HOME RANGE OVERLAP and SPATIAL-TEMPORAL INTERACTIONS in FEMALE FISHERS on the HOOPA INDIAN RESERVATION in CALIFORNIA by Kerry M

HOME RANGE OVERLAP AND SPATIAL-TEMPORAL INTERACTIONS IN

FEMALE FISHERS ON THE HOOPA INDIAN RESERVATION IN CALIFORNIA

Kerry M. Rennie

A Thesis Presented to

The Faculty of Humboldt State University

In Partial Fulfillment of the Requirements for the Degree

Master of Science in Natural Resources: Wildlife

Committee Membership

Dr. Micaela Szykman Gunther, Committee Chair

J. Mark Higley, Committee Member

Dr. Matthew Johnson, Committee Member

Dr. William J. Zielinski, Committee Member

Dr. Yvonne Everett, Graduate Coordinator

July 2015

ABSTRACT

HOME RANGE OVERLAP AND SPATIAL-TEMPORAL INTERACTIONS IN FEMALE FISHERS ON THE HOOPA INDIAN RESERVATION

Kerry M. Rennie

Spatial organization and interactions within animal populations can have important effects on population demography, social dynamics, and genetic composition. Understanding social behavior and relatedness of individuals in a population can advise species management and conservation. Previous research suggests that fishers (Pekania pennanti) are asocial and exhibit intrasexual territoriality, where home ranges rarely overlap with the same sex. On the Hoopa

Valley Indian Reservation in California, female fishers appear to overlap with members of the same sex to a greater extent than most studies report. I tested the hypothesis that female fishers tolerate some degree of spatial overlap, but that co- occurrence in the area of overlap is uncommon. I also tested the hypothesis that this overlap is mediated by habitat selection and female philopatry. I found that female fishers share a large proportion of their home ranges and overlap at a high frequency. However, dynamic interactions revealed fishers avoided each other in both space and time. Habitat used in overlap areas differed from what was available to them in non-overlapping areas but lacked a consistent pattern of ii

selection among habitat types. I used genetic samples to assess population pairwise relatedness and determined that adult female fishers were more closely related and adult male fishers less related, to each other, respectively.

Furthermore, these data showed evidence of male-biased dispersal in this population suggesting female philopatry. Information from this study is important for management of current fisher populations in order to better understand spatial patterns on the landscape needed to guide management for both source and translocation programs used for conservation of this species.

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ACKNOWLEDGEMENTS

This research was funded by the National Science Foundation Graduate Research

Fellowship (Grant Number: DGE-1049702) and in-kind contributions from Hoopa Valley

Tribal Forestry. I thank the Hoopa Tribal Council and Hoopa Valley Tribal Forestry staff for their permission and encouragement throughout this study.

I would like to sincerely thank my advisor, Dr. Micaela Szykman Gunther, for her invaluable support, guidance, and encouragement throughout my tenure as an undergraduate and graduate student at Humboldt State University. I would also like to thank my committee members J. Mark Higley, Dr. Matthew Johnson, and Dr. William

Zielinski for their expertise and suggestions on this manuscript. I extend my gratitude to

J. Mark Higley and Dr. Sean Matthews for co-founding the Hoopa Fisher Project and for their incessant faith in me to carry out this project from start to finish. I especially appreciate Dr. Sean Matthews for mentoring me and being a constant source of inspiration for me throughout many years of fisher fieldwork. I would also like to thank

Dr. Tim Bean and Angela Darnell for valuable statistical advice on the spatial-temporal component. I am indebted to Kristin Brzeski for her expertise on genetics and unwavering optimism and patience. This study would not have been possible without help from numerous field technicians in trapping efforts, especially Shannon Mendia.

Last but not least, I would like to sincerely acknowledge Caylen Cummins for all of her hard work and dedication in helping to collect telemetry and den site locations.

TABLE OF CONTENTS

ABSTRACT ...... ii

ACKNOWLEDGEMENTS ...... iv

LIST OF TABLES ...... vii

LIST OF FIGURES ...... viii

LIST OF APPENDICES ...... ix

INTRODUCTION ...... 1

METHODS ...... 9

Study Area ...... 9

Capture and Handling ...... 12

Radio telemetry ...... 13

Home Range and Overlap Estimation ...... 14

Spatial and Temporal Interaction Analysis ...... 16

Habitat Analysis ...... 20

Genetic Analysis ...... 22

RESULTS ...... 25

Home Range and Overlap ...... 25

Spatial and Temporal Interactions ...... 28

Habitat Analysis ...... 28

Genetic Analysis ...... 31

DISCUSSION ...... 38

LITERATURE CITED ...... 46

PERSONAL COMMUNICATIONS ...... 55

APPENDICES…………………………………………………………………………56

LIST OF TABLES

Table 1. Description of notation used in text for calculating spatial and temporal interactions between female fishers with overlapping home ranges on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014 ...... 18

Table 2. Mean area (km2) and percent overlap (± SE, range below) for home range (95% fixed kernel) and core use (50% isopleth) areas for pairwise and total combined overlap in annual, denning, and post-denning seasons on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014 ...... 27

Table 4. The number of alleles, observed heterozygosity (HO), expected heterozygosity (HE), tests for conformance to Hardy-Weinberg equilibrium, allele size ranges, and allele frequency ranges of 9 microsatellite loci among 127 male and female residential fishers on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014...... 33

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LIST OF FIGURES

Figure 1. Study area boundary and trap locations on the Hoopa Valley Indian Reservation for the study of female fisher home range overlap and spatial-temporal interactions in northwestern California, USA, March 2013 - March 2014 ...... 11

Figure 2. Mean within population pairwise values (R) between 46 male and 81 female resident fishers on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014. Upper and lower whiskers denote 95% confidence limits determined by 999 bootstrap resampling...... 34

Figure 3. Mean relatedness values (Queller and Goodnight’s R values from GenAlEx) within distance classes for (a) female and (b) male resident fishers on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014. Upper and lower whiskers denote 95% confidence intervals determined by 999 bootstrap resampling...... 35

Figure 4. Comparison of 46 male and 81 female AIc frequency distributions suggesting male sex-biased dispersal in fisher on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014 ...... 36

Figure 5. Mean AIc values for 46 male and 81 female resident fishers on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014. Upper and lower whiskers denote standard error bars…………………………………...... 37

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LIST OF APPENDICES

Appendix A: Fifteen Stand Structural Stages modified into five habitat classes on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014……..56

Appendix B: Annual 95% fixed kernel home ranges for 22 resident female fishers with a total of 38 pairwise combinations on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014..………….……………………………………………57

Appendix C: Spatial distribution representing (a) pairwise overlap between animal A and animal B and (b) total combined home range overlap of animal A on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014……………………..58

INTRODUCTION

Occupation and defense of space by individuals can have powerful influences on spatial and demographic structure in carnivore populations. Spatial patterns within animal populations are influenced by ecological factors as well as dispersal characteristics of males and females (Crook 1970; Crook et al. 1976) and have important effects on population demography, social structure and dynamics, and genetic composition

(Cheepko-Sade and Halpin 1987; Stenseth and Lidicker 1992). Knowledge of intraspecific spatial and temporal interactions is necessary to understand social behavior among individuals and important for successful management of the species.

Spacing patterns of animals are the result of strategies used by individual animals to maximize reproductive success and survival (Sandell 1989). Animals compete for resources, such as food, shelter, and mates (Maher and Lott 1995). One way to compete is to exclude potential competitors from the area containing the resources (i.e., being territorial; Brown and Orians 1970; Davies 1978). A territory is a particular region within an animal’s home range in which the animal has exclusive, or priority use (Powell 2012) and which may include all, or part, of the animal’s entire home range.

Animals maintain territories only when there is a limited resource, that is, a critical resource that is in short supply and limits reproduction or survival and, thus, population growth (Brown 1969). The ultimate regulator of a population of territorial animals is the limiting resource that stimulates the territorial behavior (Powell 2000). The most common limiting resource is food and, for territorial individuals, territory size tends

2 to vary inversely with food availability (Rogers 1987; Hixon 1980; Sandell 1989;

Palomares and Delibes 1994; Powers and McKee 1994; Powell et al. 1997). Territorial behavior can be predicted from variation in the availability and productivity of food

(Powell et al. 1997). Individuals may abandon territorial behavior if the limiting resource increases or is not sufficiently abundant. Such flexibility exists because a territory must be economically defensible (Brown 1969; Gorman et al. 2006) or economical to maintain

(Spencer 2012). If a limited resource is uniformly distributed and abundant, or if a resource is patchily distributed and sparsely available, the cost of defending that resource may be greater than the benefit in defending that resource. In these cases, animals should tolerate extensive home range overlap, even among members of the same sex (Powell

1993). Therefore, territorial behavior is economical primarily at intermediate levels of productivity of the limiting resource (Carpenter and MacMillen 1976, Powell 1994).

Animals establish territories usually during adolescence but sometimes into early adulthood. If a territory is occupied, young animals may need to disperse to suitable, unoccupied habitat. Natal dispersal, defined as the permanent movement an individual makes from its birth site to a place where it potentially reproduces, occurs in most birds and mammals and tends to be sex-biased, whereby one sex disperses and the other remains mostly philopatric to its natal site (Pusey1987; Perrin and Mazalov 2000;

Lebigre et al. 2008). The differential propensity of males and females to disperse is a life history trait closely related to the mating system of a species (Greenwood 1980) and is influenced by the relative costs and benefits of dispersal to each sex (Pusey 1987).

Greenwood (1980) suggests that male-biased dispersal in mammals is associated with polygyny and male defense of mates. In polygyny, parental involvement by males is minimal and their reproductive success is limited by the number of females they encounter and with whom they successfully breed (Trivers 1972). Dispersal by juvenile males reduces competition with male relatives for mating opportunities (Dobson 1982) and increases the probability of mating with unrelated females (Waser and Jones 1983), which avoids inbreeding. In addition, dispersal may expose males to a larger number of available female mating partners (Trivers 1972). In contrast, resources are more important to females, who bear the cost of raising the offspring (Greenwood 1980;

Johnson 1986), and in which reproductive success is proportional to the amount of energy available to rear young (Trivers 1972; Sandell 1989). As a result, philopatric females would benefit most from familiarity with a territory, thereby securing necessary resources to meet the demands of reproduction by maintaining close associations with female kin and remaining near natal areas, while reducing the high risks associated with dispersal.

A consequence of natal philopatry for mammals, especially among long-lived species, is that several generations of females may live close together (Waser and Jones

1983; Johnson 1986). A system of overlapping home ranges among females suggests that portions of maternal home ranges are being shared with relatives, such as offspring, and may suggest philopatric behavior (Waser and Jones 1983). Overlap may also be influenced by high prey densities, which can diminish territoriality. Long-lived territorial carnivores offer opportunities to evaluate costs and benefits of philopatry which may be facilitated by the abundance and distribution of food (Marino et al. 2012).

Individuals that have overlapping home ranges are responding to both resources and to each other, which makes measures of overlap important (Powell 2012). Areas of overlap are not used uniformly and vary in probability of use. For example, two individuals may share a large area of overlap used rarely by each, or a small but critical area used intensively. Therefore, overall area of overlap can be a poor measure of the intensity or significance of overlap. As a result, it is important to study both the static and dynamic interactions between a pair of overlapping individuals, where static interaction is the spatial overlap between two home ranges and dynamic interactions are the combined interdependent spatial and temporal movements of the two individuals whose home ranges overlap (Doncaster 1990).

Carnivore social organization can vary and be quite complex; spatial patterns range widely from overlapping and loosely defended home ranges, to exclusive, heavily defended territories occupied by individuals, pairs, or cooperatively defended packs, or groups of related individuals. Most studies have been conducted on stable social groups, such as wolves (Canis lupus) or lions (Panthera leo), which have received more attention than solitary carnivores (Macdonald and Moehlman 1982). However, a majority of carnivores are solitary, having very little contact with conspecifics (Sandell 1989).

Solitary carnivores are elusive and difficult to study but are an important part of understanding how ecological and evolutionary processes shape the spatial structure and level of sociality in carnivore populations.

The fisher (Pekania pennanti) is a mid-sized, forest dwelling solitary carnivore in the family Mustelidae. Fishers interact with conspecifics usually only during territorial

5 defense, mating, and while mothers raise their young (Powell 1993). Fishers are sexually dimorphic, polygynous, and generally occur at low densities (Fuller et al. 2001, Weir and

Corbould 2006, Thompson 2008). Like most mustelids (Kelly 1977, Powell 1979, Arthur et al. 1989), fishers exhibit intrasexual territoriality, in which home ranges tend not to overlap between individuals of the same sex, but tend to overlap extensively between the sexes (Powell 1993).

The historic geographic distribution of Pacific fishers included the Cascade Range of Washington, Oregon, and California, and the Sierra Nevada Range in California

(Gibilisco 1994, Powell and Zielinski 1994). Over-trapping for fur in the early 20th century and habitat fragmentation and loss primarily due to logging practices have resulted in range contractions and population declines throughout the Pacific states. In

April 2004, the U.S. Fish and Wildlife Service (USFWS) determined that the west coast population of fisher warranted protection under the Endangered Species Act (U.S. Fish and Wildlife Service 2004). In October 2014, the USFWS proposed to list the west coast distinct population of fisher as threatened under the ESA (U.S. Fish and Wildlife Service

2014). A better understanding regarding social behavior in fishers can help managers identify expected spatial movements on the landscape in order to help assist in range expansion for this species.

Fisher density and home range sizes vary widely across their range (Powell 1993,

Yaeger 2005, Lofroth et al. 2010, Matthews et al. 2011). On the Hoopa Valley Indian

Reservation (HVIR) in northwestern California, fisher density is relatively higher and home ranges smaller than in most regions within the Pacific states (Lofroth et al. 2010).

In addition, female fishers appear to overlap home ranges to a greater extent than other studies report (J. Mark Higley, pers. comm.); however, it is unknown if this overlap occurs in both space and time. Previous studies have demonstrated that the HVIR is the regional center of predicted habitat in northwestern California (Carroll et al. 1999, Davis et al. 2007) and the highest densities of fisher in the Pacific states (Matthews et al. 2013).

The high density of fishers on the HVIR provides an opportunity to study fisher ecology in a unique environment where their territorial behavior might be different than in other areas, perhaps due to differences in habitat composition.

Fisher home ranges and areas of overlap have been estimated using mark- recapture and radio-telemetry methods (Arthur et al. 1989, Fuller et al. 2001, Matthews et al. 2013). In species where home ranges overlap, relatives may be more likely to use shared areas simultaneously (Powell 2012) allowing for greater tolerance of overlap in territorial species. Assessing relatedness between individuals whose home ranges overlap can help to interpret the extent of overlap and degree of territoriality. Contemporary developments in genetic techniques offer field researchers powerful methods to evaluate relatedness and dispersal patterns from genetic data in wild populations of animals

(Aubry et al. 2004, Biek et al. 2006, Costello et al. 2008). Sex-biased dispersal has the potential to leave a strong genetic signature on the genetic structure of a population.

Genetic studies have been successfully conducted on population structure in fishers (Kyle et al. 2001, Drew et al. 2003, Wisely et al. 2004, Jordan et al. 2007, Tucker et al. 2012).

Combining radio-telemetry and genetic techniques can provide new insight into the population ecology of fishers in the Pacific region by improving our knowledge about

7 relatedness, sex-biased dispersal patterns, and how this can influence home range overlap and territoriality.

Few studies have focused on fisher spatial patterns (Aubry et al. 2004, Weir et al.

2013) and no studies have been conducted on fisher at small spatial and temporal scales

(i.e., interactions within home range overlap areas based on simultaneous locations).

Currently, translocation programs are being implemented as a management strategy to encourage fisher populations to re-colonize historical ranges from which they were extirpated. A better understanding of fisher spacing patterns, which may vary in different areas within their range, can help managers make more informed decisions regarding future translocation and advise assisted dispersal programs for both source and destination populations.

Due to the observed phenomenon of high female fisher overlap, this study focused on female fisher space utilization in home range overlap areas on the HVIR.

Using radio telemetry to examine patterns in space use, I aimed to document: 1) frequency, area, and size of home range overlap, 2) percent (pairwise and total combined) and volume-of-intersection of home range overlap, 3) overlap occurrence in both space and time, 4) habitat selection within the overlap areas as compared to non-overlapping areas within an individual’s home range, and 5) pairwise genetic relatedness between sexes and across geographic distances. I hypothesized that female fishers on the HVIR tolerate significant overlap with other females due to the high densities, and that this overlap is mediated by habitat selection and female philopatry. To investigate philopatric behavior, I aimed to determine pairwise genetic relatedness between same-sex fishers as

8 a function of geographic distances. I predicted that overlapping female fishers: 1) overlap at a high frequency on the HVIR, 2) avoid each other in both space and time, 3) overlap in habitat types that differ in proportion to availability within their individual home ranges, which may indicate higher prey densities in areas of overlap, and 4) are more closely related to each other than are neighboring male fishers to each other and to non- overlapping females located at greater distances. By providing insight into female fisher spatial patterns not commonly found in other parts of their range, this study can help inform future management into the flexibility of female fisher spacing behavior.

METHODS

Study Area

The study was conducted within the 366 km2 HVIR in Humboldt County in northwestern California (Figure 1). Steep slopes and rugged terrain dominate the topography with elevations ranging from between 98 and 1,170 m. Mean daily minimum and maximum temperatures range from 6 to 22°C, respectively. Mean annual precipitation is 138 cm (National Climate Data Center, March 2015).

The area is comprised of mostly forested land, bisected by the Trinity River into east and west regions. The study area included the Hostler Creek drainage in the east region and Pine Creek, Beaver Creek, Soctish Creek, and Rock Creek drainages in the west region. The study area has been used for previous fisher research where females are readily captured. The study area is composed of montane hardwood-conifer communities

(Mayer and Laudenslayer 1988) dominated by Douglas fir (Pseudotsuga menziesii), tan oak (Notholithocarpus densiflorus), Pacific madrone (Arbutus menziesii), California black oak (Quercus kelloggii), and canyon live oak (Quercus chrysolepis).

Past and current timber harvests have created a mixture of mature old-growth and early seral forests (Matthews et al. 2013). Prior to 1990, timber harvest averaged over

500 ha cut/year using clear cutting methods. Between 1994 and 2010, under the Hoopa

Tribe’s Forestry Management Plan, timber harvest averaged 150 ha annually, pre- commercial thinning of approximately 165 ha, early released of 100-175 ha, and cultural

10 burning of 6-40 hectares. Harvest is implemented using regeneration methods with green tree and snag retention in smaller stands (Higley and Matthews 2009) to maximize wildlife habitat.

Figure 1.Study area boundary and trap locations on the Hoopa Valley Indian Reservation for the study of female fisher home range overlap and spatial- temporal interactions in northwestern California, USA, March 2013 - March 2014.

Capture and Handling

Fishers were captured between January and March 2013 in Tomahawk live traps

(Model 207, Tomahawk Live Trap Company, Tomahawk, Wisconsin, USA) baited with raw chicken and modified with a wooden nest-box to provide security and shelter from elements (Wilbert 1992; Seglund 1995). Traps were placed strategically on the landscape

(Figure 1) to target adult (≥ 2 years old) and subadult (1-2 years old) resident female fishers within the study area. Juvenile (< 1 year old) fishers were processed but not included in this study. Captured fishers were immobilized with an intramuscular injection of ketamine (4.4 mg/kg) and Midazolam (2.2 mg/kg) administered by a hand- syringe.

Weight was estimated by experienced personnel prior to injection. After immobilization, fishers were marked with sterile passive integrated transponders (PIT tags, 134.2 kHz

Super Tag, Sterile, Biomark, Inc., Boise, Idaho, USA) implanted subcutaneously behind the right shoulder. An ear tissue (biopsy) sample was taken from each fisher for genetic sampling. A first upper premolar was taken from each fisher and aged by cementum annuli (Matson’s Laboratory, Milltown, Montana).

Adult and subadult female fishers were radio-collared with mortality-sensitive transmitters (Holohil model MI-2M, Holohil Systems Ltd., Carp, Ontario, Canada).

Subadult fishers that had potential to outgrow their collar or disperse out of the study area were fitted with a short piece of leather spliced to the collar to create a break-away mechanism to prevent injury or to minimize the possibility that fishers would wear collars unnecessarily in the event they could not be recaptured at the end of the study. All

13 fishers were released from their sites of capture after full recovery from anesthesia.

Capture and handling procedures were approved by the Humboldt State Animal Care and

Use Committee (protocol No. 12.13.W.42.A).

Radio telemetry

All female fishers were relocated using radio telemetry between 9 March 2013 and 9 March 2014. I used ground-based radio telemetry to obtain locations using standard techniques for triangulation (White and Garrott 1990; Samuel and Fuller 1996, Kenward

2001). Triangulations consisted of ≥ 3 bearings, a minimum of 20° apart, taken from roads or trails. Since animal movement can greatly reduce telemetry accuracy (Schmutz and White 1990), triangulations were conducted within a 20-minute period. I attempted to gather independent locations for each fisher twice a week, a minimum of 24 h apart, to ensure independence and to provide a sufficient sample size for home range estimation (>

50 locations). During the biweekly efforts, I aimed to obtain simultaneous locations between each pair of potentially overlapping females in order to assess if temporal overlap was occurring. Fisher location estimates were generated from triangulations in the program Location of a Signal (LOAS) 4.0 software (Ecological Software Solutions,

LLC, Florida, USA) using a maximum likelihood estimator, and plotted using ArcGIS

(ArcGIS version 10.1, Environmental Systems Research Institute, Redlands, California,

USA). Telemetry error was estimated by triangulating on female fishers at known locations. For this purpose, I initially used suspected den sites that were later confirmed.

Their locations were recorded as Universal Transverse Coordinates (UTM) using a

14 handheld global positioning system (GPS) with 6 m accuracy. Telemetry error was calculated as the average linear error between the estimate and the actual den location

(White and Garrott 1990).

Home Range and Overlap Estimation

Location points used to generate home range estimates included female radio telemetry locations, den and rest sites, dropped collar locations, trap sites, and incidental observations. For each fisher, annual and seasonal home range and core use areas were estimated using a bivariate normal fixed-kernel estimator in Geospatial Modeling

Environment (GME, v 0.7.2.1) with smoothing parameters calculated using a diagonal plug-in in R statistical software (v 2.15.2). Core use areas were considered areas of concentrated use within the home range in which 50 percent of locations were found.

Seasonal home range and core areas were separated into two seasons: denning (March -

August) and post-denning (September – February) in order to determine differences in home range and core area use between seasons. Fixed kernels account for differential use of area within a home range and require a large number of locations to estimate areas of core use (Seaman et al. 1999). To generate kernel home ranges a curve representing the probability density function (kernel) was placed over each location, illustrating the local area an animal might have been using when the location was obtained (Worton 1989,

Seaman and Powell 1996). A square grid was placed over all kernels and at each grid intersection the density of all overlapping kernels was averaged, resulting in a utilization distribution (UD; Seaman and Powell 1996). The UD is a 3-dimensional representation of

15 a home range that describes the relative amount of time an animal spends within its home range (Seaman and Powell 1996). The UD can then be used to generate a 2-dimensional home range estimate illustrated by isopleths (contours) surrounding the area where a specified percentage of an animal’s activity is concentrated (percent volume contours).

From the kernel density layer, I used GME to obtain 95% isopleth polygons for annual and seasonal home ranges and 50% isopleths for core use areas. I used a paired t-test to test for differences between home range size and female fisher age class since age may affect home range size. Age classes were divided into three categories: 1-3, 4-6, and 7-10 years of age. I used a paired t-test to test for differences between home range and core use area size between the two seasons (denning and post-denning).

When the home range and core areas overlapped, I used estimates of the home range to delineate the 2-dimensional and 3-dimensional regions of spatial overlap (km2) between each pair of overlapping individuals. For the 2-dimensional analysis, I determined the mean percentage of pairwise home range overlap as: mean overlap = area overlap area overlap 0.5 ( x ) (Minta 1992), where A and B represent the two different area of A area of B individuals in a pair of overlapping female fishers. Total combined home range percent overlap was calculated for each individual by dividing the total area of overlap by the total home range area and multiplying by 100. Frequency of overlap for an individual female was calculated as the total number of individual females overlapping her home range. Two fishers were removed from the pairwise and total combined overlap seasonal analyses due to a mortality event and a dropped collar. I removed five additional fishers

16 located on the periphery of the study area from the annual and seasonal total combined home range overlap analyses to minimize the confounding effects of underestimating overlap, as these peripheral females overlapped with only one other fisher and more than likely overlapped with other fishers that were not monitored during the study period. I used a paired t-test to test for differences between seasons in both pairwise and total combined overlap in home range and core use areas.

Two-dimensional overlap analysis is problematic because it does not take into account the individual’s use of space within an area of overlap and can overestimate results (Fieberg and Kochanny 2005); therefore, I also calculated 3-dimensional overlap

(volume of intersection) to provide a measure of intensity of use. I used the kernel density raster layers to obtain a volume of intersection:

∞ volume of intersection = min [UD̂ , (x,y),UD̂ , (x,y)] dxdy ∬−∞ A B where UDA and UDB are the utilization distributions (kernel density layers) for each of the individuals in the pair, respectively (Kernohan et al. 2001, Fieberg and Kochanny

2005). This resulted in an index ranging from no overlap (0) to complete overlap (1) and provided a single measure of overlap that incorporated unequal use of home range. From this method, I calculated home range overlap for each individual female as the mean volume of intersection for all females and for all occurrences of pairwise overlap within that individual’s home range.

Spatial and Temporal Interaction Analysis

I used methods proposed by Minta (1992) to test for attraction or avoidance to an area of overlap and to determine temporal interactions between pairs of fishers that shared a common area (home range overlap as previously estimated). I tested the null hypothesis that for each pair of fishers sharing an area of overlap, female fisher A and female fisher B (Table 1), each fisher would use the area spatially and temporally independently of the other (random movement) (Minta 1992, Mace and Waller 1997).

Simultaneous locations for each pair of fishers that overlapped were placed into one of the following categories of observed frequencies: 1) both individuals were simultaneously present in the overlapped area (n11); 2) only individual A was present in the overlapped area (n21); 3) only individual B was present in the overlapped area (n12);

4) both individuals were simultaneously absent from the overlapped area (n22).

Observations were considered ‘simultaneous’ if they were within 6 hours of each other, however the majority of observations occurred within a 15-minute period.

Expected frequencies (Table 1) were calculated as the proportion of overlapped area in relation to total home range area. Observed and expected frequencies of presence and absence in overlap areas were then summed across pairs of individual females. Observed frequencies were compared to expected frequencies to develop an overall chi-square

2 (χtot). If fisher A and fisher B were using the overlapping area independently, then

2 2 2 2 2 (n11−p11n) (n12−p12n) (n21−p21n) (n22−p22n) χtot = + + + p11n p12n p21n p22n where n is the total number of simultaneous locations for each pair. If fisher A and fisher

2 B were using the overlapping area independently of one another, then χtot would be small

Observed frequencies n11 Both A and B simultaneously present in overlap area n12 Only B present in overlap area n21 Only A present in overlap area n22 Both A and B simultaneously absent in overlap area Sum of observed simultaneous sightings of A and B, with A present in the n+1 overlap area (n11 + n21) Sum of observed simultaneous sightings of A and B, with A absent from the n+2 overlap area (n12 + n22) Sum of observed simultaneous sightings of A and B, with B present in the n1+ overlap area (n11 + n12) Sum of observed simultaneous sightings of A and B, with B absent from the n2+ overlap area (n21 + n22)

Expected frequencies area AB area AB Both A and B simultaneously present in overlap area x p11 area A area B area AB area AB Only B present in overlap area (1 − ) x p12 area A area B area AB area AB Only A present in overlap area x (1 − ) p21 area A area B area AB area AB Both A and B simultaneously absent in overlap area (1 − ) x (1 − ) p22 area A area B Sum of expected simultaneous sightings of A and B, with A present in the p+1 overlap area (p11 + p21) Sum of expected simultaneous sightings of A and B, with A absent from the p+2 overlap area (p12 + p22) Sum of expected simultaneous sightings of A and B, with B present in the p1+ overlap area (p11 + p12) Sum of expected simultaneous sightings of A and B, with B absent from the p2+ overlap area (p21 + p22)

2 with three degrees of freedom (df). I partitioned χtot into three parts: spatial coefficient of fisher A, spatial coefficient of fisher B, and temporal interaction (ixn) coefficient. The outcome was the combination of a spatial coefficient (L퐴:Ā or 혓퐵:퐵̅ ) or temporal

2 interaction coefficient (혓푖푥푛) and a 휒 probability value. The three partitions were defined as:

Spatial coefficient of fisher A; df = 1:

2 p+2n+1 2 (n+1+ p+1n) L퐴:Ā = ln , with 휒 = p+1n+2 퐴:Ā p+2p+1n

Spatial coefficient of fisher B; df = 1:

2 p2+n1+ 2 (n1++ p1+n) 혓퐵:퐵̅ = ln , with 휒 ̅ = p1+n2+ 퐵:퐵 p2+p1+n

Temporal interaction coefficient; df = 1:

n n n n n n 11+ 22 ( 11 + 22 + 12 + 21) 2 p11 p22 2 p11 p22 p12 p21 혓푖푥푛 = ln n12 n21 , with 휒 = 1 1 1 1 + 푖푥푛 n( + + + ) p12 p21 p11 p12 p21 p22

The spatial coefficients (L퐴:Ā and 혓퐵:퐵̅ ) for each individual in a pair of fishers, fisher A and fisher B, indicated spatial association within the overlapping area. These results translated into random use, spatial attraction, or spatial avoidance of the shared overlapping area. As the spatial coefficient approached zero, use of the shared area became random. When the spatial coefficient was greater than zero, use of the shared area was greater than expected (spatial attraction). When the spatial coefficient was less than zero, use of the shared area was less than expected (spatial avoidance). As the temporal interaction coefficient (혓푖푥푛) approached zero, simultaneous use and nonuse of the overlapped area were equal (random use). When 혓푖푥푛 was greater than zero, then

20 simultaneity was greater than solitariness (temporal attraction). When 혓푖푥푛 was less than zero, simultaneity was less than solitariness (temporal avoidance). Following Mace and

Waller (1997), spatial and temporal interaction results were further classified as symmetrical (same spatial or temporal response by the pair), asymmetrical (opposite spatial or temporal response by the pair), or singular response (only one individual showed a response).

If 혓푖푥푛 was significant, I referred to the odds ratio, or departure from expectation, of each of the four categories of observed frequencies (n11, n12, n21,n22) and expected frequencies (p11, p12, p21, p22) to interpret the magnitude of the significant interactions.

Because there is only one way to be simultaneously present and three ways to be solitary

(i.e., one or the other or both animals), the odds ratio allows for a more informative interpretation of the significant temporal interactions (Minta 1992).

Habitat Analysis

I determined habitat composition areas of overlapping areas corresponding to

Johnson’s (1980) third-order of selection which examines a fisher’s selection of habitat within its home range. I determined the habitat types using a vegetation classification and habitat typing of the HVIR’s 2012 Forest Management Plan (FMP, Higley 2012). The

FMP classified vegetation coverage by creating 1.2 ha (minimal) polygons using overlays of aerial images at a scale of 1:12,000 to 1:1,500. Unique habitat types were further stratified based on 15 “Stand Structural Stages” (SSS), modified but following Oliver and

Larson (1996). Most of the SSS strata were focused on seral stage development

21 characteristics and were developed to provide a classification of meaningful habitat classes for a number of species (Higley 2012), including the fisher. In order to meet assumptions of minimum sample sizes, I grouped the 15 SSS types into five habitat classes: MOF (old forest), YCCF (young forest), SX (stem exclusion), MCOV (sapling brushy pole), and OPEN (general open habitats and true oak woodland) (Appendix A).

I used a chi-square goodness-of-fit test (Neu et al. 1974) to examine habitat selection by comparing the number of times the different SSS types were used to the expected number of times they would be used if each fisher used habitats in proportion to their availability. For each fisher, I compared the number of observations in the home range overlap area (use) to the number of observations in the non-overlap area

(availability) for each of the five SSS, testing the hypothesis that females are using a habitat type in the overlapping areas that differ from the non-overlapping areas. When the chi-square test was significant, I determined whether a specific habitat type was avoided, selected, or used in proportion to its availability by calculating the confidence interval around the proportional use value and checked for overlap with the proportional availability value using the following formula (Neu et al. 1974):

1 1 훼 푝̂ (1−푝̂ ) 훼 푝̂ (1−푝̂ ) 푝̂ − 푧 [ 푖 푖 ]2 ≤ 푝 ≤ 푝̂ + 푧 [ 푖 푖 ]2 푖 2 푛 푖 푖 2 푛 where n is the number of used observations, 푝̂푖 is the proportional use of a habitat i, 푝푖 is the proportional availability of a habitat i, and z is the standard normal variate for a given

훼 α obtained from a z table (α = 0.05, z = 1.96). If the proportional availability value was 2 above the upper limit of the use confidence interval, then that habitat type in the overlap

22 area was used less than expected (avoided). If the proportional availability value was below the used lower confidence interval, then that habitat type was used more than expected (selected) in the overlap area. If the proportional availability value was within the confidence interval, then that habitat type was used in proportion to its availability.

Ten female fishers were removed from this analysis due to uncertainty in non- overlap areas not being shared with other females or due to limitations in sampling design, inadequate number of observations in a particular habitat type (SSS), or home range completely overlapped with other female fishers.

Genetic Analysis

Ear tissue samples were collected from male and female fishers between 23

September 2006 and 13 February 2014 and stored in 95% ethanol in individual microcentrifuge tubes at room temperature. Samples were submitted to the Rocky

Mountain Research Station Wildlife Genetics Lab in Missoula, MT for DNA extraction and molecular analysis. Whole genomic DNA was extracted from tissue samples using the QIAGEN Dneasy Tissue Kit (Qiagen, Valencia, CA, USA). Fisher tissue samples were analyzed at 10 microsatellite loci. MP0059, MP0144, MP0175, MP0197, MP0200, and MP0247 were developed from tissue samples from California (Jordan et al. 2007).

Loci MER022 (Fleming et al. 1999), GGU101, GGU16 (Duffy et al. 1998), and LUT733

(Dallas and Piertney 1998) were developed in other mustelid species [ermine (Mustela erminea), wolverine (Gulo gulo), and Eurasian otter (Lutra lutra), respectively]. The reaction volume 10 microliters (µL) contained 1.0 µL DNA, 1x reaction buffer (Applied

Biosystems, Life Technologies, Grand Island, New York, USA), 2.0 mili-molar (mM)

MgCl2, 200 micro-molar (µM) of each dNTP, 1 µM reverse primer, 1 µM dye-labeled forward primer, 1.5 mg/ml BSA, and 1 U Taq polymerase (Applied Biosystems). The

PCR profile was 94°C/5 min, [(94°C/1 min, 55°C/1 min, 72°C/30s) x 40 cycles]. The resultant products were visualized on a LI-COR DNA analyzer (LI-COR Biotechnology,

Lincoln, Nebraska, USA). All data were error checked using the program Dropout

(McKelvey and Schwartz 2005).

Pairwise relatedness (R) estimate values (Queller and Goodnight 1989) for males and females were calculated using GenAlEx, Genetic Analysis in Excel (v. 6.5, Peakall and Smouse 2006, 2012). Relatedness coefficients range from -1 to 1, where negative values indicate no relation and positive values indicate relatedness. A total of 999 permutations were completed to create mean R values and 95% confidence intervals.

Pairwise linear geographic distances between home range centroids of pairs of individuals were also calculated using GenAlEx in order to investigate dispersal patterns and relatedness as a function of distance. I assigned geographic distance between pairs of individuals into one of six distance classes; 2.5 km, 5.0 km, 7.5 km, 10 km, 12.5 km, and

15 km. Distance classes were determined based on the maximum distance between home range centroids of overlapping female pairs from this study (2.5 km) and subsequent distance classes were further categorized into 2.5 km increments. Geographic location data for male and female fishers not included in this study were used to augment the sample. Data from these individuals were collected from previous work conducted on the

HVIR between 23 September 2006 and 13 February 2014 and these data were obtained

24 either from radio-tracking (home range centroids) or trap history data. The genetic relationships from the historical data should be sufficient to hold beyond the scope of this study due to the long-lived nature of this species.

I used an assignment-based sex-biased dispersal test procedure developed by

Favre et al. (1997) and expanded by Mossman and Waser (2001) to detect genetic signatures of sex-biased dispersal in GenAlEx. For each individual, the expected genotype frequency at each locus was calculated and multiplied across multiple loci and log transformed to give a log-likelihood value. For each individual, an Assignment Index correction (AIc) was calculated from the log likelihood assignment test value as:

AIc = individual log likelihood – mean log likelihood of population

For each individual, a positive AIc value indicates that the individual is more correctly assigned to the population, while a negative AIc value indicates the opposite. The mean

AIc was calculated for males and females and tested for significance using a non- parametric Mann-Whitney U test. The genetic signal of sex-biased dispersal is indicated when there is a significant difference in frequency of distribution and mean AIc values among male and females. The least dispersing sex will have a positive mean AIc value while the most dispersing sex will have a negative mean AIc value. The conversion of negative log-likelihoods by the multiplication of -1 is not applied, hence the dispersing sex will exhibit more negative values.

RESULTS

I collected a total of 2,276 locations for 22 female fishers and obtained an average of 104 ± 3.4 (x̅ ± SE) radio telemetry locations per female (range = 64-124). Locations included 2,016 from triangulations, 177 from den sites (47 individual dens), 16 rest sites, three dropped collar sites, 58 trap sites, five visual observations, and one mortality site.

Telemetry error was 92 ± 12 m. Average age for collared female fishers was 5.1 years and ranged from 1-10 years old.

Home Range and Overlap

Overall mean 95% fixed kernel annual home range size for females was 4.94 ±

0.45 km2 (range = 2.49-10.57) and 50% core area was 1.31 ± 0.13 km2 (range = 0.53-

2.56). The mean 95% fixed kernel denning season home range size was 4.22 ± 0.34 km2

(range = 1.97-6.57) and post-denning was 5.56 ± 0.56 km2 (range = 2.99-12.39). The mean 50% core area during denning season was 1.06 ± 0.09 km2 (range = 0.42-1.85) and post-denning was 1.50 ± 0.16 km2 (range = 0.73-2.93). Home range size and core use area were significantly different between denning season and post-denning seasons (t19 =

-3.43, p = 0.003 and t19 = -3.03, p = 0.007, respectively). There was no significant difference between size of home range and core use area and age class (F = 0.12, p = 0.89 and F = 0.36, p = 0.704, respectively).

There were a total of 38 possible pairwise combinations of neighboring female fishers in which all 22 females overlapped with a minimum of one other fisher (Appendix

B). Frequency of overlap was 4.4 ± 0.36 overlapping fishers/female (range = 1-6). Post- denning season home range overlap and core use areas were significantly larger than home range overlap and core use areas during denning season (t29 = -3.36, p = 0.002, t29

= -2.62, p = 0.014, Table 2). Total home range overlap combined for all females averaged more than half of each female’s annual home range (64.5%, Table 2, Appendix C). Total combined home range overlap areas were significantly larger during post-denning season than denning season (t14 = -3.36, p = 0.005) as were core use areas (t14 = -3.03, p =

0.012, Table 2). In contrast to the high degree of overlap observed, the mean volume of intersection (3-dimensional overlap) from all occurrences of pairwise overlap was 0.09 ±

0.01 (range = 0.03-0.14) and 0.09 ± 0.01 (range = 0.02-0.34) for females overall.

Table 2. Mean area (km2) and percent overlap (± SE, range below) for home range (95% fixed kernel) and core use (50% isopleth) areas for pairwise and total combined overlap (per female) in annual, denning, and post-denning seasons on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014.

Annual Denning season Post-denning season Percent Percent Percent Area ± SE Area ± SE Area ± SE overlap n overlap n overlap n (range) (range) (range) (range) (range) (range) Pairwise overlap 38 30 30

Home range 0.95 ± 0.12 21.7 ± 1.89 0.75 ± 0.12 19.5 ± 0.03 1.11 ± 0.14 23.1 ± 0.03

0.13-3.25 3.7-78.7 0.05-2.45 0.96-61.4 0-3.15 0-71.2

Core use 0.41 ± 0.07 20.3 ± 2.95 0.33 ± 0.12 18.2 ± 0.07 0.50 ± 0.14 21.7 ± 0.09

0-1.81 0-98.9 0-1.66 0-50.8 0-1.56 0-84.9

Total combined overlap 17 15 15

Home range 2.76 ± 0.28 64.5 ± 4.83 1.96 ± 0.25 53.2 ± 6.51 2.91 ± 0.29 62.7 ± 4.91

0.96-5.65 31.3-100 0.41-3.64 1.5-95.0 1.10-5.02 25.6-100

Core use 0.70 ± 0.08 64.3 ± 5.27 0.61 ± 0.12 51.3 ± 7.45 0.93 ± 0.09 65.2 ± 5.07

0-0.96 0-99.4 0-1.75 3.7-79.1 0-1.69 25.8-100

Spatial and Temporal Interactions

Of the 22 female fishers, 38 pairs overlapped resulting in a pairwise average of

148 (range = 76-206) simultaneous locations. This resulted in a cumulative total of 5,631 locations, as a female could be simultaneous with more than one female. All 38 pairs of overlapping female fishers exhibited significant spatial and/or temporal interactions within the overlap areas. For the spatial main effect, all pairs had at least one individual that avoided the shared overlap area of which 22 (58%) pairs exhibited significant and symmetrical spatial avoidance (p<0.05). Temporal use of the home range overlap area was also significantly different from random (p<0.05) for 19 (50%) of the 38 pairs of fishers. Of the 19 pairs of fishers, 18 (95%) exhibited simultaneous temporal avoidance of the shared area, three (16%) exhibited simultaneous temporal attraction, and two

(11%) exhibited both simultaneous temporal avoidance and attraction with attraction being the stronger effect. One pair (5%) was significant for solitary use (animal B) of the overlap area greater than expected and also significant for simultaneous temporal absence, with the simultaneous temporal absence being the stronger effect. Overall simultaneous temporal absence was 1.4 to 3.6 times more frequent than expected from random use.

Habitat Analysis

I evaluated habitat selection for 12 female fishers which included 736 used

(overlap) locations and 609 available (non-overlap) locations. Overall habitat

29 composition differed from non-overlap areas for all 12 fishers, though the habitat types

2 that were selected and avoided varied among fishers (휒4 = 12.8-128.7, p ≤ 0.01). Female fishers in this study did not use habitat types in proportion to their availability. Some fishers selected for one or two of the five SSS habitat types, while other fishers avoided these (Table 3). There was no clear pattern of a particular habitat type being strongly selected or avoided in overlap areas.

Table 3. Five Stand Structural Stage habitat types selected for (+++) or against (---) in areas of overlap (use) compared to non-overlap areas (available) in 12 female fishers on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014. Fisher ID MOFa YCCFb SXc MCOVd OPENe 10054 --- +++ 10079 --- +++ 10100 +++ --- 10106 +++ --- 10125 --- +++ 10137 +++ --- +++ 10155 --- 10192 +++ ------10193 --- +++ 10213 --- 10233 ------10245 +++ a single or multi-storied old forest b young forest multi-storied, understory stem exclusion, hardwood residual c stem exclusion, dense canopy cover, single-storied stands d understory re-initiation, stand initiation, sapling brushy pole e seedling and young pole stands, non-forested areas, and true oak woodland stands

Genetic Analysis

A total of 198 fisher tissue samples (86M, 112F) were analyzed and 197 (85M,

112F) were successfully genotyped at 10 microsatellite loci. All 10 microsatellite loci were polymorphic and the number of alleles ranged from 2-5 (Table 4). Locus MP0200 was monomorphic with the exception of a single individual from the sample population

(allele size/frequency was 155/0.003 and 159/0.997, for single male and remaining population, respectively). Therefore, MP0200 was removed from the analysis as it was likely a null allele that failed to amplify. In order to detect philopatric behavior, 70 of the

197 genotypes were removed from further analysis based on the likelihood that those fishers were juveniles and had potential to disperse, leaving 127 genotypes (46M, 81F) used in the analysis. Following Hedrick (2000), I used a chi-square test in GenAlEx to test all 9 loci for Hardy-Weinberg equilibrium assumptions. All 9 loci met these assumptions; observed heterozygosity (HO) ranged 0.386-0.732 and expected heterozygosity (HE) ranged 0.385-0.659 (Table 4).

Overall mean population pairwise relatedness was -0.008, indicating lack of significant relatedness among all animals. Mean pairwise relatedness within the population was greater among females than males (Figure 2). Female fishers located more closely geographically were more related to each other than to females farther away

(Figure 3a). Female fisher mean pairwise relatedness (R) was highest in the 2.5 km distance class and relatedness (R) decreased as distance apart increased (Figure 3a). Male fishers were more closely related to each other in the 2.5 km distance class but the 95%

32 confidence interval overlapped with zero. Male fishers were less closely related in all remaining distance classes, however, 95% confidence intervals overlapped zero except for the 12.5 and 15.0 km distance classes (Figure 3b).

Frequency distributions of corrected assignment indices (AIc) for males and females indicated that males were the more likely the dispersing sex (Figure 4). There was a significant difference between the mean AIc values for males and females (Z = -

0.052, p = 0.020, Figure 5). Female fishers had positive mean AIc values that did not overlap with zero suggesting they are the least dispersing sex. Male fishers had negative mean AIc values that did not overlap with zero and suggests they are more likely to be immigrants and thus the most likely dispersing sex.

Table 4. The number of alleles, observed heterozygosity (HO), expected heterozygosity (HE), tests for conformance to Hardy-Weinberg equilibrium, allele size ranges, and allele frequency ranges of 9 microsatellite loci among 127 male and female resident fishers on the Hoopa Valley Indian Reservation in California, USA, September 2006 - February 2014. No. Allele size Locus H H P Frequency range Alleles O E (bp) range MP0059 2 0.402 0.422 0.577 153-155 0.303-0.697

MP0144 5 0.601 0.659 0.912 182-202 0.004-0.417

MP0175 5 0.732 0.634 0.157 172-192 0.028-0.272

MP0197 3 0.480 0.505 0.604 209-217 0.059-0.626

MP0247 4 0.598 0.625 0.707 121-141 0.071-0.461

GGU101 2 0.425 0.493 0.121 144-150 0.441-0.559

GGU216 2 0.409 0.392 0.618 177-179 0.268-0.732

LUT733 2 0.535 0.497 0.390 134-138 0.465-0.535

MER022 3 0.386 0.385 0.848 249-263 0.055-0.760

Mean 3.22 0.514 0.518 0.548

0.060 0.040 0.020 0.000 R -0.020 -0.040 -0.060 -0.080 -0.100 Male Female

35 a) Female fishers

0.2

0.1

R 0.0

-0.1

-0.2 2.5 5.0 7.5 10.0 12.5 15.0 Distance classes (km)

b) Male fishers

0.2

0.1

R 0.0

-0.1

-0.2 2.5 5.0 7.5 10.0 12.5 15.0 Distance classes (km)

20 15 10 5 Frequency Female 0 -5 Male -10 -15 -2.28 -1.89 -1.50 -1.11 -0.71 -0.32 0.07 0.46 0.86 1.25 1.64 Alc

0.300 0.200 0.100 0.000 Mean AIc -0.100 -0.200 -0.300 -0.400 -0.500 Male Female

DISCUSSION

Overall, female fishers overlapped at a high frequency and shared a large proportion of their home ranges to a greater extent than other studies report (Weir et al.

2013), consistent with my predictions. Despite the high degree of overlap, female fishers avoided each other in both space and time. These results supported my prediction that due to their strong territorial behavior, female fishers would exhibit temporal avoidance, and therefore not be present in overlapping areas at the same time as neighbors. Although

I expected female fishers overlap areas to differ in habitat from what was available in the non-overlapping areas, I found no consistent pattern for habitat type selection or avoidance. Finally, female fishers were more closely related to each other than males were to each other, and female fishers were more closely related to other females with home ranges closer to them than females residing further away, as predicted. These results provide evidence of philopatric behavior among females and male-biased dispersal.

Female fisher home ranges on the HVIR were smaller (4.96 km2) compared to the average (18.8 km2) across western North America (Lofroth et al. 2010). This makes the high degree of overlap even more compelling. The small home ranges of female fishers on the HVIR suggest that differences in habitat composition, and perhaps a high degree of relatedness, may allow for smaller home ranges and high degree of spatial overlap.

Although female fishers showed some degree of exclusivity within their home ranges

(non-overlapping areas), space use did not strictly adhere to the overall expected pattern

39 of intrasexual territoriality characteristic of this species and suggests flexibility in the extent of territoriality.

Despite the large number of studies evaluating the statistical properties of home range estimates and utilization distribution’s under various conditions (Worton 1989;

Boulanger and White 1990; Seaman et al. 1999), there are surprisingly few studies addressing simultaneous use of space by carnivores (Siedel 1992, Mace and Waller 1997,

Persson et al. 2010). Static interaction analyses calculate an overall measure of association during a specified time period, without considering whether animals were in close proximity at any particular point in time (Kernohan et al. 2001). Areas of overlap vary in their probability of use (Powell 2012); therefore, static interactions should be studied in conjunction with dynamic interactions. Based on static interactions alone, excluding the volume-of-intersection component, results would indicate that female fishers tolerate extensive home range overlap and do not exhibit intrasexual territoriality.

However, after I examined dynamic spatial-temporal interactions, it was apparent that female fishers avoided overlapped areas in both space and time. This suggests a social dynamic that simultaneously encourages spatial attraction, perhaps to an important resource, and temporal avoidance, to minimize contact with neighbors. The results from the 3-D volume-of-intersection analysis contradict the 2-D analyses and demonstrate the importance of concurrent study on both static and dynamic interactions. Not many similar spatial-temporal interaction analyses have been conducted on other territorial carnivores and to the best of my knowledge, this is the first study conducted on fishers.

The confounding results for the two pairs that exhibited both simultaneous temporal attraction and avoidance were likely caused by small sample sizes. The odds ratios revealed that the pairs used the overlap area simultaneously 5.5 and 4 times more than expected (respectively) compared to 1.5 and 2 times more than expected

(respectively) for simultaneous avoidance. Although the odds of simultaneous attraction were greater than the odds of simultaneous avoidance in both cases, there is no inferential test for this comparison. Although Minta (1992) recognized the limitations of assessing interactions between only 2 individuals at a time, the overall strong pattern of symmetrical avoidance (i.e., same spatial and temporal response in both members of a pair) within the areas of overlap increased confidence in the observed simultaneous interactions.

The compelling pattern of simultaneous temporal avoidance of shared overlap areas suggests that female fishers are not strongly defending the areas of overlap as territories. Territory defense is difficult to document and apparent lack of home-range overlap is often used as evidence of territoriality (Powell 2012). A territory is an area within an animal’s home range which is defended against conspecifics and may be all, or part, of an animal’s entire home range (Powell 2012). Perhaps female fishers on the

HVIR are defending territories in other areas within their home ranges (e.g., non- overlapping areas). Modern radio telemetry equipment, such as Global Positioning

Systems (GPS) and proximity sensor collars, can help researchers to better understand spatial and temporal interactions in wildlife populations. Utilizing these technologies in conjunction with emerging tools used to examine space and time, such as Time Local

Convex Hull (T LoCoHo) or Elliptical Time-Density Models, can perhaps provide insight into social behavior (Lyons et al. 2013, Wall et al. 2014).

A possible explanation for the high degree of overlap could be that food resources are mediating this behavior. As predicted by Powell (1994), it would be advantageous for an individual to defend a limited resource that is widely distributed in patches throughout its entire home range. However, in areas where food is abundant, territoriality over the food resource would diminish, and considerable overlap within home ranges would be tolerated. Small home ranges and relatively high densities of fisher on the HVIR suggest that there are high prey densities and perhaps female fishers do not need to exclude competitors due to high abundance and distribution of prey items. Although fishers have been described as generalist predators (Powell 1981; Zielinski et al. 1991; Bowman et al.

2006), studies on fisher diet composition on the HVIR and Klamath Bioregion (Golightly et al. 2006; Slauson et al. 2011) have found dusky-footed woodrats (Neotoma fuscipes) to be the most frequently consumed prey species. The dusky-footed woodrat and other rodent species are probably more available in Hoopa and the surrounding vicinity than in other regions. Availability of these prey resources may result from a combination of greater understory complexity and aridity (Carraway and Verts 1991), fairly uniform abundance and distribution of acorns from oak species in the mixed-coniferous forests, and relatively modest timber harvest management methods (particularly on the HVIR).

While the most common limiting resource for territorial animals is food, Wolff

(1989, 1993) warned that the true limiting resource may not be apparent. For some species, the limiting resource is a nest or den site (Walters et al. 1992, Doncaster and

Woodroffe 1993). Reproductive female fishers are obligate cavity users; they use cavities in large-diameter trees and snags exclusively for birthing and rearing kits (Raley et al.

2012). On the HVIR, some female fishers reuse the same reproductive den structures in subsequent years (personal observation). On several occasions, radio-tracked female fishers established home ranges on formerly occupied female home ranges that had died, and subsequently used some of the same den structures used by the former residents. The reuse patterns of den structures among breeding females strongly suggest that these preferred den structures may be a limited resource across the landscape. Perhaps exclusive territories are being maintained in areas within their home ranges that contain these den structures. Although this study did not examine this, it would be valuable to further study den site selection in relation to territoriality in fishers.

Territories can be defended without explicit aggression (e.g., scent marking) to communicate territory ownership. Fishers have plantar and anal scent glands that are used for sexual signaling and marking territory (Powell 1993). Distinct boundaries of territories may be just as difficult for researchers to identify as boundaries of undefended home ranges (Powell 2012). Territory boundaries may contain a buffer a few hundred meters wide littered with scent marks. In Hoopa, shortly after a female fisher mortality event, a neighboring female fisher will often shift or move her home range fairly quickly, either partially or entirely, onto the deceased fisher’s home range (personal observation).

These movements suggest that fishers are alerted to the opening of a territory by detecting a lack of fresh scent marking following the mortality of an individual. This may explain to some extent the high degree of overlap among females.

The lack of a pattern in selection of habitat types in the areas of overlap may have been a result of several factors. Habitat categories were combined in order to meet minimum sample sizes needed to ensure at least one expected observation occurred within each habitat category and no more than 20% of all categories contained less than five expected observations. Small sample sizes [e.g., few observations in either overlap

(use) or non-overlap (available) areas] might explain the lack of pattern in selection for or against the five SSS habitat types within the areas of overlap. Nine of the 12 fishers had

20% of all habitat categories consist of less than five expected observations. In addition, combining the 15 preliminary SSS categories down to five categories in order to meet minimal assumption requirements may have masked the ability to detect any patterns of finer resolution habitat type selection or avoidance. Furthermore, habitat types used in this study were not distinctly correlated to prey densities or rest/den site structures, and perhaps the resources within the habitat types are influencing the overlap behavior.

Finally, telemetry error may have contributed to the lack of pattern. If a fisher location was best estimated to be within a certain habitat type, but was actually in another due to triangulation error, this may further hinder the ability to detect habitat selection or avoidance within the overlap areas.

Female fishers were more closely related to each other than male fishers were to each other, which suggests limited dispersal among females compared to male fishers. In addition, female fishers with home ranges closer to one another (within 2.5 km) were more closely related to each other than females distanced further away. The results from the 2.5 km distance class were of particular interest, as this is the distance class in which

44 all female fishers that overlapped in this study occurred. The observation that females that overlap home ranges are more closely related than females who do not overlap, and that males are essentially unrelated across distance classes, in combination lends support for the theory that fishers exhibit female philopatry and male-biased dispersal (Aubry et al. 2004; Aubry and Raley 2006; Williams et al. 2000). Between 2005 and 2008, a study conducted on the HVIR marked 51 kits in their dens. Of those 51 marked kits (27 female,

24 male), 12 female and 2 males were recaptured as juveniles in subsequent seasons within or near their maternal home ranges (Matthews et al. 2013). The higher recapture rate of females suggest that females are more likely to be philopatric and low recapture rate of males suggest that male juvenile fishers either have high mortality and/or disperse prior to being recaptured.

Relatedness estimators have very large variances owing to stochastic differences among loci and to the chance of sharing alleles that are identical by decent (Blouin 2003).

Computer simulations indicate that large numbers of loci are required for reasonable relatedness estimates (e.g., ≥ 15 microsatellite loci). Although microsatellite marker variability in this study was modest (mean expected heterozygosity (HE) = 0.52) and number of loci used was low (n = 9), it appears that relatively useful and biologically informative estimates of average pairwise relatedness can be achieved. However, I recommend that more loci be included in future studies to investigate the influence of relatedness on fisher behavior.

This study supports that female fishers on the HVIR are not strictly territorial in terms of exclusion of all other females at the home range scale of resource selection

(Johnson 1980). Although this study did not reveal differences in habitat selection between overlapping and non-overlapping areas, the extensive overlap observed suggests food and other resources are widely distributed, indefensible, and not limiting. However, competition at finer scales of selection or differences in microhabitat selection may still occur. More studies are needed to quantify habitat quality (e.g., prey densities and den site availability) in relation to finer spatial and temporal scales.

Analyses of fisher spatial patterns on the landscape in relation to relatedness helps fill a gap in our understanding of fisher biology by identifying processes that affect the potential of fisher populations to expand beyond the edges of existing populations.

Spatial organization and relatedness of individuals in a population are important parameters of population structure that can advise species management and conservation.

The results from this study will help inform future fisher reintroduction and translocation efforts by better predicting spacing patterns of adult home ranges that might be expected following translocation. Male fishers may disperse into adjacent areas along the margins of fisher populations only to return in search of philopatric females to mate with or remain in areas without mates. The slow expansion of west coast fisher populations may benefit from augmentation using assisted dispersal to promote expansion into historic ranges.

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PERSONAL COMMUNICATIONS

Higley, J. M. 2012. Hoopa Tribal Forestry, Wildlife Department. One Loop Road, Hoopa, CA 95546, USA.

Appendix A: Fifteen Stand Structural Stages modified into five habitat classes on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014.

15 Stand Structural Stages Habitat Classes Old forest multi-storied MOF Old forest single-storied MOF Young forest multi-storied YCCF Understory re-initiation/stem exclusion YCCF Stem exclusion/hardwood residual YCCF Stem exclusion SX Understory re-initiation/sapling brushy pole MCOV Stand initiation/sapling brushy pole MCOV True oak woodland OPEN Young forest multi-storied/urban OPEN Understory re-initiation/seedling OPEN Stand initiation/young pole OPEN Stand initiation/seedling ÓPEN Non-forested/chaparral-brush OPEN Non-forested/open OPEN

Appendix B. Annual 95% fixed kernel home ranges for 22 resident female fishers with a total of 38 pairwise combinations on the Hoopa Valley Indian Reservation in California, USA, March 2013 - March 2014.

a) b)