(Canis Latrans Var.) in Cape Breton Highlands National Park, Nova Scotia, Canada

Home , Canso Causeway, Eastern coyote

GENETIC RELATIONSHIPS, MOVEMENT PATTERNS, SPATIAL DYNAMICS

AND DIET OF THE EASTERN COYOTE (CANIS LATRANS VAR.) IN CAPE

BRETON HIGHLANDS NATIONAL PARK

JASON WILFRED BRIAN POWER

Thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science (Biology)

Acadia University Fall Convocation 2015

This thesis by JASON WILFRED BRIAN POWER was defended successfully in an oral examination on 21 September 2015.

The examining committee for the thesis was:

______Dr. Andre Trudel, Chair

______Mr. Mark Pulsifer, External Reader

______Dr. Stephen Mockford, Internal Reader

______Dr. Søren Bondrup-Nielsen, Supervisor

______Dr. Rodger Evans, Acting Head

This thesis is accepted in its present form by the Division of Research and Graduate Studies as satisfying the thesis requirements for the degree Master of Science (Biology)

I, JASON WILFRED BRIAN POWER, grant permission to the University Librarian at Acadia University to reproduce, loan or distribute copies of my thesis in microfilm, paper or electronic formats on a non-profit basis. I, however, retain the copyright in my thesis.

______Author

______Supervisor

______Date

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TABLE OF CONTENTS

LIST OF TABLES ...... vi LIST OF FIGURES ...... x LIST OF APPENDICES ...... xiii ABSTRACT ...... xiv LIST OF ABBREVIATIONS AND SYMBOLS ...... xv ACKNOWLEDGMENTS ...... xvi CHAPTER 1: GENERAL INTRODUCTION ...... 1 BACKGROUND ...... 1 STUDY AREA ...... 6 STUDY RATIONALE ...... 7 REFERENCES ...... 9 PERSONAL COMMUNCATION ...... 14 CHAPTER 2: SPATIAL GENETIC AND BODY SIZE TRENDS IN ATLANTIC CANADA COYOTE (CANIS LATRANS) POPULATIONS ...... 16 INTRODUCTION ...... 16 METHODS ...... 19 RESULTS AND DISCUSSION ...... 21 REFERENCES ...... 28 PERSONAL COMMUNCATION ...... 32 CHAPTER 3: SPATIAL DYNAMICS OF EASTERN COYOTES (CANIS LATRANS VAR.) IN CAPE BRETON HIGHLANDS NATIONAL PARK, NOVA SCOTIA, CANADA ...... 40 INTRODUCTION ...... 40 METHODS ...... 43 RESULTS ...... 47 DISCUSSION ...... 48 REFERENCES ...... 53 PERSONAL COMMUNCATION ...... 56 CHAPTER 4: WINTER FORAGING BEHAVIOUR BY EASTERN COYOTES (CANIS LATRANS VAR.) BASED ON FRACTAL ANALYSIS OF SNOW TRACK- PATTERNS IN CAPE BRETON HIGHLANDS NATIONAL PARK, NOVA SCOTIA, CANADA ...... 64 INTRODUCTION ...... 64

METHODS ...... 66 RESULTS ...... 68 DISCUSSION ...... 69 REFERENCES ...... 74 PERSONAL COMMUNCATION ...... 76 CHAPTER 5: DIET OF EASTERN COYOTES (CANIS LATRANS VAR.) BASED ON EXAMINATION OF SCATS DURING SUMMER AND WINTER IN CAPE BRETON HIGHLANDS NATIONAL PARK, NOVA SCOTIA, CANADA ...... 82 INTRODUCTION ...... 82 METHODS ...... 87 RESULTS ...... 91 DISCUSSION ...... 95 REFERENCES ...... 101 PERSONAL COMMUNCATION ...... 106 CHAPTER 6: GENERAL CONCLUSION ...... 117 REFERENCES ...... 122

LIST OF TABLES

CHAPTER 2

Table 2.1. Summary of genetic diversity observed in the five sampled locations in

Atlantic Canada ...... 33

Table 2.2. Pairwise FST values of coyote mtDNA CR sequences for five locations in

Atlantic Canada and their associated p-values. FST values are above the diagonal; p- values are below the diagonal. The total FST value for all samples was 0.275 (p < 0.001)

...... 34

Table 2.3. Adult female morphological characteristics ± 1 SD grouped by mtDNA haplotype. All samples are from Nova Scotia. There were no statistically significant differences between groups defined by haplotype (i.e., cla28 vs. GL20) ...... 35

Table 2.4. Adult male morphological characteristics ± 1 SD grouped by mtDNA haplotype. All samples are from Nova Scotia. There were no statistically significant differences between groups defined by haplotype (i.e., cla28 vs. GL20) ...... 36

CHAPTER 3

Table 3.1. Coyotes captured in Cape Breton Highlands National Park between October

2011 and July 2013. A = adult; Y = yearling; M = male; F = female ...... 57

Table 3.2. Available and use of high human use biotopes to coyotes in Cape Breton

Highlands National Park and the community of Chéticamp from October 2011 to January

2014.Chi-square statistic shows where differences exist between low and high human presence. M = Male; F = Female; CH = Cape Breton Highlands Coyote ID ...... 58

Table 3.3. Summary of daylight and dark 2-hour movement averages for collared coyotes in Cape Breton Highlands National Park and the community of Chéticamp from October

2011 to January 2014. M = Male; F = Female; CH = Cape Breton Highlands Coyote ....59

Table 3.4. Summary of daylight and dark 2-hour movement averages for collared coyotes in Cape Breton Highlands National Park and the community of Chéticamp from October

2011 to January 2014 ...... 60

CHAPTER 4

Table 4.1. Proportion of distance travelled in each habitat ...... 77

CHAPTER 5

Table 5.1. Calendar season frequency (# scats/mean %) of occurrence of prey remains identified from coyote scats collected in Cape Breton Highlands National Park from May

2012 through August 2013. A Chi-squared test was used to test for differences between seasons of prey remains ...... 107

Table 5.2. Regional frequency (# scats/mean %) of occurrence of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Chi-squared test was used to test for differences between regions of prey remains ...... 108

Table 5.3. Calendar season percent by volume (mean% ± SD) of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Kruskal-Wallis test was used to test for differences between calendar seasons of prey remains ...... 109

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Table 5.4. Regional percent by volume (mean% ± SD) of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from

May 2012 through August 2013. A Kruskal-Wallis test was used to test for differences between regions of prey remains ...... 110

Table 5.5. Calendar season richness of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Pearson’s Chi-squared Test was used to determine if there was a statistically significant difference between calendar seasons for prey richness ...... 111

Table 5.6. Regional richness of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August

2013. A Pearson’s Chi-squared Test was used to determine if there was a statistically significant difference between regions for prey richness ...... 112

Table 5.7. Relative frequency of occurrence (%) of prey species observed within three 3 meters of both back-tracking coyotes and winter transects in CBHNP from January to

April 2013 ...... 113

Table 5.8. Moose (Alces alces) body condition at eastern coyote scavenging sites in

CBHNP from January to April 2013 ...... 114

CHAPTER 6

Table 6.1. Differences between coyotes of Cape Breton Highlands National Park and those within the community of Chéticamp ...... 120

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Table 6.2. Differences among coyotes in Cape Breton Highlands National Park and

Atlantic Canada ...... 121

LIST OF FIGURES

CHAPTER 1

FIGURE 1.1. The location of the study area, Cape Breton Highlands National Park, in northern Cape Breton Island, Nova Scotia...... 14

CHAPTER 2

Figure 2.1. Mitochondrial haplotype frequencies of eastern Coyotes from Atlantic

Canada (New Brunswick, mainland Nova Scotia, Cape Breton Island, Prince Edward

Island, and Newfoundland) ...... 37

0.30 * chest (cm), 0.34 * body length (cm), 0.31 * tail length (cm), 0.22 * front paw width (mm), 0.32 * front paw length (mm), 0.17 * back paw width (mm), 0.29 * back paw length (mm), 0.30 * shoulder (cm), 0.36 * skull width (cm), 0.36 * skull length (cm).

As all measurements load in the same direction, this component can be interpreted as representing general size differences. Boxes on this plot represent 25th and 75th percentiles, and the line represents the 50th percentile. Whiskers extend to the highest and lowest values that fall within 1.5 * the inter-quartile range. Data beyond the ends of whiskers are outliers, represented as points ...... 38

0.30 * chest (cm), 0.17 * body length (cm), 0.25 * tail length (cm), 0.34 * front paw width (mm), 0.39 * front paw length (mm), 0.37 * back paw width (mm), 0.38 * back paw length (mm), -0.02 * shoulder (cm), 0.21 * skull width (cm), 0.39 * skull length

(cm). As all measurements load in the same direction, this component can be interpreted as representing general size differences. Boxes on this plot represent 25th and 75th percentiles, and the line represents the 50th percentile. Whiskers extend to the highest and lowest values that fall within 1.5 * the inter-quartile range ...... 39

CHAPTER 3

Figure 3.1. Cumulative area of use curve (km2; 95% MCP) with sequential locations of

GPS collared coyotes in Cape Breton Highlands National Park and the community of

Chéticamp between October 2011 and January 2014 ...... 61

Figure 3.2. Cumulative area of use curve (km2; 95% MCP) with sequential locations of

CH-5 GPS-radio collared coyote in Cape Breton Highlands National Park between

November 2011 and October 2012 ...... 62

CHAPTER 4

Figure 4.1. Map of French Mountain showing the coyote paths followed and recorded in this area during the winter ...... 78

Figure 4.2. Mean (net distance travelled)2 versus the number of steps (n). The dotted black curve represents the observed data, the dotted line 95% confidence intervals, and the dashed line the expected curve from a CRW. The coyote trails are significantly straighter than a CRW………… ...... 79

Figure 4.3. Fractal analysis results versus spatial scale. (A) Fractal dimension and (B) correlation in fractal dimension between successive track segments. The dotted lines represent 95% confidence intervals. The increased Fractal dimension with scale (A) shows that coyote paths are more tortuous at larger spatial scales. The vertical gray bar in

(B) shows where the plot crosses the y-axis, going from a positive to negative correlation as scale increases...... 80

Figure 4.4. Mean fractal dimensions of coyote paths in different habitat types, measured at different spatial scales. The bars are 95% confidence intervals. For each plot, paths in habitats with the same letter did not have significantly different fractal dimensions ...... 81

CHAPTER 5

Figure 5.1. Transect locations identified for scat and prey observations in Cape Breton

Highlands National Park. Blue dots represent transect locations (18 park trails and 3 gated roads) ...... 115

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LIST OF APPENDICES

CHAPTER 3

Appendix 3.1. Reclassification of 37 biotopes from the Nova Scotia Department of

Natural Resources land cover inventory to 5 biotopes used for biotope use ...... 63

CHAPTER 5

Appendix 5.1. Description of moose (Alces alces) carcasses at eastern coyote scavenging sites in Cape Breton Highlands National Park from January to April 2013 ...... 116

xiii

ABSTRACT

The first North American adult human fatality caused directly by coyotes, coupled with seemingly high levels of aggressive coyote encounters with visitors in Cape Breton Highlands National Park, formed the rationale for this study, which has been funded by Parks Canada. The objective was to determine if coyotes within the park differ ecologically and behaviourally from other regions in northeastern North America, which might explain the heightened aggression toward humans. The eastern coyote originated from coyotes in the west, which expanded their range east and hybridized with Canis lycaon, the eastern wolf. The eastern coyote is larger than its western counterpart and has adapted to live in packs, predating large ungulates, including moose (Alces alces), possibly as a result of this hybridization. This study investigated the degree of wolf mitochondrial DNA in eastern coyotes from the Atlantic Provinces, movement patterns, use of high human use areas and diet of coyotes in Cape Breton Highlands National Park. DNA samples collected from Nova Scotia, Prince Edward Island, New Brunswick and Newfoundland coyote’s revealed mitochondrial DNA haplotypes characteristic of both C. latrans and C. lycaon with decreasing haplotype diversity consistent with sequential founder events moving from west to east and on islands. Coyotes from Nova Scotia with Eastern Wolf mitochondrial DNA are significantly larger than male eastern coyotes from the same region with coyote mitochondrial DNA. Eleven coyotes had GPS collars attached to investigate their movement patterns. Collared coyotes on the plateau of the park tended to move over larger areas than did coyotes at lower elevation. Human use habitats were avoided more in the community of Chéticamp than in the park, especially on the plateau. Daily movements differed among individuals, however coyotes in the community of Chéticamp moved significantly less during daylight hours compared to animals within the park. Fifty-seven coyote tracks were followed in the snow during the winter of 2013. Their movement paths were analysed using fractal analysis. Coyote paths were significantly straighter than correlated random walks. Path tortuosity was very low but increased with spatial scale, with no discrete breaks in tortuosity at any scale. During the winter, coyotes appeared to move along relatively straight paths scavenging on dead moose. A diet study was conducted to determine what food items coyotes consumed by season and location within the park. Coyotes appear to rely on moose throughout the year with a greater dependency during the winter months. The diversity of prey varied between the lowlands and highlands with the lowlands seeing greater diversity of prey items throughout the year. Several coyotes live within close proximity of Chéticamp. Here they likely access human food available in the form of garbage, but there have been no aggressive human-coyote encounters, and individuals show reduced daytime activity. Within the park, especially on the plateau, coyotes roam over larger areas than they do at low elevations in the vicinity of Chéticamp, they are active during the day, and rely heavily on moose for their diet. This sets the stage for aggressive encounters with humans during the high park visitation period of June to October. There is potential for visitors to directly or indirectly feed the coyotes, potentially habituating them and removing the sense of threat previously associated with interactions with humans.

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LIST OF ABBREVIATIONS AND SYMBOLS

NE North eastern mtDNA Mitochondrial DNA bp Base-pair df Degrees of freedom n Sample size SD Standard deviation U Mann-Whitney Statistic χ2 Chi-square CBHNP Cape Breton Highlands National Park KNP Kejimkujik National Park NS DNR Nova Scotia Department of Natural Resources GIS Geographic Information System GPS Global Positioning Systems CRW Correlated Random Walk VHF Very High Frequency

ACKNOWLEDGMENTS

To my supervisor, Dr. Søren Bondrup-Nielsen, thanks for providing your valuable insights to this project despite the many challenges that came our way. Dr. Donald T.

Stewart, your genetics knowledge was critical to completing this research. Thanks to

Daren Keizer, Patrick Sanderson, Wesley Pitts, Cecile Bossi, Greg Giffin, Chris

Englehart, Nathalie LeBlanc, and Seumas Nairn, for helping out in so many ways.

Thanks to Brent Patterson, Stan Gehrt, Erich Muntz, Vilis Nams, Dave Shutler, Elisabeth

Frost, Lisa Taul, Pam Mills, Dave Farrell, Emma Tichenor, Mike Boudreau, Mike

O’Brien, Gerald Bourgeois, Clarence Barrett, Archie Doucette, and Evan Wilson for various contributions to this project, both in and out in the field. Funding was provided by Parks Canada, Acadia University, Nova Scotia Department of Natural Resources-

Wildlife Division and an NSERC Discovery grant to D. T. Stewart. To my family: thanks for supporting me throughout this long and tedious journey! I couldn’t have done it without you. Finally thanks to Emily for being there for me, and encouraging me along the way.

This thesis is dedicated in memory of Emma Kate Tichenor, for her love and passion for wildlife and for her encouragement to me throughout my degree - you will be missed.

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Chapter 1: General Introduction

Background

The coyote (Canis latrans Say) has steadily expanded its North American range; populations were historically limited to the prairie region of the United States west of the

Mississippi river, portions of Mexico, and southern central Canada (Bekoff 1977; Parker

1995; Patterson 1995; Fener et al. 2005). During the latter part of the nineteenth century coyotes’ use of fragmented landscapes (primarily caused by human settlements), together with a reduction in predators and competitors (i.e., wolves and bears), made it possible for coyotes to extend their geographic range, which now spans most of North America from Newfoundland to Yukon (Canada) to Panama (Moore and Millar 1984;

Sabean 1993; Parker 1995; Hidalgo-Mihart et al. 2004; Schrecengost et al. 2009). An additional factor involved in the successful spread of coyotes is their ability to adapt to various environments and to an increased human presence. Coyotes can live in a variety of landscapes, including open grasslands (Atwood and Weeks 2003), agricultural areas

(Atwood et al. 2007; Gehrt 2007; Gehrt et al. 2009), wetlands (Atwood et al. 2004,

2007), desert terrains (Howard and DelFrate 1991), urban human settlements (Gehrt

2007; Gehrt et al. 2009) and forested landscapes (Atwood and Weeks 2003; Gosselink et al. 2003; Atwood 2006; Atwood et al. 2007).

Coyotes arrived in eastern North America via two geographically separate range expansion fronts: the “southern front” and the “northern front” (summarized by Parker

1995). The southern front originated in Louisiana and Arkansas, south of the Great

Lakes, and the northern front originated from Minnesota, Wisconsin, and Manitoba and extended eastward along the Canadian trans-continental railway during the late 1920’s to

early 1930’s (Parker 1995). Today, coyotes that live in eastern North America are known as eastern coyotes (Canis latrans var.), and have the largest body size of any existing coyote population (Thurber and Peterson 1991; Lariviere and Crete 1993; Peterson and

Thurber 1993). This larger body size may have resulted from interbreeding between the expanding coyote population and eastern wolves, Canis lycaon, (in the northeast), and red wolves Canis rufus, (in the southeast; reviewed in Parker 1995). Some (Geist 1987;

Thurber and Peterson 1991) suggest the increase in body size may have been due to selection for larger animals as a result of access to larger prey, specifically white-tailed deer (Odocoileus virginianus).

In the Canadian Atlantic Provinces, coyotes were first documented in New

Brunswick in 1958 (Squires 1968; Schrecengost et al. 2009), but their population remained low until the mid-1970’s (Moore and Millar 1986). Coyotes were likely in

Nova Scotia by the mid 1970’s, the first confirmed coyote was trapped in Guysborough

County in 1977 (O’Brien 1983; Moore and Millar 1986). First confirmed sighting in

Cape Breton Highlands National Park (CBHNP) occurred in 1980 (E. Muntz pers. comm.

2013). The Nova Scotia coyote population increased rapidly throughout the 1980’s; during the 1992-93 trapping season the harvest of coyotes was 1,270 animals (Sabean

1993), and in 2010-11 the harvest was over 2,600 animals (this increase is likely related, in part, to a trapping incentive; M. Boudreau pers. comm. 2013).

The coyote colonization in Nova Scotia has impacted a number of species.

Coyotes have been blamed for population reductions among popular game species including white-tailed deer and snowshoe hare (Lepus americanus). They have also been blamed for attacks on livestock and pets, similar to the United States (Bounds and Shaw

1994), and generalized conflict with humans (White and Gehrt 2009). More recently, in

2009, the first North American adult human fatality due to coyotes occurred in CBHNP

(CBC 2009). This was only the second coyote-caused human fatality documented in

North America; the first fatality was a child in a suburban area of Glendale California

(attack occurred in 1981; Howell 1982). Despite the extremely low incidences of human fatalities, Parks Canada has noted an increase in bold behaviour by coyotes toward humans since the 1990’s in CBHNP (E. Muntz pers. comm. 2013) and resource conservation managers have recognized a need to investigate possible causes to inform future management decisions designed to mitigate risk to park visitors.

Between 2003 and 2012 there have been eight documented incidents of fearless behaviour (i.e., a coyote that does not back away when approached by a human, or approaches a human but can be scared off), 16 acts of aggressive behaviour (i.e., a coyote that will not run away when people try to scare it, runs at people, approaches people multiple times, or circles or growls at people, but never making physical contact), and six attacks (i.e., a coyote that lunges at someone, attempts to make physical contact, or succeeds at making physical contact (E. Muntz pers. comm. 2013)). These human-coyote interactions have been primarily attributed to habituation, where coyotes exhibit diminished fear towards humans. One well documented source of coyote habituation occurs via food conditioning (Carbyn 1989; Bounds and Shaw 1994; Alexander and

Quinn 2011; Lukasik and Alexander 2011). When humans feed coyotes (directly or indirectly), coyotes’ fear of humans decrease and over time they associate humans with food (or trash and pets; Loe and Roskaft 2004). This increased aggression towards humans is what triggered Parks Canada to determine why coyotes in CBHNP seem to

behave differently than in other North American areas. Parks Canada investigation into this involves human dimensions, hazing and ecology. The study reported here focuses on coyote ecology.

The study of coyote ecology in the wild requires methodology such as scat analysis, snow tracking and radio telemetry. Radio telemetry has been a popular and convenient way to observe coyote movements (Dowd 2010; Judy 2010), biotope selection and home ranges. This form of monitoring involves trapping or snaring coyotes and immobilizing them before attaching Global Positioning System (GPS) or Very High

Frequency (VHF) radio collars. Very High Frequency collars allow coyotes to be located by triangulating via a VHF receiver, and the benefit of VHF collars is their low cost.

However, obtaining many reliable locations is time consuming, costly and requires trained personnel.

Food selection and hunting behaviours of coyotes can be quantified by indirect observations (i.e., snow tracking), identification of digestive tract contents post-mortem, or by examining individuals’ scats. In a park setting the best methods for determining diet are snow tracking and scat analysis because both methods are non-lethal. Unique hunting behaviours (revealed through snow tracking) can reveal information about prey sources; for example, during the hunt for snowshoe hare, coyotes have been observed to have short chase sequences with many sudden changes in direction, roughly following a zigzag pattern (Thibault and Ouellet 2005).

Fractal analysis is a useful way to characterize movement patterns. This technique can be used to infer changes in hunting behaviours at different spatial scales and in different vegetation types. Mandelbrot (1977) first introduced fractals to examine forms

that have a similar structure at all spatial scales, the common example being coastlines that could not be described in Euclidean forms (lines, circles or ellipses). Fractal dimension is a measure of roughness and irregularity of an object and is calculated by plotting the log of the step size (ruler size) on the x-axis and the log of the path length on the y-axis. Steeper curves indicate a more irregular or tortuous path. Animals may use various vegetation types (biotopes) within their home range, and the resulting tracks may differ depending on their activities (i.e., hunting vs. travelling).

Although coyotes are known to consume a variety of foods, they maintain a predominantly carnivorous diet (Andelt et al. 1987; Patterson 1995). Overall, coyotes are classified as opportunists, whose generalist diet varies both geographically and seasonally based on resource availability and reproductive cycles (Todd and Keith 1983;

MacCracken and Uresh 1984; Dowd 2010). Studies regarding the feeding habits of the eastern coyote have verified that white-tailed deer and snowshoe hare are primary food sources (Patterson et al. 1998; Schrecengost et al 2008). In Cape Breton, these prey items are believed to form the majority of coyote diets; however, if these food sources are absent or low in abundance, coyotes will shift their diet (Patterson 1995). Depending on the area, small mammals, fruit and berries may also be included in a coyote’s diet at certain times of the year (Fedriani et al. 2001). Coyotes may also take advantage of anthropogenic foods, although most studies suggest this is a small portion of their diets

(Moore and Millar 1986; Fedriani et al. 2001; Gehrt et al 2011; Grigione et al. 2011).

In Atlantic Canada the white-tailed deer has historically been the primary ungulate food source of coyotes (Parker 1995; Patterson 1995), but scat analysis of the eastern coyote has confirmed their consumption of moose (Alces alces). In the past, these

traces were assumed to be the result of coyotes feeding on carrion or taking occasional moose calves (Major and Sherburne 1987; Litvaitis and Harrison 1989; Dumond et al.

2001; Richer et al. 2002; Boisjoly et al. 2010; Benson and Patterson 2013). However,

Benson and Patterson (2013) found evidence of eastern coyotes and eastern coyote-wolf hybrids having killed adult moose in Ontario.

Study Area

The study area included most of CBHNP (Figure 1), a 950 km² wilderness area located in northern Cape Breton Island, Nova Scotia, Canada (46.7167 N, 60.6597 W).

An average of 150 000 people visit CBHNP each year (E. Muntz pers. comm. 2013) and over 25 000 people hike the Skyline trail – which is also the location of the 2009 fatality

(A. Boudreau pers. comm. 2013). Cape Breton Island has a cool maritime climate and the area is characterized by a rugged topography. The park is bordered by the Gulf of Saint

Lawrence in the west, where the land rises sharply from the ocean to a height of 500 m, forms a large plateau that slopes eastward to the Atlantic Ocean. There are four well- defined seasons with an average of 337 cm of snowfall and 1053 mm of rainfall per year.

The average winter temperature is approximately -5°C (but frequently drops below -

10°C) while summer brings an average temperature of 18°C (sometimes peaking above

30°C; Environment Canada 2013). An important discrepancy in snowfall and temperature has been observed between the plateau and lowlands: snowfall on the plateau is greater and persists longer, and temperature averages are usually lower year round (J. Bridgland pers. comm. 2013).

Three distinct vegetation zones exist within the park: Acadian forest, boreal forest and taiga (thick scrubby black spruce). These three forest types also provide unique

biotopes for a number of species. White-tailed deer exist in the park in low numbers and are restricted to the lowlands. Snowshoe hare are normally abundant in the park, however, hare populations cycle approximately every 10 years and during the time frame of this project population numbers were in the low phase. Moose were extirpated from

Cape Breton Island in the early 1900’s and reintroduced to CBHNP in the late 1940’s

(Bridgland et al. 2007; Smith et al. 2010). Moose could therefore have been a potential food source for incoming coyotes in the 1980’s. The population of moose in the highlands of Cape Breton is estimated at approximately 7 500 animals (J. Bridgland pers. comm. 2013; M. Smith pers. comm. 2015). Moose have altered forest regeneration in

CBHNP, thereby altering some forest landscape to grasslands. Forests have faced further alteration via outbreaks of the spruce budworm (Choristoneura fumiferana), with the last major outbreak occurring in the early 1970’s, decimating 67% of fir trees in the park (E.

Muntz pers. comm. 2012).

Study Rationale

The motivation for this study was in response to the human fatality in CBHNP and the apparent high level of coyote aggression towards humans in the park. The central question was to determine if coyote ecology in CBHNP is different from elsewhere in northeastern North America, which might make them prone to show increased aggression toward people. Thus far, ecological studies of coyotes in Nova Scotia have offered little insight into the dynamics of coyote populations in northern Cape Breton.

This study represents a cooperative effort between Parks Canada, Acadia

University and the Nova Scotia Department of Natural Resources (DNR) - Wildlife

Division. The objectives of the study were to: (1) determine the degree of coyote-wolf

hybridization in the Atlantic provinces, (2) describe differential use of space by coyotes on the CHHNP highland plateau where aggression seems to occur and lowland areas near human settlement where reports of aggression are absent, (3) determine use of different vegetation types and travel behaviour through fractal analysis of individual coyote snow tracks and (4) document the diet of coyotes within the park and surrounding area to see if there are patterns which might offer a better understanding of the recent increase in aggressive behaviour toward humans.

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Personal Communication

Allan Boudreau, Research Officer, Parks Canada, Halifax, Nova Scotia.

Michael Boudreau, Human-Wildlife Conflict Biologist, Nova Scotia Department of Natural Resources-Wildlife Division, Kentville, Nova Scotia.

James Bridgland, Park Ecologist, Parks Canada Ingonish, Nova Scotia

Erich Muntz, Project Manager, Parks Canada, Chéticamp, Nova Scotia

Figure 1.1. Study area, Cape Breton Highlands National Park, Cape Breton Island, Nova Scotia.

Chapter 2: Spatial Genetic and Body Size Trends in Atlantic Canada Coyote (Canis

latrans) Populations.1

Introduction

Coyotes, Canis latrans Say, were historically restricted to the Great Plains region of

North America west of the Mississippi River (Banfield 1974, Stains 1975) but coyotes now inhabit almost all regions between Alaska and Costa Rica (O’Brien 1983; Harrison

1992; Parker 1995; Hidalgo-Mihart et al. 2004; Fener et al. 2005). During the latter part of the 19th century, coyotes’ range expanded throughout southern Canada and the Great

Lakes region, eventually reaching New Brunswick by 1958, mainland Nova Scotia by

1977, Cape Breton Island by 1980, Prince Edward Island by 1983 and Newfoundland by

1985 (Parker 1995). Nova Scotia is connected to New Brunswick (and the rest of North

America) via the small Isthmus of Chignecto, which has served as a dispersal corridor for several large mammals into the province including coyotes (Scott and Hebda 2004).

Coyotes have demonstrated phenomenal dispersal ability at several biogeographic scales

(Bekoff 1977; Parker 1995; Patterson 1995) and have been photographed on ice flows in the Northumberland Strait and the Gulf of St. Lawrence (McGrath 2004). Cape Breton

Island, Prince Edward Island and Newfoundland are isolated islands, historically inaccessible by bridges. It is thought that coyotes arrived on these islands via ice flows from the mainland (i.e., Prince Edward Island and Cape Breton Island were colonized from a mainland source) or via ice flows from another island (i.e., Newfoundland may have been colonized from Cape Breton Island).

1 Jason W. B. Power, Nathalie LeBlanc, Søren Bondrup-Nielsen, Mike J. Boudreau, Mike S. O’Brien and Donald T. Stewart. 2015. Spatial Genetic and Body Size Trends in Atlantic Canada Coyote (Canis latrans) Populations. Northeastern Naturalist (in press).

Extensive range expansion of coyotes into eastern North America occurred after a decrease in the Canis lupus L. (gray wolf) population resulting from their local extirpation by over harvesting and habitat destruction (Thurber and Peterson 1991).

Although the phylogenetic and phylogeographic history of the genus Canis in North

America is complex, it appears that western coyotes dispersed east, where they hybridized extensively with Canis lycaon Schreber, commonly referred to as the eastern wolf (e.g., Wheeldon et al. 2010, Monzon et al. 2014), the great lakes wolf (e.g.,

Koblmuller et al. 2009; Kays et al. 2010; vonHoldt et al. 2011), or the algonquin wolf

(e.g., Chambers et al. 2012). In this paper, we use the names C. lycaon and eastern wolf for this taxon.

Eastern coyote colonization occurred at a particularly fast rate in Ontario, where they were exposed to eastern wolf populations. Their more southerly route through Ohio has been studied extensively (e.g., Leonard and Wayne 2008; Koblmuller et al. 2009;

Schwartz and Vucetich 2009; Wheeldon and White 2009; Wilson et al. 2009; Kays et al.

2010; Benson et al. 2012). In contrast to earlier hypotheses that eastern coyotes in Ohio did not hybridize with eastern wolves or with gray wolves, recent assessment of eastern coyote genomes by Monzon et al. (2014) indicates widespread admixture of eastern wolf, gray wolf, and even domestic dog (Canis lupus familiaris L.). Monzon et al. (2014) have proposed several plausible, yet presently untested, hypotheses to account for these patterns of admixture of the nuclear genomes of these various canid species.

In terms of mitochondrial DNA, there is an asymmetric pattern of matings among canid species resulting in a discordant pattern of mtDNA introgression compared to the nuclear genome (Monzon et al. 2014). Eastern coyote populations possess primarily

western coyote (C. latrans) mitochondrial DNA (mtDNA) with varying degrees of introgressed eastern wolf (C. lycaon) mtDNA (e.g., Way et al. 2010). Perhaps not surprisingly, eastern coyotes are morphologically distinct from their ancestors in the west. Eastern coyotes are the largest morphotype within C. latrans (Gompper 2002; Way and Proietto 2005; Way 2007), weighing ~15-20% more than typical western coyotes

(Blake 2006). Body size in eastern coyotes may reflect selection for eastern wolf alleles or mtDNA haplotypes that contribute to larger body size, possibly in response to an increase in predation on larger prey (e.g., white-tailed deer, Odocoileus virginianus) compared to the diet of western coyotes (Kays et al. 2010; Wheeldon et al. 2010; Monzon et al. 2014). Indeed, Benson and Patterson (2013) recently demonstrated four definitive cases of eastern coyote and (or) eastern coyote x eastern wolf hybrids killing moose

(Alces alces).

Although hybridization is a natural evolutionary process, it may also be an indicator of environmental change over time (e.g., Rutledge et al. 2010). Mitochondrial DNA, which is inherited through the maternal lineage, can be used to track such hybridization events (e.g., Nunes et al. 2010). Distinct mtDNA sequences, called haplotypes, can be used as one genetic marker in the analysis of other traits such as morphology, behaviour, health, etc. For example, Kays et al. (2010) showed that male eastern coyotes with C. lycaon mtDNA haplotypes exhibited significantly wider skulls than eastern coyotes with

C. latrans mtDNA haplotypes in northeastern North America.

With this background in mind, the objectives of this study were: (1) to determine the diversity and frequency of mtDNA control region haplotypes for coyotes in Atlantic

Canada, particularly Cape Breton Island and mainland Nova Scotia and (2) to investigate

morphological differences between groups of eastern coyotes in Nova Scotia defined by mtDNA haplotype sequences.

Methods

Sample collection and study area The majority of samples collected and analyzed in this study were from mainland

Nova Scotia (n=94) and Cape Breton Island (n=77), Canada. Samples were obtained from fur harvesters in Nova Scotia who took part in a legal harvest and voluntary coyote carcass collection program. These carcasses were brought to regional offices of the Nova

Scotia Department of Natural Resources (NSDNR) during the 2010 to 2013 harvest seasons. Additional tissue samples were obtained for New Brunswick (n=11), Prince

Edward Island (n=2), and Newfoundland (n=14) from the North American Fur Auction depot in Truro, Nova Scotia, Canada with collection data recorded from trappers upon pick up. No Canadian Council of Animal Care protocol review was required for this work as all samples were obtained from specimens that had already been killed for reasons other than this research program.

Detailed information on sex, geographic location, date and method of collection was obtained for as many animals as possible. Sampling locations were recorded to a 1 km2 grid using the Nova Scotia Atlas (Service Nova Scotia 2006). Carcasses were stored at

-20ºC until shipped to the NSDNR-Wildlife Division in Kentville, Nova Scotia, for processing. A tissue sample from each carcass was collected for genetic analysis. For carcasses that had front and back foot pads intact, we measured relaxed left foot length in mm (length from the back edge of the heel pad to the outer tip toe pad) and width in mm

(distance from the outer edges of outer toe pad); the right foot was measured in cases where the left foot was missing. Carcasses turned in with intact pelts were weighed with the pelt

on and then skinned. I weighed each skinned carcass and measured the body length (length from tip of nose to base of tail along the dorsal midline contour), chest girth (circumference around torso behind shoulders), skull length (length from the incisors to the tip of sagittal crest) and width (distance from the outer edges of the zygomatic arch). Carcass masses predominately reflect winter weight based on harvest dates (trapping season). I omitted incomplete carcasses for weight comparisons. Lower jaws were extracted from skulls and two lower canines were removed by simmering in hot water for ~15 min. Adult coyotes were identified from radiographs of canine teeth as those individuals with a pulp cavity- tooth width ratio of ≤ 0.45 as measured ~15 mm from the tip of the root (Knowlton and

Whittemore 2001) and by submitting a lower canine to Matson’s Laboratory (Montana,

USA) for cementum analysis to confirm age (Ballard et al. 1995). I measured lower canine total length and maximum width prior to submitting specimens for analysis. We identified

56 adults in our sample (25 females and 31 males).

Genetic analysis I extracted DNA from tissue samples from 199 carcasses using a DNeasy Blood and

Tissue Kit (Qiagen). For each sample, I amplified and sequenced a 343-347 base pair

(bp) fragment of the mitochondrial DNA (mtDNA) control region using published primers (AB13279: Pilgrim et al. 1998; AB13280: Wilson et al. 2000). PCR products were sent to the McGill University and Génome Québec Innovation Centre for sample sequencing. I edited mtDNA sequences to 223-228 bp in length using Jalview version 2.8

(2012). The HKY substitution model was determined as the best-fitting model of DNA evolution for these samples, using the Akaike information criterion in Modeltest (Posada and Crandall 1998). I used this model to construct a neighbor-joining tree in MEGA5

version 2.2 (Tamura et al. 2011), and assigned haplotypes, denoted as cla28, cla29 and

GL20, corresponding to previously described sequences (Kays et al. 2010).

Genetic diversity was assessed by measuring relative polymorphism levels using nucleotide diversity, haplotypic diversity (π) and θ as implemented in dnaSP 5 (Librado and Rozas 2009). We used Arlequin v.3.5 (Excoffier and Lischer 2010) to calculate overall genetic variance within and among populations using an analysis of molecular variance (AMOVA), as well as to calculate pairwise Fst values between all populations.

Morphological comparisons We compared body measurements of 56 adult eastern coyotes (25 females and 31 males) from mainland Nova Scotia and Cape Breton Island. Differences in body measurements between groups defined by haplotypes were assessed using Student’s t- tests and analysis of variance (ANOVA) as implemented in Excel and the R package aov

(R Development Core Team 2008), respectively. Morphological differences were also assessed using principal component analysis, followed by analysis of variance (ANOVA) and Tukey's Honest Significant Differences, as implemented in the R packages prcomp, aov and TukeyHSD, respectively. For each haplogroup and sex combination, missing data (which comprised 19% of total male data and 4% of total female data) were replaced by the variable mean (e.g., Pigott 2001) and variables were scaled to have unit variance.

Results and Discussion

Genetic analyses The sequences obtained in our study are consistent with previously named haplotypes

(Kays et al. 2010; Figure 2.1) and show that eastern coyote populations in Atlantic

Canada have both western coyote and eastern wolf mtDNA. The Atlantic Canada population of eastern coyotes carries one of two C. latrans haplotypes (cla28 or cla29) or

a C. lycaon haplotype (GL20; Figure 2.1), consistent with the haplotype distribution found in northeastern US populations (Kays et al. 2010). The presence of GL20 is the product of recent hybridization between C. latrans and C. lycaon and introgression of eastern wolf mtDNA into eastwardly dispersing populations of eastern coyotes (Figure

1).

Several generalizations about mitochondrial diversity in Atlantic Canada eastern coyotes can be drawn from our analysis although in some cases these observations must be considered preliminary due to small sample size (e.g., n=2 for Prince Edward Island).

Nucleotide diversity (Table 2.1) in New Brunswick was comparable to levels found previously in Ohio populations, while Nova Scotia diversity was similar to northeast populations (Kays et al. 2010). Diversity was reduced in Cape Breton samples, and

Newfoundland samples had no haplotype variation. Observed diversity levels east of

New Brunswick are consistent with populations founded by a small number of migrants.

Pairwise FST values reveal significant differences between Newfoundland and each of the following regions: Cape Breton Island, Nova Scotia, Prince Edward Island and New

Brunswick, indicating historical bottlenecks and/or restricted gene-flow between

Newfoundland and the southern provinces (Table 2.2). Similarly there is evidence of genetic differentiation between the New Brunswick and the Cape Breton Island populations (FST 0.31; p-value < 0.001). The FST value between the Nova Scotia and

Cape Breton Island populations is lower, though still significantly different from zero

(FST 0.05; p-value <0.05). There is likely some on-going genetic exchange between mainland Nova Scotia and Cape Breton Island directly or indirectly caused by the construction of the Canso Causeway connecting mainland Nova Scotia to Cape Breton

Island. As a consequence of the causeway, currents through the Strait of Canso are severely impeded leading to a build-up of ice across the strait in winter. This has facilitated the invasion of Cape Breton Island by several meso-carnivores including red fox (Vulpes vulpes) raccoon (Procyon lotor), striped skunk (Mephitis mephitis), bobcat

(Lynx rufus), coyote (Scott and Hebda 2004) and fisher (Martes pennanti), as well as other mid-sized mammals such as North American porcupine (Erethizon dorsatum; T.

Power, pers. Comm. 2015).

The FST value between Nova Scotia and New Brunswick borders on significance (FST

0.09; p-value = 0.06). Populations in these provinces likely exchange some genes as

Nova Scotia and New Brunswick are connected by the Isthmus of Chignecto, which does not appear to be a significant barrier to coyote dispersal (Scott and Hebda 2004). Other population pairs show no statistically significant genetic structure (Table 2.2). The particularly small sample size for the Prince Edward Island population makes it difficult to draw inferences about population structure in that province. The reduced nucleotide diversity observed in populations located further east, as well as the haplotypes present, supports previous hypotheses proposed for this species; i.e., that eastern coyote populations in these regions were founded by relatively few individuals who migrated from the southwest and that gene flow is likely impeded by water (e.g., Parker 1995;

Kays et al. 2010).

New Brunswick samples did not exhibit any of the rare haplotypes that Kays et al.

(2010) detected in their northeastern samples. The absence of rare alleles in our study is possibly due, in part, to sample size; we analyzed only 11 samples from New Brunswick whereas Kays et al. (2010) determined haplotypes for 453 samples from northeastern

North America. The contact zone reported by Kays et al. (2010) contains considerably more rare haplotypes compared to the more northeasterly samples, suggesting that rare haplotypes were lost the further coyotes dispersed to the east. This loss of genetic diversity is typical of dispersal, and especially dispersal with little migration between the new population and the source population (Boileau et al. 1992). Continued sampling efforts will help to further elucidate the genetic connectivity between coyote populations in the northeastern United States and New Brunswick.

The mainland Nova Scotia population of eastern coyotes has the same three haplotypes as the New Brunswick population, although the frequencies of occurrence of cla28 and cla29 are quite different. One factor contributing to this discrepancy is likely a degree of bottlenecking in the establishment of the Nova Scotia population, which is connected to New Brunswick by the small Isthmus of Chignecto. This type of sequential bottlenecking has been found previously in feral mink in Poland (Zalewski et al. 2011) and island colonizing birds (Clegg et al. 2002). Presumably a small number of individuals initially traveled this route into mainland Nova Scotia; in particular, it appears that only coyotes with the cla28 and GL20 haplotypes colonized this province. The relatively low frequency of cla29 suggests few coyotes with this haplotype made it into the province.

Interestingly, individuals with the cla29 haplotype were all collected near the Isthmus of

Chignecto, in the counties of Colchester, Cumberland and Pictou, in west-central Nova

Scotia. Therefore, it is possible that the cla29 haplotype has only recently entered Nova

Scotia. Further evidence supporting the slow spread of cla29 into Nova Scotia is the fact that the Cape Breton Island population only has two haplotypes in our sample of 77 individuals, cla28 and GL20.

The path coyotes took to get to Newfoundland is not yet understood (McGrath 2004).

It is suspected that they crossed over from the mainland on ice flows, possibly from Cape

Breton Island. Although Cape Breton Island is the closest likely source of coyotes for

Newfoundland (approximately 105 km across the Cabot Strait), it is possible they came from New Brunswick, Prince Edward Island or Québec. The first known report of coyotes on Newfoundland was from the western portion of the province in 1985 (Parker

1995). Only one or a few females carrying the GL20 haplotype may have made it across on an ice flow from Cape Breton.

Coyotes also likely dispersed onto Prince Edward Island on ice flows with the first confirmed record in 1983 (Parker 1995). The Confederation Bridge, which was completed in 1997, now links New Brunswick to Prince Edward Island. Although coyotes have been documented crossing bridges (e.g., the Golden Gate Bridge, San

Francisco, CA; Sacks et al. 2006), it is unlikely that many individuals would cross the

Confederation Bridge given its length (~11 km) and high-level of traffic. The most likely scenario again involves crossing on ice flows, limiting the number of individuals that make the crossing. Perhaps only those carrying cla28 made it to Prince Edward Island. A larger sample size will be necessary to confirm the absence of other haplotypes in these areas.

Morphological analyses We recorded skinned body weight averages for adult female and male eastern coyotes in all of our samples collected through the voluntary coyote carcass collection program.

Averages for adult female and male eastern coyotes were 12.17 kg for females (n=195) and 14.57 kg for males (n=230). Using the formula of Nelson and Lloyd (2005) to

convert skinned carcass mass to non-skinned weight gives values of 14.95 kg for females and 17.83 kg for males, respectively.

In our univariate analyses, there were no statistically significant differences in morphological variables between groups of adult male or adult female coyotes from Nova

Scotia as defined by C. latrans or C. lycaon haplotype group when each measurement was analyzed separately (Table 2.3 and 2.4). In contrast, a composite measure of overall body size based on principal components analysis did demonstrate a statistically significant difference between groups of adult male coyotes defined by haplotype.

Principal component 1 (PC1) which is generally interpreted as the “size” axis (e.g.,

Rising and Somers 1989), comprised 59% of total variation in males (Figure 2.2), and

44% of total variation in females (Figure 2.3). Male eastern coyotes that possessed GL20

(i.e., the C. lycaon haplotype) were significantly larger than male eastern coyotes that possessed western coyote haplotypes (Tukey's diff: 3.8 ± 2.9; p = 0.015). There was no significant difference in body size between groups of females defined by haplotype

(Tukey's diff: -1.5 ± 3.0; p = 0.285).

The observation that body size is correlated with mitochondrial DNA haplotype warrants further exploration. As mentioned, eastern coyotes from northeastern United

States and southeastern Canada have been shown to be fairly well genetically admixed in terms of nuclear genes from western coyote (61.73%), eastern wolf (13.58%), gray wolf

(13.62%) and domestic dog (11.07%) (Monzon et al. 2014). Although we plan to pursue additional nuclear genetic analysis (sensu Monzon et al. 2014) of our samples to determine if the morphological differences correlate with degree of eastern wolf nuclear background as well, it is possible that there is an effect on morphology conferred by the

mtDNA genome. For example, Pichaud et al. (2012) recently demonstrated that mtDNA, and not nuclear background or mito-nuclear interactions, was responsible for metabolic activity in experimental Drosophila populations. Zhang et al (2008) showed that mitochondrial mutations per se have been implicated in growth rates (and body size) in

Nanyang cattle (Bos taurus). Toews et al. (2013) showed that patterns of introgression of mtDNA in the Yellow-rumped warbler complex (Setophaga coronata spp.) can be explained by selection for mitochondrially-encoded differences in respiration rate and

ATP production efficiency. More generally, da Fonseca et al. (2008) compared 12 mitochondrial protein-coding genes from 41 mammalian species and found evidence that adaptive evolution of mitochondrial mutations was associated with metabolic requirements such as adaptations to the energy available in the diet and the body size of the species. As summarized by Galtier et al. (2009), there is growing evidence that mtDNA is not always a simple “neutral” marker of molecular diversity and it may be an important driver of functional evolutionary change.

As Kays et al. (2010) and Monzon et al. (2014) have suggested, the recent range expansion of coyotes into eastern North America and the concomitant hybridization with eastern wolves presents a fascinating case study in rapid adaptive evolution; feeding on large prey such as deer and moose may select for wolf-like traits in eastern coyotes, leading to larger body size. In future, I plan to pursue whether there is localized selection for behavioural changes in eastern coyotes in Nova Scotia, compared to their western ancestors, possibly associated with introgression of eastern wolf mitochondrial genes and selection for wolf-like behaviours.

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Personal Communication Terry Power, Regional Biologist, Nova Scotia Department of Natural Resources, Sydney, Nova Scotia.

Table 2.1. Summary of genetic diversity observed in the five sampled locations in Atlantic Canada. Total New Brunswick Nova Scotia Newfoundland Prince Edward Island Cape Breton Island Sample size 197 10 94 14 2 77

# Haplotypes 3 3 3 1 1 2

Haplotype diversity 0.465 0.733 0.48 0 0 0.267

Nucleotide diversity, π (per site) 0.01189 0.02259 0.01231 0 0 0.00661

θ (per site) 0.01189 0.01607 0.00889 0 0 0.00504

Average pairwise number of nucleotide 2.87683 5.467 2.98 0 0 1.599 differences, k

Table 2.2. Pairwise FST values of coyote mtDNA CR sequences for five locations in Atlantic Canada and their associated p-values. FST values are above the diagonal; p-values are below the diagonal. The total FST value for all samples was 0.275 (p < 0.001) New Brunswick Nova Scotia Newfoundland Prince Edward Island Cape Breton Island

New Brunswick - 0.09402 0.49083* 0.04661 0.31424*

0.06306 ± 0.0237 - 0.51736* -0.06738 0.04661* Nova Scotia Newfoundland 0.00901 ± 0.0091 0.00000 ± 0.0000 - 1 0.75539* Prince Edward Island 0.39640 ± 0.0528 0.65766 ± 0.0526 0.01802 ± 0.0121 - -0.19644

Cape Breton Island 0.00000 ± 0.0000 0.03604 ± 0.0148 0.00000 ± 0.0000 0.99099 ± 0.0030 -

* indicates that the FST value is significantly different from 0 with the associated p-value.

Haplotype Weight - Chest Body Tail Front Front Back Back Shoulder Canine Canine Skull Skull skinned girth length length paw paw paw paw height (cm) tooth tooth length width (cm) (kg) (cm) (cm) (cm) width length width length length width (cm) (mm) (mm) (mm) (mm) (mm) (mm) cla28 11.9 48.7 89.3 34.9 44.3 59.9 40.1 54.9 50.0 37.6 8.8 20.5 11.1 ±1.7 ±2.9 ±4.4 ±2.4 ±3.9 ±4.9 ±4.0 ±3.1 ±4.7 ±1.6 ±0.78 ±1.1 ±0.88

n = 19 n = 19 n = 19 n = 19 n = 17 n = 17 n = 18 n = 18 n = 16 n = 14 n = 17 n = 17 n = 17

GL20 11.5 47.3 87.2 34.0 43.2 57.2 37.7 51.8 51.5 37.0 8.9 19.3 10.1 ±1.8 ±2.3 ±2.9 ±2.8 ±3.4 ±4.7 ±5.0 ±4.4 ±1.9 ±0.97 ±0.47 ±1.3 ±0.85

n = 5 n = 5 n = 5 n = 5 n = 4 n = 4 n = 5 n = 5 n = 5 n = 3 n = 4 n = 3 n = 3

Haplotype Weight - Chest Body Tail Front Front Back Back Shoulder Canine Canine Skull Skull skinned girth length length paw paw paw paw height tooth tooth length width (kg) (cm) (cm) (cm) width length width length (cm) length width (cm) (cm) (mm) (mm) (mm) (mm) (mm) (mm) cla28 14.7 52.8 87.6 34.4 48.5 62.4 42.8 56.8 53.6 40.0 9.5 21.2 11.6 ±2.8 ±3.6 ±21.1 ±9.3 ±4.6 ±5.6 ±6.7 ±3.6 ±3.7 ±2.5 ±0.82 ±1.6 ±1.3

n = 20 n = 20 n =20 n = 20 n = 19 n = 19 n = 20 n = 20 n = 19 n = 18 n = 20 n =19 n = 19

GL20 15.1 54.4 95.9 39.0 49.0 63.2 42.3 56.9 53.6 39.0 9.4 21.3 11.9 ±2.1 ±6.3 ±5.9 ±3.9 ±4.6 ±4.9 ±1.6 ±4.8 ±3.6 ±3.3 ±0.96 ±1.4 ±0.86

n = 9 n = 10 n = 10 n = 10 n = 9 n = 9 n = 10 n = 10 n = 8 n = 6 n = 8 n = 8 n = 8

Figure 2.1. Mitochondrial haplotype frequencies of eastern coyotes from Atlantic Canada (New Brunswick, mainland Nova Scotia, Cape Breton Island, Prince Edward Island, and Newfoundland).

Figure 2.2. First principal component (PC1) values for male coyotes with one of two mtDNA haplotypes, Cla28 or GL20, using 11 morphological measurements and representing 59% of total variation. PC1 has the following loadings: 0.31 * weight (kg), 0.30 * chest (cm), 0.34 * body length (cm), 0.31 * tail length (cm), 0.22 * front paw width (mm), 0.32 * front paw length (mm), 0.17 * back paw width (mm), 0.29 * back paw length (mm), 0.30 * shoulder (cm), 0.36 * skull width (cm), 0.36 * skull length (cm). As all measurements load in the same direction, this component can be interpreted as representing general size differences. Boxes on this plot represent 25th and 75th percentiles, and the line represents the 50th percentile. Whiskers extend to the highest and lowest values that fall within 1.5 * the inter-quartile range. Data beyond the ends of whiskers are outliers, represented as points.

Figure 2.3. First principal component (PC1) values for female coyotes with one of two mtDNA haplotypes, Cla28 or GL20, using 11 morphological measurements and representing 44% of total variation. PC1 has the following loadings: 0.28 * weight (kg), 0.30 * chest (cm), 0.17 * body length (cm), 0.25 * tail length (cm), 0.34 * front paw width (mm), 0.39 * front paw length (mm), 0.37 * back paw width (mm), 0.38 * back paw length (mm), -0.02 * shoulder (cm), 0.21 * skull width (cm), 0.39 * skull length (cm). As all measurements load in the same direction, this component can be interpreted as representing general size differences. Boxes on this plot represent 25th and 75th percentiles, and the line represents the 50th percentile. Whiskers extend to the highest and lowest values that fall within 1.5 * the inter-quartile range.

Chapter 3: Spatial dynamics of eastern coyotes (Canis latrans var.) in Cape Breton

Highlands National Park, Nova Scotia, Canada

Introduction

The eastern coyote (Canis latrans var.) arrived in Nova Scotia during the late

1970’s (Sabean 1991). Coyotes were frequently classified as a nuisance species for their ability to hunt and kill livestock and pets and for damaging property (Bounds and Shaw

1994). Coyotes were also blamed for the depletion of a number of popular game species, particularly white-tailed deer (Odocoileus virginianus; Patterson 1995). Upon their arrival to Cape Breton Island in the late 1970’s and on to Cape Breton Highlands

National Park (CBHNP) in 1980’s (E. Muntz pers. comm. 2013), coyotes entered into conflict with humans. In CBHNP there have been eight fearless behaviours, 16 aggressive behaviours and six attacks by coyotes toward humans between 2003 and 2012 in CBHNP. The second human fatality (first adult) by coyotes occurred in CBHNP on 29

October 2009 (CBC News 2009); the first fatality was a 3-year old child in California in the 1980’s (Howell 1982).

Aggression by coyotes toward humans has been linked to habituation (loss of fear towards humans) or food conditioning (where coyotes associate humans or human places with food rewards; Gehrt 2012). Food conditioning can be achieved by feeding coyotes intentionally or unintentionally (Bounds and Shaw 1994; Alexander and Quinn 2011;

Lukasik and Alexander 2011). Over time, coyotes lose their fear of people and associate humans with food (garbage, pet food and pets; Loe and Roskaft 2004). When this loss of fear occurs and food becomes limited, coyotes may develop bold and aggressive behaviours, increasing the likelihood of an attack (Gehrt 2012).

The human fatality and the apparent high level of aggressive behaviour by coyotes toward humans in CBHNP initiated a study to better understand and ultimately mitigate the propensity for human/coyote interactions in the park. An important aspect of this study was to determine whether eastern coyote spatial dynamics in CBHNP increase their likelihood of coming in contact with humans.

Coyote spatial dynamics can be influenced by many factors including human activities (Atwood et al. 2004), season (Mech 1970), food availability (Dann 1981), biotope selection, and weather conditions (Andelt and Gibson 1979). Coyote eyesight is best suited for daylight and twilight conditions (Kavanau and Ramos 1975) and it has been suggested that nocturnal activity detected in many coyote populations living in human-populated areas is a direct behavioural response to avoid humans (Kavanau and

Ramos 1975; Vila et al. 1992). Atwood et al. (2004) completed a study based in a suburban to urban environment in west-central Indiana that documented coyotes as being largely nocturnal, possibly due to the fear of people which are associated with hunting and trapping activities. Kitchen et al. (2000) found coyotes to move less during diurnal hours in areas with high human activity and diurnal movement increased concurrently with cessation of human activity. McClennen et al. (2001), on the other hand, showed in the undeveloped and mainly protected (e.g., no hunting or trapping) Grand Teton

National Park which hosts human visitors, coyotes were active during both the day and night time hours. This could indicate that coyotes in a national park, where they are not hunted or trapped, learn not to fear humans and thus are equally active throughout the day and night.

Studying detailed movement activity of animals has been made feasible by

advanced radio telemetry technology involving GPS collars which are able to obtain hundreds of precise locations of wildlife species with little effort (Ganskopp 2001). GPS collars use satellites to log the geographical positions of animals up to several times per day and stores data onboard the collar until it can be downloaded remotely. This data provides insights into spatial dynamics, including extent of movement, extra territorial excursions, as well as detailed movement patterns related to time of day.

Cape Breton Highlands National Park is a remote park in Northern Cape Breton,

Nova Scotia receiving approximately 150 000 visitors per year (E. Muntz pers. comm.

2013). There is no hunting or trapping in the park at any time of the year. The adjacent community of Chéticamp on the western side of the park has just under 4 000 residents and there are approximately 300 and 500 certified trappers and hunters, respectively, in the community. There is an annual trapping and year-round hunting season for coyotes outside of the park system with about 30 coyotes harvested and marketed each year (J.

Madden pers. comm. 2015; M. Boudreau pers. comm. 2015) during the period of this study (2010-2014).

Coyote ecology has been studied throughout much of their North American range, revealing a great deal of behavioural flexibility (Bekoff 1977; Holzman et al. 1992), mainly related to prey characteristics (Bowen 1981) and biotope use (Gese et al. 1988).

Cape Breton Highlands National Park serves as a wildlife reserve and there are an estimated 50 to 100 coyotes in the park (E. Muntz pers. comm. 2013). Coyotes are naturally curious and there is potential for frequent encounters with humans, especially along high-traffic hiking trails. Specifically, managers are interested in understanding

why coyotes within the park appear to be showing high levels of aggressive behaviour towards humans compared to outside the park.

The objectives of this study were to: (1) compare the cumulative area of use by coyotes over time between different regions in CBHNP and community of Chéticamp;

(2) determine whether coyotes are attracted to high human use areas within and without the park and; (3) compare day and night time movement patterns by coyotes within the park and community of Chéticamp.

Methods

Capture and Radio Telemetry Coyotes were captured from October 2011 to July 2013, using a combination of four coiled #3 Victor soft catch leg hold traps (Oneida Victor, Inc., Ltd. Euclid, Ohio,

USA) and cable restraint devices (The Snare Shop. Lidderdale, Iowa, USA). Traps and cable restraints were opportunistically set in areas throughout the park where coyote signs

(e.g., scat and tracks) were observed; this was primarily near roads and trails but also occurred in the interior of the park. Traps and cable restraints were checked twice a day to minimize the amount of time a coyote was confined to a trap. Captured coyotes were physically restrained with a catch-pole and immobilized with an intramuscular injection of Telazol. Upon recovery from the sedative, the coyote was released at the capture site.

Parks Canada staff performed all capture and handling of coyotes. Parks Canada Animal

Care Task Force approved animal care protocols (permit number 12020).

While under sedation, coyotes were classified as being young of the year (≤12 months of age), yearling (13 to 24 months) or adult (≥24 months) based on tooth wear, tooth colouring and overall body size and shape (Parks 1979). Coyotes were sexed, weighed, and standard body measurements were collected (body length, tail length, chest

girth, front and hind pad length and width, and shoulder height). Each coyote was marked with two unique number/color combination ear tags.

Each coyote was equipped with a collar containing a GPS (Global Positioning

System) unit with a VHF beacon and with a mechanical breakaway device (Lotek

Wireless Inc, New Market, ON) set to come off 46 weeks after deployment. Collars were equipped to emit a “mortality” signal if the collar/individual was stationary for 12 hours.

GPS collars were programmed to acquire fixes every 2 hours for 12 fixes per day for the first 2 months after attachment and last 2 months of deployment and fixes every 7 hours for 3 to 4 fixes per day between these two periods. Along with geographic position, data for each fix attempt included date, time, fix status, (2 dimensional, 3 dimensional or failure), collar function status, altitude, measures of activity, dilution of precision measures related to satellite geometry, temperature and the number of accessed satellites.

Data were stored onboard the collar. In this study the GPS collar accuracy averaged approximately 3.0 m (E. Muntz pers. comm. 2012; Lotek Wireless Inc., New Market,

ON). Collared animals were located sporadically by foot, vehicle and helicopter via the

VHF signal to download GPS data.

Data editing Collar data were filtered to remove any values of zero (i.e., no coordinates taken due to poor satellite reception) for latitude or longitude. Latitude and longitude were converted to Universal Transverse Mercator (UTM). Data were removed that had a dilution of precision (DOP) value greater than 10. Dilution of precision is a measure of error. When the GPS collar uses inputs to calculate the position, the output also contains errors, and normally the output error exceeds input errors. This magnifying factor is

called DOP (a DOP between 1 and 10 has less error than >10). Date and time were converted from Greenwich Mean Time (GMT) to Local Mean Time (LMT).

With few coyotes collared, high variability among individuals and many co- variables (location of collared animals, time interval GPS collars worked correctly, and date), an individual approach was used for analyzing the data. A two-season human presence model was created to determine if coyotes were attracted to high human use areas - high human presence (15 June – 15 October, height of human visitation in the park) and low human presence (16 October – 14 June, lowest human visitation in the park).

Cumulative area of use Cumulative area of use was calculated using a 95% MCP for each individual coyote using all sequential locations. The cumulative locations increased sequentially by

50 locations up to 1000 locations and increased by 100 from 1000 to 1900 locations. A

Minimum Convex Polygon was used in the package adehabitatHR in RStudio (0.98.490) to create the total area for sequential locations (in km2).

Areas of human use A map of the study area was created using the Nova Scotia Ecological Land

Classification (2003) data from the Forestry Division of the Nova Scotia Department of

Natural Resources (DNR); this map was created in ArcMap (10.1; ESRI Canada).

Original files were used for Inverness and Victoria Counties, which consisted of 37 vegetation and land use classifications (Appendix 3.1). The urban habitat type in the original DNR layer was verified and edited in ArcMap (using air photos from 2009).

Recent (occurring since the last layers were created) residential and commercial development was added to the map of the study area. Human use areas within CBHNP

included: roads; pull offs, service roads, hiking trails, campgrounds, parking lots, beaches and golf courses. Within the park, roads and park trails were buffered with 100 m on each side and classified as human use areas.

The core area (classical home range after Burt 1943) of coyote use areas was determined by visual inspection of the data and excluded locations that may have been extraterritorial movement (areas not normally used by the coyote). The sum of human use area within each coyote’s core area of use was analysed by importing the MCP area of use from RStudio to ArcMap as polygons. These polygons were merged with the biotope layer. The data were extracted from ArcMap into an excel spreadsheet that displayed the area of the various biotopes for each coyote. The proportion of each biotope was found and converted to a percentage. Results below only report data on human use areas.

Day versus night movement patterns For assessing daylight and dark movement patterns, all GPS collar locations were edited and use was limited to those with two-hour sequential locations. The distance between each location was classified as occurring during daylight or dark hours. All data was removed for distances covered spanning the sunrise and sunset time period

(Environment Canada 2013). The distance was calculated between each location (e.g., the distance between the location taken at 10:00 and the location taken at 12:00). I calculated means of individual coyote distance traveled during daylight or dark movement periods between each 2-hour location. Means of individual coyotes’ daylight and dark movements were compared, along with daylight and dark movements for all coyotes in

CBHNP and Chéticamp.

Statistical tests Differences between high human use areas for low and high human presence were assessed using the Pearson’s Chi-Squared test. Differences between daylight and dark movements for individual coyotes were assessed using the Wilcoxon Test. Differences between daylight and dark movements for coyotes collared in CBHNP and Chéticamp were assessed using a pairwise t-test as implemented in R packages chisq.test,

Wilcox.test and t.test, respectively.

Results

Eleven coyotes were collared with GPS transmitters (9 males and 2 females). All were adults except for one male classified as a yearling (CH-19; Table 3.1). One male

(CH-7) and one female (CH-13) were collared for short durations of approximately three and two months, respectively, and thus had fewer location points compared with other individuals (Table 3.1).

The cumulative area of use based on 95% MCP of all sequential locations varied greatly among individuals (Figure 3.1). Six of the eleven collared individuals (CH-6, CH-

7, CH-11, CH-13, CH-18 and CH-19) showed stable areas of use but varying from 10 to

25 km2. CH-11’s area of use suddenly expanded after a period of no change (Figure 3.1).

CH-1 and CH-22 steady increased in cumulative area of use. CH-3 and CH-8 levelled off but not until reaching 80 km2. CH-5 had the largest area of use at almost 1200 km2

(Figure 3.2). This coyote increased its area of use steadily during the first months of collar deployment and then began levelling off (Figure 3.2). Coyotes in the community of

Chéticamp had constant area of use curves, whereas the coyotes on French/ Mackenzie

Mountains tended to increase the area of their curves (Figure 3.1). Coyotes from North

Mountain increased their area of use curve rapidly and then levelled off before a

reduction in the area of use toward the end of the collar deployment (Figure 3.1). Coyotes on the Eastern side of CBHNP increased the area of use and then levelled off (Figure 3.1;

3.2).

Coyotes in the Chéticamp area had the greatest access to human use areas and coyotes within the park had much less (Table 3.2). One of three coyotes in Chéticamp

(CH-3) was alive long enough to determine its movement within human use areas across seasons. This coyote used human use areas more during the season of high human presence compared to low human presence (Table 3.2). Two of four coyotes on

French/Mackenzie Mountains tended to use human use areas more during the season of high human presence compared to the low human presence (Table 3.2). The two coyotes on North Mountain showed no seasonal difference in their utilization of human use areas

(Table 3.2). Coyotes on the eastern side of the park used human use areas more during the season of high human presence (Table 3.2). One collared coyote (CH-1) was shot because he was circling vehicles and children and chasing bicycles; another (CH-7) was snared illegally in the community of Chéticamp and another (CH-3) was trapped legally in the community of Chéticamp.

Coyotes in CBHNP and Chéticamp showed significantly different day/night movement patterns (Table 3.4). Individually, coyotes collared in Chéticamp moved significantly less during the day compared to night (Table 3.3). Five of six coyotes within the park moved significantly more during the day compared to the night (Table 3.3).

Discussion

Only eleven coyotes were monitored with GPS collars; two adult females and nine males, all adults expect for one, a yearling. Three coyotes (two males and one

female) were centred in Chéticamp on the western coast, located southwest of CBHNP.

Two individuals were found living on the eastern side of CBHNP, near the coast. The remaining individuals were on the plateau of the park. The small sample size in this study makes generalizations about coyote spatial dynamics less reliable. However, it is clear from the analysis of cumulative area of use that individual coyotes had drastically different space use patterns. Some coyotes had relatively small, but constant, areas of use while others had much larger constant areas of use. Other coyotes were tracked moving extensively and these individuals areas of use increased for long periods or showed sudden increases. Additional data is needed to isolate more conclusive patterns of area of use as relatable to geographic conditions. Nevertheless, evaluation of the existing data still provides interesting insights into land use patterns of coyotes in Cape Breton.

The three coyotes in the Chéticamp area appear to have smaller core areas of use than coyotes within the park. Home range sizes of coyotes have been known to vary by region, food availability and reproductive season (Althoff 1978; Andelt and Gipson 1979;

Shargo 1988; Atkinson and Hackleton 1991). Coyote home range size has been documented as smaller in urban landscapes compared to more rural areas (Shargo 1988;

Atkinson and Hackleton 1991; Atwood et al 2004; Ghert et al. 2009). More broadly, red foxes also show a reduced home range size with increased urbanization (Cavillini 1996;

Goszczynski 2002). In this study there were similar trends with animals (CH-7 and CH-

13) that spent the majority of their time in the community of Chéticamp.

Only coyote (CH-3) in the Chéticamp area had enough data to compare space use across human presence seasons. This animal spent more time in high human use areas during the season of high human presence. Andelt and Andelt (1981), Roy and Dorrance

(1985) and Holzman et al. (1992) found coyotes in urban areas preferentially selected biotopes in locations that would necessitate fewer interactions with humans. Two of four coyotes on French/Mackenzie Mountain had sufficient data to compare seasonal use of human use areas, and these individuals increased their use of human use areas during high human presence, as did the two coyotes on the eastern side of the park. The two

North Mountain coyotes significantly increased their use of human use areas during high human presence. Thus, it appears that coyotes in this study do seek out human use areas during high human presence. No other study has compared coyote use of space in this way.

The three coyotes in the community of Chéticamp moved significantly less during the day compared to the night. Conversely, coyotes within CBHNP moved more during the day than night. Low level of daytime movement in human built up areas appears to be common for coyotes (Kavanau and Ramos 1975; Vila et al. 1992; Kitchen et al. 2000;

Grinder and Krausman 2001; Atwood et al. 2004). A population of coyotes in Texas not exposed to human exploitation exhibited more movement during daylight hours (Andelt

1985). Areas used during dark hours by coyotes are likely used for hunting and traveling

(Andelt and Andelt 1981; Laundre and Keller 1981; Shargo 1988; Holzman et al. 1992).

High human use areas were used more during the high human presence season by

CH-3 in Chéticamp, however his movement was highest during dark hours. This may indicate his reliance on human sources of food from restaurants and fish processing plants, while simultaneously avoiding human presence. Similarly, red foxes in Toronto

Ontario, Canada have been found to travel in human use areas, but enter them at times when there is reduced likelihood of contact with humans (Adkins and Scott 1988).

Coyotes may be avoiding people in Chéticamp due to hunting or trapping pressures. In human dominated landscapes coyotes have been found to move shorter distances during the day compared to the night (Quinn 1995; Kitchen at al. 2000). In

CBHNP coyotes may be obtaining food remains from park visitors; this familiarity with the human species may have conditioned them to more overt behaviour when humans are present. McClennen et al. (2001) showed that coyotes were active during the day and night in a mainly protected national park, which could indicate coyotes do not fear people. In the park setting, coyotes are unlikely to experience negative repercussions from humans until there is a major event (i.e., someone getting bitten). When a dangerous behaviour occurs, parks personnel will try to eliminate the offending animal, as in the case of CH-1 (E. Muntz pers. comm. 2014). Coyotes travel more during dark hours in

Chéticamp compared to within the park. Vila et al. (1992) suggested that nocturnal activity in many coyote populations living in human residential areas is a direct behavioural response to avoid humans. Way et al. (2004) showed it appeared that coyotes were more comfortable traveling through yards at night and commonly bed down 50 m away from homes during the day. Studies in New York (Bogan 2004), Massachusetts

(Way et al. 2004), Arizona (Grinder and Krausman 2001) and southern California (Riley et al. 2003) also document coyotes increase their movement during dark hours.

Although sample size is low, it appears that coyotes near Chéticamp are active at night, while in CBHNP they maintain a high level of activity during the day. Within the park, coyotes are attracted to human use areas during the season of high human presence, although the overall use of human use areas is low. In Chéticamp one coyote increased its use of human use areas during the season of high human presence but avoided daytime

movement. Thus, coyotes within the park may have a higher propensity for encountering and engaging human

References

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Personal Communication

Michael Boudreau, Human-Wildlife Conflict Biologist, Nova Scotia Department of Natural Resources-Wildlife Division, Kentville, Nova Scotia.

Jennifer Madden, Program Administrator, Nova Scotia Department of Natural Resources-Wildlife Division, Kentville, Nova Scotia.

Erich Muntz, Project Manager, Parks Canada, Chéticamp, Nova Scotia

Table 3.1. Coyotes captured in Cape Breton Highlands National Park between October 2011 and July 2013. A = adult; Y = yearling; M = male; F = female.

Location Coyote ID Length of time collared # locations Capture Sex Age (months) Date

Chéticamp CH-3 9 1479 30/10/11 M A Chéticamp CH-7 ~3 929 26/06/12 M A Chéticamp CH-13 ~2 305 15/10/12 F A French/ Mackenzie Mountain CH-11 ~5 1219 14/10/12 F A French/ Mackenzie Mountain CH-18 ~4 556 10/11/12 M A French/ Mackenzie Mountain CH-19 ~8 1386 18/05/13 M Y French/ Mackenzie Mountain CH-22 ~6 1107 01/07/13 M A North Mountain CH-1 10 1282 27/10/11 M A North Mountain CH-8 ~11 1462 18/07/12 M A Eastern CBHNP CH-5 ~9 1750 13/11/11 M A Eastern CBHNP CH-6 ~11 1854 19/11/11 M A

Table 3.2. Available and use of high human use biotopes to coyotes in Cape Breton Highlands National Park and Community of Chéticamp from October 2011 to January 2014. Chi-square statistic shows where differences exist between low and high human presence. M = Male; F = Female; CH = Cape Breton Highlands Coyote ID.

Coyote use of human use areas

Location Coyote ID Sex/Age Available Low human presence % (n) High human presence % Chi-square statistic (p-value) (%) (n)

Chéticamp CH-3 M/A 14.4 5.1 (1053) 25.3 (426) 127.2 (p ≤ 0.000) Chéticamp CH-7 M/A 35.0 - 11.4 (929) - Chéticamp CH-13 F/A 34.9 29.8 (305) - -

French/ Mackenzie CH-11 F/A 2.9 4.3 (1205) 0.0 (14) - Mountain

French/ Mackenzie CH-18 M/A 0.4 0.0 (556) - - Mountain

French/ Mackenzie CH-19 M/Y 5.3 0.8 (306) 1.0 (1080) 28.5 (p ≤ 0.000) Mountain

French/ Mackenzie CH-22 M/A 8.5 18.3 (208) 45.3 (899) 51.2 (p ≤ 0.000) Mountain

North Mountain CH-1 M/A 3.6 5.7 (1030) 6.7 (252) 0.38 (p = 0.54)

North Mountain CH-8 M/A 3.4 1.1 (734) 0.8 (728) 0.27 (p = 0.60)

Eastern CBHNP CH-5 M/A 7.9 9.5 (1128) 12.7 (622) 4.4 (p = 0.04)

Eastern CBHNP CH-6 M/A 11.4 8.5 (1091) 10.7 (763) 2.6 (p ≤ 0.000)

Table 3.3. Summary of daylight and dark 2-hour movement averages for collared coyotes in Cape Breton Highlands National Park and the community of Chéticamp from October 2011 to January 2014

Individual coyote 2-hour daily movement

Location Coyote ID Sex/Age Daylight movement (m) Darkness movement (m) Wilcoxon statistic (p-value)

Chéticamp CH-3 M/A 28.0 540.0 26416.5 (p ≤ 0.000)

Chéticamp CH-7 M/A 15.0 968.0 10520.0 (p ≤ 0.000)

Chéticamp CH-13 F/A 25.5 87.0 5990.0 (p = 0.11)

French/ Mackenzie Mountain CH-11 F/A 207.0 16.0 88632.0 (p ≤ 0.000)

French/ Mackenzie Mountain CH-18 M/A 280.5 12.0 1909.5 (p =0.005)

French/ Mackenzie Mountain CH-19 M/Y 76.5 362.0 37209.5 (p ≤ 0.000)

French/ Mackenzie Mountain CH-22 M/A 54.0 59.0 35444.0 (p = 0.23)

North Mountain CH-1 M/A 389.0 7.5 20768.0 (p ≤ 0.000)

North Mountain CH-8 M/A 179.0 75.0 33082.5 (p = 0.009)

Eastern CBHNP CH-5 M/A 203.0 164.5 75931.5 (p = 0.50)

Eastern CBHNP CH-6 M/A 310.0 69.5 93415.0 (p ≤ 0.000)

Table 3.4. Summary of daylight and dark 2-hour movement averages for collared coyotes in Cape Breton Highlands National Park and the community of Chéticamp from October 2011 to January 2014.

Location

Time of day CBHNP Mean (m) ± SD (n) Chéticamp Mean (m) ± SD (n) T-test Statistic df (p=value)

Daylight 637.0 ± 966.6 (2610) 257.4 ± 512.8 (n = 864) t = -14.8 (p ≤ 0.000) 2803.9

Dark 542.9 ± 912.0 (1903) 907.5 ± 984.8 (636) t = 8.2 (p ≤ 0.000) 1023.1

) 2

CH-1 size (km size

CH-3 CH-22

CH-11 CH-8

Cumulative area of use of Cumulativearea CH-6

CH-13 CH-18 CH-19 CH-7

Sequential locations

Figure 3.1. Cumulative area of use curve (km2; 95% MCP) with sequential locations of GPS collared coyotes in Cape Breton Highlands National Park (CH 1, 6, 8, 11, 18, 19 and 22) and the community of Chéticamp (CH 2, 7, 13) between October 2011 and January 2014.

)

size(km Concentrated area of use use of area Concentrated

Cumulative locations Figure 3.2. Cumulative area of use curve (km2; 95% MCP) with sequential locations of CH-5 GPS-radio collared coyote in Cape Breton Highlands National Park between November 2011 and October 2012.

Appendix 3.1. Reclassification of 37 biotopes from the Nova Scotia Department of Natural Resources land cover inventory to 5 biotopes used for biotope use.

ID Number Classification Name Habitat Classification

0 Natural stand Forested 1 Treated Forested 2 Burn Forested 5 Old field Human use 6 Wind throw Forested 7 Dead Forest regeneration 8 Dead – 1 Forest regeneration 9 Dead – 2 Forest regeneration 12 Treated stand Forest regeneration 13 Dead – 3 Forest regeneration 14 Dead – 4 Forest regeneration 15 Dead – 5 Forest regeneration 20 Plantation Forested 33 Brush Forest regeneration 38 Alders Forested 60 Clear cut Forest regeneration 62 Partial depletion not verified Forested 70 Wetlands general Inland and coastal water 71 Beaver flowage Inland and coastal water 72 Open bogs Bog and barren 73 Treed bogs Bog and barren 74 Coastal habitat areas Inland and coastal water 75 Lake wetland Inland and coastal water 76 Cliffs, dunes and coastal rocks Inland and coastal water 77 Inland water Inland and coastal water 78 Ocean Inland and coastal water 83 Brush Forest regeneration 84 Rock barren Bog and barren 85 Barren Bog and barren 86 Agriculture Human use 87 Urban Human use 91 Blueberries Human use 92 Miscellaneous Human use 94 Beach Human use 95 Gravel pit Human use 97 Powerline corridor Human use 98 Road corridor Human use

Chapter 4: Winter foraging behaviour by eastern coyotes (Canis latrans var.) based on fractal analysis of snow track-patterns in Cape Breton Highland National Park,

Nova Scotia, Canada.2

Introduction

The eastern coyote (Canis latrans var.) is a hybrid between the western coyote and eastern wolf (Canis lupus; review in Parker 1995) that resulted when the coyote expanded eastward in the latter part of the 19th century, leading to interbreeding (Power et al. 2015). Eastern coyotes have the largest body size of any existing coyote population

(Thurber and Peterson 1991; Lariviere and Crete 1993; Peterson and Thurber 1993), which may be the result of the described interbreeding. Some (Geist 1987; Thurber and

Peterson 1991) suggest the increase in body size may have been due to selection for larger animals as a result of larger target prey, specifically white-tailed deer (Odocoileus virginianus). Also, natural selection may favour larger bodied animals in areas where gray wolves have been extirpated (Way et al. 2010).

Coyotes are generalist predators (Lukasik and Alexander 2012) whose prey ranges in size from small rodents to large ungulates and may include livestock or domestic cats and dogs (Lukasik and Alexander 2012). Coyotes may also forage for fruit, garbage and carrion (Lukasik and Alexander 2012; Bowyer et al. 1983; Fedriani et al.

2001). The eastern coyote can prey upon adult white-tailed deer (Patterson 1995) and

2 Cécile Bossi, Jason W. B. Power, Vilis O. Nams and Søren Bondrup-Nielsen. Winter foraging behaviour by eastern coyotes (Canis latrans var.) based on fractal analysis of snow track-patterns in Cape Breton Highland National Park, Nova Scotia, Canada. Summited to PLoSONE-undergoing revisions.

more recently, coyotes have been documented killing adult moose (Alces alces) in

Ontario (Benson and Patterson 2013) and moose were found to be the dominant food item in coyote scats in Québec (Boisjoly et al. 2009).

Eastern coyotes were first caught in Nova Scotia in 1977 and known to exist on

Cape Breton Island by 1980 (O’Brien 1983; Moore and Millar 1986). In Cape Breton, coyotes occupy the highlands of Cape Breton Highlands National Park (CBHNP), a 950 km2 wilderness area mainly found at an altitude of 500 m. The population of moose in the

Highlands is estimated at 7 500 animals (J. Bridgland pers. comm. 2014). Power (2015; this thesis) has documented that moose constitute an important food source for coyotes within the park year round, but especially during the winter. In the winter, using tracks in snow cover, coyote movement patterns can be quantified to determine if they actively pursue moose as prey.

Analysis of animal movement patterns must consider spatial scale and habitat heterogeneity (Wiens 1989; Nams 2005). Movement patterns can be described at different spatial scales. At small scales, one may observe animals following tortuous paths that reflect the many movements required to successfully collect food items

(Patterson 1995). Straighter paths at a larger scale often suggest direct travel routes to a specific destination (Nams 2006). The effect of habitat heterogeneity on movement patterns at both scales must also be considered (Nams 2005).

The objective of this study was to determine how coyote movement patterns in

CBHNP vary among habitats and at different spatial scales in relation to hunting behaviour.

Methods

This study took place in CBHNP and adjacent areas from 15 January to 15 April

2013. Foraging behaviour and habitat use was based on following coyote tracks in the snow. Tracking was conducted daily (weather permitting). Coyote tracks were typically identified from the road while driving along the Cabot Trail. The coyote was then back- tracked (opposite direction of travel) to avoid interference until track were lost due to windy conditions. A Garmin Montana 650 GPS recorded fixes automatically every 10 m along the path, with an error radius of ~3 m. We also recorded locations of habitat changes, scat collections, prey observations and detection of prey remains.

We analyzed the movement patterns using fractal analysis, which is used to characterize the overall pattern of tracks of an animal in a landscape (Sugihara and May

1990). Fractal dimension is the continuous analogue to geometrical dimension. For example, the geometric dimension of a straight line is 1, and of a plane is 2; the fractal D of a line can vary from 1 (a straight line) to 2 (so tortuous that it takes up the entire two- dimensional space;([Mandelbrot 1967; Dicke and Burrough 1988)). We calculated fractal

D, and the correlation in fractal D between successive track segments, at various spatial scales. These allowed us to see how animals' movements and responses to habitat vary with scale (Nams 2005).

Tracking data was imported as a Garmin database file (.gdb) into the Garmin

MapSource computer program (version 6.16.3). To analyze movement patterns of entire paths, coordinates of every path were extracted and saved as text files (.txt) to be analyzed by the software Fractal (Nams 2013).

To evaluate movement patterns in relation to habitat, coordinates of the paths were incorporated into Excel spreadsheets and integrated into GIS software (ArcGis

10.1). A buffer of 50 m on each side of all paths was created, and habitat polygons were drawn within these buffers. Habitat identification was completed using both those waypoints collected while back-tracking and through analysis of aerial photos of the park collected in 2009 (Figure 4.1). To verify the reliability of aerial photos for photo- interpretation, 30 random points were created in a 1000 m-wide buffer along the Cabot

Trail, from Chéticamp to North Mountain. These points were verified in the field, and their respective habitat types recorded before interpreting the aerial photos. Every photo- interpreted point matched results collected during field verification; aerial photos were used to accurately determine habitat polygons.

We used three major habitat types: conifer, mixed-wood and open areas.

Coniferous stands are associated with the mature boreal landscape; mixed-wood stands are intermediate between the Acadian (hardwood stands) and boreal landscapes and are generally more open than coniferous stands because of a lack of leaves during winter; open areas are classified as an open landscape, including regions which have been extensively disturbed by the spruce budworm (thereby reducing fir stands) and bogs. The

Acadian landscape was not used for analysis because limited backtracking data were collected in this habitat. The few coyote paths back-tracked in deciduous forests (mostly on the east side of the park) were combined with those recorded in mixed-wood stand habitats, to which they are closely related. Polylines were then allocated based on habitat changes, and three different files were obtained: path segments in coniferous stands, mixed-wood stands and open areas.

Results

We followed 57 tracks from 15 January to 15 April (Figure 4.1); their lengths ranged from 150 m to >5,000 m, with a total length of 109 km. Shorter paths (<150 m) were not included in the analysis.

We first tested the hypothesis that movement patterns could be adequately described by a correlated random walk (CRW) model, by comparing the observed (net distance travelled)2 vs # of steps, to that expected under a CRW model (McCulloch and

Cain 1989). Confidence intervals for the observed values were generated by bootstrapping, treating each track as an independent sample. The expected CRW values were outside of the observed confidence intervals, signifying that coyote paths were significantly straighter than a CRW (Figure 4.2).

Then we estimated the fractal dimension vs spatial scale, for scales ranging from

10 m (the resolution of points recorded by the GPS), to 300 m, (1/16 of the longest path).

Vfractal analysis (part of the Fractal software) divides the path into segments, which are then combined into pairs, and the angle between each pair is used to estimate Fractal D

(Nams 1996).

Fractal D increased with spatial scale up to about 150 m (Figure 4.3A) as coyote paths became more and more tortuous. At larger scales the confidence intervals were too broad to determine trends. The D values were low (1.019 to 1.053), meaning coyote paths were quite straight for the considered spatial scales. The correlation plot (Figure 4.3) was positive at smaller scales, indicating heterogeneity in the path (e.g., straighter sections of path would tend to be followed by straighter sections of the path). However, as scale

increases past 135 m, the correlation plot crosses the y-axis, indicating that there are two domains of scale in coyote’s responses to their habitat, with a transition at 135 m.

Since there were two domains, further analysis was carried out in each domain.

Every path was segmented according to the habitat in which each part of the path was located (open areas n = 44, mixed-wood n = 27, coniferous n = 36). A mean D-value was calculated for each segment of path, at two different ranges of spatial scales: small (10 to

135 m) and large (≥135 m). Mean D-values of each segment of path were normalized by transforming them with log (D-1) and the means and confidence intervals were then back-transformed for final presentation. A Fligner-Killeen test was used on these residuals to test for homogeneity of variance. An ANOVA was carried out for each spatial scale to compare fractal dimensions among the three habitats and a Tukey's test was used to compare means of habitats.

Path tortuousities differed significantly among habitat types for the small scales

(F2, 104 = 3.45, p = 0.03) but not the large scales (F2,53 = 0.13, p = 0.87). At small scales paths were significantly more tortuous in the mixed habitats than the open habitats

(Figure 4.4). Coyotes also travelled further in open areas than in other types of stands

(Table 4.1).

Discussion

Coyotes do not travel randomly, but rectilinearly through their habitat. For the same number of steps, they walked longer distances than expected in the case of a CRW; their movements were oriented and the tortuosity increased slightly with spatial scale, which can be related in this case to some degree of randomness in their walking patterns increasing with spatial scale. The largest spatial scale that can be evaluated from the

output of the fractal analysis (Figure 4.3A) is 300 m; above that, the confidence intervals are too wide to facilitate conclusions. If any different movement pattern exists above this scale, it cannot be discerned using the current data.

Considering the analysis up to 300 m (Figure 4.3) and evaluating observations in the field and the shape of the whole paths, coyotes follow oriented (straight) paths. They seem to travel in one direction, swerving primarily to avoid obstacles (trees, hills, cliffs).

Only 3 cases were noted where small animal (rodents, hares) kills were encountered over the 109 km of backtracking routes; however, over the same distance, 17 moose carcasses were found which had been scavenged by coyotes (Power 2015; this thesis).

The fractal correlation analysis shows a transition in movement patterns at a spatial scale of 135 m, with heterogeneity at scales smaller than 135 m (Figure 4.3B).

Thus analyses of mean fractal D values of coyote paths through three different habitats

(mixed-wood stands, coniferous stands, open areas) were run at two different spatial scales: small (10-135 m) and large (≥135 m). Although D-values at the small scale are low, they are significantly greater in mixed-wood stands, suggesting coyotes forage more in this habitat, potentially looking for prey. Finally, coyotes travel more through open areas and mixed-wood stands during winter (Table 4.1), likely due to ease of movement in these habitats when there is snow. During periods of harsh weather, when food availability is reduced, animals need to conserve energy, a priority more difficult to achieve in coniferous stands. By travelling in open habitats, energy use is more efficient.

Coyote and moose interactions Broders (1995) studied coyote movement patterns in Kejimkujik National Park

(KNP) in 1994 and used fractal geometry to analyze data. He found coyote paths were more tortuous through forested habitats where they pursued hares, small mammals and

deer, than in open areas. Broders is the only study to use this method to analyze coyote movement, and thus remains the only work to which our study findings can be compared

(keeping in mind there was no GPS available and limited backtracking data). Coyotes in

KNP exhibited the following patterns: at the large scale (≥ 90 m) paths were significantly more tortuous in hardwood stands than open areas. At all spatial scales, paths going through mixed-wood stands always had a lower D-value than those going through coniferous and hardwood stands, suggesting coyotes search more for food in hardwood and coniferous stands than in mixed-wood stands. Also, data suggests they use open areas as travel routes, as D-values in this habitat were significantly lower than in other habitats when looking at the small scale.

The results of our fractal analysis in CBHNP show different behaviours. At large scales there is no significant difference in path tortuosity between habitats, while at the same scales coyote paths were more tortuous in mixed stands than open areas in KNP.

This difference can be reconciled by noting that in KNP coyotes are known to feed mostly on deer, primarily located at low elevations in hardwood stands. Also, snowshoe

(Lepus americanus) hare are typically found in coniferous stands. Deer are rare on the plateau in CBHNP and snowshoe hare population levels, known to be cyclical

(O'Donoghue et al. 1998), were just starting to rebound at the time of the current study.

This suggests that in this location moose were the main food source available. The homogeneity of path tortuosity between habitats at the large scale, along with the low D- values, indicates that coyotes were unlikely to have been hunting in any of the habitats at this scale. However, at the small scale, coyote paths were more tortuous in mixed-wood stands, which is also different from the results in KNP. There, D-values in mixed-wood

stands and coniferous stands were significantly lower than D-values in open areas, while the opposite was documented in CBHNP. Coyotes seem to search slightly more at a small scale in CBHNP and more at a large scale in KNP. Perhaps coyotes in CBHNP are actively foraging (moose appear to be found opportunistically) for small prey, which would be isolated on a smaller spatial scale than coyotes in KNP looking for white-tailed deer at a larger scale.

Coyotes have just recently been shown to prey upon moose (Benson and Patterson

2013), so it is difficult to say what kind of movement patterns could be expected from fractal analyses focused on moose. However, if the assumption is made that coyotes in

KNP were actively seeking deer based on their movement patterns at a large spatial scale, the hypothesis can be advanced that coyotes in CBHNP may follow the same patterns while looking for moose. There is no evidence to support this behaviour; when looking at the large spatial scale there is no significant difference in terms of tortuosity. However, coyote paths assessed for this study may have been too short and thus the ‘large scale’ considered may have been too small to reveal any pattern. When comparing the results from Broders’ thesis (1995) in KNP with results of this study (assuming the large scale used in this study was a sufficient size), it does not appear that coyotes actively hunt for moose but rather feed on carcasses that they happen to find opportunistically while moving throughout their territories.

In summary, results of fractal analysis suggest that coyote paths are mostly straight. The coyotes observed in CBHNP followed straight, linear paths to travel back and forth from moose carcasses. This movement to primary consumption of moose (over the historically more common deer and hare) may reflect low levels of both species in the

park during the study duration. Furthermore, due to severe winter conditions, it appears moose were especially vulnerable to death from natural causes, thereby becoming the most abundant food source available, and the most efficiently exploited by coyotes in terms of energy gains. However, the cause of death in moose is not conclusive and based on some recent evidence coyotes may have used various predation tactics to further exhaust moose making them easier targets. With snowshoe hare in a low in their cycle during this study period it would be necessary to document the movement patterns of coyotes after the snowshoe hare populations rebound. With less snow there could be a higher chance for moose survival. It would also be necessary to conduct movement and predator prey studies during low snow winters to fully understand predator prey relationships between moose and coyotes. Multi-year studies would also provide more data and a better understanding of how weather affects moose survival.

References

Benson, J.F., B. R. Patterson. 2013. Moose (Alces alces) predation by eastern coyotes (Canis latrans) and eastern coyote × eastern wolf (Canis latrans × Canis lycaon) hybrids. Can J Zool. 91:837–41. Boisjoly, D., J-P. Ouellet and R. Courtois. 2009. Coyote habitat selection and management implications for the Gaspésie caribou. Journal of Wildlife Management. 74:3–11. Bowyer, R.T., S. A. McKenna and M. E. Shea. 1983. Seasonal changes in coyote food habits as determined by fecal analysis. American Midland Naturalist. 109:266–73. Broders, H. G. 1995. An analysis and evaluation of methods used to determine coyote (Canis latrans) movements in relation to resource use. Acadia University, Wolfville Nova Scotia, Canada. Dicke, M. and P. A., Burrough. 1988. Using fractal dimensions for characterizing the tortuosity of animal trails. Physiological Entomology. 13:393–8. Fedriani, J. M., T. K. Fuller and R. M. Sauvajot. 2001. Does availability of anthropogenic food enhance densities of omnivorous mammals? An example with coyotes in southern California. Ecography. 24:325–31. Geist, V. 1987. Bergmann’s rule is invalid. Canadian Journal of Zoology 65:1035– 1038. Lariviere, S. and M. Crete. 1993. The size of eastern coyotes (Canis latrans): A comment. Journal of Mammalogy 74:1072–1074. Lukasik, V. and S. Alexander. 2012. Spatial and temporal variation of coyote (Canis latrans) diet in Calgary, Alberta. Cities and the Environment. 4:1-23. Mandelbrot, B. 1967. How long is the coast of Britain? Statistical self-similarity and fractional dimension. Science.156:636–8. McCulloch, C. E. and M. L. Cain. 1989. Analyzing discrete movement data as a correlated random walk. Ecology. 70:383–8. Moore, G. C. and J. S. Millar. 1986. Food habits and average weights of a fall-winter sample of eastern coyotes, Canis latrans. Canadian Field Naturalist.100:105–6. Nams, V. O. 1996. The VFractal: a new estimator for fractal dimension of animal movement paths. Landscape Ecology. 11:289–97. Nams, V. O. 2005. Using animal movement paths to measure response to spatial scale. Oecologia.143:179–88. Nams, V. O. 2006. Detecting oriented movement of animals. Animal Behaviour 72:1197–1203.

Nams, V. O. Fractal computer program, version 5.26. 2013. Available from: http://www.dal.ca/faculty/agriculture/environmental-sciences/faculty-staff/our- faculty/vilis-nams/fractal.html O’Brien, M. 1983. The coyote a new mammal, new challenges. Nova Scotia Department of Natural Resources - Wildlife Division, Kentville, Nova Scotia, Canada. O’Donoghue, M., S. Boutin, C. J. Krebs, G. Zuleta, D. L. Murray and E. J. Hofer. 1998. Functional responses of coyotes and lynx to the snowshoe hare cycle. Ecology. 79:1193–208. Paquet, P. C. 1992. Prey use strategies of sympatric wolves and coyotes in Riding Mountain National Park, Manitoba. Journal Mammalogy. 73:337–43. Parker, G. R. 1995. Eastern coyote: the story of its success. Nimbus Publishing, Halifax, Nova Scotia, Canada. Patterson, B. R. 1995. The ecology of the eastern coyote in Kejimkujik National Park Masters thesis, Acadia University, Wolfville, NS Canada. Peterson, R. O. and J. M. Thurber. 1993. The size of eastern coyotes (Canis latrans): A rebuttal. Journal of Mammalogy. 74: 1075–1076. Power, J. W., N. Weatherbee-Martin, M. Boudreau, M. O’Brien, G. Conboy and T. Smith. 2015. Survey of helminth cardiopulmonary parasites in coyotes (Canis latrans) of Nova Scotia, Canada. Journal of Compartive Parasitology. in press. Power, J. W. 2015. Genetics, movement patterns, habitat use and diet of the eastern coyote (Canis latrans var.) in Cape Breton Highlands National Park. Acadia University, Wolfville, Nova Scotia, Canada. Sugihara, G. and R. M. May. 1990. Applications of fractals in ecology. Trends in Ecology and Evolution. 5: 79–86. Thurber, J. M. and R. O. Peterson. 1991. Changes in body size associated with range expansion in the coyote (Canis latrans). Journal of Mammalogy. 72: 750–755. Way, J.G., L. Rutledge, T. Wheeldon, and B.N. White. 2010. Genetic characterization of eastern “Coyotes” in eastern Massachusetts. Noertheastern Naturalist. 17: 189-204. Wiens, J. A. 1989. Spatial scaling in ecology. Functional Ecology. 3: 385–97.

Personal Communaction James Bridgland, Park Ecologist, Parks Canada, Ingonish, Nova Scotia.

Table 4.1. Proportion of distance travelled in each habitat Habitat type Distance traveled (%) Open 47 Mixed-wood 27 Coniferous 26

Figure 4.1. Map of French Mountain showing the coyote paths followed and recorded in this area during the winter.

Figure 4.3. Fractal analysis results versus spatial scale. (A) Fractal dimension and (B) correlation in fractal dimension between successive track segments. The dotted lines represent the 95% confidence intervals. The increased Fractal dimension with scale (A) shows that coyote paths are more tortuous at larger spatial scales. The vertical gray bar in (B) shows where the plot crosses the y- axis, going from a positive to negative correlation as scale increases.

Chapter 5: Diet of eastern coyotes (Canis latrans var.) based on examination of scats during summer and winter in Cape Breton Highlands National Park, Nova Scotia,

Canada

Introduction

The coyote (Canis latrans) is native to the open plains of western Canada and the

US but has successfully expanded its range east across many different habitats and urban centers (Fox and Papouchis 2005). Coyotes arrived in New Brunswick in 1968

(Squires 1968; Schrecengost et al. 2009), reaching the mainland of Nova Scotia in 1977

(O’Brien 1983; Moore and Millar 1986) and moving to Cape Breton Island, where they were first recorded in Cape Breton Highlands National Park (CBHNP), in 1980 (E.

Muntz pers. comm. 2012). During the period of range expansion, western coyotes interbred with eastern wolves (Canis lupus lycaon) producing a new hybrid species named Canis latrans var. or the eastern coyote (Way et al. 2010). This hybridization has resulted in increased genetic diversity (Kays et al. 2010; Power et al. 2015).

Eastern coyotes have a larger body size and bigger skulls than western coyotes.

This increase in size likely contributes to their unique dietary and (potentially) behavioural characteristics (Gompper 2002). The larger skull provides additional surface area for muscle attachment, resulting in stronger jaws (Parker 1995). Notably, eastern coyotes appear to have adapted their diet to larger prey, including white-tailed deer

(Odocoileus virginianus; Kays et al. 2010) and moose (Alces alces; Benson and Patterson

2013), and seem to hunt in family groups (Parker 1995).

Coyote populations typically adhere to a generalist diet (Young and Jackson

1951; Bekoff 1977; Prugh 2005; Lukasik and Alexander 2011) and can adjust their

feeding habits on a regional basis and according to seasonal availability of prey and other food sources (Lukasik and Alexander 2011). Food selections range from small prey to large ungulates, livestock or pets, as well as foraging for fruit, eating garbage and scavenging large mammal carcasses (i.e., Bowyer et al. 1983; Fedriani et al. 2001;

Lukasik and Alexander 2011). Eastern coyotes have been known to effectively prey on adult white-tailed deer (Parker 1995; Patterson 1995), but more recently, coyotes have been documented killing adult moose in Ontario (Benson and Patterson 2013). A study in

Québec also detected moose as the dominant food item in eastern coyote scats (Boisjoly et al. 2010). This ability to consume large ungulates may indicate a shift in predatory ability.

Coyote conflicts with humans originated with attacks on farm animals in relatively rural settings, but these interactions have progressed to increasingly frequent encounters with humans in urban to suburban areas (Timm et al. 2004). The first lethal attack of a human was of a child in suburban California in 1981 (Timm et al. 2004). In

2009 in CBHNP an adult female was killed while hiking on a popular trail (E. Muntz personal comm. 2013). This was the first adult human fatality attributed to coyotes. The attack in CBHNP provided strong evidence of a deliberate act of human predation (S.

Gehrt pers. comm. 2012; B. Patterson pers. comm. 2012), although motive is unknown.

CBHNP has seen an increase in bold behaviour by coyotes toward humans since the early

1990’s. Between 2003 and 2012 there have been eight documented incidents of fearless behaviours (i.e., a coyote not backing away when approached by a person or approaching people but can be scared by people), 16 acts of aggressive behaviour (i.e., a coyote that: will not run away when people try to scare it, runs at people, approaches people multiple

times, circles or growls at people, but never makes physical contact), and six attacks (i.e., a coyote that: lunges at someone, attempts to make physical contact or succeeds at making physical contact), as defined by Parks Canada Human Wildlife Conflict protocol

(E. Muntz pers. comm. 2013). A primary motivation for the current study was the 2009 fatality to determine if coyotes in CBHNP have adopted a predatory behaviour, which may have contributed to their increased aggression towards humans.

Diet studies are used to determine which foods are utilized by wildlife and how, when and where these items are obtained (Patterson 1995). Food habit investigations contribute to a greater understanding of which foods within a particular region may become limiting to a population in times of stress (Patterson 1995). These investigations also provide insight into dynamic predator-prey systems. Dietary habits of coyotes can be studied by indirect observation such as snow tracking where predation events can be assessed along with identification of stomach contents or fecal remains. Several researchers (Lapierre 1985; Moore and Millar 1986; Backman 1987) in Nova Scotia and

New Brunswick have examined food habits of eastern coyotes based on the analysis of stomach contents. In a park system this is not possible due to the destructive nature of this sampling method. Thus, feeding behaviours in parks are best analyzed by fecal analysis and indirect observations (snow tracking). Back-tracking, especially in snow, is a convenient, non-lethal method used to study aspects of animal behaviour via indirect observations of prey capture, number of individuals, pack dynamics, movement patterns, prey remains, and collection of scats.

Coyote fecal material (scat) has been widely used to determine major dietary components for all seasons (Bowyer et al. 1983; Todd 1985; Parker 1986; Andelt et al.

1987; Toweill and Anthony 1988; Patterson et al. 1998; Dumond et al. 2001; Morey et al.

2007; Schrecengost et al. 2008; Lukasik and Alexander 2011). Bone fragments, hair, cartilage, feathers, nails, and other indigestible materials pass through the digestive system virtually unchanged, facilitating identification of prey. Many coyote scat studies document food habits in terms of percent frequency of occurrence for each prey species, although this method does not account for volume or weight of prey remains. Due to the frequent occurrence of several prey items in a single scat, overall percent occurrence is often greater than 100%. Despite its limitations, the widespread use of this method facilitates easy comparison with related regional studies.

Cape Breton Highlands National Park has a high density of moose (E. Muntz pers. comm. 2013). Thus, this study will explicitly examine the importance of moose to coyotes in CBHNP.

Environmental conditions such as heavy snowfall can negatively influence moose access to vegetation, which may lead to starvation and/or immobilization in deep snow- packs. Studies have shown that wolf-ungulate kill rates increase with snow depth (Ripple and Beschta 2004; Smith et al. 2003). When coyotes have been documented consuming large ungulates, especially moose, parts of the carcass are often left behind and one can try to piece together how the moose died (i.e., from starvation, injury or killed by predators). In addition to careful analysis of signs around a moose carcass, bone marrow fat content has been widely used as an index of body condition at the time of death for ungulates (Cheatum 1949; Baker and Leuth 1966; Neiland 1970; Franzmann and

Arneson 1976; Peterson et al. 1984; Ballard et al. 1987; Spears et al. 2003). Bone marrow fat is metabolized after other body fat reserves are depleted (Cheatum 1949; Smith and

Jones 1961). Reduction in bone marrow fat is considered a good indicator of stress, although this has been debated by some researchers (Mech and DelGiudice 1985; Ballard

1995). Marrow fat levels, collected from the femur, are frequently used to develop a generalized description about the overall condition of an individual ungulate at time of death (Cheatum 1949; Franzmann and Arneson 1976; Peterson et al. 1982; Ballard et al.

1987; Ballard 1995; Spears et al. 2003).

Feeding habits of coyotes in CBHNP have not been well studied. In the wake of the human fatality, where it is unclear if the attack was an isolated incident or the beginning of increased coyote aggression, understanding coyote diet is important. Since humans are regular visitors to the park, the objective of this study was to analyze the diet of coyotes in CBHNP and observe foraging habits to better understand predation dynamics and verify targeted prey species. Characterizing the availability/consumption of different prey items can offer evidence of regionally specific foraging patterns and may provide a potential motive for increased aggressive behaviours towards humans. More broadly, it allows for a thorough comparison of diets, predatory behaviours, and interactions between the two over a large geographic range.

Feeding behaviour was based on regular scat collection along designated transects during fall, summer, winter and spring. Dietary analysis and prey availability were studied to establish baseline information on feeding ecology of coyotes in CBHNP.

Relative frequency of occurrence of prey and prey availability was determined during the winter months, normally a challenging period for coyotes.

Methods

This study took place in CBHNP and adjacent areas from May 2012 to August

2013. Scats were collected from May 2012 to August 2013, back-tracking was conducted from January 2013 to April 2013. Prey availability and carcass remains of prey were examined from data collected while back-tracking.

Transects Scat collection transects 21 transects (Figure 5.1) were designated across the study area (18 park trails and

3 gated roads). A two km transect was randomly chosen along each park trail or gated road for study. If a trail was two km or less in length the total trail was surveyed. Prior to the formal scat surveys the 21 transects were cleared of all coyote scats to ensure collection of only new scats of known age. Transects were surveyed for scats approximately every three weeks over the fall, winter, spring and summer months.

Summer scats were collected from 21 June to 21 September (summer), 22 Septemeber to

20 December (fall), 21 December to 19 March (winter) and 20 March to 20 June (spring).

In light of the frequency and amount of snowfall over the winter months, care was taken in the early spring to determine if the scat was from the winter or spring. Scats were also separated by region; Southwestern (lowlands south west of French Mountain), Western

(plateau - French Mountain east to base of North Mountain) and Easten (hardwoods - base of North Mountain east to Alantic Ocean). Scats were also collected opportunistically (n = 52) in high traffic areas of the park while driving along the Cabot

Trail and added to the appropriate season and region. All scats collected were placed individually in sealable plastic bags with a data sheet noting tag number, general location,

date, and waypoint (Universal Transverse Mercator (UTM)). Collected scats were frozen

(-20° C) at the end of each day until further processing.

lynx (Lynx canadensis), bobcat (Lynx rufus), and red fox (Vulpes vulpes) were also present within the study area and the species of all scats were identified in the field based on diameter and length (Murie 1954). Care was taken to discard any samples that could not be positively identified as coyote.

Transect prey observations (winter) Thirty km of transects were completed while there was snow cover. For these transects, signs of potential prey species were recorded as a waypoint location and recorded as a track (single set of tracks), cluster (over three sets of tracks within a 10 m area), scat, or urine of prey species. The distance of the sign from transect was recorded as either within three m (i.e., crossing through transect) or > three m (i.e., can see from transect but is greater than three m from transect and does not cross). Relative frequency of occurance was calculated for all prey observations within three m of transects and back-tracking coyotes (see Scat Analysis for formula).

Scat processing Scats were placed in sealed nylon stockings and boiled for a minimum of 10 minutes to kill any bacteria, parasites and/or eggs. Scats were then washed to separate undigested remains from the scat matrix. Undigested material (hair, bones, etc.) was air dried, sorted and identified to species by comparison with a reference collection from the

Acadia University Wildlife Museum. Hair samples were identified by macrofeatures and microscopic identification of cuticular scale patterns, following methods outlined by

Adorjan and Kolenosky (1969). Prey species with heightened risk of improper

identification, such as small mammals and birds, were placed into general categories of:

1) small mammals (squirrels, mice, voles and shrews) and 2) birds.

Scat analysis Relative Frequency (RF) of occurrence of each prey remain was calculated based on the protocol described by Alvarez-Castaneda and Gonzalez-Quintero (2005). Relative

Frequency is represented as: RF = (fi / ∑n)(100), where RF is relative frequency, fi is the number of scats containing the prey item (frequency) and ∑n is the total number of scats in the sample. This value represents how common a prey item is, relative to the total number of scats collected in each season and elevation. Percent by volume for each prey remain was determined using the point-frame method (Chamrad and Box 1964). The percentage was obtained by adding prey remains to a plastic petri dish with a 2.5 cm grid and counting the number of squares occupied by each prey item. This number was divided by the total squares occupied by the sample and then converted to a percentage.

This method is likely less subjective than the common visual estimation of percent by volume (e.g., as in Patterson 1995 and Schrecengost et al. 2008). Prey remains that made up less than two percent of the scat volume were considered “trace” and were removed from further analysis to minimize biased emphasis (Morey et al. 2007)

Moose carcass processing If a moose carcass was discovered during back-tracking, photos were taken prior to approaching the carcass and the following information was recorded when possible: date of observation, time of observation, location (UTM), observers, general location, number of coyote tracks going in and out of the site, topography and habitat characteristics, snow depth, snow condition, body length (cm), presence of broken bones, sex, body part consumed, total portion of body consumed, jaw bone collected (yes or no),

femur collected (yes or no) and a description of the site/event. Date of death was estimated based on snow condition (recent documented storms and snow cover on the carcass). Lower jaws and femurs, when present, were collected and stored in a freezer at

-20° C.

Processing moose femurs and teeth Femurs were broken by closing a table-mounted vise on the bone. Raw bone marrow (10 cm long) from the central portion of the marrow tube was added to a pre- weighed plastic petri dish. All petri dishes with bone marrow were stored in a cool dark room, covered with cheese cloth (which did not touch the marrow) to prevent insects from accessing the marrow. Petri dishes and marrow were weighed (to the nearest 0.1 gram) every day until no further weight loss was detected, which indicated that all water content had evaporated and only the fat content remained. The final marrow weight was divided from the initial weight to determine the percent fat in the bone marrow. This measure was used as a rough measure of overall health at time of death (Leighton 2000).

Moose were aged based on tooth eruption, replacement, and wear using the lower jaws. Lower jaws that were collected from carcasses were compared to a reference collection of lower jaws from moose of known ages at the Parks Canada library. Due to cost constraints, half of the total samples were age-verified by sending two incisors to

Matson’s Laboratory (Manhattan, Montana, USA). Ages obtained from the incisors confirmed that the original estimates were accurate.

Statistical analyses Statistical analyses were conducted in R Studio (version 0.98.490; R version

3.0.2). A Pearson’s Chi-squared Test was used to determine if there was a statistically significant difference between frequency of occurrence over different seasons and regions. A Kruskal-Wallis Test was used to determine if there was a statistically significant difference for percent by volume over different seasons and regions. A

Kruskal-Wallis Test was used to determine if there was a statistically significant difference between seasons and regions in terms of prey richness.

Results

Relative frequency of occurrence A total of 294 coyote scats were collected along 966 cumulative km of trail transects. Dietary analysis of these scats indicated that moose, fruit/berries and snowshoe hare (Lepus americanus) had the highest percentage of occurrence by season and locality

(Table 5.1 and 5.2). Overall moose was the most common food item found in coyote scats (56.8%), followed by fruit/berries (33.7%) and snowshoe hare (23.5%). Small mammals (15.0%), birds (11.2%) and deer (2.0%) were less common. Lastly, anthropogenic food sources (garbage) were found in only two scats (0.70%).

Calendar Season Siginificant differences were found for relative frequency of occurrence in different calendar seasons for moose, deer, bird, small mammal and fruit (Table 5.1).

Moose was the dominant prey item for spring and winter (Table 5.1). Moose and fruit/berry remains were the highest relative frequency of occurrence for the fall season

(Table 5.1). Fruit/berries were the highest frequency of occurrence for the summer season

(Table 5.1). Deer remains were only found in the spring and winter seasons (Table 5.1).

Snowshoe hare remains were fairly consistent between calendar seasons with the hightest frequency of occurrence in the winter (Table 5.1). Bird remains were found in every season with the highest frequency of occurrence in the fall (Table 5.1). Small mammal remains were found in every season with the highest frequency of occurrence in the fall

(Table 5.1).

Region Although significant differences were found across regions for relative frequency of occurrence for moose, deer and snowshoe hare remains (Table 5.2), moose were the primary prey item for coyotes in all regions with the highest frequency of occurrence in the western region (Table 5.2). Deer remains were only found in the southwestern region

(Table 5.2). Snowshoe hare remains varied between the three regions with the highest frequency of occurrence in the eastern region (Table 5.2). Bird remains varied by region with the highest frequency in the southwestern region (Table 5.2). Small mammal remains were found in each region and had the highest frequency of occurrence in the southwestern region (Table 5.2). Fruit/berry remains were varied by region with the highest frequency in the eastern region (Table 5.2).

Percent by volume Calendar Season Siginificant differences were found for percent by volume in different calender seasons for moose, deer, bird, small mammal and fruit remains (Table 5.3). Moose remains were found above 50.0% biomass in scats for all seasons and were the highest in spring and winter (Table 5.3). Percent by volume for snowshoe hare was the highest in spring and winter. Bird remains were the highest in spring. Small mammal remains were

the highest in the fall and lowest in the summer (Table 5.3) Fruit was the highest in summer (Table 5.3).

Region Siginificant differences were found for percent by volume in different regions for moose, deer and snowshoe hare (Table 5.4). Moose remains for percent by volume were significantly less in the southwestern region compared to the eastern and western regions

(Table 5.4). Deer remains for percent by volume were only found in the southwestern region (Tabble 5.4). Snowshoe hare remains for percent by volume were significantly less in the western region compared to the southwestern and eastern regions (Table 5.4).

Bird remains for percent by volume were the lowest in the western region followed by the southwestern region and the highest in the eastern region (Table 5.4). Small mammal remains for percent by volume were lowest in the southwestern region followed by the western region and the highest in the eastern region (Table 5.4). Fruit percent by volume remains were fairly consistent between regions (Table 5.4).

Prey richness There was a significant difference (X2 = 66.87; p ≤ 0.000) between seasons for prey richness. The summer season saw the greatest number of prey items in the scat of coyotes (Table 5.5). There was no difference between prey richness for any region (X2 =

3.13; p = 0.21; Table 5.6).

Prey observations Prey availability surveys were conducted during the winter months along 116 km of back-tracking coyotes. Relative frequency of occurrence for prey observations during back-tracking coyotes were highest for tracks of snowshoe hare (32.8%), small mammals

(31.9%) and moose (12.6%); clusters were highest for snowshoe hare (6.7%) moose

(4.1%) and small mammals (3.8%); scat/urine were highest for moose (2.9%) and bird

(2.0%; Table 5.7). Prey availability surveys were conducted along 30 km of transects.

Relative frequency of occurrence for prey observations during winter transects were highest for tracks small mammal (31.9%) moose (19.0%) and snowshoe hare (15.0%).

Clusters were highest for moose (4.8%), small mammals (2.6%) and snowshoe hare

(1.8%; Table 5.7). Scat/urine was highest for bird (3.7%) and moose (2.9%; Table 5.7)

Moose carcasses A total of 17 moose carcasses, all female, were found while back tracking coyotes

(Table 5.8). The ages ranged from >1.5 to 10.5 years old; the percent of fat in the bone marrow ranged from 6.0% to 80.9%; the body length ranged from 175 to 241 cm; and the height ranged from 120 to 185 cm (Table 5.8). Only three individuals had injuries that could be detected (broken spine, broken front left leg and broken hind right leg; Table

5.4). The amount of the carcass consumed varied from minor scavenging to the majority of the carcass with just bits of meat/bone marrow remaining. Generally the body cavity was the primary portion consumed on newly scavenged carcasses (Appendix 5.1). The number of coyote tracks and scats collected around carcasses was 2-5 and 1-10, respectively (Appendix 5.1). The snow depth, snow conditions, topography and other observations varied by site (Appendix 5.1). Six moose likely died from a storm with heavy wet snow (based on the amount of snow and upright position of moose), causing moose to not be able for forage for food, thermoregulate, or move. Two moose likely died by falling off a bank and getting immobilized in heavy snow (presence of steep or rocky bank, moose lying in an odd position, and amount of snow moose were found in), three carcasses were discovered near the Cabot Trail (major highway) and may have been struck by a vehicle or scared into heavy snow. Two moose likely died by falling through

ice (thick ice with hole broken through and river or brook present) and could not get out of the snow or ice (Appendix 5.1). There was evidence of coyotes digging through snow to reach all carcasses.

Discussion

Results from this study confirm that eastern coyote is a generalist predator capable of utilizing a diverse array of food types. However no other study has documented such high annual use of moose; the frequency of moose in scats was 56.8% annually. Other studies have found moose to make up only a portion of the diet of coyotes: eastern Maine (Litvaitis and Harrison 1989; 0.63%), northwestern Wyoming

(Dowd and Gese 2012; 8.3%), southeastern Québec (Richer et al. 2002; 8%), western

Maine (Major and Sherburne 1987; 6%), and New Brunswick (Dumond et al. 2001;

8.2%). Only in eastern Québec did Boisjoly et al. (2010) report a comparable frequency of 51% moose in scats. This study documented the highest frequency of occurrence of moose in scats during the winter (77.5 %) of coyotes in CBHNP.

Numerous studies from northeastern North America have documented white- tailed deer as a primary prey item for coyotes (Hilton 1978; Harrison and Harrison 1984;

Lapierre 1985; Moore and Millar 1986; Parker 1986; Patterson et al. 1998). White-tailed deer in this study were only about 2.0% annually, compared to results by Patterson

(1995) reporting 68.4% and Litvaitis and Harrison (1989) documenting 34.9% of annual scats containing deer remains. White-tailed deer densities in the highlands of CBHNP are extremely low (Bridgland and Millette 1984), and coyotes may only have access to deer in the lowlands (E. Muntz. pers. comm. 2014).

The annual occurance of all fruit and berries was 33.7 %. Fruit and berry use were consistent between regions in the park but were consumed most during the summer and fall. Patterson (1995) and Litvaitis and Harrison (1989) saw the use of fruits by coyotes to be 20.3% and 46.2% respectively. Both Patterson (1995) and Litvaitis and Harrison

(1989) saw an increased use of fruit and berries during the summer and fall months similar to this study, which is not suprising given these are the periods when fruit typically ripens.

Snowshoe hare was the third most common food item of coyotes (23.5% annually) in CBHNP, and was consumed in every season. The annual occurance of snowshoe hare in coyote scats was lower in this study than many other diet studies in northeastern North America; Maine 30 % (Harrison and Harrison 1984), Maine 41.1 %

(Litvaitis and Harrison 1989) and southeastern New Brunswick 51.4 % (Lapierre 1985).

This study reports similar annual use of snowshoe hare (24.2 %) compared to Patterson

(1995) who reported 20.6% in Kejimkujik National Park. Snowshoe hare are known to be cyclical and have highs and lows in their population every 7-10 years (Stefan and Krebs

2001). This study and Patterson (1995) have both reported low population levels of snowshoe hare during field studies.

The annual use of small mammals in the current study was 15.0% which is on par with what Patterson (1995) found (10.7%) in KNP and Litvaitis and Harrison (1989) found (15.8%) in Maine. The current study saw an increased use of small mammals by coyotes in the fall and summer, similar to results reported by Litvaitis and Harrison

(1989) but Patterson (1995) found small mammal use to be consistent during spring, summer and fall with reduced use in winter.

Birds in the current study had an annual use by coyotes of 11.2% which is higher than Patterson (1995) documented in KNP (2.8%) and higher than Litvaitis and

Harrison’s (1989) findings in Maine (4.7%). Patterson (1995) found bird use by coyotes to increase in spring and fall and Litvaitis and Harrison (1989) had similar results to the current study.

Diversity of prey items in the diet of coyotes was similar throughout the year and region, however there was an increase in diversity during the summer season. Litvaitis and Harrison (1989) in Maine show similar results where diversity is consistent throughout the year with a slight decrease in diversity noted in summer and winter seasons. Patterson (1995) in KNP saw an increase in prey diversity in the fall season while Dowd and Gese (2012) saw an increase in diversity in the spring season.

The high dependance on moose, especially during the winter, is unique but may not be surprising. There are an estimated 7,500 moose in CBHNP with most living on the highlands (J. Bridgland pers. comm. 2014; M. Smith pers. comm. 2015). Do coyotes hunt and kill moose or do they scavenge moose that died from starvation or got caught in heavy snow?

Recently Benson and Patterson (2013) suggest it is possible for eastern coyotes to kill moose, and they cite evidence of a group of eastern coyotes killing a 20-month old moose in Ontario. Eastern coyotes have been know to form packs to cooperatively hunt or defend carcasses (Bowen 1981). However, the majority of moose carcasses examined in this study were adults that appeared to have died from causes other than coyote attacks.

It is possible that coyotes in CBHNP are killing moose, however the data from this study does not support this idea, but rather suggests that coyotes scavenge moose carcasses

where the animals were already dead (possibility from starvation, broken limbs or after becoming trapped in deep snow). In fact, Boisjoly et al. (2010) demonstrated that most moose consumed by coyotes in Québec were scavenged.

Death by starvation can be detected in moose carcasses by examining the percentage of marrow fat. Franzmann and Arneson (1976) and Peterson et al. (1984) classified calf and adult moose with marrow fat levels from femurs of <10% and <20%, respectively, to be near death from starvation. Other authors have reported marrow fat levels >10% for calves that have starved to death (Franzmann and Arneson 1976; Ballard et al. 1987). In the current study there were a number of compromised moose carcasses

(either deemed to be near starvation or with broken bones). Five moose carcasses that had bone marrow fat content <10% likely died from exhaustion while walking through deep snow or starved to death. The other deaths could be related to a heavy wet snow storm

(20-23 March 2013) resulting in moose getting permanently stuck, rendering them unable to thermoregulate or to access food and water. Winters in CBHNP have sigificant levels of snow, making it hazardous for moose to travel (Bridgland et al. 2007). From 20-23

March 2013, more than 60 cm of snow accumulated at CBHNP. This, combined with the already signficant levels of snowfall that had accumulated over the winter, likely trapped many moose throughout the park and their movement could be severely restricted for several days to weeks.

In several cases, moose carcasses were found in atypical locations in the park, including the steep slope of a cliff or in the middle of a river. It has been suggested (E.

Muntz pers. comm. 2013; S. Gehrt pers. comm 2013; B. Paterson pers. comm. 2013) that instead of chasing and killing moose, coyotes might only follow and tire them, purposely

guiding them toward a specific area where the likelihood of the moose being trapped/ultimately dying is high. A NS DNR helictoper pilot observed coyotes on both side of an ice bridge with a moose in the middle (D. Farrell pers. comm. 2014). After exhausting the moose, coyotes could move in for the kill. This form of predation is almost impossible to demonstrate without actually witnessing the event, and no tracks in snow have been found during the winter that would provide more direct evidence for this kind of behaviour. It is possible that coyote tracks indicating a chase may have been erased by new fallen snow, as none of the moose carcasses were found immediately after death. Thus, the findings of this study suggest coyote feeding on moose in CBHNP are primarily opportunistic.

Moose may have been the most biomass rich food source available to coyotes, especially on the highlands, during this study. A moose carcass may be able to sustain a coyote group for days or weeks. Moose contain a high level of protein and energy rich content; furthermore, less energy may be required to consume a moose during winter and spring months than would be expended in the hunt for small mammals. Coyote may rely on moose in CBHNP because of their apparent availability and lack of other prey, especially in the more northern regions and on the plateau regions of the park (E. Muntz pers. comm. 2014). Snowshoe hare (E. Muntz pers. comm. 2014) populations were recovering from a low in their cycle, possibly causing coyotes to rely on moose for winter survival.

The findings of this study still leave critical questions remaining in terms of the implications for park management and strategies for reducing risk to human visitors to park settings. Does a heavy dependance on moose somehow make coyotes in CBHNP

more likely to view humans as prey and thus act more aggressively? While this may be an unlikely conclusion, at the very least it may suggest that coyotes on the highlands are sensitive to the boom-and-bust fluctuations that occur within the CBHNP in terms of reduced prey diversity (compared to the community of Chéticamp and Kejimkujik

National Park; Patterson 1995) and limited avaivability of small and medimum sized prey species (i.e., snowshoe hare and white-tailed deer).

Cape Breton Highlands National Park recieves over 150 000 vistors each year (E.

Muntz pers. comm. 2013) and the most heavily used trail hosts over 25 000 visitors a year (A. Boudreau pers. comm. 2012). Coyotes in CBHNP are active during the day (see

Chapter 4, this thesis), making it highly probable that they will encounter people. There was no strong evidence for consumption of human food (no significant occurance in scats), but it remains likely that human food could be left behind on trails which could lead to habituation, especially in cases of limited food availability (Schmidt and Timm

2007). A multi-year diet study is necessary to provide a better understanding of how prey diversity, winter survivorship of moose and weather affects coyote foraging behaviour.

Stable istope work is underway as part of another study and may offer more concrete evidence of human garbage in the diet of coyotes.

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Personal Communaction

Allan Boudreau, Research Officer, Parks Canada, Halifax, Nova Scotia.

Michael Boudreau, Human-Wildlife Conflict Biologist, Nova Scotia Department of Natural Resources-Wildlife Division, Kentville, Nova Scotia.

James Bridgland, Park Ecologist, Parks Canada Ingonish, Nova Scotia.

Dave, Farrell, Helicopter Pilot, Nova Scotia Nova Scotia Department of Natural Resources-Air Services Division, Shubenacadie, Nova Scotia.

Stan Gehrt, Associate Professor, Ohio State University, Columbus, Ohio

Erich Muntz, Project Manager, Parks Canada, Chéticamp, Nova Scotia.

Mathew Smith, Park Ecologist, Parks Canada, Maitland Bridge, Nova Scotia.

Brent Patterson, Adjunct Professor, Trent University/Ontario Ministry of Natural Resources, Peterborough, Ontario.

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Table 5.1. Calendar season frequency (# scats/mean %) of occurrence of prey remains identified from coyote scats collected in Cape Breton Highlands National Park from May 2012 through August 2013. A Chi-squared test was use to test for differences between seasons of prey remains.

Calendar season

Fall Spring Summer Winter # of scats (n = 40) (n = 64) (n = 110) (n = 80) X2 p-value

13.69 Moose 18/45.0% 49/77.0% 38/34.5% 62/77.5% 72 p = 0.003348

12.99 Deer 0/0.0% 5/7.8% 0/0.0% 1/1.3% 31 p = 0.004652

1.651 Snowshoe hare 11/27.5% 13/20.3% 22/20.0% 23/28.8% 4 p = 0.6478

16.45 Bird 8/20.0% 1/1.6% 21/19.1% 3/3.8% 8 p = 0.0009133

25.01 Small mammal 12/30.0% 2/3.1% 28/25.5% 2/2.5% 04 p ≤ 0.0001

79.58 Fruit 18/45.0% 0/0.0% 81/73.6% 0/0.0% 62 p ≤ 0.0001

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Table 5.2. Regional frequency (# scats/mean %) of occurrence of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Chi-squared test was use to test for differences between regions of prey remains.

Region

East Southwest West # scats (n = 64) (n = 95) (n = 135) X2 p - value

Moose 30/47.0% 41/43.0% 96/71.0% 5.7747 p = 0.05572

Deer 0/0.0% 6/6.3% 0/0.0% 12.063 p = 0.002402

Snowshoe hare 19/30.0% 30/31.6% 20/14.8% 6.5887 p = 0.03709

Bird 6/9.4% 14/14.7% 13/9.6% 1.3674 p = 0.5047 Small mammal 11/17.2% 19/20.0% 14/10.4% 3.2467 p = 0.1972

Fruit 25/39.0% 34/36.0% 40/30.0% 0.9957 p = 0.6078

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Table 5.3. Calendar season percent by volume (mean% ± SD) of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Kruskal-Wallis Test was used to test for differences between calendar seasons of prey remains.

Calendar season Fall Spring Summer Winter 2 Prey remains (% ± SD) (% ± SD) (%± SD) (%± SD) X p - value Moose 23.3 ± 39.4 70.9 ± 44.5 18.0 ± 35.8 71.2 ± 44.5 78.9827 p ≤ 0.000

Deer - 7.8 ± 27.0 - 1.2 ± 11.3 13.9919 p = 0.002916

Snowshoe hare 16.2 ± 34.6 18.1 ± 37.5 11.8 ± 30.4 24.8 ± 42.2 3.5261 p = 0.3174

Bird 8.6 ± 24.1 1.6 ± 12.5 1.9 ± 6.4 0.9 ± 5.8 19.7626 p ≤ 0.00 Small mammal 22.9 ± 39.6 1.6 ± 12.1 12.0 ± 27.6 1.9 ± 12.1 33.4998 p ≤ 0.000

Fruit 29.1 ± 41.9 - 56.3 ± 43.1 - 147.2538 p ≤ 0.000

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Table 5.4. Regional percent by volume (mean% ± SD) of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Kruskal-Wallis Test was used to test for differences between regions of prey remains.

Region East Southwest West Prey remains (% ± SD) (% ± SD) (% ± SD) X2 p - value

Moose 41.4 ± 48.4 26.4 ± 43.0 59.2 ± 47.1 26.537 p ≤ 0.000

Deer - 6.3 ± 24.5 - 12.7866 p = 0.001673 Snowshoe hare 21.6 ± 39.8 25.7 ± 42.0 9.4 ± 27.8 11.4092 p = 0.003331

Bird 2.6 ± 13.7 3.2 ± 16.7 1.9 ± 9.5 1.7684 p = 0.413 Small mammal 10.8 ± 28.1 10.1 ± 25.8 6.2 ± 22.5 4.1708 p = 0.124

Fruit 23.6 ± 38.8 28.2 ± 41.7 23.4 ± 39.6 1.3241 p = 0.516

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Table 5.5. Calendar season richness of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Kruskal-Wallis Test was used to determine if there was a statistically significant difference between calendar seasons for prey richness.

Calendar season Fall ± SD Spring ± SD Summer ± SD Winter ± SD X2 p - value

Prey remain 1.7 ± 0.90 1.1 ± 0.30 1.7 ± 0.72 1.1 ± 0.40 66.87 p ≤ 0.000 richness

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Table 5.6. Regional richness of prey remains identified from coyote scats collected on trail transects in Cape Breton Highlands National Park from May 2012 through August 2013. A Kruskal-Wallis Test was used to determine if there was a statistically significant difference between regions for prey richness.

Region East ± SD Southwest ± SD West ± SD X2 p - value

Prey remain 1.4 ± 0.70 1.5 ± 0.71 1.4 ± 0.60 3.13 p = 0.21 richness

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Table 5.7. Relative frequency of occurrence (%) of prey species observed within three meters of both back-tracking coyotes and winter transects in Cape Breton Highlands National Park from January to April 2013.

Collection method km Moose Snowshoe hare Deer Bird observations Small mammal surveyed observations observations observations observations

Back-tracking 116 19.6 40.3 0.5 3.9 35.7 Transects 30 26.7 34.4 0.0 4.4 34.4

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Table 5.8. Moose (Alces alces) body condition at eastern coyote scavenging sites in CBHNP from January to April 2013.

General location Age Sex Fat in original bone Body length (cm) Body height (cm) Date of Death Broken limbs or bones marrow (%)

North Mountain 10.5 Female 50.2 180 - Feb. 2013 Broken spine

Lower Mary-Anne Falls road 9.5 Female 33.0 203 183 Mar. 2013 None apparent

Branch pond trail <1.5 Female 80.9 241 - Mar. 2013 None apparent

Sugar Brook lake area 10.5 Female 9.8 240 160 Feb. 2013 None apparent

North Mountain 4.5 Female 7.1 206 160 Mar. 2013 None apparent

Lower Chéticamp river 4.5 Female 40.5 210 175 Mar. 2013 Left front leg

Near Green Cove 7.5 Female 49.3 203 160 Mar. 2013 None apparent

Near Jigging Cove trail head 2.5 Female 55.6 213 180 Mar.2013 None apparent

Near Broad Cove Campground <1.5 Female 9.0 175 144 Mar. 2013 None apparent

Sunday Lake area 8.5 Unknown 31.5 - - Feb. 2013 None apparent

Upper Mary-Anne Falls road 8.5 Female 12.3 230 185 Mar. 2013 Right hind leg

French Mountain 4.5 Female 7.4 - - Jan. 2013 None apparent

Salmon Pools trail 2.5 Female 49.6 220 120 Feb. 2013 None apparent

Paquette lake area 5.5 Female 6.0 - - Mar. 2013 None apparent

French Mountain - Unknown 65.7 - - Feb. 2013 None apparent

French Mountain 9.5 Female - - - Mar. to Apr. 2013 None apparent

Old Black Brook Campground - Female - - - Mar. to Apr. 2013 None apparent

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Figure 5.1. Transect locations identified for scat and prey observations in Cape Breton Highlands National Park. Blue dots represent transect locations (18 park trails and three gated roads).

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Appendix 5.1. Description of moose carcasses at eastern coyote scavenging sites in Cape Breton Highlands National Park from January to April 2013.

General location Amount Portion Consumed Visible coyote scats Pack Snow Depth (cm) Snow Conditions Site Characteristics Comments Consumed (%) size /Topography North Mountain 50 LH 3 3 to 4 60 I; HCS MF; SB (50%) FOB Lower Mary-Anne Falls 40 BC 1 3 40 HCS; I; 5cm FS MF; OA; SG (3-5%) SWHS; MHS road Branch pond trail 30 to 40 HTQ; BC 1 5 214 HCS; 5cm FS OA: SW; SG (2-3%) SWHS; MHS Sugar Brook lake area 60 to 70 LJ; FQ; BC 1 3 120 HCS; 5cm FS EB; SB (30%) FOB North Mountain 50 RH 2 3 to 4 70 I; HCS OA; SW; FT SWHS; MHS Lower Chéticamp river >10 RFJ 5 3 to 4 100 (30 ice) I; HCS OA; RFP; SG (2-3%) EWC; URB; FTI Near Green Cove 20 Head; neck; RS 1 2 46 WS SRB; SG (5-10%) 40m from CT; FTI Near Jigging Cove trail 30 to 40 FLS; BC 4 3 to 4 16 WS OA; MF; FT 300m from CT head Near Broad Cove 10 to 20 HTQ; BC 1 2 90 I; HCS SW; SG (3-5%) 30m from CT; MHS Campground Sunday Lake area 60 to 70 BC; HL 5 3 to 4 305 HCS SW; EB; FT SWHS; MHS Upper Mary-Anne Falls >10 BRH 3 3 to 4 85 I; HCS OA; SW; FT SWHS; MHS road French Mountain 60 to 70 Everything but legs; 2 4 80 I; HCS OA; SW; FT MHS BM Salmon Pools trail 10 Nose; FLL; URC 3 4 120 I; W DBH; SBB; SG (3-5%) FTI; MHS Paquette lake area 95 Everything BM 3 4 46 WS OA; SW; FT SWHS; MHS French Mountain - BM remaining 3 2 90 FS SWT; FT Only found two leg bones French Mountain <95 Everything except 10 - - - OB; FT DDW senu Old Black Brook <95 Everything except 2 - - - OA; SW:FT DDW; SSE Campground senu Portion Consumed: Left half (LH), Body cavity (BC), Hind top quarter (HTQ), Lower jaw (LJ), Front quarter (FQ), Right half (RH), Right front joint (RFJ), Right shoulder (RS), Front left shoulder (FLS), Hind legs (HL), Back right hip (BRH), Bone marrow (BM), Front left leg (FLL), Upper rib cage (URC), Left hind quarter (LHQ), Head, Nose, Neck, Leg, and Senu. Snow Conditions: Water (W), Hard crusty snow (HCS), Fresh snow (FS), Ice (I), and Wet snow (WS). Site Characteristics\topography: Open area (OA), Steep bank (SB), Slight grade (SG), Edge of bog (EB), Mixed forest (MF), Soft wood (SW), Open bog (OB), Deep brook hole (DBH), Soft brook bottom (SBB), Bog wet hole (BWH), Flat terrain (FT), Shallow running brook (SRB), and River flood plain (RFP). Comments: Storm with wet heavy snow (SWHS), Moose may have been stuck (MHS), Died during the winter (DDW), Smell was still evident (SSE), Moose may have fell off bank (FOB), Cabot trail (CT), Fallen through ice (FTI), Extreme water currents (EWC), and Unstable rocky bottom (URB).

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Chapter 6: General Conclusions

This study examined the ecology of eastern coyotes in Cape Breton Highlands

National Park (CBHNP) and surrounding vicinity (Chéticamp) to determine if the ecosystems in the park could induce coyotes to become more aggressive towards human visitors. Since 2003 there have been 30 documented aggressive interactions between coyotes and people within the park. Concurrently, in Chéticamp, there have been no such reports despite the presence of coyotes (Table 6.1). This study specifically examined genetic characteristics (197 individuals from the Atlantic Provinces), movement patterns

(based on winter snow tracking and 11 GPS collared animals) and diet (scats) of coyotes.

Coyotes in Cape Breton are genetically different from the rest of Atlantic Canada.

Coyotes in Nova Scotia have both western coyote and eastern wolf mtDNA. Mainland

Nova Scotia has the same three haplotypes (cla28, cla29 and GL20) as coyote populations in New Brunswick. However, in Cape Breton, diversity of haplotypes was reduced, with individuals possessing only the cla28 and GL20 haplotypes. These results are consistent with Kays et al. (2010) which suggests haplotype diversity decreases as coyote populations move east. Male eastern coyotes that also have eastern wolf mtDNA are significantly larger than male coyotes with western coyote haplotypes.

This study has a number of parallels to research conducted by Patterson (1995) in

Kejimkujik National Park (KNP), but coyotes in Cape Breton have larger cumulated areas of use in relation to KNP. Coyotes collared in the community of Chéticamp tend to avoid human use biotopes (CH-7; CH-3), where some individuals in CBHNP do not.

Daily movement patterns demonstrate coyotes in CBHNP move more than coyotes in the

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community of Chéticamp during daylight hours. In general, coyotes in and around

Chéticamp were found to avoid movement during the day.

Fractal analysis revealed that coyotes in CBHNP move differently through the landscape compared to KNP. Coyotes in CBHNP tended to travel in straight lines, likely from one moose carcass to another and thereby conserving energy, by using open landscapes. In KNP coyotes tended to hunt more for prey and had irregularly movement patterns through the landscape.

Diet analysis showed coyotes in CBHNP rely on moose as a stable food item.

Prey diversity was highest in Chéticamp; the limited prey diversity in CBHNP especially on the highlands suggests coyotes in this area may have been more food-stressed. During winter back-tracking in 2013 coyotes fed on 17 moose carcasses, although causes of mortality were likely to be physiological and environmental (e.g., broken limbs, poor body condition or entrapment in deep snow) rather than a result of direct predation by coyotes (Table 6.1).

Coyotes from CBHNP tend to have larger area of concentrated use, do not avoid areas of high human use, move farther during daylight hours than dark hours, have less diversity of prey and are exposed to less negative association from humans (Table 6.1). It appears coyotes in CBHNP do not avoid humans as much as coyotes in the community of

Chéticamp, likely because they do not fear humans. Park animals may perceive humans as non-threatening, whereas in Chéticamp and vicinity coyotes are actively trapped and hunted (Table 6.1).

The eastern coyote is a permanent resident in Nova Scotia and will continue to be an important member of the ecosystems of Cape Breton Highlands National Park. Given

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the unique environment where prey diversity is limited and moose is the primary large mammalian prey species and coyotes show reduced genetic haplotypes, mangers will need to continue to monitor coyote behaviour and educate the public regarding human/wildlife conflict.

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Table 6.1. Differences between coyotes of Cape Breton Highlands National Park and those within the community of Chéticamp. Chéticamp Cape Breton Highlands National Park Food diversity High Low Negative human interactions High Low Coyote aggression Low High Human food source High Probably some

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Table 6.2. Differences among coyotes in Cape Bretion Highlands National Park and Atlantic Canada.

Item Atlantic Canada Cape Breton Highlands

Home range Small Large

Prey abundance High Low

Prey diversity High Low Travel patterns Irregular Straight line

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References

Patterson, B. R. 1995. The ecology of the eastern coyote in Kejimkujik National Park. MSc thesis, Acadia University, Wolfville, Nova Scotia, Canada.

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