BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors nonprofit publishers academic institutions researchlibraries and research funders in the common goal of maximizing access to critical research

LIFE HISTORY AND ECOLOGY OF THE WOLF SPIDER PARDOSA SIERRABANKS (ARANEAE LYCOSIDAE) IN SOUTHEASTERN ARIZONAAuthor(s) Fred Punzo and Chris FarmerSource The Southwestern Naturalist 51(3)310-319 2006Published By Southwestern Association of NaturalistsDOI httpdxdoiorg1018940038-4909(2006)51[310LHAEOT]20CO2URL httpwwwbiooneorgdoifull1018940038-490928200629515B3103ALHAEOT5D20CO3B2

BioOne (wwwbiooneorg) is a nonprofit online aggregation of core research in the biological ecological andenvironmental sciences BioOne provides a sustainable online platform for over 170 journals and books publishedby nonprofit societies associations museums institutions and presses

Your use of this PDF the BioOne Web site and all posted and associated content indicates your acceptance ofBioOnersquos Terms of Use available at wwwbiooneorgpageterms_of_use

Usage of BioOne content is strictly limited to personal educational and non-commercial use Commercial inquiriesor rights and permissions requests should be directed to the individual publisher as copyright holder

THE SOUTHWESTERN NATURALIST 51(3)310ndash319 SEPTEMBER 2006

LIFE HISTORY AND ECOLOGY OF THE WOLF SPIDER PARDOSA SIERRABANKS (ARANEAE LYCOSIDAE) IN SOUTHEASTERN ARIZONA

FRED PUNZO AND CHRIS FARMER

Department of Biology Box 5F University of Tampa 401 West Kennedy Boulevard Tampa FL 33606-1490Correspondent fpunzoutedu

ABSTRACT In 2004 we studied the life history and ecology of a population of Pardosa sierra(Araneae Lycosidae) from Cave Creek Canyon (Chiricahua Mountains Arizona) Peak abundancefor immatures occurred in July September and October About 64 of all adult males and femaleswere observed in April and May Females were observed carrying egg sacs between 5 April and 3August Diet composition consisted primarily of beetles (184) flies (153) and ants (122)Spiders were active during the day as well as at night Immatures and adults preferred sandy orrocky substrates respectively Under controlled laboratory conditions the life cycle consisted of 9instars and mean carapace width increased from 066 mm (instar 1) to 226 mm (instar 9 adultmales and females) Clutch size ranged from 35 to 92 (mean 6781) Gestation period rangedfrom 22 to 27 days (mean 2406) Significant positive correlations were found between femalemass and size offspring mass and size female size and clutch size and female size and clutchmass Mean total clutch mass was 349 of mean female mass

RESUMEN En el 2004 estudiamos la historia de vida y la ecologıa de una poblacion de aranalobo (Araneae Lycosidae) de Cave Creek Canyon (Chiricahua Mountains Arizona) La abundan-cia maxima de aranas juveniles ocurrio en julio septiembre y octubre Alrededor de 64 detodos los adultos machos y hembras se observaron en abril y mayo Se observo a las hembrasllevando sacos de huevos entre el 5 de mayo y el 3 de agosto La composicion de la dieta consistioprincipalmente de escarabajos (184) moscas (153) y hormigas (122) Las aranas estuvi-eron activas durante el dıa y la noche Los juveniles y adultos prefirieron suelo arenoso o rocosorespectivamente En condiciones controladas en el laboratorio el ciclo de vida consistio de nueveestados y el promedio de anchura de la caparazon aumento desde 066 mm (estado uno) a 226mm (estado nueve adultos machos y hembras) El tamano de la nidada vario desde 35 a 92(promedio 6781) El tiempo de gestacion vario desde 22 a 27 dıas (promedio 2406) Seencontraron correlaciones positivas significativas entre la masa de las hembras y su tamano masade los neonatos y su tamano tamano de las hembras y tamano de la puesta y tamano de lashembras y masa de la puesta El promedio de la masa total de la puesta fue 349 de la mediade la masa de las hembras

Although few empirical data are available ongeographic variation in spider populationsvariation can be observed with respect togrowth rate phenology adult size and repro-duction (Whitcomb 1967 Foelix 1996 Hen-dricks and Maelfait 2003 Punzo 2003) Wolfspiders (Araneae Lycosidae) are cursorialhunters that can be found in almost all terres-trial habitats Upon eclosion spiderlings climbonto the back of their mother where they re-main for a period of time that can range from2 to 10 days depending on the species(Gertsch 1979) After the first post-eclosionmolt these spiderlings disperse and embark ona vagrant existence (Punzo 2003)

Although life histories of some wolf spiders(Lycosidae) have been studied in detail in-cluding species in Lycosa (Miyashita 1969)Pardosa (Eason 1969 Buddle 2000) Hogna(Punzo 2003) and Pirata (Hendricks andMaelfait 2003) data are lacking for many oth-ers In addition little is known for some spe-cies concerning geographic variation in lifehistory traits ecology and behavior from allo-patric populations (Punzo 2000)

Little detailed information is available onthe natural history ecology and behavior ofPardosa sierra Banks 1898 This wolf spider canbe found in a variety of microhabitats includ-ing rocky grasslands desertscrub along the

September 2006 311Punzo and FarmermdashLife history of Pardosa sierra

margins of rocky stream banks and gently slop-ing alluvial fans (Banks 1898 Whitcomb1967) In a study on the life cycle of P sierrafrom Mill Creek California Van Dyke andLowrie (1975) found that the time required tomature from egg to adulthood in captivityranged from 152 to 210 days depending onwhether adulthood was reached in instar 7 8or 9 Adult males and females began to appearin field samples in December reaching maxi-mum abundance in February and March Nomales were collected after September and per-cent composition of females was 10 Peakabundance for immatures was observed fromJune through September (80 to 94 of all in-dividuals) declining rapidly to 62 by Januaryand to 25 by February Females were ob-served carrying egg sacs from late Februarythrough mid September Immatures preferredopen areas with sandy soil whereas adults weremost frequently found among pebbles androcks along streams

The purpose of our study was to investigatethe life cycle of an allopatric population of Psierra from a higher elevation (upper CaveCreek Canyon elevation 1790 m) in south-eastern Arizona and compare it to what hasbeen reported by Van Dyke and Lowrie (1975)for P sierra from Mill Creek California (ele-vation 825 m)

METHODS Description of Study Site We conductedfield studies in 2004 along an intermittent tributaryof the North Fork of Cave Creek Canyon in the Chir-icahua Mountains of southeastern Arizona (CochiseCounty) In this area P sierra was found in rockywoodland habitats near intermittent streams Thecreek bed contained and was surrounded by boul-ders Ground surfaces along margins of the creekbed were covered with leaf litter Dominant vegeta-tion included Arizona white oak (Quercus arizonica)Apache pine (Pinus engelmannii) alligator juniper(Juniperus deppeana) sycamore (Platanus wrightii)cliff fendler (Fendlera rupicola) rabbitbrush (Chryso-thamnus pulchellus) and scattered clumps of grasses(Bouteloua Muhlenbergia Stipa Tridens) Cave Creekis typically dry in May and June but flows intermit-tently during the rainy season ( July and August)Mean monthly precipitation in 2004 ranged from alow of 060 cm (May) to a high of 1215 cm (August)with an annual mean of 4892 cm

Collection of Animals Phenology Diel Periodicity andDiet Composition in the Field Spiders were collectedmanually by walking along the creek bed and itsmargins following a linear transect approximately

085 km in length and 12 to 30 m in width search-ing under leaf litter and loose rocks and watchingfor wandering spiders visible at the surface Spidersfrom these locations were collected using an aspi-rator (BioQuip Model 1135C Gardena California)A vacuum pump (Insect Vac BioQuip Model2820A) was used to collect arthropods present onthe leaves of shrubs and within grasses Samplingoccurred every third day at 4-h intervals starting at0800 h Voucher specimens were deposited in theInvertebrate Collection at the University of Tampa

Spiders also were collected using pitfall traps(BioQuip Model 2838A) Each trap consisted of awhite plastic bucket (12 cm in depth 15 cm in di-ameter) filled to a depth of 3 to 4 cm with ethyleneglycol and fitted with a plastic lid to deflect rainfalland provide shade The mouth of the trap was po-sitioned flush with the ground surface Traps wereset within 15 rectangular transects located along theedges of the creek bed Each transect measured 20m 9 m Traps were set along 3 linear axes (19trapsaxis) separated by a width of 3 m and eachtrap within a linear axis was separated by a distanceof 1 m Traps were checked on a weekly basisthroughout the year Spiders collected manually orfrom traps were counted to determine seasonalabundance of immatures and adults

Data from collection of spiders allowed us to de-termine seasonal abundance patterns and temporalpatterns of activity (diel periodicity) Only spidersthat were observed to be active at the surface wereused to analyze diel periodicity

After collection spiders were placed in 70 eth-anol for subsequent species identification sex deter-mination for adults and measurement of carapacewidth Carapace width for immatures and adults wasmeasured to the nearest 01 mm by using a stereo-microscope with an ocular micrometer Using cara-pace width individuals of P sierra were classified asfollows according to previous studies by Banks(1898) and Van Dyke and Lewis (1975) based onanalysis of pedipalps immatures (08 to 16 mm)adults (18 to 23 mm)

Some adult spiders were collected with a prey itemin their chelicerae (n 65 37 males 28 females)In these cases the prey item was removed andplaced in 70 ethanol for subsequent identificationto order or family depending on the degree of di-gestion

Microhabitat Preferences We recorded the specificmicrohabitats associated with adults and immaturespiders collected in the field to determine possiblemicrohabitat preferences Data were categorized forspiders collected during daylight (0730 to 1600 h n 714 adults 761 immatures) and at night (2100 to0400 h n 202 and 187) The following microhab-itats were delineated for spiders collected duringdaylight 1) collected in open rocky areas with sparse

312 vol 51 no 3The Southwestern Naturalist

vegetation and exposed to sunlight (rock sun) 2) inopen shaded rocky areas (rock shade) 3) on sandin sunlight with sparse plant cover (sand sun) 4)on sand in shade (sand shade) 5) in grass exposedto sunlight (grass sun) 6) in shaded grass (grassshade) 7) under a bush and exposed to sunlight(bush sun) 8) under a shaded bush (bush shade)9) under a tree exposed to the sun (tree sun) 10)under a shaded tree (tree shade) 11) on groundunder leaf litter or plant debris 12) on surface oflitter and 13) in bushes or on trees off the ground(off ground) For spiders collected at night micro-habitats were designated as 1) rocky substrate(rock) 2) sandy substrate (sand) 3) in grass 4) un-der bush 5) under tree 6) on surface of leaf litter7) under leaf litter or plant debris and 8) up inbushes or on trees These microhabitats were basedon a familiarity with the study site acquired over sev-eral years of field research at this location

Chi-square contingency tests were used to com-pare percentages of spiders found in specific micro-habitats (resource states) Microhabitat nichebreadth was obtained by first calculating Levinrsquos in-dex (B) B 1 pj

2 where pj proportion of in-dividuals found in a resource state (Krebs 1989) Toaccount for the possibility that resources might varyin abundance Levinrsquos standardized niche breadth(BA) was calculated BA B 1n 1 where B Levinrsquos index and n number of resource statesValues for BA can vary from 0 (narrowest nichebreadth only one resource category is used) to 10(resource categories equally represented)

Maintenance of Spiders in the Laboratory Spiderscollected in the field were brought to the laboratoryImmatures and adults were housed individually inplastic vials (3 cm in diameter 13 cm in length) orcages (13 12 10 cm) respectively and main-tained in a climate-controlled room (24 04C 65to 75 relative humidity 12L12D photoperiod) Vi-als holding immatures were placed on their side togive immatures more room for movement and theywere provided with a strip of paper toweling as asubstrate Floors of adult cages were covered withvermiculite to a depth of 3 to 4 cm Immatures werefed daily on a mixed diet of fruit flies (Drosophilamelanogaster) small mealworms (Tenebrio molitor 4mm in length) and small cricket nymphs (Acheta do-mesticus 5 mm) One prey item was given to eachspider for each feeding bout with different prey spe-cies being alternated each day (ie fruit fly meal-worm cricket etc) A similar feeding regime wasused for adults who received one adult cricket larg-er mealworm (7 to 8 mm) or cockroach nymph(Periplaneta americana 7 to 8 mm) every other dayAll spiders were provided with water ad libitum (wa-ter-soaked cotton wads) Pilot studies had shown thatthese feeding schedules allowed P sierra to reach

adulthood and breed successfully in captivity (Pun-zo unpubl data)

Gestation Period Clutch Size and Mass and GrowthRate Seventy-three pairs of males and females werehoused in plastic cages (one paircage) and servedas a pool of subjects for mating observations Matedfemales (n 31) were removed and placed sepa-rately in plastic cages housed under identical cli-matic conditions as described previously and fed anidentical diet We recorded the amount of time (indays) that elapsed between copulation and construc-tion of egg sacs

We observed 44 females construct egg sacs afterseveral days to 3 weeks in captivity Females with eggsacs were maintained separately in glass dishes (10cm in diameter 8 cm in depth) with cheese clothcovering the open top of the dish under identicalclimatic conditions as described above Hatchlingsemerged from 35 of the 44 egg sacs These egg sacswere used to determine length of gestation periodwhich was defined as the length of time that elapsedbetween construction of egg sac and when spider-lings first emerged (Canard 1987) Clutch size wasdefined as the number of spiderlings emerging fromeach egg sac (Punzo 2003)

Following dispersal of all offspring from the backof their mother females (n 40) were weighed tothe nearest 01 mg using an electronic analytical bal-ance In addition all live dispersed spiderlings fromeach female were weighed together to obtain totalclutch mass (TCM) (Enders 1976) Mean offspringmass was determined by dividing TCM by clutch size(Roff 1992) To obtain a measurement of invest-ment in reproduction relative to female size we alsocalculated relative clutch mass (RCM) by dividingTCM by female mass (Roff 1992)

Females that had egg sacs when collected in thefield were brought into the laboratory placed indi-vidually in plastic cages as described above foradults maintained under identical climatic condi-tions and given an identical feeding schedule If aprey item had not been attacked or eaten within a24-h period it was removed from the cage Uponhatching spiderlings climbed onto the back of theirmother and remained there until dispersal aftertheir first post-eclosion molt The dispersed second-instar spiderlings were also placed individually inplastic vials and maintained as described aboveThey were fed on the immature feeding regime forinstars 2 through 5 thereafter receiving the adultfeeding regime as described above

Growth rate was determined from the first instarhatchlings by removing young from egg sacs as de-scribed by Dondale (1961) and then allowing themaccess to their mother after appropriate measure-ments were taken Measurements were taken on car-apace width to the nearest 01 mm After dispersalfrom their mother spiderlings were housed as de-

September 2006 313Punzo and FarmermdashLife history of Pardosa sierra

FIG 1 Frequency distribution (number of indi-viduals) for immatures and adults of Pardosa sierrafrom Cave Creek Canyon Arizona Squares im-matures circles adult females triangles adultmales

scribed above and carapace width measured for alllife cycle stages Carapace width has been used as areliable index of growth in spiders (Hagstrom1971)

Statistical Procedures All statistical analyses fol-lowed procedures outlined by Sokal and Rohlf(1995) Pearsonrsquos correlation (r) and least-squareslinear regression on log-transformed data were usedto assess relationships between female mass andbody size (carapace width) offspring size and massfemale size and clutch size and female size and off-spring size Differences in proportions of adult ver-sus immature spiders were analyzed by an R C testof independence Size differences were analyzed us-ing an analysis of variance (ANOVA) when assump-tions for normality and homogeneity of variancewere met Otherwise a Kruskal-Wallis nonparamet-ric ANOVA was used Multiple comparisons of dif-ferent size groups were performed using Scheffetests Chi-square contingency tests were used to com-pare differences associated with diel periodicity

RESULTS Phenology and Diel Periodicity Thenumbers of immatures and adults collectedfrom the field during various months of theyear are shown in Fig 1 Of a total of 2242spiders collected during 2004 486 (n 1090) were immatures 261 (586) adultmales and 252 (566) adult females Moreimmatures were collected during July (145)September (133) and October (157)while none were collected during Decemberand January Numbers of immatures increasedmarkedly in March and declined rapidly byNovember Adult males and females first ap-peared in field samples on 16 February and 28

January respectively although in small num-bers About 64 of all adult males and femaleswere collected in April and May

Females were observed carrying egg sacs be-tween 5 April and 3 August The majority offemales with egg sacs were observed in lateJune (28) and early July (23)

Although P sierra was active during daylightas well as at night and no differences werefound between the sexes (2 207 P 05)most spiders were active during the day (2 2005 P 001) We observed 714 of 916 adultmales (78) and 604 of 788 adult females(77) between 0730 and 1600 h (diurnal)and 761 of 948 immatures (80) were activeduring the day

Diet Composition Prey items found in chelic-erae of adult spiders (Table 1) indicate that Psierra is a generalist predator that feeds on avariety of ground-dwelling insects and arach-nids Beetles accounted for 184 of preyitems followed by flies (153) ants (123)springtails (92) caterpillars (92) orthop-terans (92) spiders (92) diplurans(46) termites (46) bugs (31) harvest-men (31) and thrips (15) Interestingly43 of the 65 (662) spiders observed with aprey item were not found on open sandy orrocky microhabitats with little to no vegetationbut rather on or under plant debris or undershrubs Immature or adult stages of arthropods(13 mm) with well known chemical defensesthat were common at this site such as velvetants (Mutillidae) stink bugs (Pentatomidae)blister beetles (Meloidae) and millipedes (Di-plopoda) were not observed in the cheliceraeof P sierra

Microhabitat Preferences Microhabitat prefer-ences of P sierra during daylight hours areshown in Table 2 No significant differenceswere found between sexes for daylight (2 311 P 005) and night (2 194 P 005)samples so data were pooled for adult malesand females Of 714 adults 553 (774) wereassociated with rocky substrates along the mar-gins of the creek bed or in streamside rubbleas compared to only 76 that were found onsandy substrate With respect to immatures540 of 761 (709) were on sandy substratesas compared to 71 on rocky substratesThese data are indicative of a narrow micro-habitat niche breadth for immatures (BA 02404) and adults (BA 02146) and also

314 vol 51 no 3The Southwestern Naturalist

TABLE 1mdashNumber and percent of prey itemsfound in chelicerae of 65 adult Pardosa sierra col-lected at Cave Creek Canyon Cochise County Ari-zona from 19 March to 23 July 2004 No significantdifferences were found between sexes (chi-square P 005) Data listed as the number (n) of spiders(total 65 37 males 28 females) and percent ()of all spiders found with a particular prey item Lifecycle stage of prey A adult I immature

Prey n

Insecta

Collembola (springtails) (A) 6 92Coleoptera (beetles)

Carabidae (A) 4 61Tenebrionidae (A) 2 31Unidentified (A) 6 92

Diplura (diplurans) (A) 3 46Diptera

Asilidae (robber flies) (A) 1 15Tephritidae (fruitflies) (A) 2 31Drosophilidae (vinegarflies) (A) 5 76Unidentified (A) 2 31

Hemiptera (bugs) (I) 2 31Hymenoptera

Formicidae (ants) (A) 8 123Isoptera (A) 3 46Lepidoptera

Heterocera (moths)Unidentified (I) 3 46

Rhopalocena (butterflies)Hesperiidae (I) 1 15

Unidentified (I) 2 31Orthoptera

Acrididae (grasshoppers) (I) 4 61Unidentified (I) 2 31

Thysanoptera (thrips) (A) 1 15

ArachnidaAraneae (spiders)

Araneidae (I) 3 46Lycosidae (I) 2 31Unidentified (I) 1 15

Opoliones (harvestmen) (A) 2 31Total prey items 65

show that immatures and adults prefer differ-ent microhabitats

Data for habitat preference for P sierra atnight are shown in Table 3 Again adults andimmatures showed a marked preference forrocky and sandy substrates respectively Wefound 57 of all adults on rocks or streamsiderubble compared to 118 on sand (2 1494 P 005) For immatures 47 were on

sandy substrate compared to 96 on rockysubstrates (2 1298 P 005)

No individuals were observed to climb offthe ground and into bushes shrubs or treesAt night 155 of immatures were under leaflitter or other surface plant debris comparedto only 39 of adults (2 1104 P 005)In contrast during daylight fewer immatures(57) and adults (16) were found underlitter

Gestation Period Growth Rate Clutch Size Ob-servation of 35 egg sacs from day of sac con-struction to emergence of spiderlings showedthat gestation period under controlled labora-tory conditions ranged from 22 to 27 days(mean 2406 177 SE) Clutch size rangedfrom 35 to 92 (mean 6781 681 SE)

The surface of the cephalothorax of newlyemerged hatchlings referred to as first post-embryos (first instar) (Vachon 1957) had ayellow-orange color the rest of the embryo wascolorless Upon hatching first post-embryoswere still covered by a thin membrane After aperiod of pulsating movements of legs and ab-domen lasting from 44 to 82 sec (mean 6403 972 SE) this membrane split andlegs cephalothorax and abdomen were pulledfree followed by the mouth and spinnerets Af-ter this first molt spiderlings are now referredto as second post-embryos (second instar)(Foelix 1996)

Growth rate as measured by carapace widthis shown in Table 4 Under controlled labora-tory conditions all P sierra males and femalesattained adulthood by the ninth instar Imma-ture males and females doubled their size bythe sixth instar

Duration of first instar ranged from 14 to 21h Durations of instars 2 through 9 are shownin Table 5 The average time required to ma-ture from egg to adulthood was 21797 daysThere were overall significant differences inthe duration of instars (F 3074 P 001)Longest duration was observed for instars 6and 7 and the shortest for instars 3 and 4 Al-though adult females lived significantly longerthan males (t 2587 P 001) there was nosignificant difference in overall average life-span from instar 1 through adulthood for fe-males and males (t 381 P 005)

A summary of pertinent life history data forP sierra is shown in Table 6 Female mass andsize were positively correlated (r 2 00705 P

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

September 2006 311Punzo and FarmermdashLife history of Pardosa sierra

margins of rocky stream banks and gently slop-ing alluvial fans (Banks 1898 Whitcomb1967) In a study on the life cycle of P sierrafrom Mill Creek California Van Dyke andLowrie (1975) found that the time required tomature from egg to adulthood in captivityranged from 152 to 210 days depending onwhether adulthood was reached in instar 7 8or 9 Adult males and females began to appearin field samples in December reaching maxi-mum abundance in February and March Nomales were collected after September and per-cent composition of females was 10 Peakabundance for immatures was observed fromJune through September (80 to 94 of all in-dividuals) declining rapidly to 62 by Januaryand to 25 by February Females were ob-served carrying egg sacs from late Februarythrough mid September Immatures preferredopen areas with sandy soil whereas adults weremost frequently found among pebbles androcks along streams

The purpose of our study was to investigatethe life cycle of an allopatric population of Psierra from a higher elevation (upper CaveCreek Canyon elevation 1790 m) in south-eastern Arizona and compare it to what hasbeen reported by Van Dyke and Lowrie (1975)for P sierra from Mill Creek California (ele-vation 825 m)

METHODS Description of Study Site We conductedfield studies in 2004 along an intermittent tributaryof the North Fork of Cave Creek Canyon in the Chir-icahua Mountains of southeastern Arizona (CochiseCounty) In this area P sierra was found in rockywoodland habitats near intermittent streams Thecreek bed contained and was surrounded by boul-ders Ground surfaces along margins of the creekbed were covered with leaf litter Dominant vegeta-tion included Arizona white oak (Quercus arizonica)Apache pine (Pinus engelmannii) alligator juniper(Juniperus deppeana) sycamore (Platanus wrightii)cliff fendler (Fendlera rupicola) rabbitbrush (Chryso-thamnus pulchellus) and scattered clumps of grasses(Bouteloua Muhlenbergia Stipa Tridens) Cave Creekis typically dry in May and June but flows intermit-tently during the rainy season ( July and August)Mean monthly precipitation in 2004 ranged from alow of 060 cm (May) to a high of 1215 cm (August)with an annual mean of 4892 cm

Collection of Animals Phenology Diel Periodicity andDiet Composition in the Field Spiders were collectedmanually by walking along the creek bed and itsmargins following a linear transect approximately

085 km in length and 12 to 30 m in width search-ing under leaf litter and loose rocks and watchingfor wandering spiders visible at the surface Spidersfrom these locations were collected using an aspi-rator (BioQuip Model 1135C Gardena California)A vacuum pump (Insect Vac BioQuip Model2820A) was used to collect arthropods present onthe leaves of shrubs and within grasses Samplingoccurred every third day at 4-h intervals starting at0800 h Voucher specimens were deposited in theInvertebrate Collection at the University of Tampa

Spiders also were collected using pitfall traps(BioQuip Model 2838A) Each trap consisted of awhite plastic bucket (12 cm in depth 15 cm in di-ameter) filled to a depth of 3 to 4 cm with ethyleneglycol and fitted with a plastic lid to deflect rainfalland provide shade The mouth of the trap was po-sitioned flush with the ground surface Traps wereset within 15 rectangular transects located along theedges of the creek bed Each transect measured 20m 9 m Traps were set along 3 linear axes (19trapsaxis) separated by a width of 3 m and eachtrap within a linear axis was separated by a distanceof 1 m Traps were checked on a weekly basisthroughout the year Spiders collected manually orfrom traps were counted to determine seasonalabundance of immatures and adults

Data from collection of spiders allowed us to de-termine seasonal abundance patterns and temporalpatterns of activity (diel periodicity) Only spidersthat were observed to be active at the surface wereused to analyze diel periodicity

After collection spiders were placed in 70 eth-anol for subsequent species identification sex deter-mination for adults and measurement of carapacewidth Carapace width for immatures and adults wasmeasured to the nearest 01 mm by using a stereo-microscope with an ocular micrometer Using cara-pace width individuals of P sierra were classified asfollows according to previous studies by Banks(1898) and Van Dyke and Lewis (1975) based onanalysis of pedipalps immatures (08 to 16 mm)adults (18 to 23 mm)

Some adult spiders were collected with a prey itemin their chelicerae (n 65 37 males 28 females)In these cases the prey item was removed andplaced in 70 ethanol for subsequent identificationto order or family depending on the degree of di-gestion

Microhabitat Preferences We recorded the specificmicrohabitats associated with adults and immaturespiders collected in the field to determine possiblemicrohabitat preferences Data were categorized forspiders collected during daylight (0730 to 1600 h n 714 adults 761 immatures) and at night (2100 to0400 h n 202 and 187) The following microhab-itats were delineated for spiders collected duringdaylight 1) collected in open rocky areas with sparse

312 vol 51 no 3The Southwestern Naturalist

vegetation and exposed to sunlight (rock sun) 2) inopen shaded rocky areas (rock shade) 3) on sandin sunlight with sparse plant cover (sand sun) 4)on sand in shade (sand shade) 5) in grass exposedto sunlight (grass sun) 6) in shaded grass (grassshade) 7) under a bush and exposed to sunlight(bush sun) 8) under a shaded bush (bush shade)9) under a tree exposed to the sun (tree sun) 10)under a shaded tree (tree shade) 11) on groundunder leaf litter or plant debris 12) on surface oflitter and 13) in bushes or on trees off the ground(off ground) For spiders collected at night micro-habitats were designated as 1) rocky substrate(rock) 2) sandy substrate (sand) 3) in grass 4) un-der bush 5) under tree 6) on surface of leaf litter7) under leaf litter or plant debris and 8) up inbushes or on trees These microhabitats were basedon a familiarity with the study site acquired over sev-eral years of field research at this location

Chi-square contingency tests were used to com-pare percentages of spiders found in specific micro-habitats (resource states) Microhabitat nichebreadth was obtained by first calculating Levinrsquos in-dex (B) B 1 pj

2 where pj proportion of in-dividuals found in a resource state (Krebs 1989) Toaccount for the possibility that resources might varyin abundance Levinrsquos standardized niche breadth(BA) was calculated BA B 1n 1 where B Levinrsquos index and n number of resource statesValues for BA can vary from 0 (narrowest nichebreadth only one resource category is used) to 10(resource categories equally represented)

Maintenance of Spiders in the Laboratory Spiderscollected in the field were brought to the laboratoryImmatures and adults were housed individually inplastic vials (3 cm in diameter 13 cm in length) orcages (13 12 10 cm) respectively and main-tained in a climate-controlled room (24 04C 65to 75 relative humidity 12L12D photoperiod) Vi-als holding immatures were placed on their side togive immatures more room for movement and theywere provided with a strip of paper toweling as asubstrate Floors of adult cages were covered withvermiculite to a depth of 3 to 4 cm Immatures werefed daily on a mixed diet of fruit flies (Drosophilamelanogaster) small mealworms (Tenebrio molitor 4mm in length) and small cricket nymphs (Acheta do-mesticus 5 mm) One prey item was given to eachspider for each feeding bout with different prey spe-cies being alternated each day (ie fruit fly meal-worm cricket etc) A similar feeding regime wasused for adults who received one adult cricket larg-er mealworm (7 to 8 mm) or cockroach nymph(Periplaneta americana 7 to 8 mm) every other dayAll spiders were provided with water ad libitum (wa-ter-soaked cotton wads) Pilot studies had shown thatthese feeding schedules allowed P sierra to reach

adulthood and breed successfully in captivity (Pun-zo unpubl data)

Gestation Period Clutch Size and Mass and GrowthRate Seventy-three pairs of males and females werehoused in plastic cages (one paircage) and servedas a pool of subjects for mating observations Matedfemales (n 31) were removed and placed sepa-rately in plastic cages housed under identical cli-matic conditions as described previously and fed anidentical diet We recorded the amount of time (indays) that elapsed between copulation and construc-tion of egg sacs

We observed 44 females construct egg sacs afterseveral days to 3 weeks in captivity Females with eggsacs were maintained separately in glass dishes (10cm in diameter 8 cm in depth) with cheese clothcovering the open top of the dish under identicalclimatic conditions as described above Hatchlingsemerged from 35 of the 44 egg sacs These egg sacswere used to determine length of gestation periodwhich was defined as the length of time that elapsedbetween construction of egg sac and when spider-lings first emerged (Canard 1987) Clutch size wasdefined as the number of spiderlings emerging fromeach egg sac (Punzo 2003)

Following dispersal of all offspring from the backof their mother females (n 40) were weighed tothe nearest 01 mg using an electronic analytical bal-ance In addition all live dispersed spiderlings fromeach female were weighed together to obtain totalclutch mass (TCM) (Enders 1976) Mean offspringmass was determined by dividing TCM by clutch size(Roff 1992) To obtain a measurement of invest-ment in reproduction relative to female size we alsocalculated relative clutch mass (RCM) by dividingTCM by female mass (Roff 1992)

Females that had egg sacs when collected in thefield were brought into the laboratory placed indi-vidually in plastic cages as described above foradults maintained under identical climatic condi-tions and given an identical feeding schedule If aprey item had not been attacked or eaten within a24-h period it was removed from the cage Uponhatching spiderlings climbed onto the back of theirmother and remained there until dispersal aftertheir first post-eclosion molt The dispersed second-instar spiderlings were also placed individually inplastic vials and maintained as described aboveThey were fed on the immature feeding regime forinstars 2 through 5 thereafter receiving the adultfeeding regime as described above

Growth rate was determined from the first instarhatchlings by removing young from egg sacs as de-scribed by Dondale (1961) and then allowing themaccess to their mother after appropriate measure-ments were taken Measurements were taken on car-apace width to the nearest 01 mm After dispersalfrom their mother spiderlings were housed as de-

September 2006 313Punzo and FarmermdashLife history of Pardosa sierra

FIG 1 Frequency distribution (number of indi-viduals) for immatures and adults of Pardosa sierrafrom Cave Creek Canyon Arizona Squares im-matures circles adult females triangles adultmales

scribed above and carapace width measured for alllife cycle stages Carapace width has been used as areliable index of growth in spiders (Hagstrom1971)

Statistical Procedures All statistical analyses fol-lowed procedures outlined by Sokal and Rohlf(1995) Pearsonrsquos correlation (r) and least-squareslinear regression on log-transformed data were usedto assess relationships between female mass andbody size (carapace width) offspring size and massfemale size and clutch size and female size and off-spring size Differences in proportions of adult ver-sus immature spiders were analyzed by an R C testof independence Size differences were analyzed us-ing an analysis of variance (ANOVA) when assump-tions for normality and homogeneity of variancewere met Otherwise a Kruskal-Wallis nonparamet-ric ANOVA was used Multiple comparisons of dif-ferent size groups were performed using Scheffetests Chi-square contingency tests were used to com-pare differences associated with diel periodicity

RESULTS Phenology and Diel Periodicity Thenumbers of immatures and adults collectedfrom the field during various months of theyear are shown in Fig 1 Of a total of 2242spiders collected during 2004 486 (n 1090) were immatures 261 (586) adultmales and 252 (566) adult females Moreimmatures were collected during July (145)September (133) and October (157)while none were collected during Decemberand January Numbers of immatures increasedmarkedly in March and declined rapidly byNovember Adult males and females first ap-peared in field samples on 16 February and 28

January respectively although in small num-bers About 64 of all adult males and femaleswere collected in April and May

Females were observed carrying egg sacs be-tween 5 April and 3 August The majority offemales with egg sacs were observed in lateJune (28) and early July (23)

Although P sierra was active during daylightas well as at night and no differences werefound between the sexes (2 207 P 05)most spiders were active during the day (2 2005 P 001) We observed 714 of 916 adultmales (78) and 604 of 788 adult females(77) between 0730 and 1600 h (diurnal)and 761 of 948 immatures (80) were activeduring the day

Diet Composition Prey items found in chelic-erae of adult spiders (Table 1) indicate that Psierra is a generalist predator that feeds on avariety of ground-dwelling insects and arach-nids Beetles accounted for 184 of preyitems followed by flies (153) ants (123)springtails (92) caterpillars (92) orthop-terans (92) spiders (92) diplurans(46) termites (46) bugs (31) harvest-men (31) and thrips (15) Interestingly43 of the 65 (662) spiders observed with aprey item were not found on open sandy orrocky microhabitats with little to no vegetationbut rather on or under plant debris or undershrubs Immature or adult stages of arthropods(13 mm) with well known chemical defensesthat were common at this site such as velvetants (Mutillidae) stink bugs (Pentatomidae)blister beetles (Meloidae) and millipedes (Di-plopoda) were not observed in the cheliceraeof P sierra

Microhabitat Preferences Microhabitat prefer-ences of P sierra during daylight hours areshown in Table 2 No significant differenceswere found between sexes for daylight (2 311 P 005) and night (2 194 P 005)samples so data were pooled for adult malesand females Of 714 adults 553 (774) wereassociated with rocky substrates along the mar-gins of the creek bed or in streamside rubbleas compared to only 76 that were found onsandy substrate With respect to immatures540 of 761 (709) were on sandy substratesas compared to 71 on rocky substratesThese data are indicative of a narrow micro-habitat niche breadth for immatures (BA 02404) and adults (BA 02146) and also

314 vol 51 no 3The Southwestern Naturalist

TABLE 1mdashNumber and percent of prey itemsfound in chelicerae of 65 adult Pardosa sierra col-lected at Cave Creek Canyon Cochise County Ari-zona from 19 March to 23 July 2004 No significantdifferences were found between sexes (chi-square P 005) Data listed as the number (n) of spiders(total 65 37 males 28 females) and percent ()of all spiders found with a particular prey item Lifecycle stage of prey A adult I immature

Prey n

Insecta

Collembola (springtails) (A) 6 92Coleoptera (beetles)

Carabidae (A) 4 61Tenebrionidae (A) 2 31Unidentified (A) 6 92

Diplura (diplurans) (A) 3 46Diptera

Asilidae (robber flies) (A) 1 15Tephritidae (fruitflies) (A) 2 31Drosophilidae (vinegarflies) (A) 5 76Unidentified (A) 2 31

Hemiptera (bugs) (I) 2 31Hymenoptera

Formicidae (ants) (A) 8 123Isoptera (A) 3 46Lepidoptera

Heterocera (moths)Unidentified (I) 3 46

Rhopalocena (butterflies)Hesperiidae (I) 1 15

Unidentified (I) 2 31Orthoptera

Acrididae (grasshoppers) (I) 4 61Unidentified (I) 2 31

Thysanoptera (thrips) (A) 1 15

ArachnidaAraneae (spiders)

Araneidae (I) 3 46Lycosidae (I) 2 31Unidentified (I) 1 15

Opoliones (harvestmen) (A) 2 31Total prey items 65

show that immatures and adults prefer differ-ent microhabitats

Data for habitat preference for P sierra atnight are shown in Table 3 Again adults andimmatures showed a marked preference forrocky and sandy substrates respectively Wefound 57 of all adults on rocks or streamsiderubble compared to 118 on sand (2 1494 P 005) For immatures 47 were on

sandy substrate compared to 96 on rockysubstrates (2 1298 P 005)

No individuals were observed to climb offthe ground and into bushes shrubs or treesAt night 155 of immatures were under leaflitter or other surface plant debris comparedto only 39 of adults (2 1104 P 005)In contrast during daylight fewer immatures(57) and adults (16) were found underlitter

Gestation Period Growth Rate Clutch Size Ob-servation of 35 egg sacs from day of sac con-struction to emergence of spiderlings showedthat gestation period under controlled labora-tory conditions ranged from 22 to 27 days(mean 2406 177 SE) Clutch size rangedfrom 35 to 92 (mean 6781 681 SE)

The surface of the cephalothorax of newlyemerged hatchlings referred to as first post-embryos (first instar) (Vachon 1957) had ayellow-orange color the rest of the embryo wascolorless Upon hatching first post-embryoswere still covered by a thin membrane After aperiod of pulsating movements of legs and ab-domen lasting from 44 to 82 sec (mean 6403 972 SE) this membrane split andlegs cephalothorax and abdomen were pulledfree followed by the mouth and spinnerets Af-ter this first molt spiderlings are now referredto as second post-embryos (second instar)(Foelix 1996)

Growth rate as measured by carapace widthis shown in Table 4 Under controlled labora-tory conditions all P sierra males and femalesattained adulthood by the ninth instar Imma-ture males and females doubled their size bythe sixth instar

Duration of first instar ranged from 14 to 21h Durations of instars 2 through 9 are shownin Table 5 The average time required to ma-ture from egg to adulthood was 21797 daysThere were overall significant differences inthe duration of instars (F 3074 P 001)Longest duration was observed for instars 6and 7 and the shortest for instars 3 and 4 Al-though adult females lived significantly longerthan males (t 2587 P 001) there was nosignificant difference in overall average life-span from instar 1 through adulthood for fe-males and males (t 381 P 005)

A summary of pertinent life history data forP sierra is shown in Table 6 Female mass andsize were positively correlated (r 2 00705 P

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

312 vol 51 no 3The Southwestern Naturalist

vegetation and exposed to sunlight (rock sun) 2) inopen shaded rocky areas (rock shade) 3) on sandin sunlight with sparse plant cover (sand sun) 4)on sand in shade (sand shade) 5) in grass exposedto sunlight (grass sun) 6) in shaded grass (grassshade) 7) under a bush and exposed to sunlight(bush sun) 8) under a shaded bush (bush shade)9) under a tree exposed to the sun (tree sun) 10)under a shaded tree (tree shade) 11) on groundunder leaf litter or plant debris 12) on surface oflitter and 13) in bushes or on trees off the ground(off ground) For spiders collected at night micro-habitats were designated as 1) rocky substrate(rock) 2) sandy substrate (sand) 3) in grass 4) un-der bush 5) under tree 6) on surface of leaf litter7) under leaf litter or plant debris and 8) up inbushes or on trees These microhabitats were basedon a familiarity with the study site acquired over sev-eral years of field research at this location

Chi-square contingency tests were used to com-pare percentages of spiders found in specific micro-habitats (resource states) Microhabitat nichebreadth was obtained by first calculating Levinrsquos in-dex (B) B 1 pj

2 where pj proportion of in-dividuals found in a resource state (Krebs 1989) Toaccount for the possibility that resources might varyin abundance Levinrsquos standardized niche breadth(BA) was calculated BA B 1n 1 where B Levinrsquos index and n number of resource statesValues for BA can vary from 0 (narrowest nichebreadth only one resource category is used) to 10(resource categories equally represented)

Maintenance of Spiders in the Laboratory Spiderscollected in the field were brought to the laboratoryImmatures and adults were housed individually inplastic vials (3 cm in diameter 13 cm in length) orcages (13 12 10 cm) respectively and main-tained in a climate-controlled room (24 04C 65to 75 relative humidity 12L12D photoperiod) Vi-als holding immatures were placed on their side togive immatures more room for movement and theywere provided with a strip of paper toweling as asubstrate Floors of adult cages were covered withvermiculite to a depth of 3 to 4 cm Immatures werefed daily on a mixed diet of fruit flies (Drosophilamelanogaster) small mealworms (Tenebrio molitor 4mm in length) and small cricket nymphs (Acheta do-mesticus 5 mm) One prey item was given to eachspider for each feeding bout with different prey spe-cies being alternated each day (ie fruit fly meal-worm cricket etc) A similar feeding regime wasused for adults who received one adult cricket larg-er mealworm (7 to 8 mm) or cockroach nymph(Periplaneta americana 7 to 8 mm) every other dayAll spiders were provided with water ad libitum (wa-ter-soaked cotton wads) Pilot studies had shown thatthese feeding schedules allowed P sierra to reach

adulthood and breed successfully in captivity (Pun-zo unpubl data)

Gestation Period Clutch Size and Mass and GrowthRate Seventy-three pairs of males and females werehoused in plastic cages (one paircage) and servedas a pool of subjects for mating observations Matedfemales (n 31) were removed and placed sepa-rately in plastic cages housed under identical cli-matic conditions as described previously and fed anidentical diet We recorded the amount of time (indays) that elapsed between copulation and construc-tion of egg sacs

We observed 44 females construct egg sacs afterseveral days to 3 weeks in captivity Females with eggsacs were maintained separately in glass dishes (10cm in diameter 8 cm in depth) with cheese clothcovering the open top of the dish under identicalclimatic conditions as described above Hatchlingsemerged from 35 of the 44 egg sacs These egg sacswere used to determine length of gestation periodwhich was defined as the length of time that elapsedbetween construction of egg sac and when spider-lings first emerged (Canard 1987) Clutch size wasdefined as the number of spiderlings emerging fromeach egg sac (Punzo 2003)

Following dispersal of all offspring from the backof their mother females (n 40) were weighed tothe nearest 01 mg using an electronic analytical bal-ance In addition all live dispersed spiderlings fromeach female were weighed together to obtain totalclutch mass (TCM) (Enders 1976) Mean offspringmass was determined by dividing TCM by clutch size(Roff 1992) To obtain a measurement of invest-ment in reproduction relative to female size we alsocalculated relative clutch mass (RCM) by dividingTCM by female mass (Roff 1992)

Females that had egg sacs when collected in thefield were brought into the laboratory placed indi-vidually in plastic cages as described above foradults maintained under identical climatic condi-tions and given an identical feeding schedule If aprey item had not been attacked or eaten within a24-h period it was removed from the cage Uponhatching spiderlings climbed onto the back of theirmother and remained there until dispersal aftertheir first post-eclosion molt The dispersed second-instar spiderlings were also placed individually inplastic vials and maintained as described aboveThey were fed on the immature feeding regime forinstars 2 through 5 thereafter receiving the adultfeeding regime as described above

Growth rate was determined from the first instarhatchlings by removing young from egg sacs as de-scribed by Dondale (1961) and then allowing themaccess to their mother after appropriate measure-ments were taken Measurements were taken on car-apace width to the nearest 01 mm After dispersalfrom their mother spiderlings were housed as de-

September 2006 313Punzo and FarmermdashLife history of Pardosa sierra

FIG 1 Frequency distribution (number of indi-viduals) for immatures and adults of Pardosa sierrafrom Cave Creek Canyon Arizona Squares im-matures circles adult females triangles adultmales

scribed above and carapace width measured for alllife cycle stages Carapace width has been used as areliable index of growth in spiders (Hagstrom1971)

Statistical Procedures All statistical analyses fol-lowed procedures outlined by Sokal and Rohlf(1995) Pearsonrsquos correlation (r) and least-squareslinear regression on log-transformed data were usedto assess relationships between female mass andbody size (carapace width) offspring size and massfemale size and clutch size and female size and off-spring size Differences in proportions of adult ver-sus immature spiders were analyzed by an R C testof independence Size differences were analyzed us-ing an analysis of variance (ANOVA) when assump-tions for normality and homogeneity of variancewere met Otherwise a Kruskal-Wallis nonparamet-ric ANOVA was used Multiple comparisons of dif-ferent size groups were performed using Scheffetests Chi-square contingency tests were used to com-pare differences associated with diel periodicity

RESULTS Phenology and Diel Periodicity Thenumbers of immatures and adults collectedfrom the field during various months of theyear are shown in Fig 1 Of a total of 2242spiders collected during 2004 486 (n 1090) were immatures 261 (586) adultmales and 252 (566) adult females Moreimmatures were collected during July (145)September (133) and October (157)while none were collected during Decemberand January Numbers of immatures increasedmarkedly in March and declined rapidly byNovember Adult males and females first ap-peared in field samples on 16 February and 28

January respectively although in small num-bers About 64 of all adult males and femaleswere collected in April and May

Females were observed carrying egg sacs be-tween 5 April and 3 August The majority offemales with egg sacs were observed in lateJune (28) and early July (23)

Although P sierra was active during daylightas well as at night and no differences werefound between the sexes (2 207 P 05)most spiders were active during the day (2 2005 P 001) We observed 714 of 916 adultmales (78) and 604 of 788 adult females(77) between 0730 and 1600 h (diurnal)and 761 of 948 immatures (80) were activeduring the day

Diet Composition Prey items found in chelic-erae of adult spiders (Table 1) indicate that Psierra is a generalist predator that feeds on avariety of ground-dwelling insects and arach-nids Beetles accounted for 184 of preyitems followed by flies (153) ants (123)springtails (92) caterpillars (92) orthop-terans (92) spiders (92) diplurans(46) termites (46) bugs (31) harvest-men (31) and thrips (15) Interestingly43 of the 65 (662) spiders observed with aprey item were not found on open sandy orrocky microhabitats with little to no vegetationbut rather on or under plant debris or undershrubs Immature or adult stages of arthropods(13 mm) with well known chemical defensesthat were common at this site such as velvetants (Mutillidae) stink bugs (Pentatomidae)blister beetles (Meloidae) and millipedes (Di-plopoda) were not observed in the cheliceraeof P sierra

Microhabitat Preferences Microhabitat prefer-ences of P sierra during daylight hours areshown in Table 2 No significant differenceswere found between sexes for daylight (2 311 P 005) and night (2 194 P 005)samples so data were pooled for adult malesand females Of 714 adults 553 (774) wereassociated with rocky substrates along the mar-gins of the creek bed or in streamside rubbleas compared to only 76 that were found onsandy substrate With respect to immatures540 of 761 (709) were on sandy substratesas compared to 71 on rocky substratesThese data are indicative of a narrow micro-habitat niche breadth for immatures (BA 02404) and adults (BA 02146) and also

314 vol 51 no 3The Southwestern Naturalist

TABLE 1mdashNumber and percent of prey itemsfound in chelicerae of 65 adult Pardosa sierra col-lected at Cave Creek Canyon Cochise County Ari-zona from 19 March to 23 July 2004 No significantdifferences were found between sexes (chi-square P 005) Data listed as the number (n) of spiders(total 65 37 males 28 females) and percent ()of all spiders found with a particular prey item Lifecycle stage of prey A adult I immature

Prey n

Insecta

Collembola (springtails) (A) 6 92Coleoptera (beetles)

Carabidae (A) 4 61Tenebrionidae (A) 2 31Unidentified (A) 6 92

Diplura (diplurans) (A) 3 46Diptera

Asilidae (robber flies) (A) 1 15Tephritidae (fruitflies) (A) 2 31Drosophilidae (vinegarflies) (A) 5 76Unidentified (A) 2 31

Hemiptera (bugs) (I) 2 31Hymenoptera

Formicidae (ants) (A) 8 123Isoptera (A) 3 46Lepidoptera

Heterocera (moths)Unidentified (I) 3 46

Rhopalocena (butterflies)Hesperiidae (I) 1 15

Unidentified (I) 2 31Orthoptera

Acrididae (grasshoppers) (I) 4 61Unidentified (I) 2 31

Thysanoptera (thrips) (A) 1 15

ArachnidaAraneae (spiders)

Araneidae (I) 3 46Lycosidae (I) 2 31Unidentified (I) 1 15

Opoliones (harvestmen) (A) 2 31Total prey items 65

show that immatures and adults prefer differ-ent microhabitats

Data for habitat preference for P sierra atnight are shown in Table 3 Again adults andimmatures showed a marked preference forrocky and sandy substrates respectively Wefound 57 of all adults on rocks or streamsiderubble compared to 118 on sand (2 1494 P 005) For immatures 47 were on

sandy substrate compared to 96 on rockysubstrates (2 1298 P 005)

No individuals were observed to climb offthe ground and into bushes shrubs or treesAt night 155 of immatures were under leaflitter or other surface plant debris comparedto only 39 of adults (2 1104 P 005)In contrast during daylight fewer immatures(57) and adults (16) were found underlitter

Gestation Period Growth Rate Clutch Size Ob-servation of 35 egg sacs from day of sac con-struction to emergence of spiderlings showedthat gestation period under controlled labora-tory conditions ranged from 22 to 27 days(mean 2406 177 SE) Clutch size rangedfrom 35 to 92 (mean 6781 681 SE)

The surface of the cephalothorax of newlyemerged hatchlings referred to as first post-embryos (first instar) (Vachon 1957) had ayellow-orange color the rest of the embryo wascolorless Upon hatching first post-embryoswere still covered by a thin membrane After aperiod of pulsating movements of legs and ab-domen lasting from 44 to 82 sec (mean 6403 972 SE) this membrane split andlegs cephalothorax and abdomen were pulledfree followed by the mouth and spinnerets Af-ter this first molt spiderlings are now referredto as second post-embryos (second instar)(Foelix 1996)

Growth rate as measured by carapace widthis shown in Table 4 Under controlled labora-tory conditions all P sierra males and femalesattained adulthood by the ninth instar Imma-ture males and females doubled their size bythe sixth instar

Duration of first instar ranged from 14 to 21h Durations of instars 2 through 9 are shownin Table 5 The average time required to ma-ture from egg to adulthood was 21797 daysThere were overall significant differences inthe duration of instars (F 3074 P 001)Longest duration was observed for instars 6and 7 and the shortest for instars 3 and 4 Al-though adult females lived significantly longerthan males (t 2587 P 001) there was nosignificant difference in overall average life-span from instar 1 through adulthood for fe-males and males (t 381 P 005)

A summary of pertinent life history data forP sierra is shown in Table 6 Female mass andsize were positively correlated (r 2 00705 P

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

September 2006 313Punzo and FarmermdashLife history of Pardosa sierra

FIG 1 Frequency distribution (number of indi-viduals) for immatures and adults of Pardosa sierrafrom Cave Creek Canyon Arizona Squares im-matures circles adult females triangles adultmales

scribed above and carapace width measured for alllife cycle stages Carapace width has been used as areliable index of growth in spiders (Hagstrom1971)

Statistical Procedures All statistical analyses fol-lowed procedures outlined by Sokal and Rohlf(1995) Pearsonrsquos correlation (r) and least-squareslinear regression on log-transformed data were usedto assess relationships between female mass andbody size (carapace width) offspring size and massfemale size and clutch size and female size and off-spring size Differences in proportions of adult ver-sus immature spiders were analyzed by an R C testof independence Size differences were analyzed us-ing an analysis of variance (ANOVA) when assump-tions for normality and homogeneity of variancewere met Otherwise a Kruskal-Wallis nonparamet-ric ANOVA was used Multiple comparisons of dif-ferent size groups were performed using Scheffetests Chi-square contingency tests were used to com-pare differences associated with diel periodicity

RESULTS Phenology and Diel Periodicity Thenumbers of immatures and adults collectedfrom the field during various months of theyear are shown in Fig 1 Of a total of 2242spiders collected during 2004 486 (n 1090) were immatures 261 (586) adultmales and 252 (566) adult females Moreimmatures were collected during July (145)September (133) and October (157)while none were collected during Decemberand January Numbers of immatures increasedmarkedly in March and declined rapidly byNovember Adult males and females first ap-peared in field samples on 16 February and 28

January respectively although in small num-bers About 64 of all adult males and femaleswere collected in April and May

Females were observed carrying egg sacs be-tween 5 April and 3 August The majority offemales with egg sacs were observed in lateJune (28) and early July (23)

Although P sierra was active during daylightas well as at night and no differences werefound between the sexes (2 207 P 05)most spiders were active during the day (2 2005 P 001) We observed 714 of 916 adultmales (78) and 604 of 788 adult females(77) between 0730 and 1600 h (diurnal)and 761 of 948 immatures (80) were activeduring the day

Diet Composition Prey items found in chelic-erae of adult spiders (Table 1) indicate that Psierra is a generalist predator that feeds on avariety of ground-dwelling insects and arach-nids Beetles accounted for 184 of preyitems followed by flies (153) ants (123)springtails (92) caterpillars (92) orthop-terans (92) spiders (92) diplurans(46) termites (46) bugs (31) harvest-men (31) and thrips (15) Interestingly43 of the 65 (662) spiders observed with aprey item were not found on open sandy orrocky microhabitats with little to no vegetationbut rather on or under plant debris or undershrubs Immature or adult stages of arthropods(13 mm) with well known chemical defensesthat were common at this site such as velvetants (Mutillidae) stink bugs (Pentatomidae)blister beetles (Meloidae) and millipedes (Di-plopoda) were not observed in the cheliceraeof P sierra

Microhabitat Preferences Microhabitat prefer-ences of P sierra during daylight hours areshown in Table 2 No significant differenceswere found between sexes for daylight (2 311 P 005) and night (2 194 P 005)samples so data were pooled for adult malesand females Of 714 adults 553 (774) wereassociated with rocky substrates along the mar-gins of the creek bed or in streamside rubbleas compared to only 76 that were found onsandy substrate With respect to immatures540 of 761 (709) were on sandy substratesas compared to 71 on rocky substratesThese data are indicative of a narrow micro-habitat niche breadth for immatures (BA 02404) and adults (BA 02146) and also

314 vol 51 no 3The Southwestern Naturalist

TABLE 1mdashNumber and percent of prey itemsfound in chelicerae of 65 adult Pardosa sierra col-lected at Cave Creek Canyon Cochise County Ari-zona from 19 March to 23 July 2004 No significantdifferences were found between sexes (chi-square P 005) Data listed as the number (n) of spiders(total 65 37 males 28 females) and percent ()of all spiders found with a particular prey item Lifecycle stage of prey A adult I immature

Prey n

Insecta

Collembola (springtails) (A) 6 92Coleoptera (beetles)

Carabidae (A) 4 61Tenebrionidae (A) 2 31Unidentified (A) 6 92

Diplura (diplurans) (A) 3 46Diptera

Asilidae (robber flies) (A) 1 15Tephritidae (fruitflies) (A) 2 31Drosophilidae (vinegarflies) (A) 5 76Unidentified (A) 2 31

Hemiptera (bugs) (I) 2 31Hymenoptera

Formicidae (ants) (A) 8 123Isoptera (A) 3 46Lepidoptera

Heterocera (moths)Unidentified (I) 3 46

Rhopalocena (butterflies)Hesperiidae (I) 1 15

Unidentified (I) 2 31Orthoptera

Acrididae (grasshoppers) (I) 4 61Unidentified (I) 2 31

Thysanoptera (thrips) (A) 1 15

ArachnidaAraneae (spiders)

Araneidae (I) 3 46Lycosidae (I) 2 31Unidentified (I) 1 15

Opoliones (harvestmen) (A) 2 31Total prey items 65

show that immatures and adults prefer differ-ent microhabitats

Data for habitat preference for P sierra atnight are shown in Table 3 Again adults andimmatures showed a marked preference forrocky and sandy substrates respectively Wefound 57 of all adults on rocks or streamsiderubble compared to 118 on sand (2 1494 P 005) For immatures 47 were on

sandy substrate compared to 96 on rockysubstrates (2 1298 P 005)

No individuals were observed to climb offthe ground and into bushes shrubs or treesAt night 155 of immatures were under leaflitter or other surface plant debris comparedto only 39 of adults (2 1104 P 005)In contrast during daylight fewer immatures(57) and adults (16) were found underlitter

Gestation Period Growth Rate Clutch Size Ob-servation of 35 egg sacs from day of sac con-struction to emergence of spiderlings showedthat gestation period under controlled labora-tory conditions ranged from 22 to 27 days(mean 2406 177 SE) Clutch size rangedfrom 35 to 92 (mean 6781 681 SE)

The surface of the cephalothorax of newlyemerged hatchlings referred to as first post-embryos (first instar) (Vachon 1957) had ayellow-orange color the rest of the embryo wascolorless Upon hatching first post-embryoswere still covered by a thin membrane After aperiod of pulsating movements of legs and ab-domen lasting from 44 to 82 sec (mean 6403 972 SE) this membrane split andlegs cephalothorax and abdomen were pulledfree followed by the mouth and spinnerets Af-ter this first molt spiderlings are now referredto as second post-embryos (second instar)(Foelix 1996)

Growth rate as measured by carapace widthis shown in Table 4 Under controlled labora-tory conditions all P sierra males and femalesattained adulthood by the ninth instar Imma-ture males and females doubled their size bythe sixth instar

Duration of first instar ranged from 14 to 21h Durations of instars 2 through 9 are shownin Table 5 The average time required to ma-ture from egg to adulthood was 21797 daysThere were overall significant differences inthe duration of instars (F 3074 P 001)Longest duration was observed for instars 6and 7 and the shortest for instars 3 and 4 Al-though adult females lived significantly longerthan males (t 2587 P 001) there was nosignificant difference in overall average life-span from instar 1 through adulthood for fe-males and males (t 381 P 005)

A summary of pertinent life history data forP sierra is shown in Table 6 Female mass andsize were positively correlated (r 2 00705 P

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

314 vol 51 no 3The Southwestern Naturalist

TABLE 1mdashNumber and percent of prey itemsfound in chelicerae of 65 adult Pardosa sierra col-lected at Cave Creek Canyon Cochise County Ari-zona from 19 March to 23 July 2004 No significantdifferences were found between sexes (chi-square P 005) Data listed as the number (n) of spiders(total 65 37 males 28 females) and percent ()of all spiders found with a particular prey item Lifecycle stage of prey A adult I immature

Prey n

Insecta

Collembola (springtails) (A) 6 92Coleoptera (beetles)

Carabidae (A) 4 61Tenebrionidae (A) 2 31Unidentified (A) 6 92

Diplura (diplurans) (A) 3 46Diptera

Asilidae (robber flies) (A) 1 15Tephritidae (fruitflies) (A) 2 31Drosophilidae (vinegarflies) (A) 5 76Unidentified (A) 2 31

Hemiptera (bugs) (I) 2 31Hymenoptera

Formicidae (ants) (A) 8 123Isoptera (A) 3 46Lepidoptera

Heterocera (moths)Unidentified (I) 3 46

Rhopalocena (butterflies)Hesperiidae (I) 1 15

Unidentified (I) 2 31Orthoptera

Acrididae (grasshoppers) (I) 4 61Unidentified (I) 2 31

Thysanoptera (thrips) (A) 1 15

ArachnidaAraneae (spiders)

Araneidae (I) 3 46Lycosidae (I) 2 31Unidentified (I) 1 15

Opoliones (harvestmen) (A) 2 31Total prey items 65

show that immatures and adults prefer differ-ent microhabitats

Data for habitat preference for P sierra atnight are shown in Table 3 Again adults andimmatures showed a marked preference forrocky and sandy substrates respectively Wefound 57 of all adults on rocks or streamsiderubble compared to 118 on sand (2 1494 P 005) For immatures 47 were on

sandy substrate compared to 96 on rockysubstrates (2 1298 P 005)

No individuals were observed to climb offthe ground and into bushes shrubs or treesAt night 155 of immatures were under leaflitter or other surface plant debris comparedto only 39 of adults (2 1104 P 005)In contrast during daylight fewer immatures(57) and adults (16) were found underlitter

Gestation Period Growth Rate Clutch Size Ob-servation of 35 egg sacs from day of sac con-struction to emergence of spiderlings showedthat gestation period under controlled labora-tory conditions ranged from 22 to 27 days(mean 2406 177 SE) Clutch size rangedfrom 35 to 92 (mean 6781 681 SE)

The surface of the cephalothorax of newlyemerged hatchlings referred to as first post-embryos (first instar) (Vachon 1957) had ayellow-orange color the rest of the embryo wascolorless Upon hatching first post-embryoswere still covered by a thin membrane After aperiod of pulsating movements of legs and ab-domen lasting from 44 to 82 sec (mean 6403 972 SE) this membrane split andlegs cephalothorax and abdomen were pulledfree followed by the mouth and spinnerets Af-ter this first molt spiderlings are now referredto as second post-embryos (second instar)(Foelix 1996)

Growth rate as measured by carapace widthis shown in Table 4 Under controlled labora-tory conditions all P sierra males and femalesattained adulthood by the ninth instar Imma-ture males and females doubled their size bythe sixth instar

Duration of first instar ranged from 14 to 21h Durations of instars 2 through 9 are shownin Table 5 The average time required to ma-ture from egg to adulthood was 21797 daysThere were overall significant differences inthe duration of instars (F 3074 P 001)Longest duration was observed for instars 6and 7 and the shortest for instars 3 and 4 Al-though adult females lived significantly longerthan males (t 2587 P 001) there was nosignificant difference in overall average life-span from instar 1 through adulthood for fe-males and males (t 381 P 005)

A summary of pertinent life history data forP sierra is shown in Table 6 Female mass andsize were positively correlated (r 2 00705 P

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

September 2006 315Punzo and FarmermdashLife history of Pardosa sierra

TABLE 2mdashMicrohabitats used by Pardosa sierra during daylight (0730 to 1600 h) Data based on a sampleof 714 adults (carapace width 18 to 22 mm) and 761 immatures (carapace width 08 to 16 mm) during2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standardized index for microhabitatniche breadth

MicrohabitatNumberof adults

Percentof adults

Number ofimmatures

Percent ofimmatures

Rock sun 211 295 22 28Rock shade 342 478 32 42Sand sun 36 50 236 310Sand shade 19 26 304 399Grass sun 11 15 27 35Grass shade 24 33 19 24Bush sun 3 04 21 27Bush shade 9 12 30 39Tree sun 0 0 5 06Tree shade 16 22 2 02On leaf litterplant debris 31 43 19 24Under litterdebris 12 16 44 57In busheson trees 0 0Total 714 761BA 02146 02404

TABLE 3mdashMicrohabitats used by Pardosa sierra during night (2100 to 0400 h) Data based on a sample of202 adult (A) and subadult (SA) spiders (carapace width 18 to 22 mm) and 187 immatures (carapacewidth 08 to 16 mm) during 2004 at Cave Creek Canyon Cochise County Arizona BA Levinrsquos standard-ized index for microhabitat niche breadth

MicrohabitatNumber ofA and SA

Percent ofA and SA

Number ofimmatures

Percent ofimmatures

Rock 117 579 18 96Sand 24 118 88 471In grass 9 44 15 80Under bush 13 64 21 112Under tree 5 24 9 48On leaf litterplant debris 26 128 7 37Under litterdebris 8 39 29 155In busheson tree 0 0 0 0Total 202 187BA 03749 03607

001) as were offspring mass and size (r 2 00799 P 001) Larger females producedlarger clutches (r 2 00811 P 001) andheavier clutches (r 2 06278 P 005) Therewas no correlation between clutch size and off-spring size or size at hatching and develop-ment time (P 060) Mean TCM was 349of mean female mass

DISCUSSION This paper provides the firstdetailed information on the life cycle and ecol-ogy of P sierra for a population outside of Cal-

ifornia (Cave Creek Canyon Arizona) andfrom a higher elevation Comparisons of vari-ous life cycle parameters for P sierra from these2 populations are summarized in Table 7 (sta-tistical comparisons based on summary infor-mation) Adult size was similar in both popu-lations and values were in agreement with therange of body sizes reported by Banks (1898)for P sierra from Baja California and northernMexico

Despite similarities in body size femalesfrom the higher-elevation Cave Creek Canyon

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

316 vol 51 no 3The Southwestern Naturalist

TABLE 4mdashGrowth rate measured as width of carapace (in mm) for male and female instars of Pardosasierra Values expressed as means ( SE) Values in columns followed by a different letter are statisticallysignificant (Scheffe tests repeated measures ANOVA P 005)

Instar Males Females

Mean carapace width

Males FemalesRange forboth sexes

1 181 133 064a (003) 072a (004) 058 to 0752 213 166 078a (007) 081a (004) 073 to 0863 145 107 096b (017) 098b (011) 092 to 1044 128 143 119b (011) 122b (016) 112 to 1295 74 59 148c (024) 151c (009) 138 to 1586 91 77 158c (016) 159c (005) 151 to 1647 104 74 191d (027) 208d (021) 182 to 2198 67 85 216d (009) 228d (022) 199 to 2419 121 117 221d (018) 231d (026) 213 to 239

TABLE 5mdashDuration (in days) of instars for Pardosa sierra reared under controlled laboratory conditionsValues for duration expressed as means ( SE) and range Values in columns followed by a different letterare statistically significant (Scheffe test P 005)

Instar

Males

n Duration

Females

n Duration

2 168 2565a (303) 21 to 29 179 2616a (288) 22 to 283 132 1602b (283) 14 to 19 84 1715b (314) 15 to 214 155 1745b (299) 15 to 20 109 1794b (187) 16 to 195 77 2587a (209) 23 to 26 58 2499a (287) 22 to 276 129 3266c (498) 28 to 35 113 3307c (402) 30 to 367 118 3305c (389) 30 to 35 74 3292c (501) 29 to 368 108 2566a (388) 22 to 28 87 2588a (451) 23 to 289 120 1755b (244) 11 to 23 91 3232c (419) 18 to 42

population exhibited a significantly smallerclutch size than females from Mill Creek Cal-ifornia This might be associated with coolerambient temperatures that characterize higherelevations in the Chiricahua Mountains In ec-totherms cooler temperatures are often asso-ciated with lower basal metabolic rate result-ing in a decrease in energy allocation towardreproduction (Punzo and Olson 2005) A sim-ilar trend toward lower clutch sizes in popula-tions that inhabit higher elevations has beenreported for some species of lizards (Vitt1981) insects (Skinner 1985) solifuges (Pun-zo 1998) and other species of spiders (Punzo1991a) In contrast for some species there iseither no significant effect of elevation onclutch size (Godfray et al 1991) or popula-tions from higher elevations have a higher fe-cundity (Roff 2002) However clutch size inspiders can be affected by caloric intake (Kes-

sler 1971) and nutritional quality of foods(Punzo and Henderson 1999) Future studiesshould analyze caloric value and nutrient con-tent of common prey species from Mill Creekand Cave Creek Canyon to determine if thereare any significant differences in the percentcomposition of fat protein ash and water aswell as vitamin content of prey items

Although temperature relative humidityand photoperiod regimes were not specifiedVan Dyke and Lowrie (1975) reported a rangeof 7 to 9 instars to reach adulthood under lab-oratory conditions for P sierra from Mill CreekIn contrast all spiders derived from the CaveCreek Canyon population required 9 instars toattain adulthood Numerous species of wolfspiders including those in Lycosa (Miyashita1969) Pirata (Dondale 1961 Brown et al2003 Hendricks and Maelfait 2003) Pardosa(Eason 1969 Buddle 2000) Hogna (Gertsch

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

September 2006 317Punzo and FarmermdashLife history of Pardosa sierra

TABLE 6mdashSummary of life history data for Pardosa sierra from Cave Creek Canyon Cochise County Ari-zona Data obtained from laboratory-reared spiders

Parameter n Range Mean (SE)

Male mass adult (mg) 74 4442 to 4905 4667 (301)Male carapace width (mm) 195 204 to 231 221 (019)Female mass adult (mg) 155 4956 to 5493 5274 (296)Female carapace width (mm) 187 218 to 255 231 (022)Offspring mass (mg) 301 025 to 031 027 (003)Offspring carapace width (mm) 228 075 to 082 079 (003)Clutch sizea 117 35 to 92 6781 (681)Total clutch mass (mg) 117 1683 to 2003 1844 (187)Relative clutch mass 032 to 038 035 (008)

a Data for first egg sac produced by females clutch size number of spiderlingsegg sac

TABLE 7mdashComparison of some life cycle parameters for Pardosa sierra from Mill Creek California (ele-vation 875 m) and Cave Creek Canyon Arizona (1790 m) Data for Mill Creek population taken from VanDyke and Lowrie (1975) Values expressed as range with means in parentheses Values in rows with differentletters are statistically significant (t-test P 005)

Parameter Cave Creek Canyon Mill Creek

Adult carapace width (mm)Males 204 to 231 (221) 21 to 23 (22)Females 218 to 255 (231) 22 to 26 (24)

Clutch size 35 to 92 (678)a 44 to 107 (786)bGestation period (days) 22 to 27 (2406)a 18 to 22 (203)bNumber of instars 9 9Instar at which adulthood was attained 9 7 to 9Time to mature from egg to adulthood (days) 194 to 231 (21797)a 150 to 200 (1814)bPeak abundance (immatures) July to October September to NovemberPeak abundance (adults) April to May February to MarchAdult males first appear mid February DecemberAdult females first appear late January JanuaryLife span (both sexes days) 194 to 211 (2031) 178 to 210 (1974)Females first observed carrying egg sacs early April late FebruaryMicrohabitat preference

Immatures sand substrate sand substrateAdults rocky substrate rocky substrate

1979 Cloudsley-Thompson 1995 Punzo2003) and Schizocosa (Dondale 1961 Whit-comb 1967) have been shown to have 7 to 10instars Furthermore the instar at which adult-hood is attained can vary according to nutri-tional status and ambient temperature (Punzo1991b Foelix 1996)

Immatures and adults from Mill Creek andCave Creek Canyon populations showed astrong preference for sandy and rocky sub-strates respectively with little or no vegetationIndividuals of P sierra have long legs and ex-hibit faster running speeds than other species

of Pardosa (Van Dyke and Lowrie 1975) a traitthat would be advantageous especially onopen sandy areas with little vegetation and few-er shelter sites When disturbed spiders onsand quickly fled until they found refuge un-der a rock or plant debris Spiders on rockysubstrates sought shelter under rocks

Van Dyke and Lowrie (1975) did not studydiet composition of spiders from the MillCreek population However in the laboratoryP sierra fed on a variety of Diptera Hymenop-tera and Hemiptera In this study P sierra wasa generalist predator feeding on a variety of

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

318 vol 51 no 3The Southwestern Naturalist

ground-dwelling insects and arachnids Al-though most spiders were located on opensandy or rocky substrates with sparse plant cov-er most of those found with a prey item werelocated on or under shrubs or plant debrisOne possible explanation for this is that thesespiders move to a different microhabitat(patch) when hunting This has been reportedfor other species of cursorial spiders (Gertsch1979 Punzo 2000) Another explanation isthat once a prey item has been captured thesespiders move to sites where there might bemore shade or cover if rocks are unavailable

Peak abundance for immatures from MillCreek occurred later in the season from Sep-tember through November (Van Dyke andLowrie 1975) whereas for P sierra immaturesfrom Cave Creek Canyon peak abundance oc-curred as early as July This spider overwintersin the fourth or fifth instar at Cave Creek Can-yon Because immatures of P sierra from MillCreek do not appear until later in the seasonmost overwinter in the third instar Adultsfrom Mill Creek were most abundant earlier inthe season from February through March ascompared to April through May at Cave CreekCanyon Adult males and females from MillCreek were first observed in December andJanuary respectively (Van Dyke and Lowrie1975) as compared to mid February and lateJanuary respectively for the Cave Creek Can-yon population

In conclusion immatures of P sierra fromCave Creek Canyon in Arizona exhibit peakabundance in late summer and early fall Mostadult activity occurs in April and May Femalesare most likely to be found carrying egg sacsfrom early April through early August Thesespiders are diurnal and nocturnal in their dielperiodicity and they feed primarily on beetlesflies and ants Immatures and adults prefersandy or rocky substrates respectively with few-er individuals inhabiting leaf litter or decayinglogs Clutch size ranges from 35 to 92 with agestation period of 22 to 27 days

We thank L Ludwig T Ferraioli J Bottrell andL Summers for assistance in collecting and observ-ing animals in the field L Ludwig and M Parkerfor assistance in rearing animals in the laboratoryB Garman for consultation on statistical analysesand C Bradford and anonymous reviewers for com-ments on an earlier draft of the manuscript This

research was supported by a Faculty Research Grantto FP from the University of Tampa Field studieswere conducted with the permission and assistanceof D Bennett (US Forest Service Coronado Na-tional Forest Permit DOU0104) and the ArizonaGame and Fish Department Phoenix (Permit24204)

LITERATURE CITED

BANKS N 1898 Arachnida from Baja California andother parts of Mexico Proceedings of the Cali-fornia Academy of Science 1273ndash306

BROWN C A B M SANFORD AND R SWERDON 2003Clutch size and offspring size in the wolf spiderPirata sedentarius (Araneae Lycosidae) Journalof Arachnology 31285ndash296

BUDDLE C M 2000 Life history of Pardosa moestaand Pardosa mackenziana (Araneae Lycosidae) inCentral Alberta Canada Journal of Arachnology28319ndash328

CANARD A 1987 Analyse nouvelle de developpe-ment postembryonnaire des araignees Revue Ar-achnologique 791ndash106

CLOUDSLEY-THOMPSON J L 1995 Ecophysiology ofdesert reptiles and arthropods Springer Heidel-berg Germany

DONDALE C D 1961 Life histories of some commonspiders from trees and shrubs in Nova Scotia Ca-nadian Journal of Zoology 39777ndash787

EASON R R 1969 Life history and behavior of Par-dosa lapidicina Emerton (Araneae Lycosidae)Journal of the Kansas Entomological Society 42339ndash360

ENDERS F 1976 Clutch size related to hunting man-ner of spider species Annals of the Entomologi-cal Society of America 69991ndash998

FOELIX R F 1996 The biology of spiders OxfordUniversity Press New York

GERTSCH W J 1979 American spiders second edi-tion Van Nostrand New York

GODFRAY H C J L PARTRIDGE AND P H HARVEY1991 Clutch size Annual Review of Ecology andSystematics 22409ndash429

HAGSTROM D 1971 Carapace width as a tool forevaluating rate of development of spiders in thelaboratory and field Annals of the Entomologi-cal Society of America 64757ndash760

HENDRICKS F AND J MAELFAIT 2003 Life cycle re-productive patterns and their year-to-year varia-tion in a field population of the wolf spider Piratapiraticus (Araneae Lycosidae) Journal of Arach-nology 31331ndash339

KESSLER A 1971 Relationship between egg produc-tion and food consumption in species of the ge-nus Pardosa (Lycosidae Araneae) under experi-mental conditions of food abundance and foodshortage Oecologia 893ndash109

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook

September 2006 319Punzo and FarmermdashLife history of Pardosa sierra

KREBS C J 1989 Ecological methodology Harperand Row New York

MIYASHITA K 1969 Seasonal changes of populationdensity and some characteristics of overwinteringnymph of Lycosa t-insignita (Araneae Lycosidae)Applied Entomology and Zoology 41ndash8

PUNZO F 1991a Intraspecific variation to thermalstress in the tarantula Dugesiella echina Chamber-lin (Orthognatha Theraphosidae) Bulletin ofthe British Arachnological Society 8277ndash283

PUNZO F 1991b The effects of temperature andmoisture on survival capacity cuticular perme-ability hemolymph osmoregulation and metab-olism in Centruoides hentzi (Banks) (ScorpionesButhidae) Comparative Biochemistry and Physi-ology 100A833ndash837

PUNZO F 1998 The biology of camel-spiders (Arach-nida Solifugae) Kluwer Academic PublishersNorwell Massachusetts

PUNZO F 2000 Desert arthropods life history vari-ations Springer Heidelberg Germany

PUNZO F 2003 Observations on the natural historyand ecology of the wolf spider Hogna carolinensisWalckenaer (Araneae Lycosidae) in the north-ern Chihuahuan Desert Bulletin of the BritishArachnological Society 12399ndash405

PUNZO F AND L HENDERSON 1999 Aspects of thenatural history and behavioural ecology of thetarantula spider Aphonopelma hentzi (Orthogna-tha Theraphosidae) Bulletin of the BritishArachnological Society 11121ndash128

PUNZO F AND S OLSON 2005 Temperature and wa-

ter relations of the giant whipscorpion Mastigo-proctus giganteus (Lucas) (Arachnida Uropygi)Bulletin of the British Arachnological Society 13206ndash212

ROFF D A 1992 The evolution of life historiesChapman and Hall New York

ROFF D A 2002 Life history evolution Sinauer As-sociates Sunderland Massachusetts

SKINNER S W 1985 Clutch size as an optimal for-aging problem for insects Behavioral Ecologyand Sociobiology 17231ndash238

SOKAL B F AND F J ROHLF 1995 Biometry thirdedition W H Freeman New York

VACHON M 1957 Contribution a Lrsquoetude du devel-oppement post-embryonnaire des araignees Pre-miere note genera lites et nomencalture desstades Bulletin Societe Zoologique Francais 82337ndash354

VAN DYKE D AND D C LOWRIE 1975 Comparativelife histories of the wolf spiders Pardosa ramulosaand P sierra (Araneae Lycosidae) SouthwesternNaturalist 2029ndash44

VITT L J 1981 Lizard reproduction habitat speci-ficity and constraints on relative clutch massAmerican Naturalist 117506ndash514

WHITCOMB W H 1967 Wolf and lynx spider lifehistories Terminal Report for National ScienceFoundation University of Arkansas Departmentof Entomology Pages 35ndash39 42ndash56

Submitted 7 October 2005 Accepted 11 January 2006Associate Editor was Jerry Cook