3 3 3 4 3 2 2 1 1 4 2 phloxphbxspsp 2 2 piceafzceaspsp I1 I1 1 5 3 polygonum sp I1 populus cf fremonfremontiifremontiacffremontiitittiifit I1 pseudotsuga menziemenziesnnwnziesiisn 5 3 i1 1 purshiapnrshiaparshia trifi dentataidentataedentata I1 2 2 quercus gambellgambehigambeln 1 3 5 5 3 3 3 4 5 4 5 5 3 2 1 quercus sp 1 rhusRIMS afonaronafoarontaticaaromaticaarotaticamatica var trifrilobata I1 1 1 3 1 1 1 1 2 3 3 ribes cf cereuecereum 2 rosawoodsiirosa woodsiiwoodsie I1 2 2 1 1 5 5 5 salix sp I1 sclerocactus sp 2 shepherdiashephershephei biadia rotundifoliarotundifolia i1 i1 smilacinaspSmilacinsmilacinaaspsp 2 2 1 2 2 1 sphaeralcea sp I1 I1 toxicodendron rydbergrydbergnrydbergiiii 2 trifolium sp 1 wyethia sp 1 1
chronosequencechronosequence fig 3 some ofthe variability though the species persist in other cases nearby seen in the sequence is probably an artifact of today sampling and site to site differences rather than the more xerophytic species in the sequence a real change in vegetation however the general are prickly pear sagebrush saltbush skunk- trend toward decreasing abundance of meso- bush blackbrushblackbrush and indian ricegrassricegrass skunk- phytic species is obvious douglas fir spruce bush persists in the record until 9180 yr BPBY and currant and fir do not appear in the fossil record was most abundant in samples that date prior to after 11690 yr BP willow is also absent from 12000 yr BP it is common and abundant in the the record after 11690 yr BPBY but it is common canyons today prickly pear was most abundant in areas with permanent water in the canyons in the sample from ca 12100 yr BP and is very today rose bigbigtoothtooth maple single leaf ash and common in the canyons today indian ricegrassricegrass mountain mahogany persist until 8520 yr BPBE sagebrush saltbush and blackbrushblackbrush show little while box elder persists until 7510 yr BPBY these variability in abundance throughout the time species were not as abundant in samples from spanned by the record although they are not after 11000 yr BPBY as they werewere in samples prior found in every sample today indian ricegrassricegrass to that time hackberry and water birch become is the most widespread species in the canyons while the others more abundant in the samples after 11000 yr are frequently encountered in drier habitats BP hackberry box elder and single leaf ash are found restricted to the more mesic habitats in the canyons today oak shows little variability discussion in abundance throughout the sequence the lack of deposits of any kind dating after 7510 yr plant communities that can be described BPBF suggests that the disappearance of oak from from fossils recovered from the alcoves are the alcoves by that time was a real event al much like those found in the canyons today 156 GREAT BASIN naturalist volume 53
tp 0 44 tp ov r A j b o f AAyj YEARS M0 ajA f v 0 B P f f 7000
7510 160
8000-
8520 80- 8830190 9 00 9180100-
9730170 oooo0000-
10630110
ooo 0001-
1 690 t 120 2 olo010 llo110 21301140
12690180 13000 0 4 0 3 0 5 0 4 0 4 0 3 0 5 0 4 0 2 0 3 0 5 0 3 0 2 0 3 0 2 0 2 0 2 0 2 0 2
fig 3 fossil abundances rating 0 to 5 of selected plant species from all sites see text for discussion dotted lines refer to inferred previous or known presence today of some species
major differences lie in the abundance ofofmesomeso were located in its present position since at best phytic relictualrelicrevictualtual species and elevationallyelevation ally de- a steep embankment must be climbed and at pressed extraextralocalextrafocallocal species dating from the late worst 10 m ofvertical sandstone must be scaled pleistocene and early holocene the most 2 the existence of oak haven this active striking similarity is the abundance of gambel alcove is wet and is the only alcove currently oak in the fossil record and its abundance in supporting gambel oak we believe that condi- some areas of the canyons today while more tions at oak haven are analogous to conditions xerophytic species such as indian ricegrassricegrass and in other alcoves during the late pleistocene 3 prickly pear have shown changes in abundance all fossil deposits are located behind the their distributions appear to have changed little triplinedriplinedripdriplineline away from direct effects of precipita- in the last 13000 years tion in dry section of the alcoves this suggests inferences regarding changing precipitation that precipitation was never directly responsible regimes based on paleovegetationalpaleovegetational changes for maintenance of plant communities within seen in the fossil record are complicated by the the alcoves but that plants were dependent on distinct possibility that the communities were availability of groundwater flowing from seeps much nearer the stream in the past than the springs and the intermittent canyon stream fossil deposits are today three facts suggest that gambel oak is a definitive indicator of cli- this was true 1 bison bison camel cf mate because it is sensitive to narrow limits of camelouscamelopsCamelops and mammoth frequented alcoves moisture and temperature grover et al 1970 in the system as recently as ca 12000 yr BPBY neilson and wullstein 1983 its presence mead and agenbroad 1989 1992 it seems throughout the time spanned by this assemblage highly unlikely that any large animal would be makes it suitable for making analogies at least able to gain entrance to the alcoves ifthe stream for temperature it is found only in areas with 199311993 quaternary vegetation ON COLORADO PLATEAU 157
mean annual temperatures of 7 10cloc harper piesples is rarely found in areas with mean annual et al 1985 it is indicative of at least 450 mm of temperatures below 6cac betancourt 1984 annual rainfall and no less than 250 minmm of gambel oak also very common or abundant in winter precipitation but its proximity to a the fossil assemblages is found only in areas source ofgroundwater such as a stream modifies with mean annual temperatures of 7 10cloc100c these requirements grover et al 1970 the harper et al 1985 mean annual temperature extraextralocalextrafocallocal species will also be used as important in the escalante basin today is 10 12c webb indicators of changing temperature and mois 1985 it is unlikely that cooling in the escalante ture regimes basin during the late pleistocene exceeded the based on the plant taxa identified from the 3 4cac postulated by betancourt 1984 for samples table 2 we have designated three higher elevation sites in the paradox basin we stages late pleistocene which includes sam- interpret that mean annual temperature ex- ples dating 11000 yr BP including ohaoh3 tremes in the escalante river area may have early holocene between 11000 and 8000 yr been little different from those of today cold BP and terminal early holocene between air drainage from the aquarius plateau and in- 8000 and 7000 yr BPBY creased local water availability may have been responsible for the persistence of montane spe- paleoenvironmental chronoloChchronologyronology of Paleoenvironmental change cies in the canyons LATE pleistocene environment although temperatures in the escalante ba- 11000 YR BP elevationallyElevation ally depressed sin may have been little different from what they montane species are the hallmark of late pleis- are today it is apparent that the alcoves were tocene plant communities in forty mile can- much wetter we hypothesize that this was at- yon andwillow gulch spruce occurred as much tributable to increased groundwater rather than as 900 inm lower than today while water birch was increased local rainfall during the late pleis- up to 80 in lower douglas fir currant and tocene these canyons contained much more val- mountain mahogany exhibited depressions be- ley fill than they do today and stream base level tween 260 and 300 in and bigbigtoothtooth maple was was up to 50 in higher as much as 140 in lower all these species with stream entrenchment can take place during the possible exception of bigtoothbigtooth maple are moist periods when plant cover is abundant or extraextralocalextrafocallocal and probably occur no closer than the plentiful in the upper reaches of the stream aquarius plateau about 70 km to the north antevs 1955 if rainfall was greater at higher relictualrevictualRelictual mesophytic species were abun- elevations during the late pleistocene as is dant during this time species that appear to suggested by most workers phillips 197719841977 1984 have been common in the late pleistocene spaulding and van desenderdevenderDeseddersenderveDder 1980 betancourt such as rose and box elder are rare or buncomuncom 1984 davis et al 1984 then forty mile and mon in the canyons today where they do occur willow creeks probably began to become en- they are restricted to streamside or other shady trenched during the late pleistocene because wet situations where seasonal droughts do not of entrenchment the erosion of valley fill con- occur while gambel oak is common is some tinued and groundwater levels began to de- areas ofthe canyons it is also restricted to more crease mesic habitats the only important xerophytic EARLY HOLOCENE environment 11000 plant in the assemblages is prickly pear the TO 8000 YR BP the major reorganization of plant community of the earlier time can be the plant communities during this time noted by interpreted as representing a somewhat cooler betancourt 1984 and other workers on the environment with more available moisture than colorado plateau is apparent in this assemblage is found in the alcoves today whether these also the only extralocalextraextrafocallocal species found in the were just seasonal differences cannot be stated fossil assemblages were bigtoothbigtooth maple and at this time mountain mahogany while still at lower eleva- the escalante river served as a route by tions than they are found today elevational de- which high elevation species such as douglas fir pressionspressions decreased to 30 in and 200 m and spruce were able to disperse into the lower respectively fossils of these and other relictualrelicrevictualtual canyons mean annual temperature in the mesophytic species such as rose persisted but aquarius plateau today is about 2cac however were less abundant than in the previous stage prickly pear which is common in the fossil sam even gambel oak decreased slightly in some of 158 creatgrestGREATGRFAT BASIN naturalist volume 53
the samples there were no apparent increases less reliable or completely dried up coupled in the abundance of xerophytic plants but de- with warmer seasonal temperatures this drying creasing abundances of mesophytic plants are resulted in the disappearance of most meso- evidence of changes in both temperature and phytic species by the end of our stage moisture regimes TERMINAL EARLY HOLOCENE environ- data for forty mile canyon and willow ment 8000 TO 7000 YR BPBE the one sample gulch appear to indicate two shifts in the flora from this time period is from drobotgrobot grotto that are similar to those seen in the paradox the fact that there is only one sample in itself basin Extraextralocalextrafocallocal except rose drop from suggests a major change in the environment species 00 maiormalorJ 0 the record prior to ca 10600 yr BP bigtoothBigtooth after ca 8500 yr BPBY the sample contains only maple rose and mountain mahogany recover gambel oak and prickly pear in any quantity no for a short time between ca 10000 and 8500 yr plant macromicrofossilsmacrofossilsfossils are found above this layer in BPBY before dropping from the record gambel drobotgrobot grotto and none above layers dating a oak increased in abundance and hackberryhackhaekberiyberly be- thousand years earlier in shrub ox alcove this came common during this time indicating that sample appears to mark the beginning of ex- summer moisture may have become more reli- tremely dry environments in the alcoves and the able with the establishment of the monsoonal establishment of the present environmental boundary regime shifts in the floras seen in the escalante because of its ability to reproduce vegeta- basin during the early holocene indicate that tively cottam et al 1959 gambel oak is able prior to 10600 yr BP temperatures or tempera- to persist in areas where seedlings are unable to ture extremes began to increase and moisture become established the stand of oaks that lived availability was less reliable this resulted in in drobotgrobot grotto until 7510 yr BPBY likely repre- decreased abundances of mesophytic plants and sents the last hold outs before the local water the upslope retreat of douglas fir and spruce table became so low that even they died the after 10600 yr BPBY the recovery of mesophytic disappearance of gambel oak and all plants species such as bigtoothbigtooth maple indicated that from the alcoves is interpreted as representing moisture availability became more reliable warmer and at least seasonally drier climates again although most species do not appear to and therefore the entrenching ofthe streamstreambedbed have been as abundant as in the latest pleisto- stream base level was probably very near its cene this moisture is still more attributable to present position by this time leaving the alcoves increased groundwater due to higher stream high and dry without groundwater resources to base level groundwater levels and rate of feed the seeps and springs that had previously stream entrenchment fluctuated during this supported the plant communities stage in response to fluctuating rainfall amounts and paleoclimatic as the summer monsoon moved to its present biogeographic position considerations A last date ofofcaca 8700 yr BP on gambel oak the escalante river appears to have served leaves at cowboy cave is interpreted as marking as a major migrational route for high elevation the inception of a more xeric climate spaulding and mesophytic species during the late pleisto and van devender 1980 the sample that dates cene it is likely that stream base level was as ca 8500 yr BPBY from shrub ox alcove contains much as 50 in higher than at present at the abundant gambel oak anthers gambel oak on lower elevational limits ofa plant high tempera extremely xeric sites often fails to produce ma- tures and deficient soil moisture produce tran- ture female flowers but produces an abundance spiration stress in established individuals and of male catkinsbatkins freeman et al 1981 we be- reduce the potential for germination and seed- lieve this radiocarbon date marks the termina- ling establishment relaxation of these controls tion of mesic environments in the alcoves with can be accomplished by lowering summer tem- the stream base level no more than 10 in above peratureperature extremes which would result in in- its present position the alcoves werewe re still wetter creased effective moisture or by increasing than they are today for most of the stage but precipitation betancourt 1984 since drought continued stream entrenchment culminated in was not a limiting factor the elevational depres drier alcove environments by the end of the sion of montane species observed in this area early holocene the seeps and springs became was probably primarily the result ofcooler summer 199311993 quaternary vegetation ON COLORADO PLATEAU 159 temperatures possibly facilitated by cold air sexual reproduction in oaks stopped by the end drainage from the aquarius plateau of the stage as evidenced by the abundance of southerly displacement of the polar jet male catkinsbatkins in samples from shrub ox alcove stream during the late pleistocene has been the polar jet stream retracted to the north as proposed based on paleovegetationpaleovegetation van de- the summer monsoonal boundary approached vender and spaulding 1979 and modem plant its current position this would result in higher distributions neilson and wullstein 1983 in annual temperatures and summer extremes southeastern utah this would result in milder less available moisture more reliable summer wetter winters and cooler drier summers an precipitation and a prolonged spring drought equable climate betancourt 1984 this inter- the senescence and death of oak trees and pretationpretation corresponds well with the interpreta- the disappearance of all vegetation from the tion of the assemblages from our study as well alcoves in the canyons by ca 7500 yr BPBY are as with those of most other workers except interpreted as the result of wannerwarmer tempera- cole on the colorado plateau gambel oak was tures and the establishment ofstream base level at low elevation sites in these canyons and at near its present position most other workers on cowboy cave and bechan cave today its the colorado plateau have interpreted their re- northern limit coincides with the polar jet cords as reflecting greater effective moisture stream at 41 N latitude the fossil distribution during this time however our record reflects of gambel oak suggests that the polar jet stream more xeric conditions as a result of decreased was displaced into the central plateau at about groundwater and possibly seasonally drier con- 38 N latitude during the late pleistocene ditditionsions the disappearance ofgambelofgambel oak from the terminal date for woodland communi- low elevation sites in the central colorado pla- ties in the southwest is consistently younger teau may represent a northward retraction than the traditional date ofcaof ca 1100011 000ooo yr BBPP for andor upslope retreat of the species as condi- the end of the pleistocene however in north tions at lower elevations became too hot and dry america this boundary is generally believed to conversely the xerophytic shrub live oak whose be time transgressive watson and wright northern boundary currently coincides with the 1980 and appears to vary with latitude transi- monsoonal boundary and pinus adulisedulis ex- tional communities with mesic characteristics panded northward in response to hotter drier persisted in many areas until the end of the conditions similar to antevs altithennalaltithermalAltithermal liv- early holocene ca 7800 yr BPBY when wood- ing populations of relict hybrids between the land species disappeared from modem desert two oak species as well as between P adulisedulis and areas van devender 1977 the changes in P nwnophyllamonomonophyllaphylla occur at the middle holocene vegetation during this time involved a gradual northern limit of shrub oak and P edulisadulis and the decrease in the abundance and number of southern limit of cambelgambel oak and P monophyllanwnophyllamonophylla woodland species and a relative increase in the lanner 1974 neilson andandwullsteinwullstein 1983 to- importance of desert species many of which day shrub live oak and P adulisedulis reach their were alreadypresent phillips 1977 in the east- northern limits far south of the relict hybrid ern grand canyon peak values for vegetation populations change species flux were recorded between cole 1981 suggests that a northward shift 12000 and 8000 yr BPBY cole 1985 believes of the summer monsoon and polar jet stream that wisconsin species tended to disappear would explain his xeric record for the eastern prior to the establishment of modem species grand canyon while the same scenario is in- the major reorganization of vegetation zones voked to explain the occurrence of the hybrids during this time precludes the use ofelevational and is consistent with a recent study of modem analogs betancourt 1984 and fossil distributions ofapomictic and sexually in our study we postulate that warmer tem- reproducing populations of muttongrassmutton grass poa peraturesperatures and increased drought stress due to fendlerianafendleriana on the colorado plateau soreng fluctuating groundwater levels resulted in the and van devender 1989 it would also explain upslope retreat of montane conifers and de- the dry record for the escalante basin those creased abundance of mesophytic species dur- records from the central plateau that are inter- ing the early holocene mesophytic species preted as wet are from higher elevation sites showed a brief recovery late in our stage before where wetter cooler situations could have disappearing from the alcoves ca 8500 yr BP persisted 160 GREAT BASIN naturalist volume 53
SUMMARY literature CITED
the record from forty mile canyon and ALBFFALBEE B J 1979 A guide to the identification of seeds used willow gulch reflects at least seasonally andor by prehistoric indians pages 283321 in J D jen- nings ed swallow shelter university of utah anthro- locally wetter more equable climates in the late pology papers 103 pleistocene the macromicrofossilsmacrofossilsfossils record gradual antevsANTFVS E 1955 geologic climatic dating in the west warming and drying which became extreme american antiquity 20 317 335 sometime after 8500 yr BRB P so that all vegetation betancourt J L 1984 late quaternary plant zonation and climate in southeastern utah great basin natu- disappeareddisapdigappeared from the alcoves by 7500 yr BPBY in ralist 44 1351 35 the increased dryness is directly attributable to 1990 late quaternary biogeography of the colorado falling stream base levels and decreased ground- plateau pages 259 292 in J L betancourt I1 R van water and only indirectly to changing amounts devender and PR S martin eds packratpankrat biddensmiddens the seasonality of last 40000years40000 years ofofbioticbiotic change university of arizona andor precipitation press tucson 467 appp our record is in general agreement with COLE K L 1981 late quaternary environments in the other colorado plateau records except that it eastern grand canyon vegetational gradients over the appears to reflect at least seasonally drier condi- last 25000 years unpublished doctoral dissertation university of arizona tucson 170 appp tions during the middle holocene is prob- this 1985 past rates of change species richness and a ably because our sites are at lower elevations model of vegetational inertia in the grand canyon than the others further studies of similar mac arizona american naturalist 125 289 303 robotanical deposits in the surrounding area if COIFAMCOTTAM W P J M TUCKERTUCKFR AND R DROBNICK 1959 someS great they exist are order of this record ome clues to basin postpostpluvialpluvial climates pro- in expansion vided by oak distributions ecology 40 361377361 377 could delineate the of the changing position DAVIS 0 K L D AGENBROADAGFNBROAD P S marlinmarilnMARIINMARTIN AND J I1 northern limit of cambelgambel oak and by extension MFADMEAD 1984 the pleistocene dung blanket of bechan the geographic positions of both the monsoon cave utah pages 267 282 in H H genoways and and polarjetpolar stream throughout the late quater- M dawson eds contributions in quaternary verte- fetjet brate paleontology a volume in memorial to john E nary additional data from this area would also guilday special publications of the carnegie museum help refine the geographic and temporal of natural history 8 boundaries of both wet and dry thermalsaltithermalsaltitbermalsalti delorndeloro R J 1970 illustrated taxonomy manual of weed additional paleobotanical studies in the area are seeds agronomy publication river falls wisconsin 175 appp if late paleoecology necessary the quaternary of EOREelmoreELMORF F H 1976 trees and shrubs of the southwest the central colorado plateau is to be adequately uplands southwest parks and monuments associa- understood tion tucson 214 appp FRFFMANFREEMAN D CEC E D mcartiiurmcaimiur K I1 harpenHARPFRHARPER ANDANDCC BLAUERBLAUFR 1981 the influence of environment on the floral sex ratio of monoecious plants evolution 35 acknowledgments 194 197 COULDGOULD F W 1951 grasses of southwestern united states umvsityunivsityUnivsity of arizona press tucson 352 appp we thank louella holter bilby research GROVFR B LLEE A richardson AND A R SOUTSOUTHARDliardilardfIARD center for the we 1970 quercus gambelngambeangarnbelii as an indicator of climatic typing manuscript appreci- means proceedings of the utah academy of science ate the following people for their help in the 4718747 ist187 191 field larry agenbroad liz moskybarnoskyBa john ben- HANSENHANSFN R 1980 late pleistocene plant fragments in the jamin brian and karen cluer and emilee dung of herbherbivoresivores at cowboy cave pages 179 189 in mead emilee provided all photography and J D jennings ed cowboy cave university of utah anthropology papers 104 drafting rocky mountain region national the HharperHARPFRARPER K TTFJF J WAGSTAFFwagsiaffandlAND L M KUNLFRKUNZLER 1985 park service denver office and glen canyon biology and management of the gambel oak vegeta- national recreation area provided logistic and tive type a literature review USDA INT 179 inter- curation assistance we appreciate the special mountain forest and range experiment station ogden utah and continued help of adrienne anderson and LANNER R M 1974 natural hybridization between pinus bob schiller national park service funding adulisedulis and finusrinuspinus monophyllamonophyllcimonophylla in the american south- for this project was provided by national sci- west silvae Genegeneticsgeneticatica 23 108 116 ence foundation grants EAR 8708287 and mantinMARTINmarlmaniMAPI IN A CANDCANDWDW D bankleyBARKLEYBARKLFY 1961 seed identifica- manual of press 8845217 to mead and L D agenbroad and tion university california berkeley MFADMEAD J I1 AND L D AGENBROADAGFNBROAD 1989 pleistocene national park service contract CS 1200 A 4062 dung and the extinct herbivoresherbivores of the colorado pla- to agenbroad teau southwestern USA cranium 6 29294444 199311993 quaternary vegetation ON COLORADO PLATEAU 161
1992 isotope dating of pleistocene dung deposits pages 159 162 in J D jennings ed cowboy cave from the colorado plateau arizona and utah radio- university of utah anthropology papers 104 carbon 34 1 19 TANNER V M 1940 A biotic study of the kaiparowits MEAD J I1 L D AGENBROAD 0 K DAVIS AND P S region of utah great basin naturalist 1 97 126 MARTIN 1986 dung of mammuthusMammuthus in the and south- VAN devenderDFVFNDFRDEVENDFR T R 1973 late pleistocene plants and west north america quaternary research 25 121 animals of the sonoran desert a survey of ancient 127 pankratpackrat biddensmiddens in southwestern arizona unpub- MORRIS M SJS J E SCHMAUTZ AND P F STICKNFY 1962 lished doctoral dissertation university of arizona tuc- winter field key to the native shrubs of montana son 179 appp montana forest and conservation experiment sta- 1977 holocene woodlands in the southwestern tion montana state university and intermountain deserts science 198 189 192 forest and range experiment station USFS USDA VAN DEVENDERDFVENDFK T R AND W G SPAULDINCSPAULDING 1979 de- NEILSON R PPANDLAND L H WULLSTEIN 1983 biogeogra- velopment of vegetation and climate in the southwest- phy of two southwest american oaks in relation to ern united states science 204 701 710 atmospheric dynamics journal of biogeography 10 WATSON R A AND H E wrichtWRIGHTWRIGIIT JR 1980 the end of 275 297 the pleistocene a general critique of chronostrati PHILLIPS A M III111 1977 Pacpackratspankratskrats plants and the pleisto- graphic classification boreas 9 153 163 cene in the lower grand canyon unpublished doc- WEBBWFBB R H 1985 late holocene flooding on the escalante toral dissertation university of arizona tucson 123 river south central utah unpublished doctoral dis- sertationsertation university of arizona tucson 204 appp PIPP 1984 shasta ground sloth extinction fossil pankratpackrat WELSH S L 1986 quercus fagaceae in the utah flora creatgigreateat basin 107 111 midden evidence from the westeinwestern grand canyon naturalist 46 WELSH S L N D AIWOOD AND R MURDOCK pages 148 158 in P S martin and R G klemkleinkiem eds LN atwoodalwood andlANDJJ 1978 kaiparoadtskaiparowits flora great basin naturalist 38 125 179 quaternary extinctextinctionsions a prehistoric revolution uni- WELSH S N ATWOOD S versity of arizona press tucson 892 appp L D GOOURICH aniAND L C HIGCINSHIGGINS 1987 A utah flora great basin naturalist SORENG R JANDTT R VAN DFDEVENDERVENDER 1989 latequalate qua- JAND memoir 9 brigham young university provo 894 ternary fossils of poafendleriana muttongrassmutton grass holo- utah cene of apomictsapornicts southwestern naturalist PP expansions WITHERS K 1989 34 35 45 late quaternary vegetation and climate of forty mile canyon and willow gulch in the central SPAULDING WWGANDKG AND K L PETERSEN 1980 pleis- late colorado plateau unpublished mastermasterss thesis north tocene and early holocene paleoecology of cowboy emern arizona university flagstaff 71 appp cave pages 163 177 inm J D jennings ed cowboy cave university of utah anthropology papers 104 SPAULDING W G AND T R VAN DFVFNDFRDEVENDER 1980 late received 2 december 1991 pleistocene montane conifers in southeastern utah accepted 7 december 1992 great basin naturalist 532 appp 162 167
BOLE VOLUME GROWTH IN STEMS OF QUERCUS GAMBELIICAMBELII
warren P claryI1 and arthur R tiedemann 2
AABS13strac1 kaclkaoiKACIr shrubshi abub formtorm and tree form gambel oak quercus gamberigambehigarnbehigambebiehi stands contain a potentially significant fuelwood resource information on their growth characteristics can form a basis for future stand management stem analyses showed that height growth of shrub form stems essentially ceased after age 50 while tree form stems continued to increase in height until approximately age 100 both stem forms continued to increase in basal area and volume at a relatively constant rate as thetlletile stems increased in age and size increases in all size measures were substantially greater in tree form stems than in shrub form stems mean bole volume for tree form stems at age 100 was over 16 times that of shrub form stems sprouts from tree formfoifol m stands would reach minimum siesize for futuelwood marketing in approximately 45 years
ketikeifkey wordmordwordsmonds cambelgambel oak quercus gamberigambehi shrub formforinnorinporm tree form height growth volume growth
gambel oak quercus gambwmbeliigamberigambehiehiehl is a species gambel oak is particularly desirable as fuel important for wildlife habitat watershed pro- wood because of its heatbeat yielding quahqualitiesties ap- tectiontection and fuelwood it is found in many areas proximately 141.4 times greater than ponderosa of arizona colorado new mexico and utah pine barger and ffolliott 1972 the superior in utah the optimum elevations are 1700 heat producing qualities of this species and its 2300 m where gambel oak is a dominant in the proximity to several major population centers mountain brush or mountain mahogany oak have generated considerable interest in man- shrub potential natural vegetation zone kuch- agement and use of gambel oak for fuelwood ler 1964 harper et al 1985 west 1989 harper et al 1985 betters 1986 retail prices gambel oak has a variable growth form nor- reflect the heatbeat producing value of gambel oak mally a tall shrub or small tree it can be found it is typically sold for 10 more per ton than any as dense shrubby patches 1 m tall or as widely other species johnson and grosjean 1980 spaced trees up to 23 m tall clary and tiede- some information is available on projected mann 1986 this morphological variation growth characteristics of gambel oak based pri- prompted early taxonomists to recognize as marily on diameter increments wagstaffwagstaffl9841984 many as eight additional species within popula- however no information is known to be avail- tions now considered gambel oak harper et al able on the incremental growth of gambel oak 1985 the variability may have an environ- bole volumes because of this we conducted mental source neilson and wullstein 1983 a this study to determine the volume growth char- genetic source pendleton et al 1985 or both acteristicsacteristics of gambel oak stems to assist in fu- sexual reproduction is sporadic generally ture management of this often ignored but with limited success cottam et al 1959 neil- locally important species son and wullstein 1983 wullstein and neilson 1985 however the species has a high regen- METHODS erative capacity from adventitious buds situated on the lignotubers and rhizomesrhizomes of existing FIELD METHODS the plant materials for clones muller 1951 tiedemann et al 1987 this study were collected as part of earlier stud- these buds give rise to numerous sprouts par- ies of standing crop biomass clary and tiede- ticularlyticul arly if fire herbicidesherbicides woodcutting or mann 1986 1987 eight small tree and chaining has killed the aboveground stem shrub form plots were sampled within typical engle etalet al 1983 stands on bald mountain near ephraim utah
inte nnountiintain kosckcsc11 earaearmuclilucli station forporfoiestst serviservice US dqarfinentdepiitincnt of agricultureofagrictilture boise idiho83702idaho 83702 correspoiidathC on esponddespond with this authoratautmthoramthorhoratit hispresenthis presentpregent address forestry1 OK stistif y sdsosscicnu s Laboratorylaboratoryabiboiiboaoratoryitoizitoiy 316 R1 myrtlmyrtimyrtlemyrdlemyddie st boisekoise idahoid iboiho 83702 21adfklulic dortlynortlynitliwstst lleeavhKS iillibilli station forest serviseausemu US deputmlntofasiiciiltnrcdepartrneiit of agriculture lnndanndla braidegraideGraidealde oregon 97850978 W
162 199311993 BOLE GROWTH OF QUERCUSQULRCUS cambelliGAMBELIICAMBELII 163
TABLF 1 growth curve coefficients and RsR
variablesvan ablesabies coefficientcoeffic lent termteiter m stem 2 form Y X A B C D Rr2ra shrub basal area age 814992814 992 439716 001586 355110355 iioilo110 056 tree basal aleaarea age 330944 762575 022552 56105656 1056 058 shrub volume age 574601 846456 002810 542712542 712 068 tree volume age 169008 546096 011471 76904976 9049 055 combined volume basal area 223504 386965 002732 173616 094 shrub height age 411932 109578 026754 272519 078 tree height age 9 71913971913 819354 037815 19070119 0701 085 shrub annual inermer volume 1363821363.821363 82 053418 000016 3426153426 15 075 tree annual inermer volume 2409612409 61 242472 000009 243278 080 modomoddmodel Y adA d B 1exp CX dilD 13 a b 600 600
480 480
360 360
LU ir LU 1 5 240 1 240 0
co co 120 120-
06 5 50 75 loo100 125 150 6 5 0 iai5 idoigo 1 5 lio160 AGE YEARS AGE YEARS
fig 1 basal area cm2 versus age years a shrub form stems b tree form stems these are hereafter referred to as shrub form and foliage where the tree bole forked the the sample stands were on slopes of up to 40 largest fork was selected as the main bole these the plots sampled varied in size from 3 X 3 m boles were partitioned into 60 cm sections con for high densities of small stems to 9 X 9 m for tinuingtinging upward until stem diameter outside plots of less dense stems large gambel oak bark had decreased to approximately I1 cm the trees were represented by five stands in the last sections were therefore of variable length cascade springs area of the uinta national for- A 10 cm length was removed from the base of est utah these are referred to as tree form each section for tree ring analysis by the late tree form stands were visibly distinct from sur dr C wes ferguson and associates labora- rounding vegetation and occupied concave tory of tree ring research university ofariofardof ari- slope positions where soil depth and moisture zona tucson favored tree growth stands had to be of suffi- laboratory METHODS the basic ap- cient size to accommodate a 100loom mm2ma plot plots proach ofring count dating was augmented in this were square when possible otherwise rectangular study by the use of dendrochronological tech- stems greater than I1 m high were counted niques in instances where the ring pattern was numbered and measured for diameter at a obscured or distorted two types of controls were height of4 cm three in tree form plots or five used to reconstruct the radial tree ring sequence in shrub form plots trees were selected at ran- first a comparison was made with other areas of dom for sampling stem boles were cut 4 cm the cross section with other sections from the above ground line and separated from branches same tree braithoraithor with other trees the second using 164 GREAT BASIN naturalist volume 53 a b 160 160igo
140 140
120 120
100 loo100 I1 uj S 80 W 80- 0 0 tat7 c 60 0 60-
40 40 q 20 20 201 RL 7 0 EO 1 f 160 1 1 0 25 50 75iai5 100 5 150 0 25 50 75 loo100 5 AGE YEARS AGE YEARS
fig 2 volume cmcm3 versus age years a shrub form stems b tree form stems dendrochronological principles was to date all assumed to be equally spaced throughout the or a portion thereof of the radius in comparison section patterns ofvolume change were exam- with two relatively nearby established tree ring ined by graphic and regression methods using chronologies nine mile canyon and emory periods or height segments within trees as sam- link several notable reference points were a ple units to illustrate growth trends all regres- pair of large ringsnngsangs at 1957 and 1958 and in some sion fits were made using the richards growth instances a wide band of vessels occurred in the curve model richards 1959 1919 ring some data had to be reconstructed for individual bole sections because of tree dam RESULTS age from fire or other mury distortion due to whorls etc the diameter increments were de- little data overlap occurred between the two terterminedmined to the nearest millimeter by decades populations above age 30 in the basal area versus ege g 1980 1971 progressing from the outer ring age relationships figs la ib table 1 at age of the stem toward the pith partial decade 30 tree form stems had mean basal area values growth was recorded when the beginning or nearly 10 fold those of shrub form stems simi- ending of the section growth record fell within lar relationships occurred with volume versus a decade As the stems tended to be asymmet- age figs 2aaa ab2b table 1 at age 100 mean stem rical the longest and shortest inside bark radii volumes were 4049 cmcm3 and 65808 cm3 for were recorded shrub form and tree form respectively or a dif- cross sectional area and volume calculations ference exceeding 16 fold were made by using spreadsheet software on a the relationship of volume to basal area was personal computer diameter and volume val- more consistent between stem forms than in the ues were calculated on an air dry inside bark previously described relationships A single basis the cross sectional area for a given period function fit the full range of data for both popu- was determined for both ends of each section lations combined fig 3 table I1 using the longest and shortest radii and assum relationships of height to age varied be- ing an elliptical shape section volume for each tween the two populations rates of height period usually decadal was calculated from the growth were not greatly different among popu- top and bottom cross sectional areas for the lations for the first 20 years after 50 years period and the section length using the parab- however little additional height increment oc- oloid method period usually decadal volumes curred on shrub form stems fig 4aaa table 1 were summed across sections to give stem vol- maximum height averaged 414.1 m tree form ume totals per period bole heights were deter- stems continued growth after age 50 at substan- mined by summing the section lengths heights tial although slowly decreasing rates until ap- at a given age were estimated in the manner of proximately 939.3 rn in height was attained at age lenhart 1972 wherein annual growth tips are 100 fig ab4b table 1 199319931 BOLE GROWTH OF QUERCUS GAMBELII 165
the relationship between annual volume in- 160igoIDU crement and age was not strong for either popu-opu ra values 293829.29 A hitbitit 140 lation r2R were 2938.293829 38 betbbebbbetter fit was obtained between annual volume increment 120 and total volume rar2R 7580757580.758075.75 80so80.80 annual volume 5 a 100 increment as a function of existing volume was 25 0 0 iS greater at all volumes in tree form stems than in 80 5 0 shrub form stems illustrating more vigorous 60- 7 growth 5aaa ab5b 1 0 bw figs table 40 discussion 20 mip 0 1 a this 0 ido160igo 200 33000 400 500 600goo sampling in study was limited to central BASAL AREA CM 2 utah but stem sizes encountered were deprerepre tentativesentativesentative ofsizes across the distribution ofofgamcamgam fig 3 volume cmcm3 versus basal area cmcm2 for com- bel oak mean basal diameters of the stands in bined stem forms this study varied from 363.6 11711.7 cm in shrub form stems to lsi15115.1 24624.6 cm in tree fonnform stems clary and tiedemann 1986 1987 our shrub a form stems therefore corresponded to the av- erage 76 cm stump height diameters in western 14 colorado brown 1958 our tree form stems were similar in diameter to the larger stems in 12 north central arizona barger and ffolliott 10 1972 limited information has been available con- 8- cerning direct volume measures or growth char- 6- acteristicsacte ristics cambelgambelofofgambel oak A volume table based on a technique of visually estimated volume is 4 available for colorado chojnacky 1985 and one has been used in arizona that was devel- 2 oped by modifying a composite volume table for trees in the great lakes vicinity barger and 0 25 do50 75iai5 160igoloo100 1 5 150 AGEYEARSAGE YEARS ffolliott 1972 barger and ffolliott 1972 found that annual stand volume growth in ari- b zona averaged 0240.24 mha or about a 2 incre- ment A similar percentage increment was 14 found in utah for individual older trees wag- 1984 s 12 staff wagstaffwagstaffswagstaffeWagstaffs 1984 data showed that diameter growth in tree form stems slowed lit- 10 tle in older trees thus the rate of basal area accumulation increased with age in this study 8 P our estimates of annual growth in older tree form stems were similar to those of wagstaff although differences in magnitude between 4 shrub form and tree form stems were striking nearly all data collected basal versus 2 in areaareaversus age volume versus age height versus age and an- 0- nual volume increments in relation to total vol- 0 25 do50 iai575 loo100160 1 5 ido160 AGEYEARSAGE YEARS ume were different between stem forms volume versus basal area was the only relation- fig 4 height m versus age years a shrub form ship examined that appeared similar between stems b tree form stems stem forms 166 GREAT BASIN naturalist volume 53 c1ca b 2500- 2500
2000 2000
Z 1500 Z 1500 lu li
2 LU2 F lu ir 0ir 0 Z 1000- 9 1000 a j 1 a D Z Z Z Z 500 500-
0 160 1 0 40 60 800 100 120 140 1160igo0 0 0 do do 10 0 luu VOLUMFCMVOLUME CM 3 VOLUME CM 3 thousands thousands
fig 5 annual volume increment cm 5 versus volume cm3 a shrub form stems b tree form stems
A massive underground structure which irha averaged across broad clone occupied supports rapid and normally voluminous sprout and non clonal areas barger and ffolliott ing following top removal provides a reliable 1972 the retail value on a landscape basis reproduction strategy that should fit well into a therefore would be 740ha 1983 dollars if all coppice fuelwood management cycle of harvest harvestableharvestable volume were removed wagstaff and regrowth clary and tiedemann 1986 1984 tiedemann et al 1987 this would be espe- gambel oak is marketable when average di- cially true on the more productive sites where ameters are relatively small wagstaff 1984 clones of tree form stems or larger shrub fonnform reported that stems are salable as fuelwood stems are available while we can offer no direct when the basal diameter reaches about 9 cm evidence that tree fonnform stands will coppice to basal area of 64 cmemi this diameter based on new treeftreetreel formf rm stands rather than to shrub form our stem analyses would be attained in 45 years stands circumstantial evidence suggests this is formhormbormn stands A few 00 in our unmanaged tree fon so tree formmorm stands in this study were sepa- shrub form stems would reach marketable size rated by a distance of several kilometers yet in 90 to 100 years but a projected 170 years most oftheodtheof the stems oftheseof these stands were estab- would be required in our average unmanaged lished within a 3 year period the most likely shrub form stands cause would be sprouting following a wide our current mature tree form stands with spread hot wildfire sprouting following such marketable volumes of 1506150.6 6046604.6 mha events typically results in high stem densities As would be worth 11144 44740 per hectare of the new stand ages a natural thinning occurs clone wagstaff 1984 clary and tiedemann this is reflected in old stem scars on lignotubers 1987 marketing of the resulting sprout growth and rhizomesrhizomes tiedemann et al 1987 scars of could occur in approximately 45 years although previous stems and the underground intercon- volumes would be much less than the original nectedness of gambel oak clones suggest that harvest estimated volume at age 45 would be generations of stems arise repeatedly from the only 25 of that attained at age 100 underground structures these stems would re only one of our shrub form stands had aver- fleet the same genetic makeup as the previous age stem diameters of marketable size although stems and would be growing on the same site four of the eight stands had some stems that revenue potential of mature stands near cit- exceeded the 9gemcm diameter requirement the ies and towns is substantial maximum retail stands had mean bole volumes of 46646.6 94194.1 values can approach 55000ha55000 ha of oak clone if nhamha and no apparent correlation between vol- individual very high volume utah sites are com- ume and stand density although lower density pletely harvested wagstaff 1984 arizona for- stands tended to have larger stems thus values ests have marketable gambel oak volumes of 16 for those stands that have attained marketable 199311993 BOLE GROWTH OF QUERCUS GAMBELII 167 diameters could be 3448 6963 per hectare of forest and range experiment station ogden utah clone clary and tiedemann 1986 wagstaff 31 appp JOIINSONJOHNSON C M AND M J GROSJFAN 1980 wood for 1984 any estimate of marketability and value heatingbeating utah homes an asset or liability publication for a specific oak stand would however have to EL 95 cooperative extension service utah state be determined on site university logan 9 appp fullwoodfuelwoodFu sales can provide a valuable tool KUCHLFH A W 1964 manual to accompany the map elwood potential natural vegetation of the conterminous for oak stand management elwoodfullwoodFufuelwood cutting united states american geographical society special can generate revenue while achieving various publication 36 new york new york 39 appp stand modification goals such as modifying LFNHARILENHART D J 1972 an alternative procedure for improv- wildlife habitat conditions reynolds et al 1970 ing heightageheigh tage data from stem analysis forest science 18 332 for or stimulating sprouts in overmatureover mature stands mullerMULLFR C H 1951 the significance of vegetative repro- future fuelwood production duction in quercus Madromadronomadroniofionionno 11 129 137 nellsonNEILSON R P AND L H WULLSIFINWULLSTEIN 1983 biogeogra- phy of two southwest american oaks in i elationrelation to literature CITED atmosphericic dynamics journal of biogeography 10 275 297 bargeltBARGEBAIUFRii R L AND P F FFOLLIOIIFFOLLIOTT 1972 physical char PENDLFIONPENDLETON R L S C SANDIRSONSANDERSON AND E D acteactensticsacteristicsristics and utilization of malormajor woodland tree spe mcarthur 1985 morphologic and enzymatic vari- cies in arizona USDA forest service research paper ability among gambel oak clones in northcentralnorth central utah RM 83 rocky mountain forest and range experi- pages 19 28 in K L johnson ed proceedings of the ment station fort collins colorado 80 appp third utah shrub ecology workshop college of natu- behersbenersBFHFRSBETTERs D R 1986 gambel oak in coloradocoloradoss front ral resources utah state university logan range pages 25 30so30.30 inm P F ffolliott and W T swank REYNOLDS H G W PR CLARY AND P F FTOLLIOH 1970 eds potentials of noncommercial forestfoiest biomass for gambel oak fortor southwestern wildlife journaljoul nalnai otof for- energy technical bulletin 256 school of renewable estry 68 545 547 natural resources university of arizona tucson fficllallRICHARDSDS FFJJ 1959 A flexible growth function for empiri- BROWN H E 1958 gambel oak in west centialcentral colorado cal use journal of experimental botany 10 290 300 ecology 39 317 327 tiedemann A R W P CLARY AND R J BARBOUR 1987 CHOJNACKY D C 1985 pinyon juniper volume equations underground systems of gambel oak quercus gaingam for the central rocky mountain states USDA forest beinbelubehl in central utah american journal of botany 74 service research paper incINT 339 intermountain for 1065 1071 est and range experiment station ogden utah 27 WAGSTAFFWAGS raffFAFF F J 1984 economic considerations in use and appp managermanagementnent of gambel oak for fuelwood USDA foifolfor CLARY W P AND A R tiedeTIFDFMANNMANN 1986 distribution of est service general technical report INTINF 165 inter- biomass within small tree and shrub form quercus mountain forest and range experiment station gamberigambehi stands forest science 32 234 242 ogden utah 8 appp 1987 elwoodfullwoodFufuelwood potential in large tree quercus WFSI N E 1989 vegetation types of utah pages 18 56 in gamberigambehi stands westeinwestern journal of applied forestry K L johnson ed rangeland resources of utah co- 2 87 90 operative extension service utah state university COTTAMCO namHAM W P J M tuckerTUCKFK AND R DROBNIC K 1959 logan some clues to great basin postpostpluvialpluvial climates pro WULLSTEIN L H AND R P nellsonNEILSONaninni SON 1985 seedling aidedvided by oak distributions ecology 40 361361377377 survival and biogeography of gambel oak quercus encleENGLEENCLF D M C D BONIIAMBONHAM AND L E BARTFL 1983 gambgamgomgamhelngambeliigambellibeinhelneliieill in northern utah pages 1 3amk3mkin K L johnson ecological characteristics and control of cambelgambel oak ed proceedings of the third utah shrub ecology journal of range management 36 363 365 workshop college of natural resources utah state HARPEHARPFRii K T F J WAGSTAFFwaoWAC STAFF AND L M KUNLFRKUNZLER university logan 1985 biology and management of the gambel oak vegetative type a literature review USDA forest serv received 25 august 1992 iceleeiee general technical report I1NTINT 179 intermountain accepted 6 january6januanj 1993 great basin naturalist 532 appp 168 179
AQUATIC HABITATS LIFE HISTORY observations AND zoogeographic considerations OF THE SPOTTED FROG RANA PRETIOSA IN TULE VALLEY UTAH
peter hovingh
AABS13sritac1 kaclkaoiKACIr foulfour populations of the spotted frog rana pretiosapreciosapretiosa occur in western bonneville basin only the tule valley populations occupy aquatic habitats associated with warm 28c28oc and slightly saline 1700 2700 nmhoscmtlmboscm springs the spotted flogfrog in tule valley breeds in coldwatercoidcoldeold watelwater portions of the peripheral wetlands which exhibit maximum temperature variations 1 25c25oc maximum conductivity up to 3200 mhoscmmhormboscmscmsem and maximum ph values up to 979 7 adult frogs are found in habitats with temperatures ofof29c29oc conductivity of 4700 trnhoscrnimmhoscm and ph above 909 0 in the summer the increased summer salinity and ph in frog habitats returns to lower values by the next breeding season due to underground recharge breedingbleeding in tule valley occurs earlier than in other bonneville locations because of the warm water sources spatial and temporaltempoiallallai distribution of the spotted frog since the regression of lake bonneville 15000 years ago and threats to present habitats are discussed
key wordsspottedfrogworth spottedfrog rana pretiosapreciosapretiosa life history ecology paleozoology
the spotted frog rana pretiosapretpreciosaiosa is an aquatic lake period finally I1 will discuss some features ranid occurring in northwestern north america of tule valley that have contributed to the long extending southward to nevada and utah in survival of spotted frogs in this valley while the western utah some of the sites occupied by species has declined in the wasatch mountains spotted frogs were flooded by lake bonneville of eastern bonneville basin in northern utah 15000 years ago currey et al 1983 curreyandCurrecurreyyandand this report is part of a continuing aquatic survey oviatt 1985 tuletulctuiethlethie valley one such valley of mollusks leeches and amphibians in the flooded by lake bonneville became a closed great basin basin and separate from lake bonneville 14200 years ago sack 1990 this valley contains nu- METHODS merous artesian springs on the basin floor prob ably associated with faults and fractures tulethletuie valley is located in western millard and stephens 1977 wilberg and stolp 1985 un- juab counties in west central utah fig 1 like artesian springs in adjacent valleys tule hydrologically tule valley is a part of the bon valley artesian contain neither fish nor springs neville basin of western utah as such it has mollusk surface water impounded in saline mud flats or the spotted frog was first noted in tulethletuie val- terminal lakes and does not drain to the ocean ley in 1980 in a study of the distribution of the the springs are located 77 km west of delta spadefoot toad hovingh et al 1985 hovingh utah and occur in a north south trend for 15 1986 high conductivity 1000 3000 mhoscmlmhoscmmhoscmsem km figs 1 2 in the middle of tule valley the and temperatures 19 31c of tule valley ar springs consist of a source usually in bulrushesbulrushes tesianatesian springs suggested special life history ad- scirpus americanosamericanusanwficanusameric anus a flowing outlet in bul aptations of the spotted frog to this habitat this rushes and a terminus wetlands in open ponds pappaperwillpapererwillwillwili describe some of the physical features with or without saltgrasssaltgrass distichlis spicataspicatespicata of these springs in relation to spotted frog life the springs are surrounded by greasewood history in addition I1 will postulate movements sarcobatus venniculatusvermiculatus and pickleweed of the spotted frog to the present hydrologically allenrofeaallenrolfea occidentalisoccidentoccidentalistalis communities fautin closed basin during the lake bonneville paleo 1946 and are similar to those described for fish
721 amondsmondst ond avAM IIIK saltsait laklakilake cityityibbiby ljtdutcili 84103
168 199311993 SPOTTED FROG IN TULE VALLEY 169
GREAT L LAKE- W 1520
c36
40 C
faf1 H dulgul SPRINGS FLAT
LH SEVIER TUL lu GSM 40 0 co DESERT KC SC 3 TW NT 5 ieltabelta SNAKE ST 6 so
S 39 19 AKER
1520
WAH w WAH
0 0 kiloneterskilometersKILONETERS 50
114 113
fig 1 southwestern bonneville basin showing taletuletuie valley snake valley and the sevier desert within tule valley the basin artesian spring complexes are shown from south to north as south SO south tule ST north tule NT south central SQSC north central NC and coyote CO as well as the mountain range springs 1 6 spotted frogs have occurred in snake valley in twin springs TW gandy salt marsh springs GSM and leland harris springs LH and along deep creek upper left communities of delta and callao utah and baker nevada are shown lines are contour lines representing 1520 m 5000 feet labeled 2290 m and 3050 m 10000 feet the inset shows the region in relation to the bonneville basin of utah eastern nevada and southern idaho as well as to the great basin in western united states springs bolen 1964 these springs were stolp 1985 sack 1990 5 north central com- grouped and named in this report 1 south plex and 6 coyote complex 35353.5 km north of complex 5 km southeast of south tule springs north central springs and not shown in figure 2 2 south tule each of the larger springs was marked dis- complex 3 north tulethletuie complex 4 south tally from its source by stakes at irregular inter- central complex consisting of three isolated vals measurements of temperature conductiv- springs referred to as willow north willow and ity and ph were taken at these locations A south willow stephens 1977 wilberg and diversity of locations was selected so that areas 170 grestgrearGREATgrexr BASIN naturalist volume 53
026 coyote N
25 4 3 22 north 121 central
7
15 south 14 central 13
P 12
C north 9 tul e 7
5 sou h 4 t 3 tul e 0 3 S K I1 LO ME T E R
fig 2 detailed map showing artesian springs in tule valley the two springs in the south complex are not shown the line associated with the springs is a vehicular tiacktracktlack that can join the antelope springs gandy road with a road to coyote complex the shaded area represents saline mud flats not shown are the extensive sahnesaline mud flats on the east side of sparse and dense cover active stream flow for ph analysis were taken in the field and read and standing open waters were sampled con within 24 h with a beckman model 3560 digital ductilityductivity and temperature measurements were ph meter in the laboratory field ph determi taken in the months of february may and au- nations were made with the cole palmer model gust december in 19819811 whereas ph was meas 5985905985 90 ph meter aredured in december 1981 and june 1982 tran- complete chemical analyses such as those sects were established in two springs to performed for drinking water were carried out determine temperature variations with time of by utah water research laboratory utah state day and season university logan the US geological survey conductivity was measured in the field using analyzed the water from north tule spring and a portable meter yellow springs instruments coyote springs stephens 1977 wilberg and no 33 and corrected to 25c water samples stolp 1985 and this investigation 5 august 199319931 SPOTTED FROG IN TULE VALLEY 171
1984 examined the source and two other loca- scaphiopus interinterniontanusintermontanusmontanus which breeds in dis- tions in south tule springs tal reaches of this spring hovingh et al 1985 sizes of the springs and wetlands were esti- the absence of ranicsranids in this spring suggests a mated values obtained from tracings of aerial paleozoological explanation prior to the in- photographs courtesy of the bureau of land creased spring flows from the high precipitation management tracings of the springs and con- years of 1980 84 first expressed in 1983 north trol areas of known size were cut out and central complex contained the only stands of weighed on an electronic balance the smaller typha domingensisdomingensis 23 and scirpus acutisacutus the wetland the less accurate are the values 19 in tule valley suggesting a different the advantage of this method is that aerial pho- aquatic habitat or history tographsto show irregularities ofthe springs in the coyote complex 26 consisting of four dark due to the contrast ofbulbulrushesrushes against the springs feeding a common wetland is the largest andaridarld carbonate soils spring in tule valley this amoeboid shaped snout vent lengths SVL of metamor- spring had an outside perimeter of 5 km in 1981 phosed frogs and tadpoles were measured by the spring sources contained a higher conducbonduc ruler estimates of populations were obtained tivitttivity than other springs with spotted frogs 2700 by counting egg masses in march egg mass versus 1800 lahoslmhosJimjimhoshos and during the summer numbers represent minimal breeding adult fe- this conductivity in distal reaches of the spring male 1 numbers since egg masses sometimes hadvalueshad values of 1900 6600 lahoslmhosjlmhos with the higher sink to the substrate and the eggs disperse and 00 1 values occurring in standing open waters table hence are not counted 2 breeding may occur 1 adult frogs were observed on 22 may 19 over two months all and not egg masses are june 8 august and 19 september in areas that counted and 3 sometimes eggs are deposited had a temperature range of 17 and a con in thick bulrushesbulrushes and 29c are not observed esti ductilityductivity range of 2000 4700 rohosrmhosmhos 5000 mates of metamorphosed frog chos populations were pihospmhosjlinhos approximates 0250.25 salinity egg attempted by individually marking frogs toe by masses were found in areas with a temperature clipping of up to three phalanges on each of the range of 10 14c and conductivity range of front legs turner 1960 petersen the lincoln 2500 3600 lahoslmhosjimhos on 7 march figure 3 shows estimator white et al 1982 of population size size ranges of temperature and conductivity was utilized but in this case the first and second throughout the year at south tule complex 6 samplings were not discontinuous samples and temperature distal to the source was less than occurred over two years the source whereas conductivity was higher and lower than the source in these same locations RESULTS the increase in conductivity in summer in standing open waters and the subsequent de- distribution of spotted frogs in tule valley crease in autumn and winter suggest that evapo- figure 2 shows the locations of artesian ration of the waters contributes to this increase springs in tule valley and table I1 lists the con- the decrease of conductivity in autumn and ductivity temperature and area of these winter from the summer values suggests that springs spotted frogs were found in south tule water repercolates into subsurface howsflows water complex 5 6 north tule complex 7 9 repercolation into subsurface flows was highly 11 south central complex 1415 and coy visible in springs 6 7 9 11 and 15 where ote springs complex 26 see fig 2 during water poured into a hole that resulted from the this survey springspying 5 was colonized from spring weight of cattle hooves on the aquatic habitat 6 with the frogs traversing a distance of 4 in one holebolehoieboie measured 60 cm deep the natural over carbonate soils slow recharge ofwater into the soils contributes the absence of spotted frogs in south com- to the maintenance of more or less constant plex and north central complex might be ex- conductivity values in these distal reaches plained by the high temperature of spring 1 through the years south complex the small size and distance since conductivity increased in standing from other inhabited springs south complex open water ph measurements were made or the salinity north central complex north these values increased from 757.5ts spring source central complex 23 contains suitable habitat to 959.5 in the distal reaches of south tule com- as indicated by its use by the spadefoot toad plex 6 in the summer fig 4 spotted frogs 172 GREAT BASIN naturalist volume 53
taniTABITABLE I1 1 physical characteristicscharacter isticsristics oftuleof tuletuie valley artesiana springs
source1 ce tempetemperatureraturegrature rangrangeocrange ecoC temp conductivity conductivity surface area 2 spring ICC rohosrmhosjimjimhoshos dec 3 aug 8 range lahoslmhosiimhos m
south complex 1 2 31 1900 12 32 18 32 1800 2300 nd 2 2 29 3200 5300 ndndend0
south tule complex 3 2 26 2400 4000 50 4 11 24 1700 2100 70 5 17 27 1100 2100 600 6 14 28 1800 2 29 19 29 1300 3000 19000 north tule complex 7 10 27 1800 0 28 16 28 1100 4000 4700 8 28 1700 500 9 8 25 27 1700 1800 0 27 24 30 1100 2800 7800 10 2 25 1700 18 26 27 28 1400 2200 1500 11 7 28 1800 7 28 25 30 1600 2100 11000 12 3 22 1800 0 22 23 26 1100 2500 4800
south central complex willow springs 13 23 28 1400 2000 1400 14 4 26 1900 3 26 22 27 1100 2300 3100 15 5 11 1700 2 11 24 28 1600 2500 4700
north central complex 16 2 2 27 1500 4300 4500 17 3 26 8700 16000 nd 18 4 19 3700 7600 500 19 3 31 49000 81000 nd 20 7 23 1800 2800 1000 21 5 23 39000 61000 nd 22 4304 30 3000 6100 400 23 7 16 29 1900 4100 3 16 24 34 1700 5300 49000 24 3 3 30 1200 6800 3 9 21 26 7100 25 2 7 22 1900 9200 5200
Ccoyoteoyotecoyote complex 26 32 28 2700 0 28 20 31 1900 6600 97000
2 not dettrinineddt h namednnmed butlint less1 ss thantilin 5m11TO in numbdnuniberNunumbedniber in pparendiesesiithcscsdc notes mimhciinintber ofsitesof sitessltes sampled atit the spring conlconicomplexplex theI1 he range ofteinperatureof tempeiaturetemperature and conductivity inin these easescases represents die trentesextremesex ohneasurenientof incdmik nienic lit iiomaiom 8 to 1 1 visits at crich samplesampie site inin 14811981 springspringe9 had two sources flflangflowingang into ait eooncooncommoneo on pool mdand spring 26 had four sources flowing intoiiltoiit anionunioncommonnnion poolpooi occupied these distal reaches in summer and alexesplexes of note is the observation that coyote egg masses were found here in spring water complex contains more sodium and chloride analyses at south taletaietuletuie complex 6 at the than south tule and north tule complexes source after the water flowed through bul water originating from coyote complex 26 rushes site I1 fig 4 and in standing open contained over 1400 eglmgl dissolved solids water site M fig 4 showed a decline in cal stephens 1977 in the summer conductivity clumelum and bicarbonate in the standing open doubled in some portions of the wetlands sug- water table 2 A slight decline in sodium and gesting that dissolved solids increase in these sulfate compared to the source occurred after areas fig 3 it is unknown whether spotted the flows passed the bulrushesbulrushes table 2 shows frogs avoid areas with high dissolved solids but these results and compares these values of refugia do occur within the springs that actually south tule complex 6 with previously pub- have less conductivity than the spring sources lished values for north tule and coyote com fig 3 frogs can likewise move to areas with 199319931 SPOTTED FROG IN TULE VALLEY 173
1 3- source 0 A 0oeasteast arm 0 west arm LO 0 t 2 aj4j Q 0fu 9 o 0 8 0 E 0 P U rpdr11 ad4d onioxlomi 0 S01-4aaa i g0 0 3 0 0 s 0 qiinz E 0 0 4 0
J F M A M J J A S 0 N D
i i i i
B H 1 l 0 .0 0 0 e 0 ij 0 24 0 00 0 WUJ 8 0 IQ 9 0 0 8 0 0tat 0 1 0 itOS 8 0 tf16 0 0 0 t- z 0 telmej 0 0 8 0 8 S j0 J F M A M J J A S 0 N D M 0 N T H
fig 3 conductivity mhoscmmholmhoscmscmsem corrected to 25c A upper figure and temperature oqC 13 lower figure with time of year at south tule complex 6 A line connects the measurements of the source data represent measurements taken at sites shown in figure 4 lower ph but it appears that they tolerate ph of flowed into a previously dry east arm and south 959.5gs as indicated by the presence ofadults in such tule 5 flowed into south tule 6 besides these habitats increases in flows typha domingensisdomingensis became As a response to the wet years of 1980 84 established in areas previously occupied by flows of the springs increased in 1983 and were greasewood and typha latifolialatifolia became estab- still expanded in 1990 many springs in the lished in coyote complex although spotted north central complex coalesced north tule frogs now bred in arms of coyote springs that 9 flowed into north tule 11 north tule 11 previously were uninhabited by frogs they did 174 GREAT BASIN naturalist volume 53
not inhabit the catcattailscoattailstails of south tulethletuie spring 6 spotted frogs moved from the south arm of north tule 11 greatly reduced by an outflow into a hole to the newly flooded east arm spotted frog life history observations the spotted frog is described as inhabiting L cold permanent waters or the peripheral shal- K low waters which often have large daily tem- peraperatureture fluctuations in british columbia licht 1969 and wyoming turner 1960 turner and dumas 1972 tule valley is one of the south- ernmost localities inhabited by the spotted frog and has habitats more saline than normal tem- peraperaturestures at the water sources are higher than those at which spotted frogs initiate movement to other sites 26c turner 1960 and are at the lethal range for larval development 28c licht 1971 the frog is found in distal portions of the 0 springs in tule valley where temperature con- ductivity and ph show the greatest variation breeding occurs in open shallow pools ofwater surrounded by bulbulrushesrushes in these distal reaches or in the cattle impacted region between the andaridarld land and bulrushesbulrushes in south tule complex 6 frogs bred from B to F west arm and H to 9 J east arm see fig 4 in the summer adults occurred at the edges of wetlands in places where they were protected by bulrushesbulrushes upon disturbance no breeding occurred in the cat IL tail dominated areas that were formed as a re- a sult of the 1980 84 wet cycle table 3 summarizes the life history of the spotted frog in tule valley adults emerge in early march or if the season is warm in late february males emerge before females as indi- 7 cated at south central complex 15 where 24 frogs all males were captured on 2 march 1990 at this time no eveyevvyegeyegg masses were found emer i aj4j gence in south central complex 15 was later 1 S A 0 C G D H E I F J K than in other springs on 12 march 1988 when S I1 T E no frogs or egg masses were seen at 15 egg masses and hatchlingshatchlings were observed in south fig 4 an illustration of south tule complex showingsh g tule complex 6 this delay could be a result locations which weiewelewere sampled tipper figure the source is 1 labeled S the mtennptedinterrupted line distal to M represents the of the lower spring source temperature of ilcI PC aleaarea hoodedflooded as a resultresuit oftheodtheof the high watelwater years cattailseatcatcattciilscoattailstails grow at 15 versus the higher temperatures of 28cgsg heiehelehere wheree gigreasewoodeasewood previouslypievionslygrewgrew no measurements at other springs on 13 march 1988 candygandy salt we lieilo lower shows wereie taken hereie tietletheviefhe figure the ph measure- marsh springs and twintvan springs see I1 for ments at the sampled solid lines summer dotted lines fig sites locations snake still whiterwhlter laigelarge diamonds west arinarm in winter squares east in adjacent valley were aimalmarm in winter open chelescircles west armarrn in surnsummermer and solid under ice again suggesting the warm spring circles east aimalmarin in suisulsummerniner sources in tule valley accelerate the breeding 199311993 SPOTTED FROG IN TULE VALLEY 175
TABLETABLP 2 major ionlon analysis of water in selected springs in tule valley
north tule springs coyote springs south tulethletuiethie spiSpringsingsingswetlandsspringswetlandswetlands
1 2 1 2 2 source site I1 site M year analyzed 1974 1983 1974 1981 1983 1984 1984 1984 specific conductance mhoscmmholmhoscmscmsem 1590 1620 2400 2380 2500 1700 1590 1430 total dissolved solids eglmgl 992 910 1430 1450 1460 910 930 880 ph 77 73 82 77 76 93 calcium smolarmmolarmM olar 180 173 1781 78 1801 80 178 1461 46 14444 0920.920 92 magnesium smolarmmolarmM olar 169 144 156 1731 73 169 1241 24 1291 29 1241 24 sodium smolarmmolarmM olar 874 870 152 162 165 8938 93 8788 78 8788 78 potassium smolarmMmmolarolar 051 046 095 077 087 0450 45 0490 49 0490 49 bicarbonate smolarmMmmolarolar 4074 07 4364 36 4514 51 4464 46 2562 56 sulfate smolarmmolarmM olar 273 250 344 323 354 2362 36 2142 14 2162.162 16 chloride smolarmMmmolarolar 648 676 127 130 132 6486 48 6596 59 659gsg6 59
1 dataD it i fronfromhromerom stevem 1977 2 dataD iti froromii wilberg inciinclI stollstolp 1985198.5 season eggs were laid under conditions in in early morning to 7 12c in late afternoon which water freezes at night once entrapping temperatures ranged from 1 3cac in early breeding males in the ice 21 march 1982 morning to 4 7cac in late afternoon in south breeding can be interrupted by a recurrence of central complex 15 under breeding condi- cold weather as suggested by observations on 27 tions this compares with breeding tempera march 1983 and 23 march 1985 when breeding tures of 6cac in british columbia licht 1969 frogs and tadpoles were concurrently observed and 14c in provo utah morris and tanner egg deposition and chorusingchorusing frogs occurred 1969 by june temperatures in 11 and 15 in until early april at 15 4 april 1981 although these same locations varied from 14 18c in only minor fluctuations of temperature 2 3cac early morning to 19 23c in late afternoon occurred at a given location throughout the day temperatures remained below lethal levels to where the source current flowed in march the developing tadpoles as determined by licht temperature could fluctuate at least 8cac during 1971 and turner 1960 the day in the peripheral locations where breed- adult breeding size measured during the ing occurred eggs were observed to hatch in breeding season ranged from 43 to 66 mm for late march and early april and tadpoles SVL females N 179 with the largest size class 12 mm were observed until early may in 1981 being 55 mm and from 40 to 59 mm for males after which no tadpoles were found after N 105 with the largest size class being breeding adults could not be observed until the 45 mm female size in tulethletuie valley is 5 mm end of may as indicated by the observation of 5 smaller and maximum adult sizes are also adults between the third week in april and the smaller than at other locations turner 1960 third week in may and 139 adults between the morris and tanner 1969 turner and dumas fourth week of may and the third week ofjuneofjune 1972 licht 1975 young of the year length less than 40 minmm SVL several studies have marked spotted frogs to based on the observation that breeding adults determine population numbers growth rates were larger than 40 mm were found from mid and movement turner 1960 carpenter 1954 june to late september one juvenile observed at south central complex 15 31 adult frogs on 2 may 1987 possibly metamorphosed the were marked and one frog was recovered in the previous year adults were seen until the end of subsequent year after 99 had been examined september the percent recovery of frogs was 3 com- spotted frogs typically avoided the warm pared to 40 from a total of 54 marked in waters ofthe springs two adult frogs were seen jackson hole wyoming in a single season car in waters with temperature ofof2828 C the breed penter 1954 and 27 from a total of 1433 ing habitat at north tule complex 11 in marked in a four year study in yellowstone march had temperatures thatvaried from 1 8cac turner 1960 use of the pederson lincoln 176 GREAT BASIN naturalist volume 53
TABLETABLF 3 life history summary of the spotted frog in tule valley
month februaryfebr uary marchmetreitreti AFaprilriiril mayMay lunejuneJ ie Jlulyjulyly augustAugiist septembersepteapte mbier
week 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4
adults i1 mating adults fresh eggs embryonic eggs tadpoles immature 2
iadalada 111 adult frogs wenew UKrare iiiliiin april and may with 5 adults seen from thediedle third week inin april to the thudthird weekk in may as winmincomparedpared to 139 frogs seen from the fointhfourth week inin may to the thudthird weekk inin june 9immature iininatine frogs weieweicwelmwerm defined ais havinghavinga a shontsnoirtsnont vent length less than 40 mm based on the observation that the SVL of breedingofhreeding frogfrogs was greatelgreater than 40 mm
index yielded an estimate of some 3000 frogs at within the north tule complex in a wet cycle spring 15 however this estimate is specula- like that occurring in 1980 84 a cycle that oc- tive since only one frog was recovered it would curred only once historically between 1865 and be necessary to mark many more frogs for popu- 1880 gwynn 1989 lation estimates to be reliable this one marked predation female frog measured 44 mm on 8 march 1981 upon spotted frogs and 53 mm on 12 june 1982 and thus grew 9 two aquatic insect larvae were observed on mm in 15 months this is comparable to growth egg masses predaceous diving beetle dytisci- rates reported by licht 1975 and much faster dae and crane fly tipulidae one other than those reported by turner 1960 it may aquatic insect belostoma bakerii belostomati- reflect warmer climatic conditions and longer dae giant water bug was found in the spring growing season source as well as the peripheral habitats and has population numbers can be estimated by egg a potential for preying on tadpoles the leech mass counts table 4 these counts reflect the haemopis marmorata was also observed on egg breeding population with each egg mass repre- masses and another leech erpobdellaErpobdella punctatapunctatepunctata senting a minimum of one female of note here has the potential of devouring tadpoles occa- is the tally for south central complex 14 sionallysionally partially devoured spotted frogs were this spring contains a small population of spot- observed in one case three regurgitated and ted frogs in 1981 there were 33 egg masses in partiallyartiartlallyaily devoured spotted frogs on land in leadsead one 1982 2 egg masses and in 1991 11 egg masses dead undevoured frog in the water and one live table 4 south central complex 14 contains frog several meters on land were observed this a sustainable population egg mass numbers observation suggested predation by a coyote suggest the largest populations of spotted frogs canis latranslalatranotrans birds such as brebesgrebes podicipedi- occur in coyote complex 26 south central dae herons ardeidae and ibises threskior- complex 15 and south tule complex 6 nithidaenithidae were rare in these aquatic habitats direct movements of spotted frogs in yel- mallards anas platyrhynchosplatyrhynchous cinnamon lowstonelowstone extended up to 1290 in in 92 days and teals anas cyanoptera and american coots 700 in in 23 days turner 1960 in jackson hole fulica americana were common hovingh the maximum total direct distance was 723 in 1992 predation on spotted frogs by these wa- over 24 days with some movements of 13 in per terfowl is unknown in tule valley northern day carpenter 1954 in tule valley the move- harrier circus cyaneuscycyaneousaneus and common ravens ments were confined to areas within springs and corvus c0raxboraxcorax are both common and potential wetlands as the land between springs consists of predators open spaces of carbonate soils among grease- wood in one year only movement occurred discussion from spring 6 to spring 5 for breeding a distance of about 4 in over carbonate soils habitats of the spotted frog in tule valley movements could possibly occur during the wet include the coldwatercold water portion of thermal season of april and may but adults were not springs this habitat contrasts with those in brit seen at this time movement could only happen ishisb columbia where spotted frogs occupy 199319931 SPOTSPOTTEDSPOTFEDFED FROG IN TULE VALLEY 177
TABLETABLF 4 egg mass census of spotted frogs in tuletnietuie valleyvailey
numbernumbers of egg masmassessesges date coyote north willow willow nortnorthnornnonnh tule complexcorncormcommplex south tule 26 15 14 11 10 7 6
7 march 1981 278 59 0 67 62 0 439 21 march 1981 a 33 h 21 march 1982 2 40 23 march 1985 0 5 2 march 1990 462 5 0 0 32 15 march 1991 11 43 46 26 additional egg masses that could not be quantiquantihedquantizedqowitifiedhed bdidbaiddid not survey shallow warm water regions of cold streams presently tule valley spotted frog habitats licht 1971 in distal reaches of the spring are very isolated from each other as the dry source in tule valley conductivity ph and tem- carbonate soils do not allow movement among peratureperature show the greatest variations in british springs the exception is in south tule complex columbia habitat showed the greatest tem- where spotted frogs moved from 6 to the un- peraperatureture variations thus the spotted frog util occupied 5 and now breed therein the wet izes habitats with widely varying physical and cycle of 1980 84 coalesced two springs in north chemical parameters tulethletuie complex 9 flowed into 11 that could A unique feature oftule valley aquatic habi- allow an exchange of frogs historically the wet tats is the high summer values of conductivity cycle has occurred twice 1980 84 and 1865 80 and ph that decline to pre summer levels in the in both periods the great salt lake reached an autumn and winter this water purification elevation of 1284 m gwynn 1989 during the the great can be attributed to another unique feature 0of holocene salt lake reached an eleva- these artesian springs much of the flow from tion of 1287 m currey 1990 but it is unknown this the springs is absorbed into the ground ground whether wet cycle allowed spotted frog water recharge and probably surfaces via seep- movement among the springs the wet cycles that caused the formation of gilbert shoreline age springs at the lateral edges of saline mud elevation 1295 m 10900 10300 years ago flats on each side of the artesian springs arte- currey 1990 and the gunnison sesersevier sian springs are on a small ridge 3 8 m above lake basinbasin drainage into the great salt desert these saline mud flats this elevational differ- lake viavla the old river bed 13000 11000 years ago ence allows the subsurface flows seepage oviatt 1988 may have affected distribution of springs adjacent to saline mud flats have high the spotted frog within tule valley aquatic habi- conductivity suggesting that groundwater sup- tats both north tule 11 and south tule 6 plying these follows springs an underground have extended dry outlets that once drained saline layer that underlies central basin is the it these springs to saline mud flats west of the in this morphology that cattle hooves can punc- springsspnngs holes of the wetlands thus ture in many and since aquatic habitats in tulethletuie valley were cause water to pour as miniature waterfalls back under extensive amounts of water during the into the ground thus these tule valley springs lake bonneville times 15000 years ago the have a unique morphology that has allowed the spotted frog had to emigrate to these new spotted frog to survive for some 13000 years in habitats on the valley floor such an emigration a saline environment and to utilize the distal occurred in tule valley springs during regres- portions of the springs without an increase in sive stages of lake bonneville 14500 years ago salinity the adjacent snake valley springs are a both fish and mollusk were exterminated during contrast to the tulethletuie valley springs in that the this time three models are suggested to explain low conductivity outflows to the wetlands drain the presence of spotted frogs in tule vallleyvalivailvavalleleyVallilleyley 1 into the gandy salt marsh pond thus maintain- spotted frogs were always in tule valley and as ing more constant conductivity and ph the lake rose these spotted frogs could always 178 GREATGRFAT BASIN naturalist volume 53 find new habitats in the periphery of the basin to protect the aquatic habitats from livestock 2 spotted frogs could have migrated from ad- and the carbonate soils the spring source of jacent snake valley dunngduangduring high water times in water and the groundwater recharge of the snake valley the frogs occupied habitats above water will protect frogs from anthropogenic air the high shoreline this would suggest the pres- and land pollution ence of spotted frogs in lake creek drainage spotted frog habitats in tule valley thus see fig 1 in southern snake valley however stand in sharp contrast to spotted frog habitats there are no frogs in southern snake valley along the wasatch front in eastern bonneville today and spotted frogs now occupy habitats in basin along the wasatch front the spotted frog snake valley that were flooded by lake bon is no longer found in many historic locations neville 3 all spotted frogsflogs lived min the deep suggested impacts include 1 fragmentation of creek drainage see fig 1 above lake bon habitat by highway cuculvertslverts dams reservoirs neville in western bonneville basin where they and urbanization 2 destruction of habitats by occur today after the lake stopped rising these reservoirs channeling ofofnversrivers diversion of wa- frogs moved along the shorelines of lake bon- ters for irrigation and preventing flood plains neville the ideal time for this movement was from being flooded and 3 impacts of man on during the relatively stable provo level of lake these habitats by utilization of livestock in nipar bonneville which lasted some 300 years after ian and wetland zones and introduction of rac bass the high water threshold broke in idaho ben- coons bullfrogsbullfrogs crayfish and trout to son et al 1990 the shoreline distance from the these habitats again the unique aspects oftulethletuie the frog deep creek refugia to tule valley viavla the provoprove valley namely locations of spotted the floor the saline of level approximates 340 km which includes habitats on valley nature movement around the northern end of the fish the habitats and the isolated nature of taletuletuie itself have much ofthe habitat springs range this distance is feasible under valley prevented destruction that has occurred along the wasatch conditions of movement in the wyoming turner 1960 carpenter 1954 during wet front years shorter routes over low passes might be feasible acknowledgments in view of the above discussion of the unique hydrology of tule valley in which spotted frogs I1 thank M hovingh and W hovingh for survived for 14000 years one might ask Is their some assistance in the field M hovingh for survival assured min the future presently tule computer assistance A kelner for assistance green valley springs are in a comparatively natural with maps and david and two anony- state water is pumped for livestock and for- mous reviewers for constructive comments on merly for oil well exploration from coyote the manuscript complex 26 and north tulethletuie 11 bulrushesBulrushes have been burned annually by livestock opera- literature CITED tors limited livestock grazing occurs resulting in holes punctured in the aquatic habitats which BFNSONBENSON L V D R CURREYCURRPY R I1 DORN K R LAJOIELAJOIF greatly dimmishdiminish the size of the habitats this is C G OVIAIIOVIATT S W ROBINSON G I1 SMHIISMITIL AND S scineslSTINEinelneINF 1990 chronology ofexpansionof expansion and contraction the most severe impact on the aquatic habitats of four great basin lake systems during the past 35000 members of the centiarchidaecentrarcbidaecentrarchidae sunfish were years paleogeography paleochmatology paleoecology unsuccessfully introduced in north tule 11 7824178 241 286 BOLENBOLFN 1964 plant ecology ofspringoxspringof fed salt marshes and 7 willows once grew in south central 15 E G spring in western utah ecological monographs 34 143 166 11 1946 and 14 and in north tule fautin carpenierCARPENTER C C 1954 A study of amphibian movement and now dead stubs and trunks exist A mining in the jackson hole wildlife park copela 1954 197 company claimed north tule complex for in-in 200 currryCURREYgurrey paleolakeslakes in evolution dustrialdu strial military low flying super D R 1990 quaternary paleo in the processing of semidesert basins with special emphasis on lake sonicsonie flights occur within taletuletuie valley and bonneville and the great basin USA paleogeography adjacent valleys however the effect of sonicsonie paleoclimatology paleoecology 76 189 214 ma- booms on amphibians is unknown on the posi- CURKIYCURREY D RGR G ATWOODAIVVOOD AND D R MABEYMABFY 1983 en- joror levels of great salt lake and lake bonneville utah tive side the bureau of land management geological mineral survey map 73 closed the springs south central dis41515 north curbeyCURREYCURRFYcurney D pannPANDPANDCC G OVIAIOVIATT 1 1985 durations average tule 12 and south tule 4 5 6 within fences rates and probable causes of lake bonneville espanexpan 199319931 SPOTTED FROG IN TULE VALLEY 179
sions stillstandsstillstands and contractions during the last deep momusMORRIS R L AND WWWW TANNER 1969 thehe ecology of lake cycle 32000 to 10000 years ago pages 9 24 in the western spotted hofhogbogfrog rana prefpretiosapretwwpretpreciosaiosalosa pretlowpret iosa bandbaird P A kay and H FE diaz eds problems olandof and pros- and girard a life history study gleatgreat basin naturalist pects for predicting great salt lake levels central 29 45 81 public affairs administration university of utah salt ovianovlanONITATT C G 1988 late pleistocene and liolloiioloceneholoceneIIolocene lake lake city fluctuations in the sevier lake basin utah USA jour- paulinpautinFAUIINFAUTIN R W 1946 biotic communities of northern desert nal of paleolimnology 1 9 21 shrub biome inm western utah ecological monographs SACK D 1990 quaternary geology thletniethieonieouie valley west cen 1625116 251310251.310251951 310 trai utah utah geological mineral survey open file GkGWYNNNYNN J W 1989 the saline resources of utah utah release 143 60 appp geological mineral survey notes 23 21 31 STMIENS J C 1977 hydrologic leconleeonreconnaissancenaissance ofot the HovINGHOVINGII11 P 1986 biogeographic aspects of leeches mol- tule valley drainage basin juab and millard counties lusks and amphibians in the intermountain region utah state ofofutaliutah department of natural resources publication great basin naturalist 46 736 744 technical 56 37 appp TURNTuRNE ii F B 1992 avifauna of tule valley western bonneville turnelltunn FB 1960 population structure and dynamics of the western frog rana basin great basin naturalist 52 278 283 spotted p pretiosapreciosapretwwprefpretiosalosa baird and cuardchardcirardguardgirard in yellowstone paliparipaikparkpalk wyoming ecological HovINGHOVINCII11 P B benlonBFNIONBENTON AND D 1985 bornholdt monographs 30 251 278 aquatic parameters and life history observations of the turnenTuRNETURNERii FFBBANDAND P C duniasDUMAS 1972 ranaranapretwsapretiosapretpreciosaiosalosa bandbaird great basin spadefoot toad in utah great basin natu bann and cirardguardgirard spotted frog pages ilg1191191 1 ilg119 4 in ralistcalistralist4545 22 30 1194 in cata- logue of american amphibians and leptepreptilestiles licaitlleliello iliIII111 L E 1969 comparative breeding behavior of the willWHITEIF G C D R ANDERSONANDFKSON K PR BUHNIIAM AND red legged frog rana aurora and the aurora western D L oilsarisoris 1982 capture recapture and removal spotted frog rana pretiosapreciosapret iosa pretiosapreciosapretiosa in southwestern methods for sampling closed populations LA 8787 british columbia canadian journal of zoology 47 NERP los alamos national laboilaboratoriesatones los alamos 1287 1299 new mexico 235 appp 1971 breeding habits and embryonic thermal re- WILBI RC D E AND 13 J SIOLPSTOLP 1985 physical charac ments of the quirequirements frogs rana auroraa aurora and rana te ristics and chemical quality of selected springs inhi pretiosapreciosapretiosapretiosapreciosapretiosa in the pacific northwest ecology 52 partspaitsofjuabofjuab millard tooelethoele and utah counties utah 116 124 USGS water resource investigations repoitreportrepoint 85432485 4324 1975 comparative life history features of the west- 39 appp einern spotted frog rana pretiosapreciosapretiosa from low and high elevation populations canadian journal of zoology 53 received I1 I1 farnaryFArtehruanjcarnarynafynarynarg 1992 1254 1257 accepted 7 Decedecemberniber 1992 great basin naturalist 532 appp 180 185
GROWTH OF smallmouth BASS micropterus DOLOMIEU IN FLAMING GORGE RESERVOIR WYOMING UTAH
I1 I1 scott A mullner andwayneaand wayneWayneA A huberthuberti
AIJS 1 KACI growth ai3sntacrallsalis raol of allmouthsinsmallmouth bass micropterus dolomieudolomieu was described for three widely spaced areas progressing downstream in in flaming gorge reservoir wyoming utah significant differences among areas were detected only for age 1 fish growth in the reservoir was comparedaithcompared Aithwith that ofpopulations in other lentic habitats ofvarying growing season lengths in north america slow growth in the reservoir was related to thetiletiietlle relatively short growing season less than 90 days
key mordwordwords mautnouthmaUtsinallinouthnouth bass micropterus dolomieudolo mieu growth population structure reservoir
the native range of smallmouth bass mi- 1949 coble 1967 and hubert 1975 shorter cropteruscroptemsterustenus dolomieudolomieu extended from ontario growing seasons limit the time that fish have to canada south to northern georgia and ala- actively feed and grow each year published bama west to oklahoma and southeastern kan- assessments of smallmouth bass growth rates sas and northeast to south dakota since the are from areas where the agricultural growing mid 1800s this species has been introduced to season is 90 d the western united states and canada greatly lack of smallmouth bass population dynam- expanding its range robbinsbobbins and maccrimmon ics information in flaming gorge reservoir and 1974 lee et al 1980 other waters in the western united states although not native to wyoming and utah prompted this study we assessed growth in the smallmouth bass has also established there three widely spaced areas in flaming gorge baxter and simon 1970 in 1967 it was intro- reservoir to determine if differences occur we duced to flaming gorge reservoir wyoming hypothesized that growth in the reservoir might utah pettengill et al 1983 flaming corgorgegor e differ longitudinally due to the decline in bio- reservoir 1841 m MSL when filled has an logical productivity from headwaters to the dam agricultural growing season of 90 d koss et al marleyvarleynarleywarley 1967 wiley and varley 1978 we also 1988 natural recruitment of micropterus sppapp assessed the influence of elevation and growing has not been observed 1900 m MSL in wyo- season on growth relative to other populations ming hubert 1988 short growing seasons and we hypothesized that growth in flaming above this elevation probably lead to winter gorge reservoir would be relatively slow mortality of young of year shuter et al 1980 flaming gorge reservoir is at the highest ele- STUDY AREA vation in wyoming where successful small- mouth bass reproduction is known to occur the green river a tributary to the colorado the northern limit of the native range of river was impounded in 1962 to create flaming smallmouth bass occurs where the growing sea- gorge reservoir in southwestern wyoming son is about 120 d robbins and maccrimmon sweetwater county and northeastern utah 1974 and populations there have relatively daggett county at full pool the reservoir is 147 slow growth doan 1940 watson 1955 and km long and has a surface area of 17000 ha turner and maccrimmon 1970 growth rate of two distinct topographic regions border the the species varies over its natural and intro- reservoir the upstream two thirds of the im duced range and is at least partially related to poundmentpoundment lies on an open plateau of sage growing season length bennett 1938 tate brush covered rollinerolling hillsbills the lower one third
I1 and wildlif11 kesegichiehichleh wyoming coopratioopeiative fish md wddhte reselkeselkeseiichKeseieardeara unit uniwrsityuniveisityuniversity of wyoming laramie wyomingwyomiwyoma ng 82071 the unit isis supported by the university ofot game 1 wyoming wyoniingwyoniiug gam indand fish dqwfindtptutinenttandditluitl USU S fishfislipish aind wildlifwildlitiwildlifeWild liflitiilfe selvleeSCIVILC
180 199319931 smallmouth BASS GROWTH 181
t blacks fork green METHODS river river smallmouth bass were collected 5 20 june INFLOW 1991 from one sampling area in the inflow re- REGION gion and two sampling areas in the open hills inflow sampling area region fig 1 using boat mounted electrofish ing gear total length TL was measured and scales were collected from each fish following the methods ofjearaldofjearald 1983 scales were read 19x using a micromicrofichefiche reader and age of each fish was estimated from scale annuli fish length OPEN upper open hills at each annulus was back calculated using a 11 HILLS sampling area 35 value REGION mm intercept as suggested by carlan der1982der 1982 mean TLs at each annulus were assessed for
0 10 differences among sampling areas using one kilometers lower open hills way ANOVA sokal and rohlf 1981 A chi sampling area square test for heterogeneity was used to WYOMING determine differences in age structure among UTAH sampling areas lilinearneafnear regression analysis was used to assess the relation between mean TLs of age 4 small- oreengreen mouth bass and agricultural growing season length for 31 lakes and reservoirs across north america A sample of 25 smallmouth bass from CANYON REGION gorge flaming dam the downstream end of flaming gorge reser- was also al fig 1 flaming gorge reservoir showing the locations of voir included pettengill et 1983 the three areas from which smallmouth bass were sampled age 4 fish were compared because this age group occurred in all populations described in the literature we used and back calculated is in a wooded area of high mountains and a lengths are less affected by the method of calcu- deeply cut gorge varley 1979 lation at age 4 than at younger ages the growing the reservoirreservoii is generally separated into season length for lakes and reservoirs was deter- three regions fig 1 based on topography ge- mined using koss et al 1988 lakes were ology and hydrographic features varley 1967 classified in 30 d intervals and the upper limit wiley and varley 1978 and varley and livesay of each interval was used as the independent 1976 water quality grades from eutrophic at variable in the regression analysis the headwaters to almost oligotrophic near the statistical calculations were performed using dam varley 1967 the inflow region north- STATISTIX 313.1 analytical software 1990 and ernmost 32 km is influenced by the green and significance was determined at P 05os05.05 blackblaekblackss fork rivers water temperature there is warmer than downstream an oxygen deficient RESULTS zone sometimes occurs and maximum depth is 24 in the open hills region extends 48 km mean back calculated TLs at each age for downstream from the inflow region it is char- the smallmouth bass collected from each of the acterized by extensive bays and littoral shelves three sampling areas in the reservoir were com- maximum depth is 61 in thermal stratification pared table 1 ANOVA indicated a significant is not prevalent because of wind action the difference P 011 among locations only for canyon region extends 39 km from the open hills agelage 1 fish age structure of smallmouth bass region to the dam this region is narrow with from the three areas was computed table 1 sheer walls water is well oxygenated and ther and a chi square test indicated no significant mally stratified and maximum depth is 122 in differences in age distributions among areas 182 GRFAT BASIN naturalist volume 53
tahltahleTAHITABLE 1 calculated meantotalmean total lengths at annulus formationfOrination of smallmouthofsinallinouth bass collected fromfroin three sampling areasarcas ininflowflow INF upper open hills UOII and loweropenlowelower openropen hillshilis LOII in Flflamingarning gorge reservoir wyoming utah
mean total length minmm at each annulus year sample class site siesize I1 II11 III111 IV V VI vilVII villVIII IX X XI XII
1990 INF 13 95 UOIUOII1 3 97 lelilellhohhonLOIILOH 5 86 1989 INF 17 99 136 UOIUOII1 3 102 149 lolLOILOII1 9 85 122 1988 INF 25 97 128 157 UOII 2 99 134 182 LOII 16 89 128 159 1987 INFINIT 13 106 142 178 201 UOII 4 98 142 178 222 lolloillollLOILOII1 17 96 136 169 199 1986 INFINIT 12 104 14114 176 206 209 UOIUOII1 6 102 142 186 221 263 LOII 18 106 142 180 215 247 1985 INF 10 121 152 182 211 239 263 UOII 6 108 143 171 201 230 263 lolLOILOII1 15 102 131 157 190 219 252 1984 INF 12 100 130 160 190 219 24241 270 UOIUOII1 8 106 141 178 213 245 274 258 10LOII11 9 Hiiiili111 147 174 208 243 279 296 1983 INF 14 99 135 166 199 228 259 262 285 UOII 3 104 134 160 188 226 259 293 329 loliloil1011 2 112 149 177 208 243 279 296 316 1982 INF 7 103 139 166 193 219 239 265 293 317 UOII 2 102 143 177 210 250 288 323 347 368 LOII11011 4 100 133 ifo160igo 188 22221 246 203 226 242 1981 INF 9 104 135 163 193 223 250 277 303 330 350 UOII 1 115 163 216 269 306 338 354 365 375 391 loilLOIILOJI 2 105 134 167 215 253 282 302 325 353 381 1980 INF 1 100 129 149 164 194 214 234 259 288 318 348
UOII 1 100 133 176 214 236 258 285 328 356 372 394 LOII11011 3 104 140 186 211 244 273 304 329 350 372 394 1979 INF 1 104 134 163 188 208 227 247 267 301 326 341 361 UOII 0 LOII 0 grand INF 134 103 137 166 194 217 242 259 281 302 331 344 361 mean UOII 39 103 142 180 217 251 280 302 342 366 382 394 LOII 100 100 136 170 204 239 268 281 299 315 377 394
back calculated TLs of smallmouth bass where days is the upper limit of the 30 d interval from all sampling areas were pooled to evaluate in which the various populations occurred fig the influence ofgrowingof growing season on growth of 2 As growing season length increased mean smallmouth bass A significant Vrar2 3030.30 P TL of smallmouth bass at age 4 increased the 0012.0012 regression equation was obtained two flaming gorge reservoir studies were the only ones where a 90 d growing season oc- mean length mm at age 4 187 07770.777 days curred and they had the smallest fish at age 4 199311993 smallmouth BASS GROWTH 183
600
500 t CD 400 CU
300
200 cc
100
0 90 120 150 180 210 240 270 growing season days
fig 2 mean total lengths at age 4 for smallmouth bass from 31 lakes and reservoirs with laiyingvaiyingvarying growing seasons all lakes and reservoirs with growing seasons within a 30 day interval were plotted at the upper limit of the interval the lakes and reservoirs included the following 90 d flaming gorge reservoir wyoming present study pettengill et al 1983 120 d owen lake wisconsin bennett 1938 shadehillShadehill reservoir south dakota willis et al 1990 big lake maine watson 1955 and weber lake wisconsin bennett 1938 150 d tedenac lake ontario tinnerturner and maccinnmonmaccrimrnon 1970 lake Opeopeongoongo ontario doan 1940 winola lake pennsylvania carlandercallandercallanCailandeider 1977 cayuga lake new yolkyork webster 1954 oneida lake new york forney 196iggi19611 lewis and clark reservoirReservon south dakota willis et aal 1990 lake mcconaughy nebraska mccarraher et al 1971 hiwassiehiwasseeiiiwassee reservoir north carolina stroud 1949 and claytor lake virginia roseberry 1951 180 d santeehlah lake north carolinacalCai olina messer 1961 clarke lake pennsylvania carlandercallanCailandeidel 1977 maleomMalcommalcomsenserlSs pond illinois bennett and chiichildersChildeisdels 1957 connowingoConnowingo pond maryland heisey et al 1980 clearwater lake missouri patriarche and campbell 1958 quabbin lake maine mccaig and mullan 1960 noinorrisus reservoir tennessee stroud 1948 allerton lake illinois bennett and childers 1957 lock raven reservoir maryland carlander 1977 and Centecenterhillrhill reservoir tennessee hargis 1965 221010 d little lake oklahoma finnellfmnelletFmnelletet al 1956 pickwick reservoir aiaAlaalabamabarna hubert 1975 lake eucha oklahoma jackson 1966 and onachachitaouachachita lake alkansasarkansas hulsey and stevenson 1958 240 d fort gibson lake oklahoma carlander 1977 and P olsonoisonfolson lake california tharratt 1966 270 d pine flat lake california emig 1966
discussion at the northern edge of their native range smallmouth bass grow considerably faster than in flaming gorge reservoir probably due to among the three sampling areas the mean mean differences in growing season length fig 2 length at each age did not differ significantly for example growth rates were much faster in except for age I1 smallmouth bass the smallest tedenac lake northern ontario canada 1 fish the agelage I were in most downstream sam- turner and maccrimmon 1970 and big lake pling area which was least eutrophic young maine watson 1955 than in flaming gorge smallmouth bass in lentic habitats begin feeding reservoir smallmouth bass in southern reser on zooplankton then insects become important doirsvoirs such as pickwick tennessee hubert in the diet and finally crayfish and fish coble 1975 centerhillCente rhill tennessee hargis 1965 and 1975 possibly food resources for small fish folsom california tharratt 1966 increased in were less abundant in the most downstream TL at twice the rate observed in flaming gorge sampling area lack of significant differences in reservoir TLs ofofageofagoage 2 and older fish maybe due to similar smallmouth bass in southern waters grew abundance of crayfish and forage fishes among faster bennett 1938 coble 1967 but stroud the sampling areas or to the ability of larger fish 1948 concluded the faster growth in warmer to move among areas waters also contributed to earlier death small 184 GREAT BASIN naturalist volume 53
mouth bass in flaming gorge reservoir grew DOAN K H 1940 studies of the smallmouth bass journal slowly and fish up to age 12 were recorded the of wildlife management 4 241 266 EMIG J W 1966 smallmouth bass pages 354 366 in A maximum recorded age for smallmouth bass is calhoun ed inland fisheries management california 15 years stone et al 1954 scott and crossman department of fish and game sacramento 1973 when the population structure of small faf1finnellFINNFLLNNFLL J CRC R M JENKINSJFNKINS ANDGAN D G E HALL 1956 the mouth bass in flaming gorge reservoir is com fishery resources ofthe little river system mccurtainMcCurtam county oklahoma oklahoma fisheries research pared with that in other waters it is evident that laboratory report 55 oklahoma department ofwild- fish in flaming gorge reservoir live longer than life oklahoma city those in most other waters FORNEYFORNFY J L 1961 growth movement and survival of despite the elevation of flaming gorge res- smallmouth bass micropterus loloimeudolomieudolololoimieumeu in oneida lake new york new york fish and game journal 8 ervoir the smallmouth bass population is self 88 105 sustaining however growth rate is among the 1972 biology and management of smallmouth bass slowest recorded many factors could cause this in oneida lake new york new york fish and game brown 1960 coble 1967 keating 1970 and journal 19 132 154 haiwisHA R gisCIS H L 1965 age and growth of micropterus sal fomeyforneybomey 1972 and our analysis indicates that a hairis moilesmoldes micropterus dolomieudolomieu and micropterus buncpunc short growing season is a significant limiting tulatus in center hill reservoir tennessee factor the smallmouth bass population in unpublished mastermasterss thesis tennessee polytechnic flaming gorge reservoir may have the shortest institute cookvilleCookville iirisryheiseyhelsey P G D MAIIIUR AND N C MAGNUSSON 1980 season of self popula- growing any sustaining accelerated growth of smallmouth bass in a pumped tion and the slowest growth on record for a storage system transactions olof01 the american fisheries lake dwelling population society 109 371 377 huberthuberihunertHUBFRI W A 1975 age and growth of three black bass species in pickwick reservoir proceedings of thetlletile an- acknowledgments nual conference of the southeastern association of game and fish commissioners 29 126 134 sampling and for this 1988 altitude as the determinant of distribution of field funding study largemouthlaigemouthlaigemouth and smaUsmallmouthmouth bass in wyoming north were provided by the wyoming game and fish americanamerlean journal of fisheries management 8 386 department we appreciate the field assistance 387 ofmof M fowden K johnson and W wengert and HULSEYHUISFYAA II11 AND J II11 SIFVINSON 1958 comparison of of fish reviews of K P marsh growth rates game in lake catherine lake manuscript ofkoak carlander J hamilton and lake ouachitaOu achita arkansas proceedings zuboy and an anonymous referee of the arkansas academy of science 12 17 3311 JACKSON S WJRW JR 1966 summary of fishery management activities on lakes eucha and spavinawSpavinspavmawaw oklahoma literature CITED 1951 1964 proceedings of the annual conference of the southeastern association of game and fish com- analyticalANAIYIKAI SOIIWAHISOFTWARE 1990 STATISTIX manual anal- missioneismissiomissionersneisners 19 315 343 ytical software st paul minnesota JFARALDJEARALD A JR 1983 age determination pages 301 324 BAXIIKBAXTER G T AND J R SIMON 1970 wyoming fishes in L A nielsen and D L johnson eds fisheries bulletin 4 wyoming cainecalnegainegame and fish department techniques american fisheries society bethesda cheyennecheGheyenne maryland bfinniiibl NNI I1 I1 G W 1938 growth of the smallmouthedsmallsmail mouthed black kea7rlnKJAIINC J PF 1970 growth rates and food habits of small- bass microptenimicroptents iolomieudoloiniett lacepedeLace pede in wisconsin mouth bass in the snake clearwater and salmon riv-riv mateiswateiswaters copela 19384 157 170 ers idaho 1965 1967 unpublished mastermasterss thesis benniabennijbl NNI iii G W AND W F CHILDEIRSCIIIIDI HS 1957 the small- university of idaho moscow mouth bass micropterwmicropterus iolomieudoloinieu in warm water koss W J J R OVVFNBY P M STEURER AND D S ponds journaljoin nalnai of wildlife management 21 414 424 ezellEFLL january 1988 climatology of the united BROWNBKOWN E II11 JR 1960 little miami river headwater states number 20 supplement 1 national climate streaminvestigationsstream investigations federal aid project FIRfirF I R ohio data center asheville north carolina department of naturalofnatural resources columbus lreleeLE E D SSCC R GIGILBFRIL B F RT C H hocinhocunhoguttCUTT R E JElenkinsjenkinsJFNKINSN K I1 N S CARLANCARLANDIDiRK K D 1977 handbook of freshwater fishery D E McALLISTMALLISIFRFR AND J R slaufffrstaufff11 JR 1980 liobiologyiioilo logy vol 2 iowa state university press ames atlas of north american freshwater fishes publication 1982 standard intercepts foifor calculating lengths 1980121980 12 north carolina state museum of natural from scale measurements for some centrarchid and history raleigh percid fishes transactions of the american fisheries MCAIM cauCAR R S AND J W MULIANMULLAN 1960 growth of eight society 111 332 336 species of fishes in quabbin reservoir massachusetts coincolnCOBLE r 1D W 1967 relationship of temperature to total in relation to age of reservoir and introduction of smelt annual growth in adult smallmouth bass journal of the transactions of the american fisheriesFishenes society 89 fisheries researchReseaichsealch board of canada 24 87 99 272711273131 1975 Smallismallmouthsmallinouthnouth bass pages 213321 33 in H clepper mccarrauirmccarraiierMCCARRA ilerllerIIER D BBMM L MADSENMADSFN AND R E THOMAS eablackkablackedea blackblaek bass biology and management sport fishing 1971 ecology and fishery management of institute washington DCD C mcconaughy reservoir nebraska pages 299 311 in 199319931 smallmouth BASS GROWTH 185
G E hall ed reservoir fisheries and limnology tafetareTAFFTATE W H 1949 growth and food habit studies ofsmallof small- american fisheries society special publication 8 mouth black bass in some iowa streams iowa state american fisheries society bethesda maryland college journal of science 23 343 354 MESSFRJmesserMFSSFR J 1961 tennessee river drainage reservoirs fed TutharratttliarrarrtuarrattARRATT R C 1966 the age and growth ofeentrarchidof central ehidchid eral aid project fsr job completion report 1 north fishes in folsom lake california california fish and carolina wildlife resources commission asheville game 52452324 4 16 patriarche M HHANDRAND R S CAMPBELL 1958 the de- TURNFR G E AND 11 R maccrimmonMACRIMMON 1970 reprocepro velopment of the fish population in a new flood control duction and growth of smallmouth bass microptemimicroptenis reservoir in missouri 1948 to 1954 transactions of the dolomieudolomieu min a precambrian lake journal of the fish- american fisheries society 87 240 258 erieserles research board of canada 27 395 400 PFTTFNILL T D S BRAYIONBRAYTON J J JOHNSON D DUFFK VARLFY J D 1967 plankton periodicity as related to the W WENGERT AND M SNIGG 1983 flaming gorge chemical physical and biological environment of reservoir fisheries investigations 1983 annual report flaming gorge reservoir publication 67667 6 utah publication di- 840784 07 utah department of natural res- vision of wildlife resources salt lake city sou salt ourcesleesices lake city 1979 the influence of rainbow trout grazingglazing on ROBBINS W H AND H R maccrimmonmatMAC CRIMMON 1974 the zooplankton in flaming gorge reservoir publication black bass in america and overseas biomanagementBiomanagement 75575 5 utah division of wildlife resources salt lake and research enterprises sault saint mariemarlemane ontario city ROSFBFRRY D roseberry A 1951 fishery management of claytor VARLEYVARLFY J D AND J C livesayLIVFSAY 1976 utah ecology and an the new lake impoundment on river in virginia life history of the utah chub cliagila affatratrariaadranaatranaariaafia in flaming transactions of the american fisheries society 80 gorge reservoir wyoming utah publication 761676 16 194 209 utah division of wildlifeofwildlife resources salt lake city sconSCOTT W BAND E CROSSMAN WBANDE J 1973 freshwater fishes walsonWAISONWATSON J E 1955 the maine smallmouth fisheries re- of canada fisheries research board of canada bulle- search bulletin 3 maine department of inland fish tin 184 and game augusta SIIUIFRSHUTER B FRY J J A MALLEANMACLEAN F E J AND H A WFIFRWEBSTER D A 1954 allmouthsmallmouthsinallmouthSin bass microptenis RFIGFRRFIGER 1980 stochastic simulation of weier reiger temperature dolomiettdolomieudulodoloduiomiettmieu in cayuga lake part 1 life history and effects on first year survival of smallmouth bass trans environment cornell university agricultural experi- actions of the american fisheries society 109 1 34 ment station memoir 327 ithaca new yolkyork SOKAL R R AND F J ROIILF 1981 and2nd ed biometry WILEYWILFY R WWANDJAND J D varleyVARLFYVA RLEY 1978 diet of rainbow and W H san pianplan wandl freeman franciscoFianelseocisco california brown tiouttrout from flaming gorge reservoirRe servon 1964 SIONFSTONE U B D G PASKO AND R ROECKERROFC sioneslone M rofo KER 1954 A through 1969 fisheries research report monograph of study lake ontario saint lawrence river small series 1 wyoming game and fish department chey- mouth bass new york fish and game journal 1 1 26 enne SIROUDSTROUD R 11 1948 growth of the basses black and crap- WILLIS D W C L MILEWSKIMILFWSKI AND C S GUY 1990 in norris reservoir tennessee journal of the pie ten- growth of largemouth and smallmouth bass in south nessee academy of 23 31 99 science dakota waters prairiepi ainealne naturalist 22 265 269 1949 rate of growth and condition of game and panfish in cherokee and douglas reservoirs tennes- see and hiwassiehiwassee reservoir north carolina journal received 19 may 1992 of the tennessee academy of science 24 60 74 accepted 10 novemberNovembeibel 19927992 gigreateat basin naturalistnatuimatui ahstabst 532 appp 186 189
AUTUMN AND WINTER FOODS OF THE LESSER PRAIRIE CHICKEN tympanuchus pallidicinctus galliformes tetraonidae
1 terrytenyztenyaZ riley charles A davis2 and randall A smithsmithasmith33
absurda13siiiai iescriptiotisdescuptions of lessellesser prairieprame chicken tyinpanuchusrijmpanucjius pallidicinctuspalhdicinctiis foods in new mexico have not included comparisoneomcom panson between autumn and winter seasons we analyzed and compared prairie chicken crop contents in autumn 1976 n 9 and 1977 n 177 and winter 1977 n 4 and 1978 n 2 in a shinnery oak quercus havardii grassland in southeastern new mexico autumn foods weiewelewere a mixture of seeds x 43 vegetative material x 39 and insects x 18 especially shinnery oak acorns and insect gallsgaus x 49 short horned grasshoppers acrididae x 155 alsoweiealsoaiso weiewelewere inan important food winter foods were shinnery oak acorns x 69 and wild buckwheat eriogonum annnannuiiniuinnannnuntuhtuhlunt x 14 use of vegetative material and insects decreased fromfroin autumn to winter whereas use of acorns inciinclincreasedeased high plains bluestembluestein subtype in the southern mixed plaineprairie isis an important habitat that provides many of the foods eaten bypiameby prairie chickens therefore broad scaleseale disturbances of this community should be avoided
keyK i wordwordswory apodfiodfood bedinfeedingeedin lesser prairie chicken new mexico shinneryshinneiyhinnerioakoak tympanuchus pallidicmctuspallidicinctus
lesser praipralaraineprainerie chickens tympanuchus pal tween autumn and winter the purpose of this micinctusfidicin6tim occupy semiarid grasslands that study was to provide a description of autumn typically include a large component of shrubs and winter foods of lesser Pprairierairalrie chielenschickens eitheleitherelthelelther shinnery oak Querciquercusis havbavardhavardarditii or sand the objectives were to collect crops of prairie sagebrush artemisiafififoliaartemisia filifoha A description of chickens during autumn and winter to analyze foods used by prairiepran le chickens in the shinnery crop contents to determine the type and amount oak grasslands is incomplete Davidavisdavisetalsetalet al 1981 offoods consumed and to compare and contrast compared spring and summer diets in shinnery diets between seasons and years oak grasslands of eastern new mexico they found that prairie chickens feed on green leafy STUDY AREA vegetation in spring but change to insects in summer frary 1957 found that insects aiealeare the study area is approximately 15500 ha of important in early autumn sep oct in eastern bureau of land management lands in chaves new mexico crawford and bolen 1976 evalu- county in southeastern new mexico topogra- ated autumn diets of 90 lesser frainefrairieprairie chickens phy is gently undulating climate is semiarid collected in mid october from shinnery oak with distinct seasons and wide ranges of diurnal habitats of west texas despite the fact that their and annual temperatures nearly 75 of the study area included cultivated grains shinnery annual precipitation 30 year xX 345 amyrmmyr oak and insects were the principal natural foods falls during the growing season may through A seasonal description and comparison of october mainly from brief but often intense foods used by lesser prairie chickens in shin thunderstorms US department of com- nery oak grasslands through the entire annual merce 19761977 cycle could be used to assist land managers in the study area is in the southern mixed manipulating habitats to provide food resources prairie type where the high plains bluestem throughout the year however published de subtype grades westward into the desert prai- scriptscnphonsscriptionsionslons of lesser prairie chicken foods in rie subtype holechek et al 198979 most of shinnery oak grasslands do not differentiate be the study area 89 is on deep sandy soils
1lowilpiitiiitiitofa dcpartnent of Naturalnitin tl llonlion11ktsoinccson charitcharliinntonannton nsebesaRSCresa itdliditu station ruralburalburairiinlhoutiromerowe 1 1-box 203209 cllintoncclintonCllinton 1-loolow i 500450049 lp4tiiiiitoldcpartnentof I1 isliuits nidhiduld wildlifeVild lilill111 suniolomiobomio N mimtmlostthM i state univusityuni isittisity lasis cricrticesCrtcorticesHILLSiceslees newnowN- MCMLO 88x3 af3fh lislleslllslt hrs USDA coronado01011 idonnationalitionil flor1 utstoosthutstt300410 WV congvsstmsononicss I1 acsonncson ailaariaarlaailariai 10111a 85701
186 199311993 PRAIRIE CHICKEN FOODS IN AUTUMN AND WINTER 187
where vegetation is dominated by various com- trapping were similar to those collected by other binationsbinations ofbluestem grasses andropogon hal methods within the same season uiiiiliilil and A scoparius and shinnery oak that between year differences were noted for characterize the high plains bluestembluestein subtype autumn diets table 1 use of mast and seeds the remaining 11 of the area comprises scat primarily shinnery oak acobsacorns in 1976 diets x teredcered inclusions oftighter soils where vegetation 65 was significantly greater P 0505.05 than is dominated by short grasses especially grama in 1977 x 21 insects x 30 primarily bouteloua sppapp and buffalograssbuffalograss buchloe dac short horned grasshoppers and a variety of tytyloidesloides characteristic of the desert prairie vegetative material x 49 comprised a subtype snakeweed xanthocephalusxanthocephaluinxanthocephalum greater P 05os05.05 proportion ofdietsof diets in 1977 sarothraesarothrae and mesquite prosopis glandulosaglandulose than in 1976 animal material xX 7 vegetative are conspicuous invaders ofthis subtype in some material x 28 parts of the study area winter samples were combined between years because of the small sample size 1977 METHODS n 4419781978 n 2 foods consumed by lesser Prairieraidleprairie chickens in winter primarily consisted crop contents were from birds collected of shinnery oak acorns x 69 with lesser during autumn oct dec 1976 n 9 and 1977 amounts of green vegetation x 26 and n 17 and winter jan feb 1977 and 1978 n insects x 5 table 1 no differences P 6 mostly by shotgun or small caliber rifle 05os05.05 were detected in crop contents between two were taken from birds that died as a result autumn 1976 and winter 1977 78 shinnery oak of trapping in autumn 1977 and two were do- acorn composition of winter crops x 69 nated by hunters in autumn 1978 data from was greater P 0505.05 than in autumn 1977 crops empty crops were not used in the analysis we use ofvegetative material and insects resulting stored the frozen crops in loomi100 mlm1ma plastic storage primarily from lack of short horned grasshop bottles until analysis contents from each crop pers in winter was lower P 05os05.05 in winter were analyzed separately foods were measured crops than in crops collected in autumn 1977 by volumetric displacement to the nearest 010.1oi ml and items measuring oloi0010.1I1 ml werewere classified as trace composition of diet was determined discussion by the aggregate percent method martin et al 1946 we used borror and white 1970 and differences in autumn diets ofprairie chick correll and johnston 1970 to identify food ens between years might be explained by the items fact that annual precipitation was nearly 100 means and standard errors were calculated minmm 27 below normal in 1977 x 250 mm when an individual food item was detected in US department of commerce 1976 1977 two or more crops small sample sizes reduced lowelowerr than normal precipitation in 1977 might the power of statistical tests but we used stu- have affected the availability of food resources dentss test to test the hypothesis that there t were shinnery oak provided acobsacorns insect galls no differences in the composition of food items and leaves which together constituted 50 of between seasons and years snedecor and the autumn diet in our study and 36 of natural cochran 1989 differences were considered foods in crawford and bolenboienbolenss 1976 study significant at P 0505.05 despite the fact that crawford and bolens 1976 study area included grain fields shinnery RESULTS oak was the principal natural food in autumn in both studies shortshontshonn homedhorried grasshoppers were autumn diets primarily consisted of shin- the principal animal food in autumn frary nery oak acorns short homed grasshoppers 1957 reported crop contents from 17 lesser acrididae broom groundsel senecio spar Pprairierairalrie chickens collected in eastern new mex- tioidesnoides leaves and insect galls from shinnery ico about 45 linkmkin northeast of our study area his oak table 1 thirty different food items were sample was pooled across 6 months and so identified nearly half ofwhich were green vege seasonal comparisons with our data are not pos- tation crop contents from birds collected by sible he did find however that insects were hunters and from birds that died as a result of important in early autumn sep oct 188 GREAT BASIN naturalist volume 53
fabiTABITABLE 1 I1 mean composition otof autumn oct dec and winter jan feb crop contents of lesser prairie chickens chaves county new mexico 1976 78
autumnautaubiianiimn
1976 1977 winterWinter food item nann99 n 17 n 6 mast and seeds queraquercus havardhavaranhavardnhavardiiii acorns 61 12712 7 17 69 69 64 euphorbiaeuplwrbia sppapp 4 22 latliopennumlithosper7nuin incisuininu&uminciinclsuin 4 b Dithyditjiiraeddithyraearaea loishloishzenitvislizenizenizent tt11 total 65 130 21 68 69 64 vegetative material quercusuercim havardhavaranhavardnhavardiiii insect galls 14 82 5 17 leaves 1 2 seneciospadioidesbenecioenecio partioiiles 12 61 5 34 dalea nana 6 50 7 24 arioleriolertoonum0nuin annum 7 27 14 42 phlox sppapp 4 19 4 20 lit1wpennumlithosperutuin inuwmincisum 4 composite 4 oenothera0enotheraocnotliera sppapp 4 ttil b euplioihuieiiphorhia sppapp 3 1 tt13 1 hijmcnoihyinenoxys sppapp 2 tt11 1 xanthocephabintxantlioccphalum wrothsarothwrothraesarothraesavothrae 1 tt1ta Penrenpcmtciiionpenstenumstenum buckelhuckehibucehuckbuckelyibuge elteitehiyi tbt 2 others t1tatii tb total 28 138 49 93 26 63 animal matenalmaterialmanenal acrididae 2 14 28 82 gryllidaegryllCryllidae 3 03 lepidoptera 2 1 carabidaecarali dae tt1ta 5 36 otherss tbt1ta 2 ttb total 7 36 30 82 5 37
S E hfiaafia
similarities between autumn 1976 andainand win management implications ter 1977 78 diets probably were a result of the fact that most 46 crops collected in winter lesser prairie chickens are closely associassoni were from 1977 and the availability of acorns atedabed with the shinnery oak grassland commu- was similar between seasons differences be- nity in much oftheir occupied range within this tween autumn 1977 and winter 1977 78 diets community in new mexico lesser prairie probably resulted from the fact thatvinterthat vinterwinter diets chickens obtain most of their autumn and win- diets from a rather small number of plants were more a reflection of winter 1977 than ter and associated insects that are common in the 1978 and below normal precipitation in 1977 less grassy habitat shinnery oak is the most might have reduced acomacorn production in the heavily utilized food of prairie chickens on an and the food area increased demand on other annual basis shinnery oak acorns cacatkinsbatkinstkins 1957 sources during autumn frary found that leaves and galls in various combinations provide vegetative material is important in winter diets adult birds with 50 of their diet in autumn and that acobsacorns are important in winter jones and winter because of the importance of shin- 1963 working with prairie chickens in okla- nery oak grassland to prairie chickens for both homa showed the importance of mast and seeds food and cover broad scale eradication of this in the winter diet community should be avoided 199311993 PRAIRIE CHICKEN FOODS IN AUTUMN AND WINTER 189
acknowledgments in new mexico pages 75 80 in PR A vohs andcind F L knopf eds proceedings of the prairie grouse samposympo this research was funded in part by the US Slumsiumslum oklahoma state university stillwater FRARFRARYY L G 1957 evaluation of prairie chicken ranges bureau of land management BLM the new PRP R completion report project number W 77 R 3 mexico department of game and fish pro- new mexico department of game and fish santa fe vided collection permits we are indebted to 81 appp HOLFCIIFKHOLECIIEK L R D PIPERpipi R 11 HERBELL W J wisdom H R suminski and other for- J AND C II hr hblRBI 1989 range management principles and practices prentice mer wildlife students at new mexico state uni- hall englewood cliffs new jersey 501 appp versity for field and lab assistance this is a new lonesJONESJONFS R E 1963 identification and analysis oflesserandof lesser and mexico state university agricultural experi- creatergreater prairie chicken habitat journal of wildlife ment station journal article management 27 757 778 MARTIN A C R H GENSCII AND C P BROWN 1946 alternate methods in upland game bird food analysis literature CITED journal of wildlife management 10 8 12 SNEDECSNFDFCOKolioltOii G W AND W G CCOCHRANibanIRAN 1989 statistical methods iowa state university press ames 503 BORROR D JJANDRAND R E whireWHIFFWHITE 1970 A field guide to the appp landr US dfparimfntoidepartmentofDEPARTMENTOF GOMMFRCFCOMMERCE 1976 insects of america north of mexico houghton mifflin monthlysummamonthlysurnim nzedrizedprized co boston 404 appp station and divisional data climatological data new mexico 81 1 12 CORRELLCORRFLL D S AND M C JOHNSTON 1970 manual of JOIINSION 1977 summarized the vascular plants of texas texas research founda- monthly station and divisional data climatological data new 1 12 tion renner 1881 appp mexico 82 CRAWFORD J A AND E CG BOLFN 1976 fall diet of received 26 may 1992 lesser prairieprame chicken in west texas condor 78 142 144 accepted 10 febntar7lfcbnian 1993 DAVIS C AATT Z RILEYRILFY R A SMITH AND M J WISDOM 1981 spring summer foods of lesser prairie chickens greati eat basin naturalist 532 appp 190 193
establishment OF SHOSHONE SCULPIN COTTUS GREENEI IN A SPRING INHABITED BY MOTTLED SCULPIN C BAIRDI
1 1 derek B kuda and J S griffithgriffithagriffith1
abstractabstracrA us 1 H I1 ihetheahe shoshone sculpin cofcontusCotcottustits greenelgreeneinel is found only in springs oftheodtheof the thousand springs formation along tirtinthe snake rivelriver in idaho in 1983 a small population of shoshoneofshoshone sculpin was introduced into an unnamed springspnngspang in the thousand springs formationfoifol mationmatlon in an attempt to increase the langeiangerange oftheodtheof the species previously the only sculpin in that spring was thetiletlle mottled sculpin comCofcoffillcoffilsC ottucottufilsmils bairdihairdihabanhanbauirdidi therhe shoshone sculpin was able to establish itself and become the predominant fish within 8 years
scam keyK if luorallordluord shoshonestsiStsasi si sculpinscpm cottusottus greenel mottled sculpin cottus bandibandlbairdi syrnpatricsympatnc speciespeclpeciei species of special concern snakebuiehuiebuikiuik river idaho
As of 1982 the shoshone sculpin cottus METHODSMFTHODS rhegreereeneareeneireeneinet was found in only 25 of 40 spring sys- tems in formation in the thousand springs near shoshone sculpstulpinssculpinssculpmsins were introduced into a hagermanIIa in south central idaho wallace al ilallagennan in et small unnamed spring pond as part of an idaho 1984 the species principally inhabits springs department offishof fish and game nongame pro- the north side of from entering the snake river gram to reestablish them in portions of their rivernver kilometerkilorneter 9104910.4910glogio 4 leiaielarelativelelative to the mouth of original range griffith and daley 1984 the the snake river upriverupnver to kilometer 9504950.4950 4 spring pond referred to here as transplant of because its limited langeiangerange and the extent of spring is 15315.315 3 km upriserupriverupnver from briggs springs habitat modification the shoshone sculpin was the nearest spring inhabited by shoshone pi oposedopposedproposed as a thithl eatenedthreatened or endangered species sculpin at the time wallace et al 1984 williams 1980 it is currently a candidate transplant spring is approximately 1000 m2ma threatened or endangered species W E mar- in surface area and enters the snake river at tin USU S fish and wildlife service portland nverriver kilometer 9657965.7965 7 inm gooding county idaho oregon onalonai personalpeispels communication the ameri water flows from the spring head near the base callcailcan fisheries society considers it threatened of a basalt cliff over a 20 m long cascade into a Willwilliamsianislanis et al 1989 and idaho department of pond that is impounded by a set of cuculvertslverts the gamegarnegayne fish and considers the shoshone sculpin streainstreamstrearn drops vertically 2 inm into the snake river a priority species of special concern moseley after passing through the cuculvertslverts the dis- groves and dovesmoves 1992 charge of transplant spring is influenced by a shoshone stulpinssculpinssculpins occur sympatncallysympatricallyafth with fish hatchery water diversion near the spring mottled stulpinssculpsculpinsins coituscottus bairdi in 16 spring head systems in the thousand springs formation boulder and cobble substrate near the cas- wallace etetalal 1984 larger mottled sculpin are cade shift to gravel sand and silt at the tail of known to prey on smaller sculpin bailey 1952 the pool thelethere are dense patches of water hydoskiwydoski and whitney 1979 and are considered speedwell veronica sp and cattail typha sp a potential predator of shoshone sculpin the amphipodsamphipoda a group shown to be heavily con- purpose of this study was to assess the extent to sumed by shoshone sculpin connolly 1983 which shoshone sculpin could be successfully weiewelewere abundant 1000 5000 per inm2ma duangdunngduring the introduced into an environment that seemed study taxa such as dipterdipteransans trichoptertnchopteranstrichopteransans and physically adequate but was already occupied by oligochaetes that also are utilized by shoshone mottled sculpin sculptsculpmsculpin werewere present in densities similar to those
dcputnndepart litt olof01 BiologicbiologicilbiologicalilI suenscanuscunu s idaho statistattstate universityuniunlistyislyimly iouihloIoupowtdloloo iHlo idahoIl lilio11110 83209
190 199311993 SHOSHONE SCULPIN igi19191
TABLITABLE 1 number of stulpinssculpinssculp ins collected peipelper I1imm framefraine net andrindcind relative abundance RA electrofishing samples and percentpelper cent coituscottus greelireeneagreemireenei of total cottus sp at transplant spring idaho 1983 91 only fish 20 imnmm TL weiewelewere included on 15 august 1983 419 C greeneigreened weiewelewere introduced into transplant spring
cottus g1reeneicreenei cottu bairdi number number number number date method collected peipelper framefiame total collected peipelper frame
13 aug 1983 I111 I1 frame nets 0 0 0 30 1 6 20 nov 1983 10 fiameframe nets 4 0 2 15 23 0 6 18 feb 1984 RA I1 5 21 21 api 1984 RA 0 0 15 24 sep 1984 RA 12 27 33 3 octoct19901990 RA 20 100 0 28 sep 199iggi19911 4 frame nets 96 3 29 98 2 0 and RA
in other springs supporting dense sculpin popu- stress prior to release stulpinssculpinsSculpins less than 20 minmm lations other fish species captured in trans- TL which are ageageo 0 fish connolly 1983 were plant spring were mottled sculpin rainbow not included in the analysis because they were trout oncorhynchus mykiss and peapeamouthmouth not monitored in the 1983 and 1984 samples mylocheilusmylocheihis caurinusmaurinuscaurinus shoshone sculpinssculp 419 36 stulpinsins n mean length RESULTSRESUUFS minmm TL range 18 70 mm were dip netted and seinedheined from bickel spring at the hagerman national fish hatchery 25 kinkm downriverdownriver from on 13 august 1983 prior to the stocking of shoshone ins mottled sculpinssculpins all transplant spring on 15 august 1983 and stulpinssculpinssculp stulpins were in frame and distributed stocked at transplant spring within a few hours net samples throughout the spring pond most individuals were small or the sculpin population in the spring was moni intermediate in size 339733 97 mm TL an average tored in august 1983 prior to the introduction of 272.7 mottled stulpinssculpinssculp ins was captured per framefrarnefearne and after the introduction in november 1983 net sample griffith and daley 1984 february april and september 1984 october after the introduction of the shoshone 1990 and september 1991 1983 and 1991 the sculpin 27 individuals were collected in 1983 samples were quantitative estimates using a and 82 in 1984 table 1 mottled stulpinssculpinssculpins were frame at 4 11 random sites net the 075 m present at both the vegetated habitats and the high boxlike PVC frame has ima1 m2ma at im2 openings rocky habitats in 1983 griffith and daley 1984 the and bottom with 3 diameter mesh top mm the abundance of shoshone stulpinssculpinssculpins relative to netting attached to the sides in the 1983 frame the total number of stulpinssculpinssculpins both shoshone and net samples electrofishingelectrofishing a coffelt model BP- mottled was 15 in 1983and1983 and 16 in 1984 five IC unit producing pulsed direct current and ageageo 0 shoshone stulpinssculpinssculpins were found in septem dip nets were used simultaneously to capture ber 1984 griffith and daley 1984 indicating fish within the frame net in the 1991 frame net that some shoshone stulpinssculpinssculpins reproduced suc- samples the electrofisherelectrofisher was not employed cesscessfullyfully on 3 october 1990 20 stulpinssculpinssculpins were within the frame net instead two dip nets were collected all of which were shoshone sculpin used until both netters made three consecutive ranging from 28 to 70 minmm TL passes without capturing a fish on other sam- on 28 september 1991 100 shoshone pling dates and in areas not sampled by the stulpinssculpinssculpins were collected from frame net samples frame net the electrofisherelectrofisberelectrofisher and dip nets were and electrofishingelectrofishing table 1 in four frame sam- used to assess relative abundance of fishes ples there were 53 mature up to 80 mm TQTL stulpinssculpinsSculpins were identified and measured JLTL and 4 ageageo 0 20 mm TQTL shoshone sculpin byviewingby viewing them through a water filled plexiglas averaging 14314.3 11 mean standard devia- measuring board this aquarium like device en- tion individualsmindividualindividualismindividualsm2sm forty three other sho- abled us to discriminate these small morpho- shone sculpin were electrofishedelectrofished along the logically similar fish while minimizing handling perimeter of the pond highest shoshone 192 GREAT BASIN naturalist volume 53
sculpin densities were among veronica where peraperaturestures brown 1989 rough and shoshone one frame net captured 29 fish two mottled stulpinssculpinssculpins both utilize the unique habitat pro- stulpinssculpinssculpins were found in cobbles and boulders vided by springs and both have a limited geo- where the spring cascades into the pond graphic distribution density data from transplant spring suggest discussion that shoshone stulpinssculpsculpinsins may have been able to occupy or utilize habitat with lowerwaterlower water veloci- ties and dense vegetation more effectively than shoshone sculpin has become the predomi- mottled stulpinssculpinssculpins daley et al 1982 observed nant fish in transplant spring in less than an that shoshone stulpinssculpinssculpins rarely occupied areas 8 year period that period represents two or with surface velocities greater than 60 80 cmsamsems three generations based on typical longevity of the highest densities of shoshone sculpin typi- 343 4 years connolly 1983 reproduction was cally occur in aquatic vegetation daley et al successful in 1984 but a substantial increase in in 1982 and this report when shoshone stulpinssculpinssculpins population size was not recognized until 1990 were absent or less abundant in transplant unfortunately we have no data from 1985 to spring mottled stulpinssculpinssculpins utilized vege- 1989 to assess the rate of change net aquatic frame tation and low water velocity areas griffith and sampling was probably more thorough in 1991 daley 1984 apparently however they were than methods used in 1983 which have may displaced from this habitat but not from the underestimated densities although we believe cascade at the pond head by shoshone stulpinssculpinssculp ins the bias was minor mottled stulpinssculpsculpinsins primarily utilize rocky sub- A smallersmaileralleraileralier unnamed sin spring entering the strates and moderate water velocities bailey snake river 010.1oloi km downstream from trans- 1952 hydoskiwydoski and whitney 1979 page and plant spring also was colonized recently by burr 1991 mottled stulpinssculpinssculpins in north carolina shoshone stulpinssculpinssculpins nine fish were captured streams selected habitats with mean focal point there with an electrofisherelectrofisher in september 1991 velocities of 488848 88 cmsemsams and 71 of the stulpinssculpinssculpins when the spring was sampled in 1981 83 only occupied sites with overhead rocky shelters mottled sculpin and rainbow trout were found facey and grossman 1992 it appears that there griffith unpublished data we suspect shoshone and mottled stulpinssculpinssculpins may segregate that shoshone stulpinssculpinssculpins have the may migrated based partially upon water velocity short distance downstream from transplant spring shoshone stulpinssculpinssculpins introduced to transplant acknowledgments spring were able to reproduce compete and survive in the spring environment in the pres- we thank dan daley krishna merkley jim ence of the larger mottled stulpinssculpinssculpins other sym- smothers todd anderson mike black and patric stulpinssculpinssculpins show habitat segregation by rick phillipp for their field assistance gary selecting different substrates water velocities scoppettone paul marsh and an anonymous depths or temperatures in oregon streams the reviewer provided helpful comments on an ear- reticulate sculpin cottuscoituscottus perperplexesperplexusplexus occupied lier version of this manuscript this project was fifflesriffles and pools in the absence of other sculpin supposupportedrtedarted by the idaho field office of the power species finger 1982 in the of the nature conservancy idaho company presence game paiute sculpin coftuscottuscoituscottus beldingibeldingi the larger re- and the idaho department of fish and ticticulateulate sculpin used pools more frequently nongame program matheson and brooks 1983 found that mot- tled sculpin in virginia streams preferred colder literature CITED water than did the potomac sculpin coituscoftuscottus girardiirardi which occupied slowwaterslowsiow water velocity and balballeyBAILEYBAI U-Y J E 1952 life history and ecology of the sculpin gottus silty substrates in california the rough sculpin cottus bairdi pundukituspunctulcitus in southwestern montana copela 243245243 245 coituscottus asperrimus selected deeper water bbownBKOWN L R 1989 temperature preferences and oxygen 20 cm than did the pit sculpin coituscottus piten consumption of three species of sculpin cottus from sis and marbled sculpin coituscoftuscottus klatnathensisklamathensis the pit river drainage california environmental bi- ology of 26 mcicropstnacropsmclmcicrops brown 1991 rough stulpinssculpinssculpins typi- fishes 223236223 236 1991 differences habitat choice and behavior cally fed streams in occupy spring and they are among three species of sculpin cottusCWUS in artificial physiologically limited to a narrow range aftemoftemof tem stream channels copela 1991 810819810 819 199319931 SHOSHONE SCULPIN 193
CONNOLLYCON NOLLY P J 1983 life history of shoshone sculpin MOSFLFYMOSELEY RANDCR annAND C crovesGROVESGROVFS 1992 rare threatened and coituscottus greeneigreenedgreenei in southcentralsouthcentral idaho unpublished endangered plants and animals of idaho and2nd ed re- mastermasterss thesis university of idaho moscow 79 appp port of conservation data center idaho department DALEYDALFY D M J S GRIFFHHGRIFFITH R L WALLACWALLACE F AND P J of fish and game boise 38 appp CONNOLLY 1982 relative abundance and habitat facefagePAGFPAGE L M AND B M BURR 1991 A field guide to preference of the shoshone sculpin cottus greeneigreenedgreenei freshwater fishes north america north of mexico pages 601 610 in proceedings of the annual confer- houghton mifflin co boston 432 appp ence of the western association of fish and wildlife WALLACE R LJL J S GRIFFITH D M DALEYDALFY PPJJ CON agencies NOLLY AND G B BECKHAMBFCKIIAM 1984 distribution of the FACEYFACTY D E AND G D GROSSMAN 1992 the relation- shoshone sculpin cottus greeneigreened cottidae in the of great ship between water velocity energetic costs and mi hagerman valley south central idaho basin 44 324 326 crocrohabitathabitat use in four north american stream fishes naturalist 324326 WILLIAMS D 1980 and wildlife hydrobiologia 239 161 6 J endangered threatened and plants of the status of shoshone sculpin FINGERFINGFR T R 1982 interactive segregation among three review federal register 4560 19853 species of stulpinssculpsculpinssculpmsins cottus copela 1982 680694680 694 WILLIAMS J E ET AL 1989 fishes of north americaamenea GRIFFIIHGRIFFITH J SANDDS AND D M DALFYDALEY 1984 re establishment endangered threatened or of special concern 1989 of shoshone sculpin cottus gree in the hagerman greeneigreenednei in fisheries 14 2202 20 valley idaho report to idaho department of fish and WYDOSKI R SANDRS AND R R wnirnfyWHITNEY 1979 inland fishes of game 12 pp ap washington university of washington press seattle MAIHESONMATHESON BROOKS R E AND G R 1983 habitat seg- and london 220 appp regationre between cottus bairdi and cottus girardi an example of complex inter and intraspecific resource partitioning american midland naturalist 110 165- received 5 february 1992 176 accepted I1 february 1993 great basin naturalist 532 appp 194 198
USE OF BOULDER POCKET HABITAT BY RAINBOW TROUT oncorhynchus MYKISS IN FALL RIVER IDAHO
1 1 2 daniel N streubelstreubel1 andaandjand J S giffithGriffith 12
abstract abundance of rainbow trout oncorhynchus inymykisskiss in relation to characteristics of pockets created by boulders was studied in fall river southeastern idaho to determinedeteimme depth and surface area of pockets most selected by rainbow trout fish weiwelweree counted by snorkeling and pocket physical dimensions were measured an electivity index defined habitat selection in the following terms the most suitable habitat was 070 7 rn maximum depth 050 5 m minimum depth 2 and 3 in surlsurfsuifacesurfaceaceaee aleaarea some study reaches of fall river had more suitable pockets available for trout than were being utilizeduti lied
key uwrcswords rainbow trout oncorhynchus rnymykisskiss habitat use idaho stream rehabilitation
boulders create a major source of trout habi- cial outwash through which it flows basalt and tat in many higher gradient western rivers they ash flow tuff bedrock define the channel form create pools or pockets with increased depth sinuosity is low approaching 101.0loio1 0 and there are and provide surface turbulence that maybemay be the no meander pools overall gradient in the study only cover available to trout water depth and reach is 0640.640 64 boulder cover were important in determining within channel habitat was homogeneous density of trout in a colorado stream stewart and consisted predominantly of run habitat as 1970 boulder placement is a commonly used defined by helm 1985 little woody debris technique in stream rehabilitation rosgen and had been retained in the channel at the 14 16 fittante 1986 and may provide effective dura- rrsecm3secmusecrosec low flows of late summer 1991 the ble trout habitat lere 1982 stream margin hadpulledhad pulled awayfromaway from anyverticalany vertical this study evaluated age 1 and older wild banks formed by high flows leaving no bank rainbow trout oncorhynchus mykiss use of habitat to provide cover for larger trout the boulder pocket habitat in fall river idaho obj- study reach contained paiute sculpin cottus ectivesjectives were to determine the proportion of beldingibeldingzbelbeidingi longnoselongnose dace rhinichthysrhimchthys catalaccatarac trout using boulder pocket habitat and to assess tae and a few utah suckers catostomus the extent to which fish selected pockets of arardensaddensns and mountain whitefish rrosopiumprosopium specific surface area and depth wilmilmiiwilliaThamsommoni in addition to the wild rainbow and occasional cutthroat oncorhynchus clarki METHODS trout in august of 1990 and 1991 snorkel surveys the fall river originates in the southwest were conducted to estimate trout density portion of yellowstone national park it flows throughout the study area these indicated that east into targhee national forest idaho and density of trout larger than ageageo 0 averaged 0350.350 35 then through agricultural lands to join henrys fish100fish 100 m2ma or approximately 136 fishlinfishkmfishkrnfishkinfishken grif- fork of the snake river approximately 10 km fith unpublished data three sites repre- south of ashton in fremont county the study senting a range of boulder pocket densities area at an elevation of about 1740 m extends 7 were selected for the present study sites were km halfofwhich is within the targhee national 160 170 m long and averaged 26 46 m wide A forest and half immediately below the stream boulder was defined as 040.40 4 m diameterdiameteitel situ- channel has been shaped by coarse grained gla ated so that its top was at or above the water
duartdtartdtinitnniitofbioloplilslkikcsntofantof biologibiology I sdncessances idahoidabo state uiiiversityuniveisityuniversity patelloPpocatelloatello idahoidabo 83209 2authorhauthorAutautlioilioi choinwhointowhomto coitcsloiiliieloikspondtnn silotshouldillriitiI IKI1 addressed
194 199319931 TROUT USE OF BOULDER POCKETS 195
TABLF 1 characteristics of boulder pockets used by 83 rainbow trout in three study sections of fall river idaho suminersummer 1991
trout peper r pocket number of number of characteristic occupied pockets trout average range maximum depth m 046 0 0 0460 46 0550 55 2 3 151 5 1 2 0560 56 065ogs0 65 7 16 17 1 5 0660ogg66 0750 75 15 30 20 1 5 0760 76 0850 85 8 19 24 1 5 0860 86 0950ogs95 2 3 151 5 1 2 096ogg0 96 1051 05 2 8 40 1061.06log1 06 1151 15 1 4 40 minimum depth m 026 0 0 0 0260 26 0350 35 1 1 10 0360 36 0450 45 6 12 20 1 4 0460.460 46 0550 55 11 24 22 1 4 0560 56 0650ogs65 10 26 26 1 5 0660.66ogg0 66 0750 75 9 20 22 1 5 surface area m2 046 0 0 046 075 2 2 10 076 125 1 2 20 126 175 3 4 131 3 1 3 176 225 2 3 15 1 2 226 275 3 7 23 2233 276 325 6 13 26 1 4 326 375 5 11 22 151 5 376 450 4 8 20 131 3 451 550 2 5 25 141 4 551 650 1 2 20 651 850 2 4 20 851 1100 1 3 30 liol110111 01 135013 50 1 2 20 135113.5113 51 160016 00 1 4 40 igol160116 01 205020 50 1 3 30 205120 51 260026 00 1 4 40 260126.0126 01 300030 00 1 5 50
surface to create a pocket oflower velocitywatervelocitywater to differentiate fish by size categories under- immediately downstream the low boulder water visibility was approximately 4 m and density site LBD had 38 boulders that fit these water temperature ranged from 14 to 19 C criteria and averaged 050.5 boulders100boulders 100 m2ma of from 1000 to 1500 MDT when observations stream surface the intermediate boulder den- were made sity site JBDIBD had 60 boulders average 101.0iolo after snorkeling we recorded dimensions of boulders100boulders 100 inm2ma2 and the high boulder density all pockets in the section we demarcated the HBD site contained 84 boulders 21212.1 boul lateral margins of a pocket by the abrupt change ders100ders 100 m2ma in water velocity that occurred there initially we during the last two weeks of august 1991 used a velocity meter marsh mcbimey model boulder locations in each site were mapped and 201 on a range of pockets in each site and then trout focal point positions recorded by a snor- completed demarcation by eye water velocity keler moving slowly upstream fish larger than which ranged from 080.8os to 121.2 isecmsec along the about 15 cm were included with most 15 25 cm thalweg outside boulder pockets in all sites was and a few as large as 30 cm no effort was made generally 030.3 050.5 isecmsec within the pockets 196 GREAT BASIN naturalist volume 53
080.8uo D maximum depth 060.6og0 6 el minimum depth k 040.4 s 0 02 020.2 j f 0 Y liilir 0 01oi0.1 0.202 0 3 oubajoaj 0 5 0.606og 0.707 0.808 0.909og 101.0iolo iili111.1 02 0 06 07 08 09
02 1 020.2 I
04040.4
06og060.6
08080.8 depth Wm
fig 1 minimum depth in andcind maximum depth in of boulder pockets used by rainbow trout in falls river idaho electivitiesplectivihes are indicated 05 strong selection 025 but 05 moderate selection 0 025 no selection 050 5 buthut 00250.0250 025 moderate avoidance and 050 5 strong avoidance
to evaluate the selection by trout of the HBD reach and smaller pockets were primarily pocket parameters an electivity index D was found in the LBD and IBD reaches calculated 0 pocket surface area was partially a function D r p of boulder diameter aithwith pocket area 18811.881 X R2 st57.57 r p 2prmpr 455724.5572 boulder diameter rar2 57 N 182 for all sites combined the correlation was where r is the proportion of the resource used higher at lower boulder density sites but at the by rainbow trout and p is the proportion avail- HBD site area of an individual pocket was also able in the environment baltz and moyle affected by the presence of adjacent boulders 1985 following baltz and moyle 1985 we all trout observed in the study sites were in interpreted strong selection to be indicated by boulder pockets eighty three fish were found D 050.5os moderate selection 025 but 05os050.5 no with 001717 and 66 at sites LBD IBD and HBD selection 0 0250.25 moderate avoidance 050.5os respectively the total number ofboulder pock but 0250.25 and strong avoidance 050.5os elec- ets holding trout was 10 17 of pockets pre tivity values were calculated for maximum and sent at IBD and 27 32 ofpockets present at minimum depth and surface area of the boulder HBD A comparison of utilized pocket meas- pockets ureurementsments showed no significant difference be- tween the two sites P 0505.05 and the data were RESULTS pooled for analysis As water depth and surface area of a pocket there was a wide range of maximum and increased the number of fish present generally minimum depths and surface area of boulder increased table 1 no trout used pockets in pockets available on fall river maximum depth which minimum depth was less than 0260.26 m and among the three study sites ranged from 030.3 to maximum depth was less than 0360.36 m llli111.1 m and averaged 070.7 m minimum depth As surface area increased the number offish ranged from 020.2 to 070.7 m averaging 0450.45 m per pocket generally increased to a maximum of pocket surface area ranged from 0250.25 to 28 m2ma 5 table 1 average number ofoffishfish per pocket and averaged 242.4 m2ma the larger and intermedi was 141.4 in pockets with surface areas 2252952252.25 mm2ma ate sized pockets were primarily found in the 222.2 in surface areas of 2262.26 4504.50 m2ma 222.2 in 199311993 TROUT USE OF BOULDER POCKETS 19719
080.8 T
1 060.6og 444 4 H 444 4 444 4 040.4 0 020.2 u 0 1 0 n CU E 1ini n c LD QIQI i n 1 i r CO eslekl LO r tonhont- cooo L I inln cl inln ln ninln 0 LLL r 0 C cslasl 0cd i 02020.2 J 04040.4
06og060.6
08080.8 surface area mam2
fig 2 surfaceSurfcleecice area mam2 of boulder pockets used by lambowrainbow trout in falls river idaho electivities are indicated 05050.50 5 strong selection 0250 25 but 050 5 moderate selection 00002525 no selection 050 5 but 00250 025 moderate avoidance and 050 5 strong avoidance
surface areas between 4514.51 and 8508.50 mm2ma and 353.5 bow trout in the fall river baltz and moyle in surface areas ss85858.5 m2ma 1985 evaluated rainbow trout habitat in a the electivity index demonstrated that trout tributary of the sacramento river california were selective in the micromicrohabitathabitat they occu- and found strong selection for depths greater pied ElectiAelectivityty values for maximum depth indi than 060.6og m similar to the threshold value for our catedbated moderate selection at depths equal to or study the habitat suitability index HSI for greater than 07m0 7 m and strong selection at depths rainbowtroutrainbow trout raleigh et al 1984 indicates that greater than 090.9og m fig 1 minimum pocket depths greater than 0460.46 m have a suitability depth was not a sensitive index of trout density index value of 1 the highest value possible not minimum depths of 0.606og m and at 06 deeper there until fall river pocket depths of 07070.7 m were was moderate selection and over ot07 070.7 m strong reached was there moderate to strong selection selection 1 pockets with surface fig areas and trout moderately avoided pockets at depths equal to or exceeding 3 mm2ma were moderately or of 050.5os m thus the HSI did not accurately pre- strongly selected 2 fig dict depth selection on fall river minimum habitat for which rainbow showed trout a pocket depth appearedappeared to be a less useful indi- or moderate selection viewed strong was by cator of habitat selection for rainbow the fall river us as most suitable habitat for the study sites trout fifty of the 178 pockets in the three sites fell pocket surface area was also a factor affect- within those limits thirteen optimal pockets ing trout density only four fish were found were located within the IBD reach and none in pockets 15151.5 mm2ma and those 3 m2ma were se- within the LBD reach thirty seven were lo- lected 1969 found cated within the HBD reach and more fish lewis that surface area and depth along with volume velocity were found in that reach A total of 23 of the 50 current and cover accounted for 70 77 the optimal pockets were not occupied by trout of variation in numbers of trout in pools of little prickly creek montana discussion if surface area requirements reflect the size of territories defended by individual trout maximum water depth in boulder pockets in optimal habitat agonistic behavior by individ- strongly influenced selection of habitat by rain ual trout might serve to establish maximum 198 GREAT BASIN naturalist volume 53
density alienallenailen 1969 and grant and kramer literature CITED 1990 reviewed the literature for fluvial sal monidsrnonidsmonads though data for rainbow trout were ALLENALLFN K R 1969 limited strong similarities were found among limitations on production in salmonid populations streamsstr earnsearms pages 3 18 the salmonid in sti in T G northcoteNortheote seven species they reviewed for ed symposium on salmon and trout in streams insti- fish 15 20 cm long average territory size in tute of fisheries university of british columbia van- pools was approximately 1 5 m2ma in fall river couver canada the estimated area occupied by individual trout ballBALI D mandpbmoylrM AND R B MOYLE 9851985 microhabitatmicrobabitatmieroMicrohabitat use by an assemblage of california stream fishes developing based on our observations offishfishhishbish of abundance per criteria for instream flow determinations transactions 05 go ma pocket ranged from 050.5 to 606.0 mm2 and averaged of the american fisheries society 114 695 704 252.5 mm2ma however two thirds of the fish were GRANI J WWAA AND D L kramenKRAMERKRAMFR 1990 territory size as inhabiting areas 25252.5 mm2ma suggesting that a predictor of the upper limit to population density of smaller territories might be required boulder juvenile salmonsalmonidsids in streams canadian journal of in fisheries and aquatic sciences 47 1724 1737 pockets than from in the pools which the data of HELMHFLM W T rdED 1985 glossary of stream habitat terms grant and kramer 1990 were generated special report western division american fisheriesFishenes lack of summer holding habitat in the LBD society 34 appp reach appeared to limit trout abundance as the KFNNFUYKENNEDY G J A AND C D stranceSIRANGFSTRANGE 1982 the distri- bution of salmonidsids reach contained no and salmon in upland streams in relation to quality pockets no trout depth and gradient journal of fish biology 20 579 were present summer holding habitat did not 591 appear to limit trout numbers in the HBD and lerfLERE M E 1982 the long term effectiveness of three illiIBD1131 reaches because there were 23 pockets types of stream improvement structures installed in montana streams s with optimal dimensions that were not utilized unpublished mastermasters thesis mon- tana state university boemanbozeman 99 appp trout density in these reaches might have been lewis S L 1969 physical factorss influencing fish popula- depressed by low recruitment or factors such as tion in pools of a trout stream transactions of the winter mortality and food availability american fisheries society 98 14 19 RAL R although trout distribution is closely tied to ralrunrabrunrunruheuh F T HICKMANKMAN R C SOLOMON AND PE C neisonNELSONNFLSON 1984 habitat suitability information rain- physical habitat in fall it is clear that river ubow trout USU S fish and wildlife service publication simply adding boulders to rivers will not auto- FWSOBS 8210608210 60 64 appp matimaticallycally increase trout populations pockets ROSGENROSGFN D AND B L fittanteFITTAN Tr 1986 fish habitat struc- created by boulders must meet depth and sur- tures a selection guide using stream classification pages 163 179 G miller A arway and R face area before fish will inhabit in J J F requirements carline eds fifth trout stream habitat improvement them as shown on fall river other studies have workshop lock haven pennsylvania found that water depth alone is not the major stfvvastavvaSIFWARIin PAP A 19197070 physical factors influencing trout den- limiting factor for trout populations kennedy sity in a smallsmail stream unpublished mastermasters s thesis colorado state 78 and strange 1982 water velocity and available university fort corins appp wl C KKIIAMI1 1aaa m G M 1967 physical microhabitathabitat of trout un- also micro cover influence trout density lewis 1969 published mastermasterss thesis colorado state university these environmental requirements as well as fort coffins 42 appp other limiting factors must be understood be fore boulders are effectively used for habitat received 10 june 1992 improvement accepted I1 february 1993 great basin naturalist 532 appp 199 202
HUMORAL creatinine IN WALLEYE stizostedion VITREUM
1 harry L holloway jr and craig A shoemaker
ABSIBAC published fish blood parameterspaiapalameters are limited to commercially cultured species ege g rainbow trout on- corhynchus mykiss and channel catfish ictalurus punctatuspunct atus however as walleye stizostedion vitreum and other fish increase in value to the angler hatchery and fish managers will require data on these species blood sera were collected from live walleye in the field creatininecreatimne values were determined colorimetrically and health range values were established from these data creatinineCreatimne levels ofwalleye serum 0060 06 0720 72 madlmgdl were higher in three species but lower than recognized in a recent catfish study creatinine levels may be important in predicting diseases in which the kidney is adversely affected
key words creatininecredcreatimne walleye stizostedion vitreum health range shrumserum colorinwriccolonmetriccolon metric determination
the need for a rapid means of assessing the mcgilvery and goldstein 1979 phospho- condition offishof fishhishbish exposed to environmental deg creatine acts in muscles of vertebrates as a res- radation and disease has long been recognized ervoir of chemical energy for the phosphoryla- logically blood is a medium with which to do tion ofadpof ADP to reconstitute ATP during muscle this since no other single body tissue is more contraction since creatinine excretion depends reflective of total body metabolism since our upon phosphocreatine content creatinine can interests are in disorders of skeletal muscle we be used to assess muscle mass creatinine is elected to establish the range of serum creat- constant with about 0020.02 gmkg of body weight inine levels a nonprotein constituent in fish excreted per day thus when muscle degener- including feral walleye creatinine an end ates from paralysis or muscular dystrophy the product ofcreatineof creatine metabolism is a normal and creatinine content of urine falls mcgilvery and alkaline component of urine and blood warner goldstein 1979 because there is less muscle and williams 1977 recognized the need for an mass and phosphocreatine to spontaneously cy- indicator in assessing the health of channel cat- cle to creatinine oser 1965 reported that fish and utilized blood serum components iiee creatinine in mammals is more readily excreted metabolic products electrolytes enzymes than uric acid or urea and that even a slight field et al 1943 phillips 1958 barnhart increase in blood creatinine is evidence of im 1969 and bentinck smith et al 1987 deter- paired kidney function in contrast with ammo- mined ranges of variation in blood serum con- nia which is excreted by the gills creatinine is stituents in several fish species under different excreted by the kidney in fish smith 1929 protocols ie collecting holding rearing etc shell 1961 interpreted periods of increased according to wamermamerwarner and williams 1977 natu- creatinine concentration as indicating kidney ral variation must be established before using impairment in smallmouth bass micropterus these constituents to assess fish health hoffman dodolomiedolomitedolomieudololomieiomiemieu however he concluded that cause and effect evidence linking the two 1963 and wedemeyer and chatterton 19719711 phenomena is available defined natural variation as being two stand- not for fish as it is for mammals ard deviations of the mean value normally dis- tritributedbuted data for healthy fish populations in MATERIALS AND METHODS mammals phosphocreatine spontaneously cy- cles at a slow rate to form creatinine which is mature walleye 5 arsyrs ofage were collected excreted in urine the cycling rate depends by trap nets isls181.8 X 242.4 in and experimental gill upon the total phosphocreatine content ie nets isls181.8 x 75 in 19 25 38 51 and 64 mm muscle mass at a given temperature and ph bar measure meshes in april and may 1991
biology department university ofot north dakota box 8238 umveisityuiiiversity stationstadonstanonstatton crandgrand forks northnoi th laotadakota 5820258902 USA
199 200 GREAT BASIN naturalist volume 53
fabifahlTAHI 1 1 seiselserumnm creatinine values foifolfor selected freshwater fish species values converted from mg and mg100 ml to madlmgdl authors health range mean SD range species tat22 SD of mean madlrngdlmgdl madlmgdl field etetalal 1943 caiencaipncarp n 5 19 0056 00420 042 00870 087 brook trout n 5 19 00720 072 00510 051 00850 085oss lakes nealnear madison wisconsin phillipsplnllips19581958 brook trout n 0041 biowntioiitnbrown trouttront n 0049 lakes near cortland new york barnhartbamhait19691969 rainbow troutti out n 16 0105 00530 053 0200 20 parvin lake bellbeilbellvueBelivue Coloiacoloradodo warner and williams 1977 channel cattishcatfish n 107 000ooo0 00 0370 37 01780 its178 096 ponds at gallatin tennessee hille1982hille 1982 rairalrainbowabownbow troutti out n 0020 02 021709170 217917 review papelpaper including fish data collected in USA Coloiacoloradodo and italy bentinck smith et al 1987 channel catcatfishfisli n 376 05430 543 0399 020 2 2509502 50 rearingbearingrealbealbeai ing ponds stonevilleStoneville mississippi hollowayandIIHolloollowaywayandand shoemaker unpublished walleye lake oahebaheoalie n 15 000ooo0 00 0570 57 027609760 276976 0145 0060 06 0480 48 lake sakakawea it 15 0060 06 0640 64 03480 348 0145 ols0180 18 0720 72 merritt reservoir n 15 ooiool0010 01 0440 44 02240 224 0106 0060 06 0420 42 combined 000ooo0 00 0560 56 02830 283 0140141 0060 06 0720 72
ffromrom lake oahebahe south dakota lake sakakdakak RESULTS AND discussion abeaawea north dakota and merritt reservoir ne- braska table 1 heparinized lomi10 ml syringes creatinine values from each lake were ana- and 21 gauge needles were used to collect blood lyzed for normality using the kolmogorov smiramir from live adult walleye by cardiac puncture nov test sokal and rohlf 1981 data from each samples were coagulated clot milked and cen- lake and pooled data were nonnonnallynormallynallynaily distributed trifugedtrifuged within 24 hrs of collection sera were since all D values were not significant p collected in microcentrifugemicrocentrifuge tubes frozen on oi0101.01 means were compared using williams s dry ice for transport to the laboratory and 1974 modification ofoftukeystukey s honestly signifi- placed in a 80csoc freezer until analyzed cant difference test and no significant differ- sigma diagnostics 1989 protocol for the ences occurred among values for each reservoir colorimetric determination ofofcreatininecreatinine was util- p oi0101.01 ized A bausch and lomb spectronicSpectronic 20 was used creatinine levels of walleye blood serum to make detendeterminationsaeten ninations at 500 nm creatinine were within the range of values observed for concentration in madlmgdl was calculated using the other fish eg salvelinus fontinalis brook manufacturers formula and dilutions of creat trout salmo trutta brown trout oncorhyn- inineanine standard 15 mdi were utilized as controls chus mykiss rainbow trout cyprinus carpio to ensure that procedures were reliable carp and ictalurus punctatuspunctatus channel catfish 199311993 creatinine IN WALLEYE 201
table 1 mean creatinine concentration in wall more recent catfish study we assume these eye closely approximated the value for channel differences to be due to development of more catfish wamerwarner and williams 1977 table 1 sensitive procedures and higher chemical speci- besides specificity and ecology of the target ficity population the method of capture age offishof fishhish method of drawing blood and diet are all vari- acknowledgments ables that should be considered in accepting study specimens and evaluating results how- we acknowledge HLH principal investiga- ever bentinck smith etetalal 1987 suggested the tor use of federal aid sport fish restoration variation in serum constituents in channel cat- funds from nebraska game and parks commis- fish was due principally to variability among sion project F 77 R north dakota game and individual fish they contend that individual fish department project F 2 R and south studies generate results characteristic of the dakota game fish and parks department species being analyzed and conditions to which project F 14 R financial support was pro- the fish had been exposed the 300 difference vided to CASGAS by the academic program and between the two channel catfish studies table student awards committee biology depart- 1 is too great to be explained by improved ment university of north dakota we wish to methodology and instrumentation in such a thank christopher ottinger graduate student in short time span thus differences must be at- biology university of north dakota for his tritributed to variation in channel catfish or incon- technical assistance in the field sistencies in procedures walleye values may be useful as reference points in studies using simi- literature CITED lar fish collecting blood sampling and analysis methods in assessing the health of this species BARNHAIHBARNHART R A 1969 effects of certain variables on he- circulating levels of creatinine are used pri- matmatologicalological characteristics of rainbow trout transac- of marily as an index of renal function in man tions the american fisheries society 98 411 418 BFNIINCKBENTINCK SMITIIJSMIIII J M H bflfaubeleaupwateiistiiatpwairkslkal C S hepler 1977 high serum creatinine concen TUCKER F sillesSTILES P R bowserBOWSFK AND L A BROWNBKOWN trattrationsorationsions are encountered in cases of human re- 1987 biochemical reference ranges for commercially nal inflammation and obstruction in some reared channel catfish progressive fishpish culturist 49 108 114 instances reflecting the degree of impairment FIFLDFIELD J BCB C A elvriljfmelvriijem AND C JUDAY 1943 A study smith 1929 showed creatinine to be excreted of the blood constituents of carp and trout journal of in the urine of fish thus creatinine concenttaconcentraconcentra biological chemistry 148 261 269 tionseions may be elevated in fish with diseases af- HELPER 0 E 1977 pages 285 288 in manual of clinicalofchnical laboratory methods ath4th ed illi- fecting the bacterial thomas springfield kidney eg kidney nois disease there was no apparent pathological HILLEHILLF S 1982 A literature review of the blood chemistry evidence that the kidneys were diseased in fish of rainbow trout salimsaline gairdnetgairdnergairdnengairdneTnen rich journal of fish examined biology 20 535569535 569 HOFFMAN R G 1963 statistics in the practice ofineof medicineofinedicinedicine bentinck smith et al 1987 stated that the journal of american medical association 185 864 analytical procedure utilized must be appropri- 873 ate for each constituent being measured and McGlmcgilverymcgi LVERY R WANDGW AND G coldsteinGOLDSIFINGOLDSTEIN 1979 biochemis- that measurements must fall within the of try a functional approach W B saunders philadel- range phia pennsylvania 862 appp analytical linearity when absorbanceabsorb ance of diluted oserOSFR B L 1965 hawk s physiological chemistry 14th14tb ed standards 0 10 madlmgdl was graphed creatinine mcgraw hill book company new york 1472 appp concentration on x axis and absorbanceabsorbance on the PIIILLUsPIIILLIPS A M JRjik 1958 the organic composition otof brook and brown blood axis a straight line trout progressive fish culturculter yaxisy resulted the reference ist20ist 20 114 116 values presented for creatinine in walleye blood SHELLSIIFLL E W 1961 chemical composition ofbloodoxbloodof blood ofsmallofsmall- serum meet these requirements and are ofvalue mouth bass USU S fish and wildlife service research in establishing a range of health we consider report 57 USU S government printing office washing- ton DC 36 appp values of 0.06 1 creatinine 006 0720.72 madlmgdl table SIGMAS MA DIACNOSIIdiagnostics s 1989 creatinine procedure no 555 representative of feral walleye populations sam- st louis missouri 7 appp pled under field conditions and exhibiting no SMIIH H W 1929 the excretion of ammonia and urea by apparent disease symptoms or gross pathology the gills of fish journal of biological chemistry 81 727 742 levels for the reported walleye are much higher SOKAL R RANDR AND F J ROIII r 1981 biometry W II1114 free- 2 5 times than for other species except the man and company new york 859 appp 202 GREAT BASIN naturalist volume 53
WARNERWARN 1 M C AND R W WILLIAMS 1977 comparison WILLIAMS J D 1974 A simplified regression formulation between serum values otof pond and intensive raceway oftukeysoitukeyturkeys s test journal of experimental education 42 cultured channel catfish ictalunisictalurus punctatuspunctatus rafi 80 82 aesquenesque journal of fish biology 11 385 395 wlwedemeyejidrmryfk G AND K chkitertoncllalirklon 1971 some blood received 26 march 1992 chemistry values for juvenile coho salmon oncorhynOncorhyn accepted 10 february 1993 chus kisutchsutoh journal of the fisheries research board of canadaofcanada 28 606 608 great basin naturalist 532 appp 203 206
INFLUENCE OF PREY MOVEMENT ON THE AIM OF PREDATORY STRIKES OF THE WESTERN rattlesnake CROTALUS VIRIDIS
121 2 1 3 1 4 douglas F schmidtschmidthschmidt1 wilhamwilliam K hayes13 and floyd E hayes14
ABSIRACT the purpose of this study was to determinedeteimmedeteimme whether the western rattlesnake crotalus virvinbindivindiviridisidisdi aimsarmsalms its predatory strike at the headthoraxhead thorax region of mice primarily on the basis of cues related to direction of prey movement we hypothesized that when rattlesnakes strike at anesthetized backward moving mice most strikes would be aimed at the forward most moving region iei e the posterior abdomentailabdomentail region of prey however most micemlee were struck in the anterior headthoraxhead thorax region implying that the visual infrared image of mice rather than directional movement guides the aim of predatory strikes
key words reptiliaheptReptiha serpentes rattlesnake crotalus virvindisidis predator prey feeding behavior strike aim
rattlesnakes generally strike rodent prey in ment kardong 1986a kardong and mackessy the headthoraxhead thorax region minton 1969 kardong 1991 based on photographic studies dieffen- 1986a hayes 1986 1991 1992a kardong and bach and emslie 1971 suggested that move- mackessy 1991 which presumably reduces the ment of the mouse s head attracts the strike of risk of missing forward moving prey hayes and E climacophoraclimacophora to the anterior region hayes galusha 1984 provides the quickest means of 1986 using featureless cylindrical models of prey death kardong 1986a hayes 1992a and mice concluded that direction of movement minimizes the threat of retaliatory injury in- alone can be utilized by rattlesnakes to aim the flicted by the rodent kardong 1986a hayes strike at the forward most moving anterior re- 1992b other ophidiaophidiansns that reportedly strike gion release of the predatory strike itself ap- the headthoraxhead thorax region of mice include the rat pears dependent upon prey movement de- snakes elaphe climacophoraclinwcophoraclimacophoraphorn diefenbach and tected by visual andor infrared cues chiszar et emslie 1971 andandeE quadrimrgataquadrivirgata mori 1991 al 1983 hayes and duvall 1991 since motion- and the cottonmouth agkistrodon piscipiscivoruspiscivorousvorus lessess anesthetized and dead mice with all other kardong 1982 the rough green snake stimuli present often are not struck klauber opheodrys aesaestivusaestwusaestfivustivuswusmus also frequently strikes the 1956 dullemeijerDullemeijer 1961 cock buning et al anterior region of insects goldsmith 1986 19811981chiszaretall992chiszar et al 1992 however when feeding on lizards E quadrivir thus considering the significant contribu- gata and the puerto rican racer Alsophis por tion of prey movement to the predatory strike toricensis most often strike at the midbody we designed a test of our hypothesis that the mori 1991 rodriguez robles 1992 western rattlesnake crotalusGrocrotalusfalusfains virviridisidis aims its strike primarily on the basis of to aim the strike at the headthoraxhead thorax region orofr cues related to direction of a mouse a rattlesnake must be able to difdifferendifferedferen prey movement tiatediate the anterior and posterior ends ie longi- tudinal polarpolarityityl ofpotential prey discrimina- METHODSMFTHODS tion of these regions may be accomplished by several means including the visual andor infra- subjects were 14 adult northern pacific red image of the rodent ie gross morphologi- rattlesnakes crotalus viridis oreganusoreganus from cal features detected by the eyes and facial pits southeastern washington and two great basin respectively and the direction of prey move rattlesnakes C v lutosuslutolukosussus from southeastern
department of biological sciences walla walla college college place washington 99324 presentpresent address baptist medical center sleepsteep disorder center 9601163096011 630 exit 7 little rock arkansas 72205 31 31resentpresentresent addressaddi ess biology departmentdepal tinenttenent southern college collegedaleCollegedale tennessee 37315 antholauthorauthol to ahornwhomwhorn con espondenceesponderespondencecorrespondencencenee should be addressed 4 present address department of natural sciences loinalorna linda university loinalorna linda california 9235092330
203 204 GREAT BASIN naturalist volume 53
taniTABLETABI I1 1 sites on anesthetized mice mus musculusmuseums either anterior or posterior half where the fangs where strucistruelstinckshinck by rattlesnakes crotalus vindisvinchs and latencylat encyney of snakesofsnakes to strikesti ike sec mice were drawn by monofilainentinonofilament penetrated two mice struck exactly in the mid- stringacrossstringstung across the arena floor in eithereithelelthel a fulwardforward or backward dle were discarded for the present purpose directiondn actionection videotapes were made of most trials to confirm the of observations 1 accuracy our and to measure number ofsitesof sitessltes struck latency to strike directiondnectionofof the time sec until a strike was initiated how- movement anterior 1ostcriorPospostenortenor X SD ever the sample of videotaped trials was re- duced due to a malfunction of the video system forwardforwald 9 5 560 522 in agreement with kardong 1982 1986a vis-as4s baekBackbackwardwaidwald 10 5 470 499 ual observations were sufficiently reliable for
axa 1 1 judging the site struck by ax2X 00 1 00 the snakes amann muriiimannmuiii wlntmyuwhit y U 20 11 50 statistical tests with alpha set at 05os05.05 fol- lowed siegel 1956 oregon all were well habituated to captivity RESULTS 1 3 years and maintained on a 1212 LD cycle at 25 35c prior to the study the snakes con in table 1 the ofstrikes fumedsumed live adult laboratory mice mus muscu- data present number delivered to the anterior and halves of lus on an irregular 1 2month basis posterior the anesthetized mice no difference existed our experimental design was to present between forward moving and backward movamov rattlesnakes with anesthetized mice moving ing conditions table 1 mice were most often either forward or backward the forwardhorward mov- struck in the anterior headthoraxhead thorax region re ing mice served as controls since we expected gardgardlessgarblessless of the direction of movement when most strikes be aimed to at the anterior data were pooled to include both forward and headthoraxhead thorax region of the body kardong backward moving presentations the proportion 1986a the backward hyby for moving mice we of mice bitten in the headthoraxhead thorax region was sized that strikes pothepothesized most would be aimed at significant 66 of all observations one tailed the forward most moving region the poste ie chi square 2792.79 df 1 p 05os05.05 from the riorbior abdomentailabdomentail region of the prey videotaped trials there also was no difference in for each trial a snake was transferred to a 91 latency to strike forward n 6 or backward x 61 x 46 cm L X W x H wooden box adult n 7 moving mice table 1 thus the rattle- laboratory mice were anesthetized with 010.1oi mg snakes treated forward and backward moving vetalargVetalarg mouse and a monofilament fishing mice in a similar manner line was tied around the center of the body to field by field videotape analyses indicated manipulate direction of mouse movement dur- the total duration of successful strikes ranged ing presentation to the snake each snake was from 02670.267 to 05670.567 sec x 04 seesec latency to randomly assigned several forward or back contact the mouse was 00670.067 03330.333 sec contact ward moving presentations with a minimum of with the mouse lasted 00670.067 01670.167 secsee and re- one day between successive trials we assumed coil required 00670.067 01670.167 secsee cf van riper all strikes from a given snake were independent 1955 kardong 1986a rowe and owings 1990 no individual accounted for more than 20 of hayes 1991 1992a there were no obvious the data cf kardong 1986a snakes were not differences in the form or success of strikes at allowed to consume struck mice until after com- forward and backward moving mice pletion of the study mice were dragged for several minutes in broad sweeps at a 90 angle discussion to the snake minton 1969 to within 15 cm of the snakesnakess snout velocity of the moving mouse although rattlesnakes apparently can utilize determined from slow motion videotape re- directional movement to aim the strike at the view was approximately 50 150 cmseccusecemsee which headthoraxhead thorax region of mouse models hayes was comparable to sprint speeds of wild mice 1986 our results table 1 indicate that when peromyscus numiculatusnuiniculatus measured by similar striking at live mice the aim is guided primarily means for each presentation in which the by other cues since the snakes mostly struck at mouse was struck a given mouse was used until the anterior headthoraxhead thorax region of backward struck once we recorded the site on the mouse moving mice it seems likely that rattlesnakes 199319931 rattlesnake PREDATORY STRIKE BEHAVIOR 205
discriminate the anterior and posterior ends of literature CITED prey ie longitudinal polarity by visual andor infrared cues rather than by directional move- CIIISARCIIISZAR D R K K lerLEE C W radoRADCLIFFERADC nrrrnarr AND II11 M ment this conclusion is in disagreement with SMIIIISMITIL 1992 searching behaviors by rattlesnakes fol- our original working hypothesis lowing predatorypredatoiystnkesstrikes pages 369 382 in J A camp- bell and E D brodie jr eds biology ofot the pitviperspitvipers although the relative contribution of visual selva tyler texas and infrared cues remains unknown the CIIISARCIIISZAR D C wradciirfrRADCLIFFEW K M SCUDDHSCUDDEIJ AND D headthorax oriented strikes observed in a DUVALL 1983 strike induced chemosensorychemosensory search- congenitally blind rattlesnake by kardong and ing by rattlesnakes the role of envenomation related chemical cues in the strike pages 1991 that polarity the in post environment mackessy suggest of 125 139139mdin D muller schwarze and R M silversteinsliverSilverstem infrared image alone can be deciphered how- eds chemical signals III111 plenum press new yolkyork ever cock buning et al 1981 observed that COCK BUNINGBUNINC T DFDE R C GORIS AND S TFRASIHMA blindfolded agkistrodon blom occasionally 1981 the role of thermosensitivitythermo sensitivity in the feeding be- blomhoffiblomhoffthofft havior of the strike the infrared be- pit viper agkistrodon bloinhofftblomhofftblomhofft breutbredibredlbrevi warm spot an cue left caiduscaudus japanese journal of herpetology 9 7 27 hind when a mouse moves just before the strike dieffenbac n C OANDOANDSS G emslieEMSLIF 1971 cuesinflucues influ- and suggested that visual cues may be relied on encing the direction of prey ingestion of the japanese for aiming the strike considering the neural snake elaphe climacophorachmacophoraclimacophora colubridae serpentes Herpetologic a 27 461 466 visual integration of the and infrared systems of dullemfijfrdullemfijeh P 1961 some remarks on the feeding be- rattlesnakes hartline 1984 it is likely that both havior of rattlesnakes kon ned akad wetenschWetensch systems when functional contribute to coordi- proc ser C 64 383396383 396 nation of the strike aim GOLDSMITHGOLDS ml illlii S K 1986 feeding behavior of an arboreal insectivorous snake opheodnjsopheodrys aestivusaestivus colubri- from the slow motion videotape analyses it dae southwestern naturalist 31 246 249 appeared that most strikes showed no deviation HARTLINE P H 1984 infrared and visual senses in snake from their original course cf kardong 1986b optic tectum pages 405 420 in L bohsbobs R D keynes thus we believe strikes delivered to anterior and S H P maddrell eds comparative physiology of sensory systems cambridge university press cam- regions of moving prey were aimed on the basis bridge united kingdom of information received prior to striking ac- HAYESHAYFS W K 1986 factors influencing the release and aim cordingly we concur with kardong 1986a and of predatory strikes in the rattlesnake crotaGrotacrotaluslits vindis kardong and mackessy 1991 that rattlesnakes oreganusoreganus unpublished mastermasterss thesis walla walla college college place washington v 36 appp rely on visual andor infrared cues to distinguish 1991 ontogeny of striking prey handling and en- the most vulnerable headthoraxhead thorax region of venomationveno mation behavior of prairie rattlesnakes crotalus mice prior to the attack and aim their strike v vindis toxicon 29 867 875 accordingly 1992a factors influencing venom expenditure by prairie rattlesnakes crotalus v uitditvirvindisidis feeding on wild although rattlesnakes typically aim their mice toxicon 30 449 460 strike at the anterior region of mice the reaction 1992b striking prey handling and venom expen- of the prey may limit their success when strik- diture by prairie rattlesnakes feeding on birds and journal of 26 496 ing at unrestrained lab mice kardong 1986a mice herpetology 499 HAYESHAYFS WWKANDDK annAND D DUVALL 1991 A field study of prairie and at items manipulated by monofilament line rattlesnake predatory strikes Herpetologic a 47 78 81 anesthetized lab mice in this study and models HAYESHAYFS WKANDJW K annAND J G GALUSIIA 1984 effects of rattle- of mice in hayes 1986 snakes most frequently snake crotalus andismndisviridis oreganusoreganus envenomation upon mus bite the head thorax mobility of male wild and laboratory mice mus- headthorax region however hayes culus bulletin ofthe maryland herpetological society 1992a found that strikes by prairie rattlesnakes 201359013520 135 144 C v virviridisidis were distributed randomly along KARDONCKARDONG K V 1982 comparative study of changes in prey the axis ofwild mice peromyscus maniculatus capture behavior in the cottonmouth agkistrodon pis civorus and egyptian cobiacobra naja haje copeiacopela 1982 the discrepancy is best explained by relative 337 343 differences in mobility lab mice are much 1986a the strike behaviouibehaviour of the rattlesnake slower hayes and galusha 1984 and undoubt- crotalus vindis oreganusoreganus journal of comparativeofcomparative psy- edly less able to evade strikes than wild mice chology 100 304 314 1986b slow the predatory strike of the rattlesnake motion videotape analyses indicate that when things go amiss copela 1986 816 820 the latter can often leap clear of the strike after KARDONG K V AND S P MACKESSYmaoMAC kessyKFSSY 1991 the strike it is launched by a snake hayes 1991 1992a behavior of a congeniallycongenitally blind rattlesnake journal of thus a distinction can be made between what herpetology 25 208 211 KLAUBEKLAUBFRii L M 1956 rattlesnakes their habits life histo- the snake aims at and where the fangs success- ries and influence on mankind university of californiaCah fornia fully contact prey press berkeley 2 volumes 206 creviGREAT BASIN naturalist volume 53 mingonMINIONMINTON S A 1969 the feedingstrikefeeding strike of thetimberthe timber rattle- ground squirrels and rattlesnakes ethology 86 237 snake journal of herpetology 3 121 124 249 moniMOKI A 1991 effects of pleyprey size and type on prey han slegelSIEGELSIFGFL S 1956 nonparametricNonnonparparametricametnc statistics for the behavioral ifingdlingefinglhing behavior in elaphe quadnvirgataquadrivirgata journal of her- sciences mcgraw hill new york petpetologyology 25 160 166 VAN riperRIPFR W 1955 how a rattlesnake strikes natural RODRKUI roi3iesroiilrobil rs J A 1992 notes on the feeding be- history 64 308308311311 havior of the puerto rican racer plophisalophis portoricensis serpentes colubridae journal of herpetology 26 received 3 july 1991 loo1001 0 102 accepted 10 february 1993 ROWED M P aniAND D II11 OWINGS 1990 probing assess- ment and management during interactions between information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged unpublished manuscripts pertaining to the biolog- to designate properly prepare label and deposit ical natural history of western north america pref- high quality voucher specimens and cultures docu- erence will be given to conciseconeisenelse manuscripts ofup to menting their research in an established perma- 1212000000ooo words nent collection and to cite the repository in publi- SUBMIT manuscripts to james R barnes editor cation great basin naturalist 290 MLBM brigham references IN THE teffteaftefaTEXT are cited by author and young university provo utah 84602 A cover date eg martin 1989 or martin 1989 multiple letter accompanying the manuscript must include citations should be separated by commas and listed phone numbers of the author submitting the in chronological order use et al after name of manuscript it must also provide information de- first author for citations having more than two au- scribing the extent to which data text or illustra- thors tions have been used in other papers or books that acknowledgments under a centered main head- are published in press submittedsubmitsubmittedortedoror soon to be ing include special publication numbers when ap- submitted elsewhere authors should adhere to propriateprop riate the following guidelines manuscripts not so pre- literature CITED also under a centered main pared may be returned for revision heading lists references alphabetically in the fol- manuscript preparation consult vol 51 no 2 lowing10wing formats of this journal for specific instructioninstructionss on style and mack G D and L D flake 1980 habitat rela- format these instructions guidelines FOR manu- tiontionshipsships of waterfowl broods on south da- scripts UBMITTEDSUBMITTEDS TO THE GREAT BASIN NATURnaturalistilistvlistALIST kota stock ponds journal of wildlife man- supply greater detail than those presented here agement 4444695695 700 in condensed form the guidelines are printed at sousa W P 1985 disturbance and patch dynambynam the back oftheofodthethe issue additional copies are available lesies on rocky intertidal shores pages 101 124 from the editor upon request also check the pickett S great in S T A and P white eds the most recent issue oftheodtheof the basin naturalist ecology of natural disturbance and patch dy- for changes and refer totheto the CBE style manual namicsnamiasnainicscs academic press new york ath5th edition council of biology editors one coulson R N and J A 1984 illinois 200 111 witter forest center suite illiliiiilillii east wacker entomology ecology and management john chicago IL 60601429860601 4298 USA PHONE drive wiley and sons inc new york 669 appp 312 6160800616 0800 FAX 312 6160226616 0226 24 1 TABLES are double spaced on separate sheets and TAPETYPE AND DOUBLE SPACE all materials including designed to fit the width of either a single column literature cited table headings and figure legends or a page use lowercase letters to indicate foot- avoid hyphenated words at right hand righthand margins notes underline words to be printed in italics use stan- photocopies OF FIGURES initially dard bond 22 x 28 cm leaving cm ma on are submitted cineineln 25 marginsrginsagins with the manuscript all sides manuscript editors may suggest changes lettering on figures should be large enough to SUBMIT 3 COPIES ofthe manuscript and the originoriginal withstand reduction to one or two column width on a 5255.25 or 35 inch disk utilizing WordwordperfectPerfect originals must be no larger than 22 X 28 cm 424.2 or above number all pages and assemble each copy separately title page abstract and key words NOTES if the manuscriptinanuscript would be more appro- text acknowledgments literature cited appen- priate as a hortshorts communication or note follow the dices tables figure legends figures above instructions but do not include an abstract A CHARGE of 45 page is made TITLE PAGE includes an informative title no longer per for articles than 15 words names and addresses of authors a published the rate for subscribers will be 40 per page however manuscripts with complex tables running head of fewer than 40 letters and spaces I1 footnotes to indicate change of address and author andor numerous half tones will be assessed an to whom correspondence should be addressed if additional charge reprints may be purchased at other than the first author the time of publication an order form is sent with proofs ABSTRACT states the purpose methods results the and conclusions of the research it is followed by FINAL CHECK 6 12 key words listed in order of decreasing imim- cover letter explains any duplication of infor- portance to be used for indexing mation and provides phone numbers teffteaftefaTEXT has centered main headings printed in all 3 copies of the manuscript and WordwordperfectPerfect capital letters second level headings are centered disk in upper and lowercase letters third level head conformity with instructions ings begin paragraphs photocopies of illustrations ISSN 001736140017 3614 GREAT BASIN naturalist volvoi 53 no 2 june 1993 CONTENTS articles depletion of soil moisture by two cold desert bunchbunchgrassesgrasses and effects on photosyntheticperformancephotosynthetic performance jay E anderson and nancee L toft 97 on the relative importance offloral color shape and nectar rewards in attract- ing pollinatorspollinators to mimulus steven D sutherland and robert kvickeryKVickK vickeryery jr 107 response of a sonoran riparian forest to a loyear10 year return flood J C stromberg B D richter D T patten and L G wolden 118 habitat selection of merriam s turkey meleagris gallopavogallopavo merrimerriaminwrriamiami hens with boultspoults in the black hills south dakota mark A rumble and stanley H anderson 131134 squirrels as predators J R callahan 137 late quaternary vegetation and climate in the escalante river basin on the central colorado plateau kim withers and jim I1 mead 145 bole volume growth in stems of quercus gambegambeliigambelligambeliieillliiillili warren P clary and arthur R tiedemann 162462 aquatic habitats life history observations and zoogeographic considerations of the spotted frog rana pretpretiosapreciosaiosa in tule valley utah peter hovingh 168 growth of smallmouth bass micropterus dolomieudolomieu inin flaming gorge reser- voir wyoming utah scott A mullner and wayne A hubert 180 autumn and winter foods of the lesser prairie chicken tympanuchus pal lidicinctus galliformes tetraonidae terry Z riley I1 charles adavisA davis and randall A smith 186486 establishment of shoshone sculpin coituscottus greenelgreenei in a spring inhabited by mottled sculpin C bairdi derek B kuda and J S griffith 190490 use of boulder pocket habitat by rainbow trout oncorhynchus mykiss in fall river idaho daniel N streubel and J S griffith 494194 humoral creatinine in walleye stizostedion vitreum harry L holloway jr and craig A shoemaker 199499 influence of prey movement on the aim of predatory strikes of the western rattlesnake crotalus virviridisidis douglas F schmidt william K hayes and floyd E hayes 203