DAILY ACTIVITY PATTERNS OF MOUNTAIN ( CONCOLOR ) IN

RELATION TO THE ACTIVITY OF THEIR PREY IN SOUTHERN

ARIZONA

By

Emil B. McCain

A Thesis

Presented to

The Faculty of Humboldt State University

In Partial Fulfillment

Of the Requirements for the Degree

Master of Science in

Natural Resources: Wildlife

August, 2008

ABSTRACT

Daily Activity Patterns of Mountain Lions ( Puma concolor ) in Relation to the Activity of their Prey Species in Southern

Emil B. McCain

Food resources are not evenly distributed over space or time, and therefore, changes in prey abundance and availability may influence predator behavior both spatially and temporally. It has been suggested that mountain lions ( Puma concolor ) follow the daily activity patterns of their main prey species. In the Sonoran Desert the javelina ( Pecari tajacu ) is an important prey item for mountain lions and it has been shown that javelina shift from a diurnal activity pattern during winter months to a nocturnal pattern in the summer. I examined whether mountain lions shift their activity patterns between summer and winter following the activity of the javelina. Alternatively,

I examined whether mountain lions shifted their diet to other species that were more active during the period when mountain lions were active. I analyzed 117 mountain fecal samples to determine their diet during summer (16 April – 15 October) and winter

(16 October – 15 April) and I used the date/time stamps from 4,528 trail-camera photographs collected during March 2001-September 2006 in southern Arizona to index daily activity patterns of mountain lions and their prey species.

Mountain lions did not track the activity of one particular prey species, but appeared to shift their daily activity patterns and diet according to temperature and availability of different prey species in a given season. Coues white-tailed deer

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(Odocoileus virginianus couesi ) were the most common prey in both winter and summer, but the mountain lion diet was supplemented with seasonally abundant and vulnerable domestic calves (Bos taurus ) in summer and javelina in winter. Because mountain lion activity patterns and their diet both changed between winter and summer, it was difficult to discern exactly what drove these seasonal shifts. High temperatures may have influenced mountain lions to shift towards nocturnal activity in the summer. Coues white- tailed deer remained primarily diurnal through the summer, while the excessive daytime temperatures may have limited mountain lion movements and reduced their ability to exploit this resource. The occurrence of deer in mountain lion diet decreased in summer, and cattle increased. With the cooler daytime temperatures and the absence of calves of domestic cattle in the winter, mountain lions became more diurnal, which coincided with the activity of Coues white-tailed deer and javelina, both of which became more prevalent in the mountain lion diet at that time.

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ACKNOWLEDGEMENTS

I thank my advisory committee at Humboldt State University; Dr. T. Luke George and Dr. Richard T. Golightly of the Wildlife Department, and Dr. William Zielinski of

Redwood Sciences Laboratory. Mr. Jack L. Childs of the Borderlands Detection

Project provided tremendous field instruction, guidance, encouragement, partnership, and friendship; thank you. The Integral Ecology Research Center and G. Wengert and M.

Gabriel provided financial and logistic support and managed funding from the Arizona

Game and Fish Department, Disney Wildlife Conservation Fund, Switzer Foundation,

Woodland Park Zoo, Milwaukee Zoo, Bergin County Zoo, Coronado National Forest,

USDA Pacific Southwest Research Station, Redwood Sciences Laboratory, Buenos Aires

National Wildlife Refuge, and several individual donors, thank you all. The Bell family provided accommodations within the study site, and continuous technical and logistical support; I am forever grateful. M. Abbott, with instruction from C. Hass, conducted microscopic identification of hair remains in the diet study. M. Culver, Z. Hackle and S.

Carrillo conducted genetic analysis, with funding from the Arizona-New Jaguar

Conservation Team, its chairmen (T. Johnson and B. Van Pelt), and agency signatories.

A. Childs, L. Colvin, M. Colvin, J. Conklin, S. Bell, S. Bless, J. Brun, M. Holister, C.

McGary, J. McCain, A. Neils, Pancho, S. Pavlik, G. Paz, M. Pruss, W. Rizzo, K.

Shallcross, T. Snow, Sundog, M. Terrio, V. Walkosak, S. Walkosak, and T. Wright all assisted with field work. R. Borque contributed the site map. Last, but not least, I thank my loving parents, Jim and Roz McCain, for their support and encouragement. v

TABLE OF CONTENTS

PAGE

ABSTRACT ...... iii

ACKNOWLEDGMENTS ...... v

LIST OF TABLES ...... vii

LIST OF FIGURES ...... viii

LIST OF APPENDICES ...... x

INTRODUCTION ...... 1

MATERIALS AND METHODS ...... 5

Study Site ...... 5

Diet ...... 5

Activity Patterns...... 8

Data Analysis ...... 10

RESULTS ...... 12

Diet ...... 12

Activity Patterns...... 16

DISCUSSION ...... 23

LITERATURE CITED ...... 29

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LIST OF TABLES

Table Page

1 Prey items present in mountain lions scats (n = 117) from southern Arizona 2004-2005. Data is presented for all prey items found in scats and the most dominant prey item in each scat. Percent of occurrence is the number of times a specific item was found as a percentage of all items found. Frequency of occurrence is the percentage of total scats in which an item was found (Ackerman et al. 1984), where n = the number of scats...... 13

2 Frequency of occurrence of the dominant prey species found in 117 mountain lion scats in southern Arizona 2004-2005 during summer (16 April – 15 October) and winter (16 October – 15 April). The frequency of occurrence was the percentage of total scats in which each prey item was found. P-values are Chi-square tests for the null hypothesis of no difference in the frequency of occurrence of prey species between summer and winter. Expected values were calculated for each species as the proportion of that species in the total annual diet multiplied by the number of scats in each respective season...... 14

3 Total number of trail camera photographs taken of mountain lions and their top six prey species during summer (16 April – 15 October) and winter (16 October – 15 April) in southern Arizona from 2001 – 2006. could not be identified to species in all photographs and therefore were lumped ...... 17

4 Pearson Correlation coefficients (r) and associated P-values between daily activity patterns of mountain lion and the activity patterns of their top six prey species in southern Arizona from March 2001-September 2006. Activity was measured as the number of trail camera photographs taken of each species during each of the 8 time bins throughout the 24-h period. Skunks could not be identified to species in all photographs and therefore were lumped ...... 21

vii

LIST OF FIGURES

Figure Page

1 Topographic relief of the study area, showing Tucson, Arizona (black polygon), the U.S. / Mexico border (black line) and the major mountain ranges and valleys within the study area. The study area extended from the crest of the Baboquivari Mountains east to the San Rafael Valley and approximately 80 km north of the International border, specifically focusing within the mountain ranges (white ovals). Exact camera locations cannot be given due to the confidentiality of jaguar location data generated on this study...... 6

2 Frequency of occurrence of the top six prey species in mountain lion scat in southern Arizona from 2004-2005 during summer (16 April – 15 October) (n=67) and winter (16 October – 15 April) (n=50). Solid Bars represent observed diet and open bars show expected values for each species (calculated as the proportion of each species in the total annual diet multiplied by the number of scats from each season). Skunks could not be distinguished to species in all scats and therefore were lumped...... 15

3 Daily activity pattern of mountain lions in southern Arizona from March 2001 – September 2006 (n=696) based on percentage of trail camera photographs taken during each of eight 3-h time bins throughout the 24-h period. Solid bars represent winter (16 October – 15 April) and open bars represent summer (16 April – 15 October). Time Bins: MidNight = 2230- 0130; LateNight = 0130-0430; Dawn = 0430-0730; EarlyDay = 0730- 1030; MidDay = 1030-1330; LateDay = 1330-1630; Dusk = 1630-1930; and EarlyNight = 1930-2230 (Mountain Standard Time)...... 18

4 Percent of photographs of Coues white-tailed deer (n = 789), domestic cow (n = 365), cottontail (n = 253), javelina (n = 351), spp (n = 355), and white-nosed (n = 144) taken during eight 3-h time bins in southern Arizona. Solid bars represent winter (16 October – 15 April) and open bars represent summer (16 April – 15 October). Time Bins: MidNight = 2230-0130; LateNight = 0130-0430; Dawn = 0430-0730; EarlyDay = 0730-1030; MidDay = 1030-1330; LateDay = 1330-1630; Dusk = 1630-1930; and EarlyNight = 1930-2230 (Mountain Standard Time). Skunks could not be identified to species in all photographs and therefore were lumped ...... 19

viii

Figure Page

5 Correlations between mountain lion activity and prey activity based on trail camera photographs taken during eight time bins throughout the 24-h period in southern Arizona from March 2001 – September 2006. Lines are best fit regression lines. Skunks could not be identified to species in all photographs and therefore were lumped ...... 22

ix

LIST OF APPENDICES

Appendix Page

A True sunrise (lower line) and sunset (upper line) times for each day throughout the year at Nogales International Airport, Arizona (Mountain Standard Time)...... 34

B Average daily temperature recorded within the study area at the Nogales International Airport from 2000-2005 (averaged for all four years). The solid vertical lines separate the six hottest months (16 April – 15 October = summer) from the six coldest months (16 October – 15 April = winter)...... 35

C Species photographed at trail cameras in southern Arizona from March 2001 – September 2006. The total number of photographs was used to calculate detection rates (photographs/100 "trap" nights) for each species. Skunk and squirrel species could not be identified to species in all photographs and therefore were lumped ...... 36

x

INTRODUCTION

The foraging rate of a predator is the product of three factors: (1) the rate of prey encounter; (2) the rate of prey detection; and (3) the probability of a successful capture once prey has been detected (Taylor 1984). Food resources are not evenly distributed over space or time (Swingland and Greenwood 1983), and therefore, changes in prey activity and distribution may influence predator behavior both spatially and temporally

(Ward and Krebs 1985). When prey are active or moving, they are more easily detected by carnivores, and therefore, may be more vulnerable than when stationary (Hopkins

1979, Taylor 1984, Zielinski 2000). It has long been hypothesized that pursuit and ambush predators track the movements of their prey by concentrating hunting efforts when and where prey are most available (Curio 1976, Hopkins 1979, Zielinski 2000).

Zielinski (2000) suggested that predators which hunt during the time of day when prey are most vulnerable would benefit from higher capture rates at lower costs, than individuals that hunt at random times. The relationship between daily activity patterns of predators and prey, and specifically the behavior of predators in response to changes in prey activity, remains poorly understood in large mammalian species.

Carnivores may also respond to changes in prey availability by switching their diets to alternative prey species that become more available (Zielinski et al. 1983, Ward and Krebs 1985, Leopold and Krausman 1986, Yanez et al. 1986, O'Donaghue et al.

1998, Harveson et al. 2000, Zielinski 2000, Rosas-Rosas et al. 2003). For example, pine ( Martes americana ) shift their diets from ground-dwelling squirrels and voles

(Microtus spp ) in summer to Douglas squirrels ( Tamiasciurus douglasii ) in winter

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(Zielinski et al. 1983). This change in diet is associated with a change in activity from being primarily diurnal in summer to primarily nocturnal in winter. The authors suggested that this switch occurred to exploit nocturnally vulnerable prey species in winter and diurnally vulnerable species in summer.

Mountain lions (Puma concolor ) range from the northern Rocky Mountains of the

United States (Hornocker 1970, Seidensticker et al. 1973) and western Canada

(Spreadbury et al. 1996) to the Venezuelan llanos (Scognamillo et al. 2003) and the of Peru (Emmons 1987). Their daily activity patterns vary across their range

(Hornocker 1970, Seidensticker et al. 1973, Scognamillo et al. 2003, Emmons 1987). The daily activity patterns of mountain lions in southern (Waid 1990), Peru (Emmons

1987) and (Scognamillo et al. 2003) were found to remain constant through out the year, despite large changes in temperature. In central (Hopkins 1979) and southern (Beier et al. 1995) and southern Texas (Waid 1990) mule deer

(Odocoileus hemionus cooki ) are the single most important prey species. These authors all suggested the daily activity pattern of mountain lions is directly tied to the activity of their primary prey, mule deer, which exhibits no seasonal change in activity patterns. In

Idaho Seidensticker et al. (1973) found that mountain lions are more active in the daytime during the summer, presumably to take advantage of seasonally abundant diurnal ground squirrels, but return to hunting large ungulates during nocturnal and crepuscular periods the rest of the year. Therefore, several studies have suggested that the daily activity patterns of mountain lions are associated with the activity patterns of their prey.

Extreme seasonal temperature changes can cause prey species, such as javelina

(Pecari tajacu ), to shift their daily activity patterns to better meet energetic requirements

3

(Zervanos and Hadley 1973, Bigler 1974, Bissonette 1978, Waid 1990, Carrillo et al.

2002). Javelina restrict daily activity to early mornings and late evening during the summer but switch to more diurnal activity during winter in the Sonoran Desert

(Zervanos and Hadley 1973) and southern Texas (Waid 1990).

Within the Sonoran Desert of the southwestern United States and northern

Mexico, mountain lions prey heavily upon desert mule deer, Coues white-tailed deer ( O. virginianus couesi ) and javelina (Robinette et al. 1959, Ackerman et al. 1984, Waid 1990,

Cunningham et al. 1999, Rosas-Rosas et al. 2003), with abundant small supplementing diets locally (Leopold and Krausman 1986, Harveson et al. 2000).

Interestingly, mountain lions consistently prey upon javelina in direct proportion to their availability (Ackerman et al. 1984, Cunningham et al. 1995, Cunningham et al. 1999,

Harveson et al. 2000). The Sonoran Desert, therefore, offers an environment to examine mountain lion response to a seasonal change in the daily activity pattern of a primary prey species, the javelina.

If the daily activity patterns of their prey changes seasonally, mountain lions could respond in two ways: they may track changes in prey activity while keeping their diet constant, or they may change their diet while maintaining similar activity patterns throughout the year. Because javelina are an important prey item in mountain lion diets in the American Southwest (Harveson et al. 2000, Rosas-Rosas et al. 2003) and their daily activity patterns vary between seasons (Zervanos and Hadley 1973), they provide a system to test the hypothesis that mountain lions track the activity patterns of their prey.

I tested these two hypotheses by examining the activity pattern of mountain lions and their main prey species as well as mountain lion diet throughout the year in southern

4

Arizona. Specifically, I examined whether mountain lions shift from a crepuscular pattern in summer to a more diurnal pattern in winter following the activity of the javelina.

Alternatively, I examined whether mountain lions shift their diet to species that are more active during the period when they are active.

To assess the daily activity patterns of mountain lions and their prey species I utilized non-invasive remotely triggered trail cameras, which record a photographic image as well as the time and date of any activity of any species that passes by (Bridges et al. 2004). Aside from a short note on the timing of feeding bouts at carcasses (Pierce et al. 1998), this is the first study to examine mountain lion circadian behavior using trail cameras. Use of this method has been exercised to describe daily activity of black

( americanus ) (Bridges et al. 2004), Indian ( bengalensis ) (Vanak and

Gompper 2007), jaguar ( onca ) (McCain and Childs 2008), and

( pardalis ) (Dillon and Kelly 2007). The technique may provide a useful alternative to traditional methods of studying activity patterns. Contrary to methods using

GPS or radio telemetry with motion sensors, trail cameras do not require that individuals must be captured, handled, or collared, and the activity of multiple species may be measured simultaneously.

MATERIALS AND METHODS

Study Site

The study was conducted from the crest of the Baboquivari Mountains east to the

San Rafael Valley and approximately 80 km north of the International border. The study area encompassed portions of the Coronado National Forest, the Buenos Aires National

Wildlife Refuge, lands managed by the Bureau of Land Management, and the State of

Arizona, and several private ranches (Figure 1).

The study area included biotic communities of Madrean evergreen woodland and semi-desert scrub (Brown 1994). The oak woodland-oak grassland community was the dominant vegetation type between 1100 m and 2000 m. Below 1100 m, mesquite-Sonoran desert scrub predominated (Toolin et al. 1979, Brown and Lopez

Gonzalez 2001, Hatten et al. 2005). The climate is semiarid with approximately 400 mm annual rainfall, half of which occurs between July and September (Hass 2002).

Diet

I collected mountain lion fecal remains opportunistically (Ciucci et al. 1996) and on line transects conducted approximately every six weeks in conjunction with maintaining trail cameras. I identified scats as mountain lion by size, shape, and associated tracks and sign at the site of collection (Murie 1974, Childs 1998, Elbroch

2003). I collected all scats that appeared to be of felid origin that were > 30 mm in

5 6

Figure 1. Topographic relief of the study area, showing Tucson, Arizona (black polygon), the U.S. / Mexico border (black line) and the major mountain ranges and valleys within the study area. The study area extended from the crest of the Baboquivari Mountains east to the San Rafael Valley and approximately 80 km north of the International border, specifically focusing within the mountain ranges (white ovals). Exact camera locations cannot be given due to the confidentiality of jaguar location data generated on this study.

7 diameter (Cunningham et al. 1999, Rosas-Rosas et al. 2003, Novack et al. 2005). Each scat was assigned a unique ID number and recorded GPS coordinates at the site of collection. I estimated age of each scat to the nearest week based on weathering and state of decomposition, considering recent environmental conditions. Scats were placed in paper bags and upon returning from the field, they were frozen until analysis. Dr. M.

Culver at the University of Arizona conducted mitochondrial DNA analysis to identify the predator species and to exclude sympatric jaguar (Panthera onca ), ( rufus ), ( latrans ), and grey fox ( cinereoargenteus ) from the analyses.

Those scat samples that genetic analysis confirmed to be of mountain lion origin

(or failed to suggest otherwise) were air-dried and then sterilized in a pressure cooker for

15 min. to prevent exposure to parasites or disease (C. Hass pers. comm. - Borderland

Carnivore Studies). I dissected each scat sample to identify prey species from any available distinguishing characteristics within the remains, including : dentition, tooth characteristics, claw type, size and shape of hooves (Golightly et al. 1994, Ciucci et al.

1996, Rosas-Rosas et al. 2003, Novack et al. 2005), and macro and microscopic characteristics of the hair (Mayer 1952, Moore et al. 1974, Moore et al. 1997). I compiled a complete reference collection of known hair samples from all local mammalian species from Borderland Carnivore Studies (Vail, Arizona), University of

Arizona, Reid Park Zoo (Tucson, Arizona) and several local hunters. I first analyzed all scat samples at a macro level (Rosas-Rosas et al. 2003), and any questionable or unknown samples were then analyzed microscopically (Moore et al.1997). Prior to microscopic analysis, I soaked ≥ 5 hairs in carbon-tetrachloride for 15-25 minutes, then

8 in xylene for an additional 15-25 minutes and immediately mounted the hair on slides, which were compared with the reference collection and field manuals (Moore et al. 1997,

C. Hass pers. comm. - Borderland Carnivore Studies). I presented the contents of scats as both the frequency of occurrence (percentage of total scats in which a prey item was found) and percentage of occurrence (number of times a specific item was found as percentage of all prey items found) (Ackerman et al. 1984, Ciucci et al. 1996, Núñez et al. 2000, Rosas-Rosas et al. 2003, Novack et al. 2005). However, to address sample size assumptions for statistical analysis, I used only the dominant prey species in each scat

(Cunningham et al. 1999, Rosas-Rosas et al. 2003).

Activity Patterns

Following non-invasive sampling techniques developed for ( Panthera tigris ) (Karanth 1995, Karanth and Nichols 1998, Karanth et al. 2004), I implemented a series of systematic surveys with automatic trail cameras arranged in grids over the three mountain range complexes in the study area. I placed at least one trail camera along the most probable travel-route within each 25 km 2 grid and maintained > 1 km distance between cameras (Henschel and Ray 2003, Silver et al. 2004). The number of 25 km 2 grid cells in each mountain range complex ranged from 5-35 and the number of cameras ranged from 5-35 per mountain range complex over the course of the study. To address autocorrelation issues that occur with wide-ranging carnivores, I recorded only the first detection when more than one photograph was taken of the same species at a given camera site within 60 min (Bridges et al. 2004). Any subsequent photographic records of the same species were discarded, unless physical characteristics indicated that

9 photographs unambiguously represented two different individuals. Although cameras were placed > 1 km apart, individuals may have been able to move between camera sites in < 60 min; therefore pictures of the same species on adjacent cameras also had to be separated by at least 60 min to be included in the analysis.

I focused surveys on major wildlife travel-routes and natural funnels through the mountain ranges within my study area. I placed cameras along known mountain lion travel-routes (based on tracks, scrapes, scat, and the existing knowledge of local ranchers, hunters and biologists) along washes, trails, dirt roads, ridges, and canyon bottoms

(Henschel and Ray 2003, Karanth et al. 2004, Silver et al. 2004), and in areas where wildlife travel was naturally directed or funneled by landscape features. I used trail camera systems designed for continuous surveillance of medium to large mammals. I used two types of cameras: digital (Cuddeback Digital ® - Non Typical, Inc., Park Falls,

WI) and film (CamTrakker ® - Camtrak South, Inc., Watkinsville, GA). All cameras were fully automatic and used passive infrared motion sensors that detect heat-in-motion. I attached camera units to trees and aimed approximately 0.3 m above the level of the trail or travel-route. Cameras monitored activity 24 h per day with a five min delay between photographs to avoid multiple photographs of the same individual (Main and Richardson

2002). I serviced cameras approximately every six weeks to replace batteries and film or memory card.

I examined daily activity patterns using the time stamp on each photograph

(Bridges et al. 2004, McCain and Childs 2008). Because the cameras require heat-in- motion to be triggered, each photograph was considered to represent active behavior of the subject species. The time stamps recorded from these activity events were compiled

10 to form an index of activity times for different species (Bridges et al. 2004). I divided the

24-h day into eight 3-h time bins: MidNight = 2230-0130; LateNight = 0130-0430;

Dawn = 0430-0730; EarlyDay = 0730-1030; MidDay = 1030-1330; LateDay = 1330-

1630; Dusk = 1630-1930; and EarlyNight = 1930-2230. All times were recorded in

Mountain Standard Time. Because the camera monitoring of this study was continuous and the number of camera-trap days was large (27,416), the effort throughout the 8 time bins was assumed to be equal.

Day length changed throughout the year, and therefore sunrise and sunset times fluctuated through the seasons (Appendix A). Sunrise times varied from 0519 near the summer solstice, to 0724 near the winter solstice. Sunset times ranged from 1721 at the winter solstice to 1932 around the summer solstice. The time bins were constructed so that sunrise always occurred within the “Dawn” category, and “Dusk” encompassed the sunset except for a few days during the year.

Field methods were approved by Humboldt State University Institutional

Care and Use Committee (IACUC Numbers 04/05.W.43.A and 06/07.W.133.E).

Data Analysis

I divided the year into two seasons based on local temperature records from the

Nogales International Airport, National Oceanographic and Atmospheric Agency monitoring station approximately 40 km southeast of the center of the study site. I used average daily temperatures from 2002-2006 to delineate the hot (summer) period from 16

April to 15 October, and the cooler (winter) period from 16 October to 15 April

(Appendix B).

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I used Chi-Square Test of Independence to test for differences in frequency of occurrence of prey in mountain lion scats between summer and winter. However, to clearly define the sample size for statistical analysis, I considered only the dominant prey item per scat in the analyses (Cunningham et al. 1999, Rosas-Rosas et al. 2003). The dominant prey item in each scat was the species which had the largest percentage by volume of remains in each scat. If two species were equally represented, I selected the larger body size of the two as the dominant species because smaller prey species can bias results due to disproportionate surface area to volume ratios (Ackerman et al. 1984). I compared the dominant prey remains in mountain lion diet between summer and winter.

I also tested each prey species between seasons to determine if the proportions of each species in the diet changed with season. For the within-species comparison, the observed value was the frequency that a particular prey item was observed in a season, and the expected value was the proportion of that species in the total annual diet multiplied by the number of scats from each respective season (Rosas-Rosas et al. 2003). I used only the prey species that comprised >5% of the annual mountain lion diet for the diet analysis.

I used a Chi-squared contingency analysis to test for seasonal changes in daily activity patterns. I tested for differences in the number of photographs in each of the eight times bins for each of the two seasons. The level of association of the activity patterns between mountain lion and each prey species was compared using Pearson

Correlations. Activity was measured as the number of photographs taken during each of

8 time bins throughout the 24 h period.

RESULTS

Diet

Between March 2004 and December 2005 I collected 128 fecal samples that were identified as of probable mountain lion origin based on size, shape and associated signs

(Murie 1974, Childs 1998, Elbroch 2003). Genetic analysis at the University of Arizona confirmed 32 of those to be from mountain lion and excluded six bobcat and three coyote scats. The others were unconfirmed and assumed to be of mountain lion origin. The 117 mountain lion scat samples contained 143 prey items (24 scats contained two different prey species and four scats contained three species). Coues white-tailed deer was the most frequent prey item (36.8%), followed by domestic cattle ( Bos taurus ) (14.5%), cottontail rabbit ( Sylvilagus floridanus ) (13.7%), javelina (9.4%), skunks (skunks could not be identified to species in all scat and were therefore lumped) (8.5%), and white- nosed coati ( narica ) (7.7%) (Table 1). The remaining 13.3% of the diet was comprised of 9 different species (Table 1).

Mountain lion diet differed significantly between summer and winter (χ2= 20.95, df = 9, P = 0.013) (Table 2). Specifically, mountain lions preyed more on cattle in summer than winter (χ2= 4.73, df = 1, P = 0.037), and more on javelina (χ2= 3.74, df = 1,

P = 0.044) and Coues white-tailed deer ( χ2= 2.99, df = 1, P = 0.041) in winter than summer (Figure 2).

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Table 1. Prey items found in mountain lions scats (n = 117) from southern Arizona 2004-2005. Data are presented for all prey items found in scats and the dominant prey item in each scat. Percent of occurrence is the number of times a specific item was found as a percentage of all items found. Frequency of occurrence is the percentage of total scats in which an item was found (Ackerman et al. 1984), where n = the number of scats. All Prey Items _ Dominant Prey Item Percent of Frequency of Frequency of Common Name n Occurrence (%) Occurrence (%) n Occurrence (%) Canis spp 4 2.8 3.4 4 3.4 Coues white-tailed deer 45 31.5 38.5 43 36.8 Domestic cattle 17 11.9 14.5 17 14.5 Eastern cottontail rabbit 22 15.4 18.8 16 13.7 Javelina (Collared peccary) 12 8.4 10.3 11 9.4 spp 1 0.7 0.9 - - Mountain lion 6 4.2 5.1 5 4.3 1 0.7 0.9 1 0.9 Ringtail 1 0.7 0.9 - - Skunk spp 16 11.2 13.7 10 8.5 Small spp 3 2.1 2.6 1 0.9 Snake spp 1 0.7 0.9 - - Sonoran mud turtle 1 0.7 0.9 - - Arizona grey squirrel 1 0.7 0.9 - - White-nosed coati 12 8.4 _ 10.3 9 7.7 Total 143 100 100 117 100 13

Table 2. Frequency of occurrence of the dominant prey species found in 117 mountain lion scats in southern Arizona 2004-2005 during summer (16 April – 15 October) and winter (16 October – 15 April). The frequency of occurrence was the percentage of total scats in which each prey item was found. P-values are Chi-square tests for the null hypothesis of no difference in the frequency of occurrence of prey species between summer and winter. Expected values were calculated for each species as the proportion of that species in the total annual diet multiplied by the number of scats in each respective season.

Summer Winter Frequency of Frequency of Occurrence Occurrence Common Name Scientific Name n (%) n (%) P Canis spp Canis spp 4 6.0 - - 0.084 Coues white-tailed deer Odocoileus virginianus couesi 18 26.9 25 50.0 0.041 Domestic cattle Bos taurus 14 20.9 3 6.0 0.037 Eastern cottontail rabbit Sylvilagus floridanus 12 17.9 4 8.0 0.152 Javelina (Collared peccary) Pecari tajacu 3 4.5 8 16.0 0.044 Mountain lion Puma concolor 4 6.0 1 2.0 0.304 Raccoon lotor - - 1 2.0 0.247 Spilogale gracilis, macroura, Skunk spp M. mephitis, Conepatus leuconotus 7 10.4 3 6.0 0.416 Small mammal spp Small Mammal spp - - 1 2.0 0.247 White-nosed coati Nasua narica 5 _ 7.4 _ 4 _ 8.0 _ 0.917 Total 67 100 50 100

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15

Summer

30

25 20 15 10

5 0

Frequency of occurence (%) occurence of Frequency Coues white- Domestic Eastern Javelina Skunk spp White-nosed tailed deer cattle cottontail coati

Winter

30

25 Observed

20 Expected

15

10

5

0

Frequency of occurence (%) occurence of Frequency Coues white- Domestic Eastern Javelina Skunk spp White-nosed tailed deer cattle cottontail coati

Figure 2. Frequency of occurrence of the top six prey species in mountain lion scat in southern Arizona from 2004-2005 during summer (16 April – 15 October) (n=67) and winter (16 October – 15 April) (n=50). Solid Bars represent observed diet and open bars show expected values for each species (calculated as the proportion of each species in the total annual diet multiplied by the number of scats from each season). Skunks could not be distinguished to species in all scats and therefore were lumped.

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Activity Patterns

I operated between 8 and 41 trail cameras from March 2001 through September

2006. Cameras operated for a total of 27,416 working camera-trap days, producing 4,528 photographs of native wildlife species and domestic cattle (Appendix C). I obtained 696 independent photographs of mountain lions (Table 3). The six prey species that were found in >5% of the scats comprised 86.7% of the mountain lion diet by frequency of occurrence of the dominant prey species in each scat (Table 2). The number of photographs of the top six prey species ranged from 789 for white-tailed deer to 144 for the white-nosed coati (Table 3).

Mountain lions were generally crepuscular with peaks in activity before Dawn and just before or at Dusk (Figure 3). Coues white-tailed deer were primarily diurnal with activity peaking at PostDawn and dropping throughout the day (Figure 4). Domestic cattle were generally diurnal to crepuscular (Figure 4). The eastern cottontail rabbit was crepuscular, peaking before Dawn and during or after Dusk with minimal activity during the day (Figure 4). Javelina were also primarily crepuscular with peaks in activity in the

PostDawn and again in the Dusk and PostDusk periods (Figure 4). Skunks were almost completely nocturnal (Figure 4). The white-nosed coati was strongly diurnal, with a sharp peak in Early Day and decreasing throughout the day, with no activity by Dusk

(Figure 4).

The daily activity patterns of mountain lions and four of their top six prey species differed significantly between summer and winter. During summer mountain lion

17

Table 3. Total number of trail camera photographs taken of mountain lions and their top six prey species during summer (16 April – 15 October) and winter (16 October – 15 April) in southern Arizona from 2001 – 2006. Skunks could not be distinguished to species in all scats and therefore were lumped. Species Summer Winter Total Mountain lion 346 350 696 Coues white-tailed deer 333 456 789 Domestic cow 109 256 365 Eastern cottontail rabbit 177 76 253 Javelina 207 144 351 Skunk spp 238 117 355 White-nosed coati 86 58 144

18

30% Winter Summer 25%

20%

15%

10% % of Photographs 5%

0%

t n y ay a igh aw D usk ight D teD D N teN Mid MidNight a EarlyDay La L Early

Figure 3. Daily activity pattern of mountain lions in southern Arizona from March 2001 – September 2006 (n=696) based on percentage of trail camera photographs taken during each of eight 3-h time bins throughout the 24-h period. Solid bars represent winter (16 October – 15 April) and open bars represent summer (16 April – 15 October). Time Bins: MidNight = 2230-0130; LateNight = 0130-0430; Dawn = 0430-0730; EarlyDay = 0730- 1030; MidDay = 1030-1330; LateDay = 1330-1630; Dusk = 1630-1930; and EarlyNight = 1930-2230 (Mountain Standard Time).

=

Coues White-Tailed Deer Domestic Cow Cottontail Rabbit

Winter 45% 45% 45% 40% 40% 40% 35% 35% 35% Summer 30% 30% 30% 25% 25% 25% 20% 20% 20% 15% 15% 15% 10%

%of Photographs 10% %of Photographs %of Photographs 10% 5% 5% 5% 0% 0% 0% t ht n k ht n y y g ay ay s g t a a w D u i h ight w Day a yD idDay D N a ly dD Dusk teNi D te yN te D r arl M a Dawn idDay teDay Dusk yNig a a Mi ateD MidNight La E L arl M a l MidNigh L E L E MidNight EarlyDay L EarlyNight LateNight Ear

Javelina Skunk spp White-Nosed Coati

45% 45% 40% 45% 40% 40% 35% 35% 35% 30% 30% 30% 25% 25% 25% 20% 20% 20% 15% 15% 15%

10% Photographs of % 10% %of Photographs

%of Photographs 10% 5% 5% 5% 0% 0% 0% t t ht n y k ht t t h n ay g w a ay g h h y ht ight i a i g g sk g aw dDay N yD eD Dus i i Day u N D lyD Dusk te D t yN N N lyDa e D idNig te r Mi arl MidDay a d e Dawn t yNi M a a LateDay MidNigh La E L arl t MidDay l L E E Mi a Ear La EarlyNight L Ear

Figure 4. Percent of photographs of Coues white-tailed deer (n = 789), domestic cow (n = 365), cottontail rabbit (n = 253), javelina (n = 351), skunk spp (n = 355), and white-nosed coati (n = 144) taken during eight 3-h time bins in southern Arizona. Solid bars represent winter (16 October – 15 April) and open bars represent summer (16 April – 15 October). Time Bins: MidNight = 2230-0130; LateNight = 0130-0430; Dawn = 0430-0730; EarlyDay = 0730-1030; MidDay = 1030-1330; LateDay = 1330-1630; Dusk = 1630-1930; and EarlyNight = 1930-2230 (Mountain Standard Time). Skunks could not be identified to species in all photographs and therefore were lumped.

19

20 activity was highest at Dusk, EarlyNight, and MidNight, while winter activity increased during Dawn and MidDay, with a large peak at LateDay ( χ2= 26.43, df = 7, P < 0.001)

(Figure 3). The proportion of photographs was higher during all daytime periods (Dawn,

EarlyDay, MidDay, and LateDay) and lower during all nighttime periods (Dusk,

EarlyNight, MidNight, and LateNight) in winter as compared to summer. Coues white- tailed deer were more active at LateNight during summer and more active at Dawn and

EarlyDay in winter ( χ2= 24.47, df = 7, P = 0.001) (Figure 4). Domestic cattle were more active at LateNight in summer and active throughout the day during winter ( χ2= 31.30, df

= 7, P < 0.001) (Figure 4). Javelina were strongly crepuscular in summer and increased diurnal activity during EarlyDay, MidDay, and LateDay in winter ( χ2= 53.81, df = 7, P <

0.001) (Figure 4). Skunks were strictly nocturnal in summer and more active in

LateNight and LateDay in winter ( χ2= 31.40, df = 7, P < 0.001) (Figure 4). Activity patterns did not change with season for cottontail ( χ2= 4.51, df = 7, P = 0.719)

(Figure 4) or coati (χ2= 11.77, df = 6, P = 0.067) (Figure 4).

Mountain lion activity was significantly correlated with the activity of only one prey species (Table 4). Mountain lion activity had a strong negative correlation with white-nosed coati activity during the summer months (Table 4). were strictly diurnal (Figure 4) and mountain lions were largely nocturnal (Figure 3) resulting in a negative correlation in activity patterns for these species during the summer. Mountain lion activity was most positively correlated, although not significantly, with javelina activity (Figure 5), and that correlation was strongest during the winter (Table 4).

Table 4. Pearson Correlation coefficients (r) and associated P-values between daily activity patterns of mountain lion and the activity patterns of their top six prey species in southern Arizona from March 2001-September 2006. Activity was measured as the number of trail camera photographs taken of each species during each of the 8 time bins throughout the 24-h period. Skunks could not be distinguished to species in all scats and therefore were lumped. Coues white-tail deer Domestic cattle Cottontail rabbit Javelina Skunk spp White-nosed coati Season r P r P r P r P r P r P Winter -0.36 0.388 -0.16 0.697 0.29 0.491 0.54 0.169 0.23 0.587 -0.37 0.366 Summer -0.55 0.158 -0.50 0.203 0.61 0.108 0.49 0.220 0.58 0.130 -0.80 0.016 Overall -0.52 0.185 -0.46 0.250 0.51 0.197 0.57 0.143 0.34 0.414 -0.69 0.057

21

White-tailed Deer Domestic Cow Cottontail Rabbit

250 120 100 200 100 80 80 150 60 60 100 40 40 50 20 20 Number of Photographs of Number Number of Photographs of Number Number of Photographs of Number 0 0 0 0 50 100 150 200 0 100 200 0 50 100 150 200 Mountain Lion Mountain Lion Mountain Lion

Javelina Skunk spp White-nosed Coati

100 120 50 80 100 40 80 60 30 60 40 20 40 20 20 10 Number of Photographs of Number

0 Photographs of Number Number of Photographs of Number 0 0 0 50 100 150 200 0 50 100 150 200 0 100 200 Mountain Lion Mountain Lion Mountain Lion

Figure 5. Correlations between mountain lion activity and prey activity based on trail camera photographs taken during eight time bins throughout the 24-h period in southern Arizona from March 2001 – September 2006. Lines are best fit regression lines. Skunks could not be distinguished to species in all scats and therefore were lumped. 22

DISCUSSION

Mountain lions in southern Arizona showed significant seasonal changes in their daily activity patterns and their diet. Mountain lion activity was crepuscular overall, but lions exhibited more activity during daytime periods in winter and during nighttime periods in summer. Mountain lions could be reducing their daytime activity during the summer months in response to hot daytime temperatures. In other studies, however, mountain lions have not been observed changing their daily activity patterns across seasons. Even in southern latitudes with high diurnal summer temperatures, such as southern Texas (Waid 1990), Peru (Emmons 1987) and Venezuela (Scognamillo et al.

2003) mountain lions remained crepuscular year round and did not show any seasonal shift in activity patterns. This suggests that mountain lions in southern Arizona are either more responsive to seasonal temperature changes than lions in other areas or are responding to something else, possibly changes in the activity patterns of their prey species (Seidensticker et al. 1973, Emmons 1987, Waid 1990, Beier et al. 1995,

Scognamillo et al. 2003).

Seasonal changes in mountain lion daily activity patterns and diet may reflect the daily activity patterns of different prey species, which could ultimately affect their vulnerability and availability to predators (Taylor 1984, Emmons 1987, Sunquist and

Sunquist 1989, Scognamillo et al. 2003). Coues white-tailed deer were the most common prey species comprising 36.8% of the annual diet and 50% of the winter diet. Deer activity was diurnal, changing little between summer and winter. However, the proportion of deer in the mountain lion diet changed dramatically,

23 24 appearing significantly more in winter than in summer. This change could have been a result of increased overlap in the activity of mountain lions and deer during winter months. Mountain lions were crepuscular to nocturnal in the hot summer months, as they have been described at other sites (Emmons 1987, Waid 1990, Beier et al. 1995).

However, during the cooler winter months, mountain lions switched to more diurnal activity. Therefore, during the summer mountain lions were less active during daytime periods when deer were most active. However, with the cooler daytime temperatures during the winter, they shifted towards more daytime activity, and the presence of deer increased dramatically in their diet.

Unlike desert mule deer, which are crepuscular, Coues white-tailed deer are mostly active during daylight hours (Ockenfels and Brooks 1994, Heffelfinger 2006).

These desert-adapted Coues white-tailed deer appear to be able to withstand high daytime temperatures (especially while birthing and raising their fawns), perhaps limiting predation by mountain lions. The decreased overlap in the activity of mountain lions and

Coue’s white-tailed deer may force mountain lions to prey on alternative (nocturnal or crepuscular) prey during the summer.

Domestic cattle were most strongly represented in the mountain lion diet during summer months, but decreased during the winter. Cunningham et al. (1995) observed a similar increase in cattle in mountain lion diet during summer months in central Arizona.

The increase in the proportion of domestic cattle in mountain lion diet during the summer is likely the result of both the inaccessibility of diurnally active Coues white-tailed deer during summer and the availability of domestic cattle calves. Adult cows gave birth to calves in the spring months and calves remained in the area until the fall (D. Bell, pers.

25 comm. – ZZ Cattle Corporation). Since adult cattle average over 500 kg and are generally too large for mountain lions to kill, cattle were only truly available to mountain lions in the summer when calves were abundant (Cunningham et al. 1995, Cunningham et al. 1999). After calves were removed in the fall, cattle significantly decreased in the mountain lion diet, while deer and javelina increased.

Both white-tailed deer and cattle give birth and raise their young in the summer months. The native prey species, having evolved with mountain lions and in a desert environment, may be more adapted to enduring high daytime temperatures and, therefore, limiting predation risks to their young, than domestic cattle. Deer were strongly diurnal in the summer fawning months, despite the extreme daytime temperatures, possibly to avoid or reduce vulnerability to generally crepuscular mountain lions. Also, deer are solitary and more secretive during the fawning period and utilize the anti-predator strategy of concealing their young until fawns are large enough to evade predators (Schwede et al. 1993, Ballard et al. 1999). Deer fawns hide and remain motionless much of their young life, while domestic calves are active and move with their mother from their first day of life, making them much more vulnerable to visually- oriented predators (Cunningham et al. 1995). These differences in adaptation to desert temperatures and rearing strategies between wild and domestic species may have been responsible for the increased occurrence of cattle and decrease of deer in the mountain lion diet during summer.

Javelina appeared in mountain lion diet most prominently in the winter months.

Javelina shifted their daily activity patterns from crepuscular in summer to diurnal in winter, which is consistent with the shifts observed in other studies (Zervanos and Hadley

26

1973, Waid 1990). Perhaps due to the greater degree of overlap in activity patterns during winter, javelina became significantly more prevalent in mountain lion diet at that time.

Changes in the mountain lion diet have been observed when prey availability changes over time (Leopold and Krausman 1986, Rosas-Rosas et al. 2003). For example, mountain lions in Chile prey primarily on guanacos ( Lama guanicoe ), but supplement their diet with European hares ( Lepus capensis ) when guanacos populations were low

(Yanez et al. 1986). However, the dietary change observed in my study was on a seasonal scale, and may be linked to changes in the vulnerability of different prey species as their daily activity patterns change with the seasons. Similar to the seasonal changes observed in this study, Siedensticker (1973) found mountain lions in Idaho switching daily activity patterns to exploit a seasonally available food resource, diurnal ground squirrels, then switching back to hunting mule deer during crepuscular and nocturnal hours the rest of the year.

None of the positive correlations between mountain lion activity and the activity of their top six prey species were significant, suggesting that mountain lions are not closely tracking the activity patterns of any particular prey species. Mountain lion activity was significantly negatively correlated with coati and only in summer when mountain lions were most nocturnal and coatis were diurnal. In fact the slope of the regression line, which should indicate the general relationship between the activity patterns of predator and prey, was negative for three of the six prey species, suggesting mountain lion activity was not strongly influenced by the activity of their primary prey,

27 and emphasizing the importance of temperature fluctuations in the seasonal changes in activity patterns observed in this study.

The diet portion of the study used well-established methods and faced the same biases of other studies that examine fecal remains: that smaller prey species may be overrepresented due to their greater surface area/volume ratios (Ackerman et al. 1984,

Cunningham et al. 1999, Rosas-Rosas et al. 2003). The trail-camera aspect of this study, however, is a fairly recent approach to indexing levels of activity during different times of day (Pierce et al. 1998, Bridges et al. 2004, Dillon and Kelly 2007, McCain and Childs

2008). While trail-camera data may not provide as much information as telemetry with activity sensor or GPS technology, it is cost effective and offers constant monitoring of multiple species at multiple sites. It is not species specific or specific to the marked individuals in a given population, therefore, offering a broader view of a variety of species. Camera placement is, however, of critical importance and caution must be taken in assuming that all individuals and all species are equally detectable at all camera sites.

For example, in this study, cameras were placed at locations most likely to detect mountain lion and jaguar. Some of the prey species may not use the major trails and canyon bottoms to the same degree as the large felids (or may even avoid these areas), and thus the number of photographs of each species likely does not reflect their relative abundance in a region. However, the bias likely only affects the total number of photographs taken of a given species at a given site, not the time of day they are taken.

The activity pattern aspect of this study examined the relative proportions of activity during each of the eight time categories, and regardless of whether a camera site is a particularly productive site for a given species, that species should still have an equal

28 probability of detection during each of the eight time bins. The non-invasive methodologies used during this study allowed for the examination of both diet and activity patterns without having to capture or handle any . The combination of these methodologies provides a low-cost method of addressing interesting ecological and behavioral questions in wildlife biology.

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34

21:36

19:12

16:48

14:24

12:00

TIME 9:36

7:12

4:48

2:24

0:00 001 031 062 092 122 152 182 212 242 272 302 332 362 JULIAN DATE

Appendix A. True sunrise (lower line) and sunset (upper line) times for each day throughout the year at Nogales International Airport, Arizona (Mountain Standard Time).

35

35

30

25

20

15

10 Summer TEMPERATURE (C) 5 Winter Winter 0

y y r ay e ber b ber ruar April M June July b March ugust em Januar e A cem F October e Sept Novem D Month

Appendix B. Average daily temperature recorded within the study area at the Nogales International Airport from 2000-2005 (averaged for all four years). The solid vertical lines separate the six hottest months (16 April – 15 October = summer) from the six coldest months (16 October – 15 April = winter).

36

Appendix C. Species photographed at trail cameras in southern Arizona from March 2001 – September 2006. The total number of photographs was used to calculate detection rates (photographs/100 "trap" nights) for each species. Skunk and squirrel species could not be identified to species in all photographs and therefore were lumped. Detection Rate Number of (Photos / 100 Common Name Scientific Name Photographs "Trap" Nights) American Taxidea taxus 2 0.0097 Black bear Ursus americanus 126 0.6132 Bobcat Lynx rufus 282 1.3723 Coues white-tailed deer Odocoileus virginianus couesi 789 3.8396 Coyote Canis latrans 19 0.0925 Domestic Felis silvestris catus 2 0.0097 Domestic cattle Bos taurus 365 1.7762 Domestic Canis lupus familiaris 92 0.4477 Eastern cottontail rabbit Sylvilagus floridanus 253 1.2312 Grey fox Urocyon cinereoargenteus 449 2.1850 Jaguar Panthera onca 54 0.2628 Javelina Pecari tajacu 351 1.7081 Mountain lion Puma concolor 696 3.3870 Raccoon Procyon lotor 1 0.0049 Ringtail astutus 120 0.5840 Spilogale gracilis, Mephitis mephitis, M. macroura, and 355 1.7276 Skunk spp Conepatus leuconotus Didelphis virginiana Sonoran Opossum californica 172 0.8370 Sciurus arizonensis and 251 1.2215 Squirrel spp Spermophilus variegatus White-nosed coati Nasua narica 144 0.7008 Wild turkey Meleagris gallopavo 13 0.0633