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

The amount of time female pumas Puma concolor spend with theirkittensAuthor(s) John W Laundreacute and Lucina HernaacutendezSource Wildlife Biology 14(2)221-227 2008Published By Nordic Board for Wildlife ResearchDOI httpdxdoiorg1029810909-6396(2008)14[221TAOTFP]20CO2URL httpwwwbiooneorgdoifull1029810909-639628200829145B2213ATAOTFP5D20CO3B2

BioOne (wwwbiooneorg) is a nonprofit online aggregation of core research in the biologicalecological and environmental sciences BioOne provides a sustainable online platform for over170 journals and books published by nonprofit societies associations museums institutions andpresses

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

Usage of BioOne content is strictly limited to personal educational and non-commercial useCommercial inquiries or rights and permissions requests should be directed to the individualpublisher as copyright holder

The amount of time female pumas Puma concolor spend withtheir kittens

John W Laundre amp Lucina Hernandez

Laundre JW ampHernandez L 2008 The amount of time female pumasPuma concolor spend with their kittens - Wildl Biol 14 221-227

In the sport hunting of pumas Puma concolor most states and prov-

inces of the United States and Canada do not allow the killing of females

with kittens However female pumas can be away from their kittens

and if detected by hunters at these times can be mistakenly killed

To assess the extent to which females with kittens might mistakenly be

killed we need to have estimates of the percent of time female pumas

are with their kittens on a daily basis Previous estimates range within

52-83 but are based on simultaneous locations taken during the day

when pumas are least active To provide a more accurate assessment of

the amount of time females spend with their young we analyzed tele-

metry data collected over 24 24-hour blocks for 15 females and their

kittens We collected data from June to September during 1989-1999 in

southeastern Idaho and northwestern Utah We found that females

with 7-12 month-old kittens were within 200 m of their kittens an aver-

age 162iexcl38 (N=12) of the time These females were gt10 km from

their kittens 309iexcl67 of the time Three females with kittens in dens

were near their dens 103 122 and 23 of the time Females were

within 200 m of their kittens the least amount of time (51iexcl21

N=8) during 1100-1430 and the most amount of time during 2300-

0130 (294iexcl30 N=6) and 0700-1030 (232iexcl31) We conclude

that the probability that a hunter would encounter a female without her

kittens wasgt80

Key words dependent kittens females Idaho pumas time together USAUtah

John W Laundre amp Lucina Hernandez Instituto de Ecologıa AC Du-rango Regional Center Blvd Guardiana 123 34100 Durango DurangoMexico - e-mail addresses launjohnhotmailcom (John W Laundre)hernlucihotmailcom (Lucina Hernandez)

Corresponding author John W Laundre

Received 10 October 2005 accepted 27 May 2007

Associate Editor Urs Breitenmoser

In most western states and provinces of the UnitedStates and Canada sport hunting for pumas Pumaconcolor occurs The hunting seasons usually lastfromfall toearly springandvary in length fromstateto state To hunt pumas hunters use trained dogs to

chase and eventually corner them usually up a treeIn most states hunters rely on snow to find freshtracks of pumas that their dogs can follow To findthese tracks hunters travel the roads and trails intothe mountains early in the morning looking for

WILDLIFE BIOLOGY 142 (2008) 221

tracksmade by pumas during the previous night Inall states hunters are allowed to kill female pumaswithout dependent kittens (approximately j13-15months old Logan amp Sweanor 2001) Femalepumas can reproduce at any time of the year andkittens can stay with their mothers for up to 17-18months Thus during the hunting season there willbe females with kittens ranging in age from birth toalmost independence It is the responsibility of thesport hunter to discern if a female has kittensMosttimes hunters should be able to distinguish betweenmale and female pumas based on track size orwhenthe puma is in the tree on the presence of testesHowever a hunter can only determine if a female iswith kittens if they travel with her and leave tracksor if on occasion their dogs chase a kitten up a treenear or with its mother (JW Laundre pers obs)The problem arises in that kittens are not alwayswith their mothers When a hunter finds a singlefemale track crossing the road hedoes notknow forsure that she does not have kittens If he sees noevidence of kittens during the chase he can mis-takenlykillafemalewhohaskittensbutwasnotwiththemat the timeof the chase If orphanedkittens arelt6months old they will probably die of starvation(Cougar Management Guidelines Working Group2005)Because the loss of such orphaned individuals

can possibly affect recruitment into the populationit is desirable to determine how likely it is to en-counter a female without her kittens We can thenestimate the probability that a hunter will kill afemale puma with undetected kittens Others haveestimated the percent time females spend with theiryoung based on simultaneous radiotelemetry loca-tions (BarnhurstampLindzey 1989Maehr et al 1989Logan amp Sweanor 2001) their estimates rangewithin 53-82 However all of these studies reliedon locations taken once or twice a day or on pointlocations (LaundreampKeller 1984) usuallyobtainedduring daytime hours and mostly from airplaneFemales are more likely to be with their young atthese times (Barnhurst amp Lindzey 1989) and re-location data may overestimate the time femalesspendwith their kittens especially at timeswhen thefemales are active and more likely to cross roadsleaving tracks for the hunters to find The fact thatBarnhurstampLindzey (1989) foundkittenswith theirmothers only 19-43 of the time based on snow-tracking data seems to support this observationSnow-tracking data however may tend to under-estimate the time females spend with their kittens

especially kittens in dens because we cannot es-timate the time females and kittens are togetherbased on tracks Thus we are still uncertain aboutthe percent time a female spends with and withouther kittens over the course of 24 hours Such in-formation would be helpful in evaluating the pos-sible risk of a hunter mistakenly killing a femalepuma with kittens

During 1989-1999we intenselymonitored radio-collared pumas over 24-hour time blocks Duringthis time we had the opportunity to monitor 15females simultaneously with their kittens for alto-gether 24 24-hour sessions In this paper we pre-sent the analysis of these data relative to the per-cent time these females were near their kittens overthe 24 hours This analysis should provide us withnew information regarding the spatial and temporalaffinity between female pumas and their kittens

Methods

Study area

Our study area was in south-central Idaho andnorthwestern Utah USA (Fig 1) We chose thestudy area because it was representative of the typeof mountain physiognomy and habitat structurecharacteristic of this region The total area of2400 km2 contained approximately 1700 km2 ofpumahabitatwithinfive small semi-isolatedmoun-tain ranges (of 65-760 km2)with elevationsof 1830-3151 m asl (see Fig 1) A variety of roads andtrails made most parts of the mountains accessiblevia motorized vehicles which facilitated our tele-metry efforts Mule deer Odocoileus hemionus werethe principal prey species of pumas with only aremnant (lt50)elkCervuselaphuspopulationOtherspecies pumas occasionally preyed on during thestudyperiodincludedcoyotesCanis latransbobcatsLynx rufus and porcupines Erethizon dorsatum

Mountain ranges were internally fragmented in-to open and forested habitat patches that varied insize Forested patches consisted of various mixesof Douglas fir Pseudotsuga menziesii subalpine firAbies lasiocarpa juniper Juniperus osteosperma andJ scopulorum pinyon pine Pinus edulis quakingaspen Populus tremuloides and curl-leaf mountainmahoganyCercocarpus ledifoliusDominant shrubsin open areas included big sagebrush Artemisiatridentata gray rabbitbrush Chrysothamnus nause-osus bitterbrush Purshia tridentata and buffalo-berryShepherdia rotundifoliaThe climate ishotand

222 WILDLIFE BIOLOGY 142 (2008)

dry in the summers (20-35xC) and cold andwindy inthewinters (-25-4xC)Humidity rarely exceeds 40andprecipitation is sporadicwithanannual averageof 300 mm

Field procedures

We captured pumas primarily in thewinter with theaidof traineddogsAsdidotherhunterswecheckedroadsandtrails early in themorningto lookfor freshpuma tracks made the night before Once we foundfresh tracks we released the dogs and when theycornered a puma in a tree we tranquilized it with amixture of Ketamine hydrochloride (9 mgkg) andXylazine hydrochloride (15 mgkg) administeredintramuscularly via a dart gun We lowered eachtranquilized animal from the tree with a rope at-tached to a legWe attached a radio-collar (WildlifeMaterials Inc Murphysboro Illinois USA) toeach animal and then administered an antidote ofYohimbine in equal quantity to the XylazinehydrochlorideResearchers stayedwith the animalsuntil they recovered sufficiently For kittens weradio-collared only individualsi6months oldWe

recapturedandchangedthecollarsas neededOur capture procedureswereapprovedby theAnimalWel-fare Committee of Idaho StateUniversity

During June-September weused triangulation to monitor 24-hour movements of collared pu-mas Once we located the collaredanimalswewere to followwe stra-tegically placed two monitoringstations so as to optimize the diffe-rence between compass bearingsWedetermined the locations of thestations from USGS topographicmaps and handheld GPS units Ateach station we used a 4-elementdouble yagi null antenna systemWedetermined the direction of theantenna elements with a compassrosemounted on the antenna poleWe adjusted the rose to magneticnorth with a handheld compassand periodically checked this ad-justment during the 24 hoursTrained Earthwatch Institute vol-unteers took simultaneous read-ings each half hour from each sta-tion To insure data accuracy at

least onemember of the research staffwas present atall times and periodically checked the work of thevolunteers In addition to the telemetry stationsapproximately4-5 timesover the24hoursaresearchstaffmember collected a third compass bearingwitharooftop-mounteddoubleyaginullantennasystemFor these third bearings the staff member wouldpositionthevehicleatalocationdeterminedfromtheUSGSmaps or a handheldGPS unitWe combinedthese third bearings with the two taken simulta-neouslyatthestationsandwiththeLOCATEII(ver182)program(VNamsvnamsnsacnsca)estim-ated the animalrsquos most likely position by the max-imumlikelihoodestimatormethod(Lenth1981)Toestimate the triangulation error of our stations wemeasured thedistance fromtheLOCATEestimatedlocationtotheonebasedonlyonthetwomonitoringstationsWeusedtheaverageofthesemeasurementsas an estimate of our telemetry error for locationsbased only on the two bearings from the telemetrystations Pumas were normally in the mountains(Blum 2003) and our stations were usually gt1 kmfromtheanimalswewere locatingThusweassumed

Figure 1 Location of our study area in southern Idaho and northwestern Utah Thestudy area consisted of five small mountains (of 647-760 km2 gt2000 m asl)separatedbyflatvalleyterrainRoadsallowingaccess tothemountainsare indicatedbythedashedandsolid linesSmall towns(with5-200 inhabitants) in theareaare indicatedby the black dots The topographic lines begin at 2000 m asl and are indicated forapproximately every 300 m

WILDLIFE BIOLOGY 142 (2008) 223

aminimal impactofourpresenceontheirbehaviourConverselybecauseof the sizeof themountainsandthe accessibility to the valleys within the mountainranges we were rarely gt5 km from the animals wemonitored and were able to monitor most pumasover the 24hourswithoutmoving stationsAlsoweassumed minimal signal error because reception ofthe signal was predominately line of sight with littleproblems with signal bounceForeach24-hour sessionweentered thedata into

a program we developed to estimate the location ofthe animal at half-hour intervals We then matchedup the simultaneous locations of female pumaswiththeir kittens in an Excel worksheet and determinedthe female-kitten distance for each half-hour lo-cationWhentherewasgt1radio-collaredkittenwecalculated the distance the female was from eachkitten and then averaged these distances Once wecalculated the female-kitten distances for each halfhour we then counted the number of locationswhere the female was near her kittens Because oftelemetry error it is possible to misclassify a femaleas being away from or near her kittens when indeedshe was the opposite To reduce these errors we in-corporated our telemetry error (180iexcl187 m N=129) and assumed that if a femalersquos estimated posi-tion was within 200 m of the estimated locations ofher kittens they were probably together We alsocounted the number of locations where the femalewasgt10 km from her kittens to facilitate compar-isons with other studies We then expressed thesevalues as percents of the total number of locationsfor the 24-hour session Our sample unit was theindividual animal so for all our calculations weaveraged the sessions of females for which we hadmore than one 24-hour session to avoid pseudo-replicationWe used Sigmastat software (Systat Software

Inc Point Richmond CA USA) for all statisticalanalyses and reported allmeans asiexclstandard error

Results

Over the studyperiod our research teammonitored15adult pumaswithkittens for 24 complete 24-hoursessions For 12 of the females we were able tosimultaneouslymonitor their radio-collared kittensduring 21 monitoring sessions We had singlesessions for nine females and three four and sixsessions for theother threeThekittensranged inagewithin 7-12 months We were also able to monitor

one time each three other females whowe knew hadrecently born kittens (lt1 month old) at known densites

The distance females were from their kittensranged within 00-38 km over the 24 hours (Fig2A) The percent time females were j200 m fromtheir kittens over the 24 hours averaged 162iexcl38(N=12) Conversely females were gt10 km fromtheir kittens 309iexcl67 of the time The three fe-males with kittens in dens were near their dens 103122 and 23 of the time

Relative to time of day females were usuallyi10 km from their kittens during 1100-1600Theywere within 06-08 kmduringmost of the restof the time (see Fig 2A) There were three periodsover the 24 hours when females werej200 m fromtheir kittens an average 23-29 of the time 1500-1730 (257iexcl57 N=5) 2300-0130 (294iexcl30 N=6) and 0700-1030 (232iexcl31 N=8

Figure 2 Average distance (in km) 12 female pumas were fromtheir 7-12month-old kittens over the course of 24 hours (A) andaverage percent time these 12 females were j200 m of theirkittens (B)Averagesamplesize foreachhalf-hoursamplewas98female-kitten pairs The average standard error for the half-hourly distances between females and kittens was 02iexcl001 Therange of sunset and sunrise times over the study period is in-dicated

224 WILDLIFE BIOLOGY 142 (2008)

see Fig 2B) During 1100-1430 females were neartheir kittens the least amount of time (51iexcl21N=8 see Fig 2B)

Discussion

Basedonsimultaneous telemetry locationscollectedat varying time intervals over the year four femalesand their 7-12 month-old kittens in southern Utahwere together69ofthe time (BarnhurstampLindzey1989) Three Florida panther Puma concolor coryifemales with dependent kittens (no ages werespecified) were within 05 km of each other 45of the time andweregt10 kmapart 47of the time(Maehr et al 1991) In contrast in New Mexicokittens gt6 months old were with their mothers82 of the time (Logan amp Sweanor 2001) Thesetelemetry-based estimates are substantially higherthan the 162 we found for the 12 females within200 m of their kittens Barnhurst amp Lindzey (1989)and Logan amp Sweanor (2001) observed that takingdaytime telemetry locations when pumas are in-active (Beier et al 1995) probably overestimated thepercent time females and kittens were together Wedidfind that thepercent time femaleswerewith theirkittenschangedthroughthedaytimewithoneof thehighest average percentages being early morning(0700-1030) If telemetry flights in these studieswere in the early morning (when the air is moststable) then the estimates of Barnhurst amp Lindzey(1989) and Logan amp Sweanor (2001) were probablyoverestimates of the total time females and kittensare together However given the variability wefound in the amount of time they were close to theirkittens (0-420) some of the differences could bedue to the small sample sizes (3-4 females) of theseprevious studiesIn contrast to their telemetry based estimates

BarnhurstampLindzey (1989) found femalesand their7-12 month-old kittens together 43 of the timebasedonsnowtracksBecauseBarnhurstampLindzey(1989) recorded only tracks crossing roads theycontended that this estimate should be the mostrealistic to the percent of time hunters using thesametechnique shouldencounter femaleswiththeirkittens Their estimate however was still higherthan the 168 we found via telemetry Againbecause of the variability we found among days andanimals the higher percent of Barnhurst ampLindzey(1989) couldbe related to their limited sample sizeoftwo animals Another possibility is that the percent

time females associate with their kittens differsbetween summer when we collected our data andwinter However we currently lack the data to testthis supposition

Barnhurst amp Lindzey (1989) found that twofemales were together with their 0-6 month-oldkittens for 63 of the telemetry locations Incontrast their snow data indicated that femalestraveled with young kittens 19 of the time whichwas only slightly higher than our telemetry estimateof 23-122 for three females with lt1 month-oldkittensFemales rarelymovetheirkittensawayfromtheir birth sites before they are two months old(Logan amp Sweanor 2001) so females will be alonewhen they travel for at least two of six months afterthe kittens are born Therefore the estimate ofBarnhurst amp Lindzey (1989) from snow tracking isperhaps more applicable for kittens of 3-6 monthswhich we would expect to be higher In all casessample sizes were extremely small (2-4 animals) butconsidering our data and the snow-tracking dataofBarnhurstampLindzey (1989) females appear tobewith their 0-6 month-old kittenslt20 of the time

During the diel cycle females are least likely to benear their young from late morning to mid-after-noon (seeFig 2B) It is during this time that they arealso the furthest away (see Fig 2A) We speculatethat in late morning females separate from theiryoung perhaps to sleepundisturbed It appears thatthey rejoin their young in late afternoon but thenseparateagain towardeveningwhentheybegin tobeactive (Beier et al 1995) Over the nighttime thepatternof percent time females are near their kittensappears similar to the pattern of activity (ie dis-tance moved per hour) Beier et al (1995) reportedand forwhatwe found inour study (Fig 3)Femalesshowed a peak in activity during 1600-2200 (seeFig 3) and during this period we found them to beaway from their young gt80 of the time (see Fig2B) Shortly before midnight females seem to rejointheir kittens which corresponded to the reducedactivity we (see Fig 3) and also Beier et al (1995)noted An hour or so before sunrise we (see Fig 3)and Beier et al (1995) found that pumas increasedtheir activity and again we found that they sepa-rated from their young (see Fig 2B) The cycle com-pletes itself when females rejoin their young aroundsunrise before they separate again in late morningConsequently the duel requirements of caring fortheir youngandactivity primarily hunting appearsto produce a distinct diel pattern of association be-tween females and their young

WILDLIFE BIOLOGY 142 (2008) 225

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

tracksmade by pumas during the previous night Inall states hunters are allowed to kill female pumaswithout dependent kittens (approximately j13-15months old Logan amp Sweanor 2001) Femalepumas can reproduce at any time of the year andkittens can stay with their mothers for up to 17-18months Thus during the hunting season there willbe females with kittens ranging in age from birth toalmost independence It is the responsibility of thesport hunter to discern if a female has kittensMosttimes hunters should be able to distinguish betweenmale and female pumas based on track size orwhenthe puma is in the tree on the presence of testesHowever a hunter can only determine if a female iswith kittens if they travel with her and leave tracksor if on occasion their dogs chase a kitten up a treenear or with its mother (JW Laundre pers obs)The problem arises in that kittens are not alwayswith their mothers When a hunter finds a singlefemale track crossing the road hedoes notknow forsure that she does not have kittens If he sees noevidence of kittens during the chase he can mis-takenlykillafemalewhohaskittensbutwasnotwiththemat the timeof the chase If orphanedkittens arelt6months old they will probably die of starvation(Cougar Management Guidelines Working Group2005)Because the loss of such orphaned individuals

can possibly affect recruitment into the populationit is desirable to determine how likely it is to en-counter a female without her kittens We can thenestimate the probability that a hunter will kill afemale puma with undetected kittens Others haveestimated the percent time females spend with theiryoung based on simultaneous radiotelemetry loca-tions (BarnhurstampLindzey 1989Maehr et al 1989Logan amp Sweanor 2001) their estimates rangewithin 53-82 However all of these studies reliedon locations taken once or twice a day or on pointlocations (LaundreampKeller 1984) usuallyobtainedduring daytime hours and mostly from airplaneFemales are more likely to be with their young atthese times (Barnhurst amp Lindzey 1989) and re-location data may overestimate the time femalesspendwith their kittens especially at timeswhen thefemales are active and more likely to cross roadsleaving tracks for the hunters to find The fact thatBarnhurstampLindzey (1989) foundkittenswith theirmothers only 19-43 of the time based on snow-tracking data seems to support this observationSnow-tracking data however may tend to under-estimate the time females spend with their kittens

especially kittens in dens because we cannot es-timate the time females and kittens are togetherbased on tracks Thus we are still uncertain aboutthe percent time a female spends with and withouther kittens over the course of 24 hours Such in-formation would be helpful in evaluating the pos-sible risk of a hunter mistakenly killing a femalepuma with kittens

During 1989-1999we intenselymonitored radio-collared pumas over 24-hour time blocks Duringthis time we had the opportunity to monitor 15females simultaneously with their kittens for alto-gether 24 24-hour sessions In this paper we pre-sent the analysis of these data relative to the per-cent time these females were near their kittens overthe 24 hours This analysis should provide us withnew information regarding the spatial and temporalaffinity between female pumas and their kittens

Methods

Study area

Our study area was in south-central Idaho andnorthwestern Utah USA (Fig 1) We chose thestudy area because it was representative of the typeof mountain physiognomy and habitat structurecharacteristic of this region The total area of2400 km2 contained approximately 1700 km2 ofpumahabitatwithinfive small semi-isolatedmoun-tain ranges (of 65-760 km2)with elevationsof 1830-3151 m asl (see Fig 1) A variety of roads andtrails made most parts of the mountains accessiblevia motorized vehicles which facilitated our tele-metry efforts Mule deer Odocoileus hemionus werethe principal prey species of pumas with only aremnant (lt50)elkCervuselaphuspopulationOtherspecies pumas occasionally preyed on during thestudyperiodincludedcoyotesCanis latransbobcatsLynx rufus and porcupines Erethizon dorsatum

Mountain ranges were internally fragmented in-to open and forested habitat patches that varied insize Forested patches consisted of various mixesof Douglas fir Pseudotsuga menziesii subalpine firAbies lasiocarpa juniper Juniperus osteosperma andJ scopulorum pinyon pine Pinus edulis quakingaspen Populus tremuloides and curl-leaf mountainmahoganyCercocarpus ledifoliusDominant shrubsin open areas included big sagebrush Artemisiatridentata gray rabbitbrush Chrysothamnus nause-osus bitterbrush Purshia tridentata and buffalo-berryShepherdia rotundifoliaThe climate ishotand

222 WILDLIFE BIOLOGY 142 (2008)

dry in the summers (20-35xC) and cold andwindy inthewinters (-25-4xC)Humidity rarely exceeds 40andprecipitation is sporadicwithanannual averageof 300 mm

Field procedures

We captured pumas primarily in thewinter with theaidof traineddogsAsdidotherhunterswecheckedroadsandtrails early in themorningto lookfor freshpuma tracks made the night before Once we foundfresh tracks we released the dogs and when theycornered a puma in a tree we tranquilized it with amixture of Ketamine hydrochloride (9 mgkg) andXylazine hydrochloride (15 mgkg) administeredintramuscularly via a dart gun We lowered eachtranquilized animal from the tree with a rope at-tached to a legWe attached a radio-collar (WildlifeMaterials Inc Murphysboro Illinois USA) toeach animal and then administered an antidote ofYohimbine in equal quantity to the XylazinehydrochlorideResearchers stayedwith the animalsuntil they recovered sufficiently For kittens weradio-collared only individualsi6months oldWe

recapturedandchangedthecollarsas neededOur capture procedureswereapprovedby theAnimalWel-fare Committee of Idaho StateUniversity

During June-September weused triangulation to monitor 24-hour movements of collared pu-mas Once we located the collaredanimalswewere to followwe stra-tegically placed two monitoringstations so as to optimize the diffe-rence between compass bearingsWedetermined the locations of thestations from USGS topographicmaps and handheld GPS units Ateach station we used a 4-elementdouble yagi null antenna systemWedetermined the direction of theantenna elements with a compassrosemounted on the antenna poleWe adjusted the rose to magneticnorth with a handheld compassand periodically checked this ad-justment during the 24 hoursTrained Earthwatch Institute vol-unteers took simultaneous read-ings each half hour from each sta-tion To insure data accuracy at

least onemember of the research staffwas present atall times and periodically checked the work of thevolunteers In addition to the telemetry stationsapproximately4-5 timesover the24hoursaresearchstaffmember collected a third compass bearingwitharooftop-mounteddoubleyaginullantennasystemFor these third bearings the staff member wouldpositionthevehicleatalocationdeterminedfromtheUSGSmaps or a handheldGPS unitWe combinedthese third bearings with the two taken simulta-neouslyatthestationsandwiththeLOCATEII(ver182)program(VNamsvnamsnsacnsca)estim-ated the animalrsquos most likely position by the max-imumlikelihoodestimatormethod(Lenth1981)Toestimate the triangulation error of our stations wemeasured thedistance fromtheLOCATEestimatedlocationtotheonebasedonlyonthetwomonitoringstationsWeusedtheaverageofthesemeasurementsas an estimate of our telemetry error for locationsbased only on the two bearings from the telemetrystations Pumas were normally in the mountains(Blum 2003) and our stations were usually gt1 kmfromtheanimalswewere locatingThusweassumed

Figure 1 Location of our study area in southern Idaho and northwestern Utah Thestudy area consisted of five small mountains (of 647-760 km2 gt2000 m asl)separatedbyflatvalleyterrainRoadsallowingaccess tothemountainsare indicatedbythedashedandsolid linesSmall towns(with5-200 inhabitants) in theareaare indicatedby the black dots The topographic lines begin at 2000 m asl and are indicated forapproximately every 300 m

WILDLIFE BIOLOGY 142 (2008) 223

aminimal impactofourpresenceontheirbehaviourConverselybecauseof the sizeof themountainsandthe accessibility to the valleys within the mountainranges we were rarely gt5 km from the animals wemonitored and were able to monitor most pumasover the 24hourswithoutmoving stationsAlsoweassumed minimal signal error because reception ofthe signal was predominately line of sight with littleproblems with signal bounceForeach24-hour sessionweentered thedata into

a program we developed to estimate the location ofthe animal at half-hour intervals We then matchedup the simultaneous locations of female pumaswiththeir kittens in an Excel worksheet and determinedthe female-kitten distance for each half-hour lo-cationWhentherewasgt1radio-collaredkittenwecalculated the distance the female was from eachkitten and then averaged these distances Once wecalculated the female-kitten distances for each halfhour we then counted the number of locationswhere the female was near her kittens Because oftelemetry error it is possible to misclassify a femaleas being away from or near her kittens when indeedshe was the opposite To reduce these errors we in-corporated our telemetry error (180iexcl187 m N=129) and assumed that if a femalersquos estimated posi-tion was within 200 m of the estimated locations ofher kittens they were probably together We alsocounted the number of locations where the femalewasgt10 km from her kittens to facilitate compar-isons with other studies We then expressed thesevalues as percents of the total number of locationsfor the 24-hour session Our sample unit was theindividual animal so for all our calculations weaveraged the sessions of females for which we hadmore than one 24-hour session to avoid pseudo-replicationWe used Sigmastat software (Systat Software

Inc Point Richmond CA USA) for all statisticalanalyses and reported allmeans asiexclstandard error

Results

Over the studyperiod our research teammonitored15adult pumaswithkittens for 24 complete 24-hoursessions For 12 of the females we were able tosimultaneouslymonitor their radio-collared kittensduring 21 monitoring sessions We had singlesessions for nine females and three four and sixsessions for theother threeThekittensranged inagewithin 7-12 months We were also able to monitor

one time each three other females whowe knew hadrecently born kittens (lt1 month old) at known densites

The distance females were from their kittensranged within 00-38 km over the 24 hours (Fig2A) The percent time females were j200 m fromtheir kittens over the 24 hours averaged 162iexcl38(N=12) Conversely females were gt10 km fromtheir kittens 309iexcl67 of the time The three fe-males with kittens in dens were near their dens 103122 and 23 of the time

Relative to time of day females were usuallyi10 km from their kittens during 1100-1600Theywere within 06-08 kmduringmost of the restof the time (see Fig 2A) There were three periodsover the 24 hours when females werej200 m fromtheir kittens an average 23-29 of the time 1500-1730 (257iexcl57 N=5) 2300-0130 (294iexcl30 N=6) and 0700-1030 (232iexcl31 N=8

Figure 2 Average distance (in km) 12 female pumas were fromtheir 7-12month-old kittens over the course of 24 hours (A) andaverage percent time these 12 females were j200 m of theirkittens (B)Averagesamplesize foreachhalf-hoursamplewas98female-kitten pairs The average standard error for the half-hourly distances between females and kittens was 02iexcl001 Therange of sunset and sunrise times over the study period is in-dicated

224 WILDLIFE BIOLOGY 142 (2008)

see Fig 2B) During 1100-1430 females were neartheir kittens the least amount of time (51iexcl21N=8 see Fig 2B)

Discussion

Basedonsimultaneous telemetry locationscollectedat varying time intervals over the year four femalesand their 7-12 month-old kittens in southern Utahwere together69ofthe time (BarnhurstampLindzey1989) Three Florida panther Puma concolor coryifemales with dependent kittens (no ages werespecified) were within 05 km of each other 45of the time andweregt10 kmapart 47of the time(Maehr et al 1991) In contrast in New Mexicokittens gt6 months old were with their mothers82 of the time (Logan amp Sweanor 2001) Thesetelemetry-based estimates are substantially higherthan the 162 we found for the 12 females within200 m of their kittens Barnhurst amp Lindzey (1989)and Logan amp Sweanor (2001) observed that takingdaytime telemetry locations when pumas are in-active (Beier et al 1995) probably overestimated thepercent time females and kittens were together Wedidfind that thepercent time femaleswerewith theirkittenschangedthroughthedaytimewithoneof thehighest average percentages being early morning(0700-1030) If telemetry flights in these studieswere in the early morning (when the air is moststable) then the estimates of Barnhurst amp Lindzey(1989) and Logan amp Sweanor (2001) were probablyoverestimates of the total time females and kittensare together However given the variability wefound in the amount of time they were close to theirkittens (0-420) some of the differences could bedue to the small sample sizes (3-4 females) of theseprevious studiesIn contrast to their telemetry based estimates

BarnhurstampLindzey (1989) found femalesand their7-12 month-old kittens together 43 of the timebasedonsnowtracksBecauseBarnhurstampLindzey(1989) recorded only tracks crossing roads theycontended that this estimate should be the mostrealistic to the percent of time hunters using thesametechnique shouldencounter femaleswiththeirkittens Their estimate however was still higherthan the 168 we found via telemetry Againbecause of the variability we found among days andanimals the higher percent of Barnhurst ampLindzey(1989) couldbe related to their limited sample sizeoftwo animals Another possibility is that the percent

time females associate with their kittens differsbetween summer when we collected our data andwinter However we currently lack the data to testthis supposition

Barnhurst amp Lindzey (1989) found that twofemales were together with their 0-6 month-oldkittens for 63 of the telemetry locations Incontrast their snow data indicated that femalestraveled with young kittens 19 of the time whichwas only slightly higher than our telemetry estimateof 23-122 for three females with lt1 month-oldkittensFemales rarelymovetheirkittensawayfromtheir birth sites before they are two months old(Logan amp Sweanor 2001) so females will be alonewhen they travel for at least two of six months afterthe kittens are born Therefore the estimate ofBarnhurst amp Lindzey (1989) from snow tracking isperhaps more applicable for kittens of 3-6 monthswhich we would expect to be higher In all casessample sizes were extremely small (2-4 animals) butconsidering our data and the snow-tracking dataofBarnhurstampLindzey (1989) females appear tobewith their 0-6 month-old kittenslt20 of the time

During the diel cycle females are least likely to benear their young from late morning to mid-after-noon (seeFig 2B) It is during this time that they arealso the furthest away (see Fig 2A) We speculatethat in late morning females separate from theiryoung perhaps to sleepundisturbed It appears thatthey rejoin their young in late afternoon but thenseparateagain towardeveningwhentheybegin tobeactive (Beier et al 1995) Over the nighttime thepatternof percent time females are near their kittensappears similar to the pattern of activity (ie dis-tance moved per hour) Beier et al (1995) reportedand forwhatwe found inour study (Fig 3)Femalesshowed a peak in activity during 1600-2200 (seeFig 3) and during this period we found them to beaway from their young gt80 of the time (see Fig2B) Shortly before midnight females seem to rejointheir kittens which corresponded to the reducedactivity we (see Fig 3) and also Beier et al (1995)noted An hour or so before sunrise we (see Fig 3)and Beier et al (1995) found that pumas increasedtheir activity and again we found that they sepa-rated from their young (see Fig 2B) The cycle com-pletes itself when females rejoin their young aroundsunrise before they separate again in late morningConsequently the duel requirements of caring fortheir youngandactivity primarily hunting appearsto produce a distinct diel pattern of association be-tween females and their young

WILDLIFE BIOLOGY 142 (2008) 225

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

dry in the summers (20-35xC) and cold andwindy inthewinters (-25-4xC)Humidity rarely exceeds 40andprecipitation is sporadicwithanannual averageof 300 mm

Field procedures

We captured pumas primarily in thewinter with theaidof traineddogsAsdidotherhunterswecheckedroadsandtrails early in themorningto lookfor freshpuma tracks made the night before Once we foundfresh tracks we released the dogs and when theycornered a puma in a tree we tranquilized it with amixture of Ketamine hydrochloride (9 mgkg) andXylazine hydrochloride (15 mgkg) administeredintramuscularly via a dart gun We lowered eachtranquilized animal from the tree with a rope at-tached to a legWe attached a radio-collar (WildlifeMaterials Inc Murphysboro Illinois USA) toeach animal and then administered an antidote ofYohimbine in equal quantity to the XylazinehydrochlorideResearchers stayedwith the animalsuntil they recovered sufficiently For kittens weradio-collared only individualsi6months oldWe

recapturedandchangedthecollarsas neededOur capture procedureswereapprovedby theAnimalWel-fare Committee of Idaho StateUniversity

During June-September weused triangulation to monitor 24-hour movements of collared pu-mas Once we located the collaredanimalswewere to followwe stra-tegically placed two monitoringstations so as to optimize the diffe-rence between compass bearingsWedetermined the locations of thestations from USGS topographicmaps and handheld GPS units Ateach station we used a 4-elementdouble yagi null antenna systemWedetermined the direction of theantenna elements with a compassrosemounted on the antenna poleWe adjusted the rose to magneticnorth with a handheld compassand periodically checked this ad-justment during the 24 hoursTrained Earthwatch Institute vol-unteers took simultaneous read-ings each half hour from each sta-tion To insure data accuracy at

least onemember of the research staffwas present atall times and periodically checked the work of thevolunteers In addition to the telemetry stationsapproximately4-5 timesover the24hoursaresearchstaffmember collected a third compass bearingwitharooftop-mounteddoubleyaginullantennasystemFor these third bearings the staff member wouldpositionthevehicleatalocationdeterminedfromtheUSGSmaps or a handheldGPS unitWe combinedthese third bearings with the two taken simulta-neouslyatthestationsandwiththeLOCATEII(ver182)program(VNamsvnamsnsacnsca)estim-ated the animalrsquos most likely position by the max-imumlikelihoodestimatormethod(Lenth1981)Toestimate the triangulation error of our stations wemeasured thedistance fromtheLOCATEestimatedlocationtotheonebasedonlyonthetwomonitoringstationsWeusedtheaverageofthesemeasurementsas an estimate of our telemetry error for locationsbased only on the two bearings from the telemetrystations Pumas were normally in the mountains(Blum 2003) and our stations were usually gt1 kmfromtheanimalswewere locatingThusweassumed

Figure 1 Location of our study area in southern Idaho and northwestern Utah Thestudy area consisted of five small mountains (of 647-760 km2 gt2000 m asl)separatedbyflatvalleyterrainRoadsallowingaccess tothemountainsare indicatedbythedashedandsolid linesSmall towns(with5-200 inhabitants) in theareaare indicatedby the black dots The topographic lines begin at 2000 m asl and are indicated forapproximately every 300 m

WILDLIFE BIOLOGY 142 (2008) 223

aminimal impactofourpresenceontheirbehaviourConverselybecauseof the sizeof themountainsandthe accessibility to the valleys within the mountainranges we were rarely gt5 km from the animals wemonitored and were able to monitor most pumasover the 24hourswithoutmoving stationsAlsoweassumed minimal signal error because reception ofthe signal was predominately line of sight with littleproblems with signal bounceForeach24-hour sessionweentered thedata into

a program we developed to estimate the location ofthe animal at half-hour intervals We then matchedup the simultaneous locations of female pumaswiththeir kittens in an Excel worksheet and determinedthe female-kitten distance for each half-hour lo-cationWhentherewasgt1radio-collaredkittenwecalculated the distance the female was from eachkitten and then averaged these distances Once wecalculated the female-kitten distances for each halfhour we then counted the number of locationswhere the female was near her kittens Because oftelemetry error it is possible to misclassify a femaleas being away from or near her kittens when indeedshe was the opposite To reduce these errors we in-corporated our telemetry error (180iexcl187 m N=129) and assumed that if a femalersquos estimated posi-tion was within 200 m of the estimated locations ofher kittens they were probably together We alsocounted the number of locations where the femalewasgt10 km from her kittens to facilitate compar-isons with other studies We then expressed thesevalues as percents of the total number of locationsfor the 24-hour session Our sample unit was theindividual animal so for all our calculations weaveraged the sessions of females for which we hadmore than one 24-hour session to avoid pseudo-replicationWe used Sigmastat software (Systat Software

Inc Point Richmond CA USA) for all statisticalanalyses and reported allmeans asiexclstandard error

Results

Over the studyperiod our research teammonitored15adult pumaswithkittens for 24 complete 24-hoursessions For 12 of the females we were able tosimultaneouslymonitor their radio-collared kittensduring 21 monitoring sessions We had singlesessions for nine females and three four and sixsessions for theother threeThekittensranged inagewithin 7-12 months We were also able to monitor

one time each three other females whowe knew hadrecently born kittens (lt1 month old) at known densites

The distance females were from their kittensranged within 00-38 km over the 24 hours (Fig2A) The percent time females were j200 m fromtheir kittens over the 24 hours averaged 162iexcl38(N=12) Conversely females were gt10 km fromtheir kittens 309iexcl67 of the time The three fe-males with kittens in dens were near their dens 103122 and 23 of the time

Relative to time of day females were usuallyi10 km from their kittens during 1100-1600Theywere within 06-08 kmduringmost of the restof the time (see Fig 2A) There were three periodsover the 24 hours when females werej200 m fromtheir kittens an average 23-29 of the time 1500-1730 (257iexcl57 N=5) 2300-0130 (294iexcl30 N=6) and 0700-1030 (232iexcl31 N=8

Figure 2 Average distance (in km) 12 female pumas were fromtheir 7-12month-old kittens over the course of 24 hours (A) andaverage percent time these 12 females were j200 m of theirkittens (B)Averagesamplesize foreachhalf-hoursamplewas98female-kitten pairs The average standard error for the half-hourly distances between females and kittens was 02iexcl001 Therange of sunset and sunrise times over the study period is in-dicated

224 WILDLIFE BIOLOGY 142 (2008)

see Fig 2B) During 1100-1430 females were neartheir kittens the least amount of time (51iexcl21N=8 see Fig 2B)

Discussion

Basedonsimultaneous telemetry locationscollectedat varying time intervals over the year four femalesand their 7-12 month-old kittens in southern Utahwere together69ofthe time (BarnhurstampLindzey1989) Three Florida panther Puma concolor coryifemales with dependent kittens (no ages werespecified) were within 05 km of each other 45of the time andweregt10 kmapart 47of the time(Maehr et al 1991) In contrast in New Mexicokittens gt6 months old were with their mothers82 of the time (Logan amp Sweanor 2001) Thesetelemetry-based estimates are substantially higherthan the 162 we found for the 12 females within200 m of their kittens Barnhurst amp Lindzey (1989)and Logan amp Sweanor (2001) observed that takingdaytime telemetry locations when pumas are in-active (Beier et al 1995) probably overestimated thepercent time females and kittens were together Wedidfind that thepercent time femaleswerewith theirkittenschangedthroughthedaytimewithoneof thehighest average percentages being early morning(0700-1030) If telemetry flights in these studieswere in the early morning (when the air is moststable) then the estimates of Barnhurst amp Lindzey(1989) and Logan amp Sweanor (2001) were probablyoverestimates of the total time females and kittensare together However given the variability wefound in the amount of time they were close to theirkittens (0-420) some of the differences could bedue to the small sample sizes (3-4 females) of theseprevious studiesIn contrast to their telemetry based estimates

BarnhurstampLindzey (1989) found femalesand their7-12 month-old kittens together 43 of the timebasedonsnowtracksBecauseBarnhurstampLindzey(1989) recorded only tracks crossing roads theycontended that this estimate should be the mostrealistic to the percent of time hunters using thesametechnique shouldencounter femaleswiththeirkittens Their estimate however was still higherthan the 168 we found via telemetry Againbecause of the variability we found among days andanimals the higher percent of Barnhurst ampLindzey(1989) couldbe related to their limited sample sizeoftwo animals Another possibility is that the percent

time females associate with their kittens differsbetween summer when we collected our data andwinter However we currently lack the data to testthis supposition

Barnhurst amp Lindzey (1989) found that twofemales were together with their 0-6 month-oldkittens for 63 of the telemetry locations Incontrast their snow data indicated that femalestraveled with young kittens 19 of the time whichwas only slightly higher than our telemetry estimateof 23-122 for three females with lt1 month-oldkittensFemales rarelymovetheirkittensawayfromtheir birth sites before they are two months old(Logan amp Sweanor 2001) so females will be alonewhen they travel for at least two of six months afterthe kittens are born Therefore the estimate ofBarnhurst amp Lindzey (1989) from snow tracking isperhaps more applicable for kittens of 3-6 monthswhich we would expect to be higher In all casessample sizes were extremely small (2-4 animals) butconsidering our data and the snow-tracking dataofBarnhurstampLindzey (1989) females appear tobewith their 0-6 month-old kittenslt20 of the time

During the diel cycle females are least likely to benear their young from late morning to mid-after-noon (seeFig 2B) It is during this time that they arealso the furthest away (see Fig 2A) We speculatethat in late morning females separate from theiryoung perhaps to sleepundisturbed It appears thatthey rejoin their young in late afternoon but thenseparateagain towardeveningwhentheybegin tobeactive (Beier et al 1995) Over the nighttime thepatternof percent time females are near their kittensappears similar to the pattern of activity (ie dis-tance moved per hour) Beier et al (1995) reportedand forwhatwe found inour study (Fig 3)Femalesshowed a peak in activity during 1600-2200 (seeFig 3) and during this period we found them to beaway from their young gt80 of the time (see Fig2B) Shortly before midnight females seem to rejointheir kittens which corresponded to the reducedactivity we (see Fig 3) and also Beier et al (1995)noted An hour or so before sunrise we (see Fig 3)and Beier et al (1995) found that pumas increasedtheir activity and again we found that they sepa-rated from their young (see Fig 2B) The cycle com-pletes itself when females rejoin their young aroundsunrise before they separate again in late morningConsequently the duel requirements of caring fortheir youngandactivity primarily hunting appearsto produce a distinct diel pattern of association be-tween females and their young

WILDLIFE BIOLOGY 142 (2008) 225

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

aminimal impactofourpresenceontheirbehaviourConverselybecauseof the sizeof themountainsandthe accessibility to the valleys within the mountainranges we were rarely gt5 km from the animals wemonitored and were able to monitor most pumasover the 24hourswithoutmoving stationsAlsoweassumed minimal signal error because reception ofthe signal was predominately line of sight with littleproblems with signal bounceForeach24-hour sessionweentered thedata into

a program we developed to estimate the location ofthe animal at half-hour intervals We then matchedup the simultaneous locations of female pumaswiththeir kittens in an Excel worksheet and determinedthe female-kitten distance for each half-hour lo-cationWhentherewasgt1radio-collaredkittenwecalculated the distance the female was from eachkitten and then averaged these distances Once wecalculated the female-kitten distances for each halfhour we then counted the number of locationswhere the female was near her kittens Because oftelemetry error it is possible to misclassify a femaleas being away from or near her kittens when indeedshe was the opposite To reduce these errors we in-corporated our telemetry error (180iexcl187 m N=129) and assumed that if a femalersquos estimated posi-tion was within 200 m of the estimated locations ofher kittens they were probably together We alsocounted the number of locations where the femalewasgt10 km from her kittens to facilitate compar-isons with other studies We then expressed thesevalues as percents of the total number of locationsfor the 24-hour session Our sample unit was theindividual animal so for all our calculations weaveraged the sessions of females for which we hadmore than one 24-hour session to avoid pseudo-replicationWe used Sigmastat software (Systat Software

Inc Point Richmond CA USA) for all statisticalanalyses and reported allmeans asiexclstandard error

Results

Over the studyperiod our research teammonitored15adult pumaswithkittens for 24 complete 24-hoursessions For 12 of the females we were able tosimultaneouslymonitor their radio-collared kittensduring 21 monitoring sessions We had singlesessions for nine females and three four and sixsessions for theother threeThekittensranged inagewithin 7-12 months We were also able to monitor

one time each three other females whowe knew hadrecently born kittens (lt1 month old) at known densites

The distance females were from their kittensranged within 00-38 km over the 24 hours (Fig2A) The percent time females were j200 m fromtheir kittens over the 24 hours averaged 162iexcl38(N=12) Conversely females were gt10 km fromtheir kittens 309iexcl67 of the time The three fe-males with kittens in dens were near their dens 103122 and 23 of the time

Relative to time of day females were usuallyi10 km from their kittens during 1100-1600Theywere within 06-08 kmduringmost of the restof the time (see Fig 2A) There were three periodsover the 24 hours when females werej200 m fromtheir kittens an average 23-29 of the time 1500-1730 (257iexcl57 N=5) 2300-0130 (294iexcl30 N=6) and 0700-1030 (232iexcl31 N=8

Figure 2 Average distance (in km) 12 female pumas were fromtheir 7-12month-old kittens over the course of 24 hours (A) andaverage percent time these 12 females were j200 m of theirkittens (B)Averagesamplesize foreachhalf-hoursamplewas98female-kitten pairs The average standard error for the half-hourly distances between females and kittens was 02iexcl001 Therange of sunset and sunrise times over the study period is in-dicated

224 WILDLIFE BIOLOGY 142 (2008)

see Fig 2B) During 1100-1430 females were neartheir kittens the least amount of time (51iexcl21N=8 see Fig 2B)

Discussion

Basedonsimultaneous telemetry locationscollectedat varying time intervals over the year four femalesand their 7-12 month-old kittens in southern Utahwere together69ofthe time (BarnhurstampLindzey1989) Three Florida panther Puma concolor coryifemales with dependent kittens (no ages werespecified) were within 05 km of each other 45of the time andweregt10 kmapart 47of the time(Maehr et al 1991) In contrast in New Mexicokittens gt6 months old were with their mothers82 of the time (Logan amp Sweanor 2001) Thesetelemetry-based estimates are substantially higherthan the 162 we found for the 12 females within200 m of their kittens Barnhurst amp Lindzey (1989)and Logan amp Sweanor (2001) observed that takingdaytime telemetry locations when pumas are in-active (Beier et al 1995) probably overestimated thepercent time females and kittens were together Wedidfind that thepercent time femaleswerewith theirkittenschangedthroughthedaytimewithoneof thehighest average percentages being early morning(0700-1030) If telemetry flights in these studieswere in the early morning (when the air is moststable) then the estimates of Barnhurst amp Lindzey(1989) and Logan amp Sweanor (2001) were probablyoverestimates of the total time females and kittensare together However given the variability wefound in the amount of time they were close to theirkittens (0-420) some of the differences could bedue to the small sample sizes (3-4 females) of theseprevious studiesIn contrast to their telemetry based estimates

BarnhurstampLindzey (1989) found femalesand their7-12 month-old kittens together 43 of the timebasedonsnowtracksBecauseBarnhurstampLindzey(1989) recorded only tracks crossing roads theycontended that this estimate should be the mostrealistic to the percent of time hunters using thesametechnique shouldencounter femaleswiththeirkittens Their estimate however was still higherthan the 168 we found via telemetry Againbecause of the variability we found among days andanimals the higher percent of Barnhurst ampLindzey(1989) couldbe related to their limited sample sizeoftwo animals Another possibility is that the percent

time females associate with their kittens differsbetween summer when we collected our data andwinter However we currently lack the data to testthis supposition

Barnhurst amp Lindzey (1989) found that twofemales were together with their 0-6 month-oldkittens for 63 of the telemetry locations Incontrast their snow data indicated that femalestraveled with young kittens 19 of the time whichwas only slightly higher than our telemetry estimateof 23-122 for three females with lt1 month-oldkittensFemales rarelymovetheirkittensawayfromtheir birth sites before they are two months old(Logan amp Sweanor 2001) so females will be alonewhen they travel for at least two of six months afterthe kittens are born Therefore the estimate ofBarnhurst amp Lindzey (1989) from snow tracking isperhaps more applicable for kittens of 3-6 monthswhich we would expect to be higher In all casessample sizes were extremely small (2-4 animals) butconsidering our data and the snow-tracking dataofBarnhurstampLindzey (1989) females appear tobewith their 0-6 month-old kittenslt20 of the time

During the diel cycle females are least likely to benear their young from late morning to mid-after-noon (seeFig 2B) It is during this time that they arealso the furthest away (see Fig 2A) We speculatethat in late morning females separate from theiryoung perhaps to sleepundisturbed It appears thatthey rejoin their young in late afternoon but thenseparateagain towardeveningwhentheybegin tobeactive (Beier et al 1995) Over the nighttime thepatternof percent time females are near their kittensappears similar to the pattern of activity (ie dis-tance moved per hour) Beier et al (1995) reportedand forwhatwe found inour study (Fig 3)Femalesshowed a peak in activity during 1600-2200 (seeFig 3) and during this period we found them to beaway from their young gt80 of the time (see Fig2B) Shortly before midnight females seem to rejointheir kittens which corresponded to the reducedactivity we (see Fig 3) and also Beier et al (1995)noted An hour or so before sunrise we (see Fig 3)and Beier et al (1995) found that pumas increasedtheir activity and again we found that they sepa-rated from their young (see Fig 2B) The cycle com-pletes itself when females rejoin their young aroundsunrise before they separate again in late morningConsequently the duel requirements of caring fortheir youngandactivity primarily hunting appearsto produce a distinct diel pattern of association be-tween females and their young

WILDLIFE BIOLOGY 142 (2008) 225

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

see Fig 2B) During 1100-1430 females were neartheir kittens the least amount of time (51iexcl21N=8 see Fig 2B)

Discussion

Basedonsimultaneous telemetry locationscollectedat varying time intervals over the year four femalesand their 7-12 month-old kittens in southern Utahwere together69ofthe time (BarnhurstampLindzey1989) Three Florida panther Puma concolor coryifemales with dependent kittens (no ages werespecified) were within 05 km of each other 45of the time andweregt10 kmapart 47of the time(Maehr et al 1991) In contrast in New Mexicokittens gt6 months old were with their mothers82 of the time (Logan amp Sweanor 2001) Thesetelemetry-based estimates are substantially higherthan the 162 we found for the 12 females within200 m of their kittens Barnhurst amp Lindzey (1989)and Logan amp Sweanor (2001) observed that takingdaytime telemetry locations when pumas are in-active (Beier et al 1995) probably overestimated thepercent time females and kittens were together Wedidfind that thepercent time femaleswerewith theirkittenschangedthroughthedaytimewithoneof thehighest average percentages being early morning(0700-1030) If telemetry flights in these studieswere in the early morning (when the air is moststable) then the estimates of Barnhurst amp Lindzey(1989) and Logan amp Sweanor (2001) were probablyoverestimates of the total time females and kittensare together However given the variability wefound in the amount of time they were close to theirkittens (0-420) some of the differences could bedue to the small sample sizes (3-4 females) of theseprevious studiesIn contrast to their telemetry based estimates

BarnhurstampLindzey (1989) found femalesand their7-12 month-old kittens together 43 of the timebasedonsnowtracksBecauseBarnhurstampLindzey(1989) recorded only tracks crossing roads theycontended that this estimate should be the mostrealistic to the percent of time hunters using thesametechnique shouldencounter femaleswiththeirkittens Their estimate however was still higherthan the 168 we found via telemetry Againbecause of the variability we found among days andanimals the higher percent of Barnhurst ampLindzey(1989) couldbe related to their limited sample sizeoftwo animals Another possibility is that the percent

time females associate with their kittens differsbetween summer when we collected our data andwinter However we currently lack the data to testthis supposition

Barnhurst amp Lindzey (1989) found that twofemales were together with their 0-6 month-oldkittens for 63 of the telemetry locations Incontrast their snow data indicated that femalestraveled with young kittens 19 of the time whichwas only slightly higher than our telemetry estimateof 23-122 for three females with lt1 month-oldkittensFemales rarelymovetheirkittensawayfromtheir birth sites before they are two months old(Logan amp Sweanor 2001) so females will be alonewhen they travel for at least two of six months afterthe kittens are born Therefore the estimate ofBarnhurst amp Lindzey (1989) from snow tracking isperhaps more applicable for kittens of 3-6 monthswhich we would expect to be higher In all casessample sizes were extremely small (2-4 animals) butconsidering our data and the snow-tracking dataofBarnhurstampLindzey (1989) females appear tobewith their 0-6 month-old kittenslt20 of the time

During the diel cycle females are least likely to benear their young from late morning to mid-after-noon (seeFig 2B) It is during this time that they arealso the furthest away (see Fig 2A) We speculatethat in late morning females separate from theiryoung perhaps to sleepundisturbed It appears thatthey rejoin their young in late afternoon but thenseparateagain towardeveningwhentheybegin tobeactive (Beier et al 1995) Over the nighttime thepatternof percent time females are near their kittensappears similar to the pattern of activity (ie dis-tance moved per hour) Beier et al (1995) reportedand forwhatwe found inour study (Fig 3)Femalesshowed a peak in activity during 1600-2200 (seeFig 3) and during this period we found them to beaway from their young gt80 of the time (see Fig2B) Shortly before midnight females seem to rejointheir kittens which corresponded to the reducedactivity we (see Fig 3) and also Beier et al (1995)noted An hour or so before sunrise we (see Fig 3)and Beier et al (1995) found that pumas increasedtheir activity and again we found that they sepa-rated from their young (see Fig 2B) The cycle com-pletes itself when females rejoin their young aroundsunrise before they separate again in late morningConsequently the duel requirements of caring fortheir youngandactivity primarily hunting appearsto produce a distinct diel pattern of association be-tween females and their young

WILDLIFE BIOLOGY 142 (2008) 225

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

Given that females spend considerable amountsof time away from their young what are themanagement implications As proposed by Barn-hurstampLindzey (1989) theprobability thatahunterwill detect a female with kittens is low and even lessforfemaleswithextremelyyoungvulnerablekittensFemales are most likely to cross roads and trailsleaving tracks for hunters to find when they areactive at night Based on our data in Fig 2B it isduring these periods of activity that females are lesslikely accompanied by their young This furtherdecreases the probability that a hunter will find afemalersquos track accompanied by those of her kittensWhen a hunter releases his dogs on a single femaletrack the dogs usually run faster than the humansThus hunters rarely travel closely with their dogsduring the chase (JW Laundre pers obs) Manytimes when the dogs reach the puma and she haskittens the female goes in one direction and thekittens in another (JW Laundre pers obs)Because the dogs have been tracking the scent ofthe female they most often continue on her trailleaving the kittens behind (JW Laundre persobs) Thus even if a female had rejoined her kittensafter crossing a road by the time the dogs chase herup into a tree she is alone again Hunters normallyknow when the dogs have a puma in a tree by thechangeof the frequencyand toneof their barkTheythen travel directly to the tree missing much of thechase trail where they might have noted kittentracks Consequently the hunter would mistakenlyassume the female was without kittens and maydecide to kill her If we add to this the fact that

females are either pregnant or with dependent kit-tens approximately 80 of their adult lives (Loganamp Sweanor 2001) and that females can give birth tokittens at anytime of the year then the probabilitythat ahunterwill unknowinglykill either apregnantfemale or one with undetected dependent kittens isextremely high

One way of reducing possible orphaning of lt6month-oldkittens is to adjust thehunting seasons totimes when the minimal possible number of suchyoung kittens occurs in the population (CougarManagement Guidelines Working Group 2005)However given the frequency distribution of births(Cougar Management Guidelines Working Group2005 Laundre amp Hernandez 2007) 60 of thekittens arej6months old (born in July-December)and thus dependent on their mothers during thetraditional December-March hunting season Thisleaves little room for adjustment of the seasons Anobvious solution would be to prohibit the killing ofall females However given the current concernabout the impacts of puma predation on ungulatepopulations (Ballard et al 2001) it is unlikely thatgameagencieswouldenact sucharestrictionOntheother hand under the current hunting regulationsthe lawsprohibiting thekillingof femalepumaswithkittens would seem relatively ineffective If we con-tinue the harvest of female pumas we must acceptthat the undetected loss of kittens for future re-cruitment will occur What remains to be studied isthe impactof this lossonthepopulationdynamicsofhunted puma populations

Acknowledgements - our project was initiated in 1985 as along-term study of puma ecology behaviour and con-servation andwas conductedunder the auspices of IdahoStateUniversity and theNorthernRockies ConservationCooperative We thank the following organizations forfinancial and logistic support of thefieldworkused for thebasis of this analysis ALSAM Foundation Boone andCrockett Club Earthwatch Institute Fanwood Founda-tion Idaho State University National Rifle AssociationThe Eppley Foundation US Bureau of Land Manage-ment NorthernRockies ConservationCooperative Ida-ho Department of Fish and Game Mazamas Merrill GandEmitaEHastingFoundation Patagonia Inc SEA-CON of the Chicago Zoological Society WilliamH andMattie Wattis Harris Foundation Utah Division ofWildlife and Wiancko Charitable Trust We wouldlike to thank the many Earthwatch volunteers withoutwhose help this workwould not have been accomplishedWe also would like to thank Ken Jafek Kevin AllredJanet Loxterman BrianHolmesKellyAltendorf CarlosLopez Gonzalez Laura Heady and Scott Blum fortheir help in the field We extend a special thanks to

Figure3Averagedistance (inkm)the females inourstudymovedper hour over the 24-hour cycle The average number of animalswe used per hour is the same as in Fig 2 The range of sunset andsunrise times over the study period is indicated

226 WILDLIFE BIOLOGY 142 (2008)

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227

C Patrick and G Ordway for their support Lastly wethank Harley Shaw for his helpful comments on thismanuscript

References

BallardWBLutzDKeeganTWCarpenterLHamp

deVos JC Jr 2001 Deer-predator relationships areviewof recentNorthAmerican studieswith emphasison mule and black-tailed deer - Wildlife Society Bul-

letin 29 99-115Barnhurst D amp Lindzey FG 1989 Detecting femalemountain lions with kittens - Northwest Science 6335-37

Beier P Choate D amp Barrett RH 1995 Movementpatterns ofmountain lions during different behaviors -Journal of Mammalogy 76 1056-1070

Blum S 2003 Habitat use by mountain lions (Pumaconcolor) in the Northern Great Basin - MSc thesisIdaho State University Pocatello 76 pp

Cougar Management Guidelines Working Group 2005

Cougar Management Guidelines 1st edition- Wild-FuturesBainbridge IslandWashingtonUSA137pp

Laundre JW amp Hernandez L 2007 Do female pumas

(Puma concolor) exhibit a birth pulse- Journal ofMammalogy 88 1300-1304

Laundre JW amp Keller BL 1984 Home-range size

of coyotesA critical review - Journal ofWildlifeMan-agement 48 127-139

Lenth RV 1981 On finding the source of a signal -Technometrics 23 149-154

Logan KA amp Sweanor LL 2001 Desert puma evolu-tionary ecology and conservation of an enduring carni-vore - Island Press Washington DC 463 pp

Maehr D Land ED ampRoof JC 1991 Social ecologyof Florida panthers- National Geographic Researchand Exploration 7 414-431

Maehr DS Roof JC Land ED amp McCown JW1989 First reproduction of a panther (Felis concolorcoryi) in southwestern Florida USA - Mammalia 53129-131

WILDLIFE BIOLOGY 142 (2008) 227