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A comparative ecological study between coyotes (Canis latrans) in a protected and urban habitat

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Title:
A comparative ecological study between coyotes (Canis latrans) in a protected and urban habitat a closer look at enteric parasites and diet between Florida coyotes
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Book
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English
Creator:
Manning, Denara Lynn
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University of South Florida
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Tampa, Fla
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Subjects / Keywords:
Hymenolepis spp
Ascaris spp
Balantidium coli
Blastocystis spp
Entamoeba histolytica
Dissertations, Academic -- Environmental Science & Policy -- Masters -- USF   ( lcsh )
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bibliography   ( marcgt )
theses   ( marcgt )
non-fiction   ( marcgt )

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Abstract:
ABSTRACT: Coyotes (Canis latrans) have inhabited Florida (USA) since the 1960s and are currently found throughout the state. The purpose of the present study was to obtain information on enteric parasites and diet of Florida coyotes from two different habitat types. Seasonal variation in diet was also examined. Fresh coyote fecal samples were collected from protected and urban habitats in Pinellas County, Florida (USA; 27°54' n, 82°41'w) from may 2005 to march 2007. A standard fecal flotation examination and formalin-ethyl acetate sedimentation were utilized on fecal samples collected from the protected (n=40) and urban (n=50) habitats. Five novel (newly documented) parasites of coyotes were discovered; one cestode (Hymenolepis spp.), one nematode (Ascaris spp.), and three protozoa (Balantidium coli, Blastocystis spp., and Entamoeba histolytica).^ Novel parasites of Florida coyotes were also discovered two cestodes (diphyllobothrium latum and dipylidium caninum), two nematodes (toxocara canis and uncinaria stenocephala), one trematode (paragonimus spp.), and four protozoa (cryptosporidium spp., giardia canis, isospora spp., and sarcocystis cruzi). One cestode (Taenia spp.), three nematodes (Ancylostoma caninum, Physaloptera spp., and Trichurus vulpis), and one trematode (Alaria spp.) were also recovered, all of which have previously been documented in Florida coyotes. Diet items were identified to the lowest possible taxonomic level by gross morphological characteristics and medullary configurations of dorsal guard hairs. A poisson regression was utilized to determine the relation between diet items and habitat, season, and interaction.^ In the protected habitat (n=49), vegetative matter (96%), Insecta (53%), and Rodentia (45%) were recovered most often, as opposed to berries (56%) and Lagomorpha (32%) in the urban habitats (n=71). Overall, vegetative matter, berries, and Lagomorpha were recovered most often from Florida coyote fecal samples. Odocoileus virginianus, Lagomorpha, and berries varied the most between wet and dry seasons. It is suggested that Florida coyotes are more susceptible to reinfection by novel parasites because of their rapid range expansion and lack of acquired immunity. Rapid habitat loss in Florida (i.e., urbanization) lowers survival of adult coyotes, increases the probability of transmission of disease between wild and domestic canids, and alters the diet of coyotes by lowering biological diversity of available prey items.
Thesis:
Thesis (M.S.)--University of South Florida, 2007.
Bibliography:
Includes bibliographical references.
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Mode of access: World Wide Web.
Statement of Responsibility:
by Denara Lynn Manning.
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Title from PDF of title page.
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Document formatted into pages; contains 80 pages.

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University of South Florida
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aleph - 001930011
oclc - 213273814
usfldc doi - E14-SFE0002194
usfldc handle - e14.2194
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SFS0026512:00001


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ABSTRACT: Coyotes (Canis latrans) have inhabited Florida (USA) since the 1960s and are currently found throughout the state. The purpose of the present study was to obtain information on enteric parasites and diet of Florida coyotes from two different habitat types. Seasonal variation in diet was also examined. Fresh coyote fecal samples were collected from protected and urban habitats in Pinellas County, Florida (USA; 27¨54' n, 82¨41'w) from may 2005 to march 2007. A standard fecal flotation examination and formalin-ethyl acetate sedimentation were utilized on fecal samples collected from the protected (n=40) and urban (n=50) habitats. Five novel (newly documented) parasites of coyotes were discovered; one cestode (Hymenolepis spp.), one nematode (Ascaris spp.), and three protozoa (Balantidium coli, Blastocystis spp., and Entamoeba histolytica).^ Novel parasites of Florida coyotes were also discovered two cestodes (diphyllobothrium latum and dipylidium caninum), two nematodes (toxocara canis and uncinaria stenocephala), one trematode (paragonimus spp.), and four protozoa (cryptosporidium spp., giardia canis, isospora spp., and sarcocystis cruzi). One cestode (Taenia spp.), three nematodes (Ancylostoma caninum, Physaloptera spp., and Trichurus vulpis), and one trematode (Alaria spp.) were also recovered, all of which have previously been documented in Florida coyotes. Diet items were identified to the lowest possible taxonomic level by gross morphological characteristics and medullary configurations of dorsal guard hairs. A poisson regression was utilized to determine the relation between diet items and habitat, season, and interaction.^ In the protected habitat (n=49), vegetative matter (96%), Insecta (53%), and Rodentia (45%) were recovered most often, as opposed to berries (56%) and Lagomorpha (32%) in the urban habitats (n=71). Overall, vegetative matter, berries, and Lagomorpha were recovered most often from Florida coyote fecal samples. Odocoileus virginianus, Lagomorpha, and berries varied the most between wet and dry seasons. It is suggested that Florida coyotes are more susceptible to reinfection by novel parasites because of their rapid range expansion and lack of acquired immunity. Rapid habitat loss in Florida (i.e., urbanization) lowers survival of adult coyotes, increases the probability of transmission of disease between wild and domestic canids, and alters the diet of coyotes by lowering biological diversity of available prey items.
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Ascaris spp.
Balantidium coli.
Blastocystis spp.
Entamoeba histolytica.
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PAGE 1

A Comparative Ecological Study between Coyotes ( Canis latrans ) in a Protected and Urban Habitat: A Closer Look at Enteric Parasites and Diet between Florida Coyotes by Denara Lynn Manning A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science Department of Environmental Science and Policy College of Arts and Sciences University of South Florida Co-Major Professor: Melissa Grigione, Ph.D. Co-Major Professor: Robert Brinkman, Ph.D. Ricardo Izurieta, Ph.D. Steven Harper, Ph.D. Date of Approval: October 18, 2007 Keywords: Hymenolepis spp., Ascaris spp., Balantidium coli, Blastocystis spp., Entamoeba histolytica Copyright 2007 Denara Lynn Manning

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ACKNOWLEDGMENTS Most importantly I would like to thank the Lord Jesus Chri st for giving His life for me, for always being faithful to His promises, and for giving me guidance when I don’t know where to go or what to do. I would also lik e to thank my husband, Justin Wofford, for his continued love, support, and encouragement in everything I do. I want to thank my mom, dad, sisters (Chelsea, Daubri, Dalesha), and fam ily from the bottom of my heart. I cannot express how thankful and blessed I am to ha ve such a wonderful family who supports me every step of the way. Throughout this research my committee (Melissa Grigione, Ricardo Izurieta, and Steve Harper) has cont inued to support and guide me, for which I am most grateful. I would also like to th ank Sarah Clavio for all of her help. Our friendship is one that will trul y last a lifetime, solidified by all of those countless hours in the field. This research would not have been as intensive if it we re not for all of the volunteers who dedicated themselves and thei r time to this cause. I cannot thank Jay Jones enough for all of her help in the fi eld collecting samples and for her overall excitement, dedication, and enthusiasm. I also wish to thank Catherine Hughes and Michelle Dachsteiner for their continued a ssistance in the lab. I express utmost appreciation to Bruce Rinker a nd all of the staff at Brooker Creek Preserve for providing access into the preserve, for the use of an ATV, and for always being an integral part of this study. I am also grateful for the a ssistance of Pinellas County Animal Control (Kenny Mitchell, Welch Agnew, and Richard Stahl) with the urban study. Lastly, I sincerely thank Pinellas Count y for providing the grant whic h funded this research.

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i TABLE OF CONTENTS List of Tables................................................................................................................. .iii List of Figures................................................................................................................ .iv Abstract....................................................................................................................... ......v CHAPTER ONE: A COMPARATIVE STUDY BETWEEN ENTERIC PARASITES OF COYOTES IN A PROTECTED AND URBAN HABITAT.........1 Introduction...........................................................................................................2 Study Sites............................................................................................................6 Protected Habitat.......................................................................................6 Urban Habitat............................................................................................6 Methods.................................................................................................................8 Field Methods...........................................................................................8 Laboratory Methods..................................................................................8 Statistical Methods....................................................................................9 Results.................................................................................................................10 Discussion...........................................................................................................20 Enteric Parasites of Florida C oyotes in a Protected Habitat...................20 Enteric Parasites of Florida Coyotes in Urban Habitat...........................22 Novel Parasites of Canis latrans .............................................................24 Novel Parasites of Florida Coyotes.........................................................26 Conclusion..............................................................................................28 Implications.............................................................................................29 Literature Cited...................................................................................................31 CHAPTER TWO: A COMPARATIV E STUDY BETWEEN THE DIET OF COYOTES IN A PROTECTED AND URBAN HABITAT..............................40 Introduction.........................................................................................................41 Study Sites..........................................................................................................45 Protected Habitat.....................................................................................45 Urban Habitat..........................................................................................45 Methods...............................................................................................................47 Field Methods.........................................................................................47 Laboratory Methods................................................................................47

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ii Statistical Methods..................................................................................48 Results.................................................................................................................50 Discussion...........................................................................................................63 Diet of Coyotes in Diffe rent Habitat Types............................................63 Diet of Florida Coyotes...........................................................................68 Conclusion..............................................................................................71 Literature Cited...................................................................................................74

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iii LIST OF TABLES CHAPTER ONE Table 1. Parasites recovered from fecal samples of coyotes in the protected habitat (n=40)..........................................................................................13 Table 2. Parasites recovered from fecal samples of coyotes in urban habitats (n=50)......................................................................................................14 Table 3. Novel parasite species of C. latrans found during the course of this study and the habitat they were found in................................................15 Table 4. Novel parasite species of Florida co yotes found during the course of this study and the habitat they were found in.....................................16 Table 5. Previously documented parasite spec ies of Florida coyotes found during the course of this study and the habitat they were found in........17 CHAPTER TWO Table 1. Diet items consumed by c oyotes in protected habitat (n=49).................53 Table 2. Diet items consumed by c oyotes in urban habitats (n=71).....................54 Table 3. Diet items consumed by Fl orida coyotes (prote cted and urban habitats combined)..................................................................................55

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iv LIST OF FIGURES CHAPTER ONE Figure 1. Distribution of th e number of enteric parasite species identified per coyote fecal sample collected.................................................................18 Figure 2. Novel parasites of Canis latrans .............................................................19 CHAPTER TWO Figure 1. Comparison of diet items c onsumed between coyotes in protected and urban habitats...................................................................................56 Figure 2. Seasonal variation in the percentage of coyote fecal samples collected in a protected habitat that contain each diet item....................57 Figure 3. Seasonal variation in the percentage of coyote fecal samples collected in a protected habitat that contain each diet item (represented by the lowest possible taxonomic level)............................58 Figure 4. Seasonal variation in the percentage of coyote fecal samples collected in urban habitats th at contain each diet item...........................59 Figure 5. Seasonal variation in the percentage of coyote fecal samples collected in urban habitats th at contain each diet item (represented by the lowest possible taxonomic level)............................60 Figure 6. Seasonal variation in the percentage of coyote fecal samples collected from Florida coyotes that contain each diet item (represented by the lowest possible taxonomic level)............................61 Figure 7. Florida coyote diet (prote cted and urban habitats combined) (n=120)....................................................................................................62

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v A COMPARATIVE ECOLOGICAL ST UDY BETWEEN COYOTES (CANIS LATRANS) IN A PROTECTED AND URBAN HABITAT: A CLOSER LOOK AT ENTERIC PARASITES AND DIET BETWEEN FLORIDA COYOTES Denara Lynn Manning ABSTRACT Coyotes ( Canis latrans ) have inhabited Florida (USA) since the 1960s and are currently found throughout the state. The purpose of the present study was to obtain information on enteric parasites and diet of Florida coyotes from two different habitat types. Seasonal variation in diet was also examined. Fres h coyote fecal samples were collected from protected and urban habitats in Pinellas County, Florida (USA; 27o54’ N, 82o41’W) from May 2005 to March 2007. A standard fecal flotation examination and formalin-ethyl acetate sedimentation were utilized on fecal sa mples collected from the protected (n=40) and urban (n=50) habitats. Five novel (new ly documented) parasi tes of coyotes were discovered; one cestode ( Hymenolepis spp.), one nematode ( Ascaris spp.), and three protozoa ( Balantidium coli Blastocystis spp., and Entamoeba histolytica ). Novel parasites of Florida coyotes were also discovered; two cestodes ( Diphyllobothrium latum and Dipylidium caninum ), two nematodes ( Toxocara canis and Uncinaria stenocephala ), one trematode ( Paragonimus spp.), and four protozoa ( Cryptosporidium spp., Giardia canis Isospora spp., and Sarcocystis cruzi ). One cestode ( Taenia spp.), three nematodes ( Ancylostoma caninum Physaloptera spp., and Trichurus vulpis ), and one trematode

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vi ( Alaria spp.) were also recovered, all of which have previously been documented in Florida coyotes. Diet items were identif ied to the lowest possible taxonomic level by gross morphological characteristics and medullary configurations of dorsal guard hairs. A Poisson Regression was utilized to dete rmine the relation between diet items and habitat, season, and interaction. In the protected habitat (n =49), vegetative matter (96%), Insecta (53%), and Rodentia (45%) were r ecovered most often, as opposed to berries (56%) and Lagomorpha (32%) in the urban habita ts (n=71). Overall, vegetative matter, berries, and Lagomorpha were recovered most often from Florida coyote fecal samples. Odocoileus virginianus Lagomorpha, and berries varied the most between wet and dry seasons. It is suggested that Florida coyotes are more susceptible to reinfection by novel parasites because of their ra pid range expansion and lack of acquired immunity. Rapid habitat loss in Florida (i.e., urbanization) lowe rs survival of adult coyotes, increases the probability of transmission of disease between wild and domestic canids, and alters the diet of coyotes by lowering biological di versity of available prey items.

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1 CHAPTER ONE: A COMPARATIVE STUDY BETWEEN ENTERIC PARASITES OF COYOTES IN A PROTECTED AND URBAN HABITAT

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2 INTRODUCTION Coyotes ( Canis latrans ) have inhabited Florida (U SA) since the 1960s and are currently found throughout th e state (Wooding and Hardisky, 1990; Maehr et al., 1996; Main et al., 1999; Main et al., 2000). To date, there has been minimal research conducted on these new Florida residents to determine what ecological effects they have on the communities they inhabit. Coyotes have b een widely studied throughout North America, but due to vast differences of flora and fauna it is unclear how well these studies apply to Florida coyotes (Seesee et al ., 1983; Thurber and Peterson, 1991) Habitat loss in Florida (i.e., urbanization) lowers survival of adu lt coyotes and increases the probability of transmission of disease between wild and domestic canids (Grinder and Krausman, 2001). Furthermore, the overall health of individuals declines when heavy parasitic infections occur (Belden and Kiesecker, 2005). Documentation of enteric (intestinal) para sites of Florida coyo tes is important for a number of reasons. First, the health of the coyote populat ion is directly affected by intestinal parasitic infecti on (Lindsay et al., 1997). Second, knowledge of parasite species which infect coyotes is essential in or der to determine if any measures need to be taken to prevent transmission between coyotes and domestic animals (Arjo et al., 2003). Third, domestic animals and coyotes can act as reservoirs for infections to humans. When infected wild animals, such as coyotes, have increased interaction with areas frequented by humans, as is the case with dense human popul ations and drastic

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3 urbanization, the risk of infection to humans is increased. Humans can become infected with enteric parasites if th ey consume viable parasites or drink contaminated water (Rubel and Wisnivesky, 2005). Children, due to their close contact with the soil, are especially susceptible to consumption of parasites (Matsuo and Kamiya, 2005). Although studies have been conducted on th e intestinal para sites of coyotes elsewhere in the United States (Arther a nd Post, 1977; Conder and Loveless, 1978; Pence and Meinzer, 1979; Seesee et al., 1983; Arjo et al., 2003; Gomppe r et al., 2003), few studies have focused on Florida coyotes. C onti (1984) and Foster et al. (2003) conducted research on enteric parasites of Florid a coyotes, but both studies were based on necropsies from coyotes in less densely populated counties. This present study is the first conducted on Florida coyotes using non-inva sive fecal examination techniques. Specifically, we utilized standard fecal flota tion and formalin-ethyl acetate sedimentation to compare enteric parasite species of Flor ida coyotes between tw o different habitats, protected and urban, in the most densely populated county in Florida (USCB, 2004). The primary objectives of this study were to investigate differences in enteric parasites between coyotes from protected a nd urban habitats and to document any novel (i.e., newly documented) parasi tes. Specifically, we compar ed species richness, defined as the number of species, and composition of enteric parasites betw een study locations. Additionally, we documented any novel ente ric parasites recove red, whether novel to Canis latrans (i.e., never been documented in coyotes before) or to Florida coyotes (i.e., never been documented in Florida coyotes before).

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4 Species composition of enteric parasites: Null hypothesis : Enteric parasites of coyotes in the protected and urban habitats will not differ significantly. Alternate hypothesis : Enteric parasites of coyotes in the protected habitat will differ significantly from those parasites of coyotes in the urban habitat. It is presumed that there will be several enteric parasites found in both habitats. However, we predict that there will be si gnificant differences in parasite species composition between protected and urban habita ts. Specifically, we predict that urban coyote samples will contain more parasites commonly found in domestic animals. This prediction is based on the increased probabi lity that coyotes and domestic dogs utilize common areas in the urban environment and beca use of this coyotes in the urban habitat might have more enteric parasites that ar e normally documented in domestic dogs. This could result from an infected domestic dog def ecating in an area, thus depositing viable parasite eggs in the urban environment. If an urban coyote were to come into contact with these viable eggs and consume them, the coyote could then become infected with parasites. Novel enteric parasites: Null hypothesis : All enteric parasites in Fl orida coyotes will have been previously documented. Alternate hypothesis : Florida coyotes will be in fected with novel enteric parasites. Due to unique flora, fauna, and environmen tal conditions in Florida, it is also presumed that novel (i.e., newly documented) enteric parasites will be documented.

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5 Specifically, we presume that novel enteric parasites of Canis latrans which have not previously been documented in coyotes elsewh ere in the United States may be discovered during the course of this study. Additionall y, little research has been conducted on coyote populations in Florida. As such, little is known about what enteric parasites infect these animals. Therefore, we also predict th at this research may discover novel parasites of Florida coyotes.

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6 STUDY SITES Pinellas County is the most densely popul ated county in Florida with over 1281 people per square kilometer (USCB, 2004). Two types of habitat were compared in Pinellas County during this st udy: protected and urban. Protected Habitat Brooker Creek Preserve (BCP; 27o54’ N, 82o41’W) was used for the protected habitat (Bean et al., 2005). BCP is an 8500 acr e wilderness area that is actively managed for natural resource protection. Located in the northeast corner of the county, the boundaries of BCP are shared with densely populated residential areas. The study site consists of extensive pine flatwoods and fr eshwater swamps. Fauna include white-tailed deer ( Odocoileus virginianus ), raccoons ( Procyon lotor ), armadillo ( Dasypus novemcinctus ), bobcat ( Lynx rufus ), otter ( Lutra canadensis ), wild turkey ( Meleagris gallopavo ), red-shouldered hawks ( Buteo lineatus ), wood storks ( Mycteria americana ), bobwhite quail ( Colinus virginianus ), and gopher tortoise ( Gopherus polyphemus ). Urban Habitat Different sites throughout Pinellas Count y were used for urban habitat. Sites were determined using GIS (ArcGIS v.8) to plot existing geospatial information including land-use categories, railroads, bi ke trails, and power lines throughout the

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7 county. Subsequently, a map revealing ru ral, sub-urban, and urban areas based on residential (land-use category) population density was genera ted. Sites were identified based on criteria expected to support urban coyot es. The sites were constrained such that they had land cover of urban sites similar to that of BCP, were traversed by power lines, bike trails, or inactive railroads, and were located in urban areas.

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8 METHODS Field Methods Fresh coyote feces were collected over th e course of two years (May 2005 thru March 2007) from trails, power lines, and bike trails in the protected (n=40) and urban (n=50) habitats. Paths were traversed on f oot, bicycle, and by ATV three times a week during the course of this study. Each fecal sample was measured (length and diameter) and photographed in the field. Species origin of the fecal samples was determined by adjacent sign (tracks) and dimensions of feces. Upon confirmation of coyote scat, the sample was assigned a unique identification code and its longitude and latitude were recorded by use of a GPS unit. Finally, the sample was placed in its own paper bag. To avoid collecting bobcat feces, only those samples in excess of one inch in diameter were collected (Gompper et al., 2003). To avoi d collecting domestic dog feces, only samples which contained hair and bone fragments and/or which were accompanied by coyote tracks were collected (W ooding et al., 1984). Laboratory Methods To determine the enteric parasites present in each sample, a standard fecal flotation (specific gravit y = 1.25) examination was conducted within 12 hours of collection (Thornton et al., 1974). Two grams of the fresh sa mple was preserved in 10% formalin and stored at room temperature unti l analyzed (Gillespie et al., 2005). To allow

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9 for maximum recovery of ova, oocysts, and larv ae, a formalin ethyl-acetate sedimentation technique was also utilized on each sample (P rice, 1994). Parasites were identified based on measurements obtained by an ocular microm eter fitted to a compound microscope and review of morphological characteristics observed from photographs taken of each specimen (Zaman, 1984; Chessbrough, 1987). Statistical Methods Differences in enteric parasite compos ition between protected and urban habitats were tested using a 2 x 19 chi-squared conti ngency table. Specifica lly, the different types of enteric parasite species that infect coyotes in the protec ted habitat were compared to the types of parasite species that infect urban coyotes and a chi-square analysis was utilized to determine if the parasite species found in the different habitat types were significantly different. The contingency table utilized all parasite species that were recovered during the course of this study.

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10 RESULTS Ten helminth species cons isting of three cestodes ( Diphyllobothrium latum Hymenolepis spp., and Dipylidium caninum ), six nematodes ( Ancylostoma caninum Ascaris spp., Physaloptera spp., Toxocara canis Trichurus vulpis and Uncinaria stenocephala ), and one trematode ( Paragonimus spp.) were recovered from coyote fecal samples (n=40) collected from the protected habitat (Table 1). Seven protozoan species ( Isospora spp., Blastocystis spp., Entamoeba histolytica Sarcocystis cruzi Balantidium coli Cryptosporidium spp., and Giardia canis ) were also recovered from coyote fecal samples found in the protected habitat (Table 1). Nine helminth species, all of which are very common enteric parasites of domestic dogs, were recovered from coyote fecal samples (n = 50) collected in urban habitat (Table 2). Helminth spec ies consisted of three cestodes ( Diphyllobothrium latum Dipylidium caninum and Taenia spp.), four nematodes ( Ancylostoma caninum Trichurus vulpis Toxocara canis and Uncinaria stenocephala ), and two trematodes ( Alaria spp. and Paragonimus spp.) (Table 2). Two protozoan species, Balantidium coli and Blastocystis spp., were also recovered from the urban habitat (Table 2). Of the 40 scat samples collected from th e protected habitat, 47.5% contained three or more parasite species, while only 4.0% of the 50 scat samples collected in the urban habitat contained three or more species (Figur e 1). When infected with parasites (Figure 1; zero values excluded), coyote scat in the protected habitat had an average of 2.6

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11 parasite species/scat and thos e in the urban habitat had an average of 1.4 species/scat. The mean number of parasite species per in fected scat (zero values excluded) was significantly greater in the protected than in urban hab itat (t = 3.84 df = 49 P = 0.0003) (JMP, v.5.1.2, SAS Institute Inc.). In addition to the 2 x 19 chi-square analysis (X2 = 29 df = 18, P < 0.05), a 2 x 2 chi-square analysis (X2 = 4 df = 1, P < 0.05) was also conducted with all expected values less than 5 combined. Both analyses i ndicated that the enteric parasite species that infect coyotes in the protected habitat do differ significantly from those parasite species that infect urban coyotes. Th erefore, the null hypothesis that enteric parasites of coyotes in the protected and urban habitat woul d not differ significantly was rejected. During the course of this study 19 different parasite species were recovered from coyote feces collected from protected and urban habitats. Only 9 parasite species that were found in the protected habitat were also found in fecal samples collected from the urban habitat. The fecal sa mples collected in the protec ted habitat contained eight parasite species that were not documented in fecal samples from the urban habitat. Two parasite species ( Taenia spp. and Alaria spp.), both of which infect domestic dogs and have been previously documented for Florida coyotes, were recovered from fecal samples collected in the urban habitat that were not recovered from fecal samples collected in the protected habitat. Of the parasites found in both habitats, two ( B. coli and Blastocystis spp.) were novel to C. latrans (Table 3 and Figure 2) and five ( D. latum D. caninum T. canis U. stenocephala and Paragonimus spp.) were novel to Florida coyotes (Table 4). Therefore, the null hypothesis that all enteric parasites in Florida coyotes will have been

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12 previously documented was rejected. Only two ( A. caninum and T. vulpis ) of the parasites found in both habitats had previ ously been documented in Florida coyotes (Table 5). Three parasites found only in the protected habitat ( Hymenolepis spp., Ascaris spp., and E. histolytica ) were novel to C. latrans (Table 3 and Figure 2), four ( Cryptosporidium spp., G. canis Isospora spp., and S. cruzi ) were novel to Florida coyotes (Table 4), and one ( Physaloptera spp.) had been previously documented in Florida coyotes (Table 5).

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13 Table 1. Parasites recovered from fecal samp les of coyotes in the protected habitat (n=40). %n Cestoda Diphyllobothrium latum2135 Dipylidium caninum252 Hymenolepis spp.183 Nematoda Ancylostoma caninum 208 Ascaris spp.1208 Physaloptera spp. 52 Toxocara canis231 Trichurus vulpis 31 Uncinaria stenocephala231 Trematoda Paragonimus spp.2135 Protozoa Balantidium coli1156 Blastocystis spp.12510 Cryptosporidium spp.2135 Entamoeba histolytica1239 Giardia canis283 Isospora spp.23514 Sarcocystis cruzi 2 2081 Novel parasites in Canis latrans2 Novel parasites in Florida coyotes

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14 Table 2. Parasites recovered from fecal samp les of coyotes in urban habitats (n=50). %n Cestoda Diphyllobothrium latum242 Dipylidium caninum242 Taenia spp. 42 Nematoda Ancylostoma caninum 2412 Toxocara canis221 Trichurus vulpis 105 Uncinaria stenocephala221 Trematoda Alaria spp. 21 Paragonimus spp.221 Protozoa Balantidium coli142 Blastocystis spp. 1 421 Novel parasites in Canis latrans2 Novel parasites in Florida coyotes

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15 Table 3. Novel parasite species of C. latrans found during the course of this study and the habitat they were found in. The table shows wh ich habitat type (protected and/or urban) the coyote fecal samples were collected from. P rotecte d Habitat U r b an Habitat Cestoda Hymenolepis spp. Yes Nematoda Ascaris spp. Yes Protozoa Balantidium coli YesYes Blastocystis spp. YesYes Entamoeba histolytica Yes Novel parasite species for Canis latrans

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16 Table 4. Novel parasite species of Florida c oyotes found during the course of this study and the habitat they were found in. The table shows which habitat type (protected and/or urban) the coyote fecal samples were collected from. P rotecte d Habitat U r b an Habitat Cestoda Diphyllobothrium latum YesYes Dipylidium caninum YesYes Nematoda Toxocara canis YesYes Uncinaria stenocephala YesYes Trematoda Paragonimus spp. YesYes Protozoa Cryptosporidium spp. Yes Giardia canis Yes Isospora spp. Yes Sarcocystis cruz i Yes Novel parasite species for Florida coyotes

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17 Table 5. Previously documented parasite sp ecies of Florida coyotes found during the course of this study and the habitat they we re found in. The table shows which habitat type (protected and/or urban) the coyo te fecal samples were collected from. P rotecte d Habitat U r b an Habitat D omest i c Dog Cestoda Taenia spp. YesYes Nematoda Ancylostoma caninum YesYesYes Physaloptera spp. YesYes Trichurus vulpis YesYesYes Trematoda Al ar i a spp. YesYes Previously documented for Florida coyotes

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18 0 10 20 30 40 50 60 0123+ Number of Parasite SpeciesPercentage of Fecal Samples (%) Protected Habitat (n=40) Urban Habitat (n=50) Figure 1. Distribution of the number of ente ric parasite species identified per coyote fecal sample collected. Coyote scat collected in the protected habitat had an average of 2.6 parasite species/scat (zero va lues excluded) while scat co llected in the urban habitat had an average of 1.4 parasite spec ies/scat (zero values excluded).

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19 D E B C A Figure 2. Novel parasites of Canis latrans (A) Hymenolepis spp. (B) Ascaris spp. (C) Balantidium coli (D) Blastocystis spp. (E) Entamoeba histolytica

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20 DISCUSSION Enteric Parasites of Florida Coyotes in a Protected Habitat All novel parasite species documented during the course of this study for C. latrans and for Florida coyotes were discovered in coyote fecal samples collected in the protected habitat. Additionally, protozoa we re recovered more often from fecal samples collected from the protected habitat as opposed to those collected from the urban habitats. This is probably due to the fact that th e protected habitat was a seasonal wetland and protozoa are easily transmitted through water. While some coyote fecal samples collected in the protected habitat were infected with helminths, protozoa infected them the most. The protozoa Isospora spp., Blastocystis spp., and Entamoeba histolytica were found in 35%, 25%, and 23%, respectively, of the samples collected from the protected habitat. No other parasites were recovered more often from fecal samples co llected from the protected habitat. While it is unknown, due to the current discovery of these novel parasitic infections of coyotes, what effect Blastocystis spp. and Entamoeba histolytica have on coyotes, it has been well documented that Isospora spp. infect coyotes. Coyotes from across the United States have been known to be infected with Isospora spp. For example, coyotes in Oregon (Dunbar and Giorda no, 2003), Utah (Conder and Loveless, 1978), Colorado (Arther and Post, 1977), Texas (Thornton et al., 1974), and New York (Gompper et al., 2003) have been docu mented as being infected with Isospora spp.

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21 Isospora spp. was found in 35% of the scat collected from Florida coyotes in protected habitat and from scat samples collected fr om coyotes in New York (Gompper et al., 2003), but only 3% of the scat samples collect ed during a study of coyotes in Colorado (Arther and Post, 1977) were in fected. Based on necropsies of coyotes in Utah (Conder and Loveless, 1978) and Texa s (Thornton et al., 1974), Isospora spp. infected 18% and 66%, respectively, of the animals examined. Even though Sarcocystis spp. has not previously been documented for coyotes in Florida, it has been documented in coyotes in other states (Dunbar and Giordano, 2003). Coyotes are also known to be definitive hosts of the parasite (Dubey et al., 1989). Due to the current findings of Sarcocystis spp. in fecal samples collected from the protected habitat, the parasite has been listed as a novel parasite of Florida coyotes. Twenty percent of the fecal samples collected from Florida coyotes were infected with Sarcocystis cruzi Likewise, 20% of the fecal samples collected from coyotes in Colorado were infected with Sarcocystis cruzi (Arther and Post, 1977). Results of Sarcocystis spp. (via fecal examination) from studies conducted on coyotes in Utah, Idaho, and New York are similar to those of the present study of Florida coyotes. Fourteen percent of the fecal samples colle cted from coyotes in Utah and Idaho were found to be infected with Sarcocystis spp. (Fayer and Johnson, 1975) and 27% of the fecal samples collected from coyotes in New York were infected with Sarcocystis spp. (Gompper et al., 2003). Necropsie s preformed on coyotes in Georgia (Holzman et al., 1992) and Oklahoma (Cummings et al., 2000) show ed that of the animals examined, 6% and 4%, respectively, were infected with Sarcocystis spp.

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22 Water is a major vehicle for transmission of Cryptosporidium and the infective or viable stage of this parasite is prolonged in moist environments (Fayer, 2004). Thus, it is not surprising that 13% of the coyote fecal samples collected in the protected habitat, which supports a large number of seasonal wetlands, contained Cryptosporidium while samples collected from the urban habitat did not. Cryptosporidium is of zoonotic importance due to outbreaks in drinking wa ter and recreational water (Fayer, 2004). According to MacKenzie et al. (199 4), the “defining recognition” of Cryptosporidium as a public health concern occu rred in Milwaukee, Wiscons in in 1993, when the largest water-borne disease outbreak ever recorded occurred in the public drinking water supply and approximately 403,000 people c ontracted cryptosporidiosis. Giardia is also of zoonotic importance becau se it is known to infect humans and cause disease. Giardia was first documented to infect coyotes in 2003 (Santin et al., 2003), when it was discovered that coyotes can se rve as a reservoir for the parasite. Later that year, Gompper et al. (2003) discovered Giardia in 15% of the fecal samples collected from coyotes in New York. During the present study, Giardia was recovered in 8% of the fecal samples collected from the protected habitat. Enteric Parasites of Florida Coyotes in Urban Habitat While protozoa did comprise of some of th e enteric parasites th at infected coyotes in urban habitats, it was the helminth species that infected them the most. Helminth species, all of which were very common en teric parasites of domestic dogs, were recovered most often from coyote scat samples collected in the urban habitat. Thus, it is presumed that coyotes in the urban habitat could have received these parasites from

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23 infected domestic dogs. During the present study, protozoa may not have survived as long in the environment as helminths. Theref ore, recovery and identification of protozoa in the urban habitat may have been limited. Parasites may limit coyote population grow th in a density-dependent way. Ancylostoma caninum the common dog hookworm, for exampl e, has been suggested as a regulator of coyote populations via increased neonatal mortality (Pence et al., 1988). Twenty-four percent of all fecal samples collected from the urban habitat were infected with A. caninum No other parasite was found more frequently in the urban habitat. Additionally, twenty percent of the samples collected from the protected habitat were infected with A. caninum Approximately 20% of the coyotes examined during a study in Kansas were infected with A. caninum (Ameel, 1955). In our study, the only other parasites found more often in the protected habitat were Isospora spp. (35% of the samples were infected), Blastocystis spp. (25% of the samples were infected), and E. histolytica (23% of the samples were infected), all three of which are protozoa. The fact that dog hookworms were the most prevalent enteric pa rasites of coyotes in the urban habitat is very important for fu ture densities of urban coyotes in Pinellas County and is of zoonotic importance. Acco rding to Radomski (1989), a threshold level of only >300 hookworm larvae/kg was needed to cause mortality in coyote neonates. Thus, a study on hookworm densities within co yotes of Pinellas County would give further insight into the health of urban coyot es and provide insight into the viability of their populations. Ancylostoma caninum is of zoonotic importance as well because infective stages of this hookworm can pe netrate human skin cau sing cutaneous larva migrans (Traub et al., 2005). Thus, preventa tive measures that hinder transmission of

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24 parasites between coyote populations, domestic animals, and humans should be taken (Erickson, 1944; Traub et al., 2005). Coyote fecal samples collected from the pr otected habitat had, on average, more parasite species per sample than those samples collected from the urban habitat. These averages, especially those of the protected ha bitat, are similar to those found in a study conducted by Holmes and Podesta (1968) in Ca nada. During their study of helminths, Holmes and Podesta (1968) found that the av erage number of parasi tes that infected coyotes in Canada was 2.0 parasite species per coyote. Increased numbers of enteric parasites weakens the condition of the intestines. This is important because pathogenic activities of parasites depend primarily upon th e resistance of the host and the condition of the intestinal tract (Brown, 1975). Add itionally, the ability of a host to acquire resistance to a parasite depends on immunit y, nutritional state, a nd the condition of the intestinal tract within that host (Brown, 1975). While immunity can be built up, severe and prolonged exertion breaks down acquire d immunity and renders the animal susceptible to reinfection (Olsen, 1974). Novel Parasites of Canis latrans Newly established coyotes in Florida woul d be expected to lack resistance to novel parasites due to recent exposure. In a ddition, rapid habitat loss in Florida, mainly due to urbanization, lowers survival of adu lt coyotes and increases the probability of transmission of disease between wild and domestic canids (Grinder and Krausman, 2001). It is suggested that Florida coyotes are more sus ceptible to reinfection by novel parasites because of their rapi d range expansion and lack of acquired immunity. This is

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25 of importance for the species because the ove rall health of coyote populations declines when heavy parasitic infections occur (Belden and Kiesecker, 2005). During the course of this study, five novel enteric paras ite species were discovered which, to my knowledge, have not been previously documented in Canis latrans Of these, one cestode ( Hymenolepis spp.), one nematode ( Ascaris spp.), and three protozoa ( Balantidium coli Blastocystis spp., and Entamoeba histolytica ) were recovered, some of which are potentially pathogenic to humans (Abe et al., 2002). Biomolecular studies would need to be conducted on Hymenolepis spp., Ascaris spp., and Blastocystis spp. to determine which species were present. All of the novel parasites documented in this study for C. latrans were discovered in fecal samples collected from coyotes in the protected ha bitat. Two of these, Balantidium coli and Blastocystis spp. were also discovered in fecal samples collect ed from coyotes in the urban habitat. Blastocystis has been reported in feca l matter of domestic dogs ( Canis familiaris ) and cats ( Felis cattus ) (Duda et al., 1998) as well as in cattle ( Bos taurus ), pigs ( Sus domestica ), and zoo animals (Abe et al., 2002). While Hymenolepis has more recently been documented in domestic dogs (Traub et al., 2005), it has been well known that Entamoeba (Jordan, 1967; Northway, 1975; and Wittnich, 1976), Ascaris (Traub et al., 2005), and Balantidium (Dikmans, 1948; Das, 1999) infect them. Infection by all novel parasites of C. latrans discovered during the present study occurs through passive transmission (i.e., neithe r parasite nor host takes an active role in transmission) from contaminated sources, including soil and wa ter. Additionally, autoinfection (i.e., proglottid disintegrates in the intestine reinfecting the host) can occur with Hymenolepis (Price, 1994).

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26 While human infections of Balantidium coli are rare, Ascaris Blastocystis Entamoeba and Hymenolepis are of zoonotic importance. Ascaris spp. is a very common nematode of animals and humans throughout the world, but rarely results in death. Originally, Blastocystis spp. was considered a nonpathogenic yeast, but in 1967, it was reclassified as a protozoan (Z ierdt et al., 1967) and today it is known to be pathogenic to humans. Blastocystis spp. is frequently found in the intestinal tract s of humans (Price, 1994), causing diarrhea in immunosuppressed indi viduals (Zierdt, 1991), and reports of infection continue to increase. Blastocystis spp. and Entamoeba spp. are two of the few amoebas to infect humans (Price, 1994). Entamoeba spp. is ranked as an important parasite of humans due to its wide distri bution and pathogenic prope rties (Olsen, 1974). While each parasite is cosmopolitan, Entamoeba spp. is more commonly found in warm, moist climates (Olsen, 1974) Infection by the tapeworm Hymenolepis spp. occurs through consumption of contaminated sources. Intermediate hosts are not required for certain species of Hymenolepis but others utili ze rodents, fleas ( Ctenocephalides spp.), or cockroaches ( Periplaneta spp.) (Price, 1994). Novel Parasites of Florida Coyotes In addition to the five novel enteric parasites discovered for C. latrans nine enteric parasite species were discovered during the present st udy which have not previously been documented for Florida coyot es, but have been documented for coyotes in other states. Of the novel parasites discovered fo r Florida coyotes, two cestodes ( D. latum and D. caninum ) were found in both the protected and urban habitats. During the present

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27 study, 13% of the fecal samples collected from the protected habitat were infected with D. latum while only 4% of the samples collected from the urban habitat were infected. Five percent (5%) of the coyotes examined by Holmes and Podesta (1968) in Alberta, Canada were infected with Diphyllobothrium spp. Dipylidium caninum has been known to infect coyotes (Ameel, 1955; Butler and Grundmann, 1954), dogs, and humans throughout the United States. While D. caninum is referred to as the “dog tapeworm,” human infection can occur when the intermed iate host (usually a flea) is consumed (Brown, 1975). Two nematodes ( T. canis and U. stenocephala ) novel to Florida coyotes were discovered in both the protected and urba n habitats during the present study. T. canis recorded in the present study for Florida coyotes in protected (3%) and urban (2%) habitats is similar to that recorded for c oyotes in New York (2%; Gompper et al., 2003) and Canada (1%; Holmes and Podesta, 1968), but lower than that recorded for coyotes in Utah (6%; Conder and Loveless, 1978). In the present study, U. stenocephala infections of Florida coyotes in protected (3%) and urban (2%) habitats are lower than those documented for coyotes in Montana (18%; Se esee et al., 1983), Canada (16%; Holmes and Podesta, 1968), and New York (6%; Gompper et al., 2003). While the effect of these parasites on Florida coyotes is not definitive ly known, it is presumed that they will not routinely be pathogenic to Fl orida coyotes. Additionally, Uncinaria infections in other carnivores are usually less severe than those of the dog hookworm ( Ancylostoma caninum ) (Bowman, 1999). Paragonimus spp. is the only trematode documented in this study that is novel for Florida coyotes. Florida coyot e scat collected from the prot ected and urban habitats were

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28 infected with this trematode. While Paragonimus spp. has been reported in fox (Bekoff, 1978), dogs (Bekoff, 1978), and coyotes outside of Florida, documentation of these parasites discovered via fecal examination is limited due to the loca tion of the parasite within the host’s body. Paragonimus spp. is a fluke that resi des in the lungs of the infected animal (Brown, 1975). Detection of Paragonimus spp. in fecal samples would only result from the host swallowing parasite eggs (i.e., swallowing sputum) (Brown, 1975). Conclusion While this study has likely identified the ma jority of enteric pa rasites that infect Florida coyotes, there were significant di fferences in species composition of enteric parasites of coyotes between study locations. Some parasite species were recovered from both habitat types, but overall more protozoa were documented in the protected habitat (probably due to it being a seasonal wetla nd) and more helminths in the urban. Additionally, all of the helminths documented in the urban habitat are common parasites of domestic dogs. Another significant di fference between study locations was the number of novel parasite speci es recovered. More novel para site species were recovered from the protected habitat while more pa rasite species known to commonly infect domestic animals were found in the urban habitat. My previous prediction that parasite s recovered from the urban coyote scat samples would contain more parasites commonly found in domestic animals was supported by the findings that all of the he lminths documented in coyotes in the urban habitat are common parasites of domestic animals. Therefore, it is presumed that coyotes

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29 in the urban habitat received these parasites from infected domestic dogs. Additionally, from a wildlife management perspective, domestic animals should not be allowed in nature preserves due to the heightened risk of wildlife becoming infected with parasites known to infect domestic animals. Pets coul d introduce new parasites into the protected area resulting in wildlife (i.e., coyotes) becoming infected. While this study has shown that previ ous studies of coyotes from other geographical locations do apply to Flor ida coyotes, it has also documented novel parasites of the coyote in Fl orida. Optimum temperatures for helminths are 27 – 34oC and with Florida being closer to the tropics these temperatures are available throughout most of the year. Additionally, biodiversity of parasite species increases near the tropics. Therefore, the vast differences in climate, flora and fauna found in Florida, as opposed to other states, could be partly responsible for the recent discovery of these novel infections in the coyote. Implications One example of a preventative method that would hinder transm ission of parasites between coyote populations, domestic animals, and humans is for pet owners to be more aware of their pet’s behavior while in public areas. Specifically, pet owners should prevent their pet from coming into contact with feces previously deposited in urban areas. When infected canids (domestic dogs or coyot es) defecate in areas visited by domestic dogs, it is possible that dogs could become inf ected if they consume viable parasites. Thus, not allowing pets to come into contac t with infected feces would help prevent infection.

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30 Another preventative method that pet owners can take is to remove pet feces from urban/public areas. Removal of pet feces w ould result in fewer viable hookworms being present in these urban habita ts. In theory, this could result in fewer coyotes being infected. Assuming that proper veterinary ca re (i.e., yearly fecal exams) is given to domestic animals, the severity of enteric parasite infections would be minimal. Conversely, coyotes do not receive such care. Thus, infection would pose a greater threat to their health. A third example of a preventative method that humans should take is to perform sanitary behavior, such as fr equent hand washing, especially for children. Parasites documented in this study remain viable outsi de of the host while in water and soil. Humans can also become infected with thes e enteric parasites if they consume viable parasites or drink contaminated water (R ubel and Wisnivesky, 2005). Children, due to their close contact with soil are more susceptible to infection. These methods are especially critical due to the rapid disappearance of wild habitats. Areas frequented by wild canids domestic canids, and humans overlap when dense human populations and vast urbanization are present, as is the case in Pinellas County, Florida (Canon et al., 2004). Hence, boundaries between wildlife and domestic animals become obscured and the risk of transm ission of diseases incr eases (Tigas et al., 2002).

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31 LITERATURE CITED ABE, N., M. NAGOSHI, K. TAKAMI, Y. SAWANO, AND H. YOSHIKAWA. 2002. A Survey of Blastocystis sp. in livestock, pets, and z oo animals in Japan. Veterinary Parasitology 106:203-212. AMEEL, D.J. 1955. Parasites of the coyote Canis latrans Say, in Kansas. Journal of Parasitology 41:325. ARJO, W.M., E.M. GESE, C. BROMLEY, A. KOZLOWSKI, AND E.S. WILLIAMS. 2003. Serologic survey for diseas es in free-ranging coyotes ( Canis latrans ) from two ecologically distinct areas of Utah. Journal of Wildlife Diseases 39:449-455. ARTHER, R.G. AND G. POST. 1977. Coccidia of coyotes in Eastern Colorado. Journal of Wildlife Diseases 13:97-100. BEAN, D.L., E. ROJAS-FLORES, G.W. FOSTER, J.M. KINSELLA, AND D.J. FORRESTER. 2005. Parasitic helminths of Eurasian Collared-Doves ( Streptopelia decaocto ) from Florida. Journal of Parasitology 91:184-187.

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32 BEKOFF, M. 1978. Coyotes: biology, behavior, and management. Academic Press. New York, New York, 384pp. BELDEN, L.K. AND J.M. KIESECKER. 2005. Gl ucocorticosteroid hormone treatment of larval treefrogs increases infection by Alaria sp. trematode cercariae. Journal of Parasitology 91:686-688. BOWMAN, D.D. 1999. Geor gis’ Parasitology for Veterinarians, 7th Edition. W.B. Saunders Company. Philadelphia, Pennsylvania, 430pp. BROWN, H.W. 1975. Basic Clin ical Parasitology, 4th Edition. Appleton Century Crofts. New York, New York, 355pp. BUTLER, J.M. AND A.W. GRUNDMANN. 1954. The intestinal helminths of the Coyote Canis latrans Say, in Uta h. Journal of Parasitology 40:440-443. CANON-FRANCO, W.A., L.E.O. YAI, S.L. P. SOUZA, L.C. SANTOS, N.A.R. FARIAS, J. RUAS, F.W. ROSSI, A.A.B. GOMES, J.P. DUBEY, AND S.M. GENNARI. 2004. Detection of antibodies to Neospora caninum in two species of wild canids, Lycalopex gymnocercus and Cerdocyon thous from Brazil. Veterinary Parasitology 123:275-277.

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33 CHESSBROUGH, M. 1987. Medical Laboratory Manual for Tropical Countries. Vol. 1, 2nd Edition, University Press, Cambridge, 84 pp. CONDER, G.A. AND R.M. LOVELESS. 1978. Parasites of the coyote ( Canis latrans ) in Central Utah. Journa l of Wildlife Diseases 14:247-249. CONTI, J. A. 1984. Helminths of foxes and coyotes in Florida. Proceedings of the Helminthological Society of Washington 51:365-367. CUMMINGS, C.A., A.A. KOCA N, R.W. BARKER, AND J. P. DUBEY. 2000. Muscular sarcocystosis in coyotes from Oklahoma. Journal of Wildlife Diseases 36(4):761763. DAS, U. 1999. A case of Balantidium coli infection in a dog. The Indian Veterinary Journal 76(2):174. DIKMANS, G. 1948. The Dog, Canis familiaris a New Host of Balantidium spp. Proceedings of the Helminthological Society of Washington 15(1):40-41. DUBEY, J.P., C.A. SPEER, AND R. FAYER. 1989. Sarcocystosis of animals and man. CRC Press, Boca Raton, Florida, 215 pp.

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34 DUDA, A., D. STENZEL, AND P. BOREHAM. 1998. Detection of Blastocystis sp. in domestic dogs and cats. Vete rinary Parasitology 76:9-17. DUNBAR, M.R. AND M.R. GIOR DANO. 2003. Abundance a nd condition indices of coyotes on Hart Mountain National Ante lope Refuge, Oregon. Western North American Naturalist 62(3):341-347. ERICKSON, A. B. 1944. Helminths of Minnesota canidae in relation to food habits, and a host list and key to the species reported from North America. American Midland Naturalist 32:358-372. FAYER, R. 2004. Cryptosporidium : a water-borne zoonotic parasite. Veterinary Parasitology 126(1-2):37-56. FAYER, R. AND A.J. JOHNSON. 1975. Sarcocystis fusiformis infection in the coyote ( Canis latrans ). Journal of Infecti ous Diseases 131:189-192. FOSTER, G.W., M.B. MAIN, J.M. KINS ELLA, L.M. DIXON, S.P. TERRELL, AND D.J. FORRESTER. 2003. Parasitic helm inths and arthropods of coyotes ( Canis latrans ) from Florida, U.S.A. Comparative Parasitology 70:162-166. GILLESPIE, T.R., E.C. GREINER, AND C. A. CHAPMAN. 2005. Gastrointestinal parasites of the colobus monkeys of Uganda. Journal of Parasitology 91:569-573.

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35 GOMPPER, M.E., R.M. GOODMAN, R.W. KAYS, J.C. RAY, C.V. FIORELLO, AND S.E. WADE. 2003. A survey of the parasites of coyotes ( Canis latrans ) in New York based on fecal analysis Journal of Wildlife Diseases 39:712-717. GRINDER, M. AND P.R. KRAUSMAN. 2001. Morbidity – mortality factors and survival of an urban coyote population in Arizona. Journal of Wildlife Diseases 37:312-317. HOLMES, J.C. AND R. PODESTA. 1968. The he lminths of wolves and coyotes from the forested regions of Alberta. Canadian Journal of Zoology 46:1193-1204. HOLZMAN, S., M.J. CONROY, AND W.R. DAVIDSON. 1992. Diseas es, parasites and survival of coyotes in South-Centra l Georgia. Journal of Wildlife Diseases 28:572-580. JORDAN, H. 1967. Amebiasis ( Entamoeba histolytica ) in dog. Veterinary Medicine and Small Animal Clinician 62:61. LINDSAY, D.S., J.P. DUBEY, AND B.L. BLAGBURN. 1997. Biology of Isospora spp. from humans, nonhuman primates, and domestic animals. Clinical Microbiology Reviews 10:19-34.

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36 MAC KENZIE, W.R., N.J. HOXIE, M.E. PR OCTOR, M.S. GRADUS, K.A. BLAIR, D.E. PETERSON, J.J. KAZMERCZAK, D. G. ADDISS, K.R. FOX, J.B. ROSE AND J.P. DAVIS. 1994. A massive outbreak in Milwaukee of Cryptosporidium infection transmitted through the public water supply. New England Journal of Medicine 331:161-167. MAEHR, D.S., R.T. McBRIDE, AND J.J. MU LLAHEY. 1996. Status of coyotes in South Florida. Florida Field Naturalist 24:101-107. MAIN, M.B., S.F. COATS, AND G.M. A LLEN. 2000. Coyote distribution in Florida extends southward. Florida Field Naturalist 28:201-203. MAIN, M.B., P.B. WALSH, K.M. PORTIE R, AND S.F. COATS. 1999. Monitoring the extending range of eoyotes in Florid a: Results of the 1997-98 Statewide Scent Station Survey. Florida Fi eld Naturalist 27:150-162. MATSUO, K. AND H. KAMIYA. 2005. Modified sugar centrif ugal flotation technique for recovering Echinococcus multilocularis eggs from soil. Journal of Parasitology 91:208-209. NORTHWAY, R. 1975. Entamoeba histolytica in a dog. Veterinary Medicine and Small Animal Clinician 70:306.

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37 OLSEN, O.W. 1974. Animal Parasites: Their Life Cycles and Ecology. 3rd Edition. University Park Press. Baltimore, 352 pp. PENCE, D.B., F.F. KNOWLTON, AND L.A. WINDBERG. 1988. Transmission of Ancylostoma caninum and Alaria marcianae in coyotes ( Canis latrans ). Journal of Wildlife Diseases 26:560-563. PENCE, D.B. AND W.P. MEINZER. 1979. Helminth parasitism in the coyote, Canis latrans from the Rolling Plains of Texas. International Journal for Parasitology 9:339-344. PRICE, D.L. 1994. Procedure Manual for the Diagnosis of Intestinal Parasites. CRC Press. Boca Raton, Florida, 263 pp. RADOMSKI, A.A. 1989. Host-parasite relationshi ps of helminths in a coyote population from southern Texas with particular re ference to the dog hookworm. M.S. Thesis, Texas Tech University, Lubbock, Texas, 132 pp. RUBEL, D. AND C. WISNIVESKY. 2005. Magnit ude and distribution of canine fecal contamination and helminth eggs in two areas of different urban structure, Greater Buenos Aires, Argentina. Veterinary Parasitology 133:339-347.

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38 SANTIN, M., K. LUDWIG, R. FAYER, AND J.M. TROUT. 2003. First report of Giardia in coyotes ( Canis latrans ). Journal of Eukaryotic Microbiology 50:709. SEESEE, F.M., M.C. STERNER, AND D.E. WORLEY. 1983. Helminths of the coyote ( Canis latrans Say) in Montana. Jour nal of Wildlife Diseases 19:54-55. THORNTON, J.E., R.R. BELL, AND M.J. RE ARDON. 1974. Internal parasites of coyotes in Southern Texas. Journal of Wildlife Diseases 10:232-236. THURBER, J.M. AND R.O. PETERSON. 1991. Changes in body size associated with range expansion in the coyote (Can is latrans). Journal of Mammalogy 72:750755. TIGAS, L.A., D.H. VAN VUREN, AND R.M. SAUVAJOT. 2002. Behavioral responses of bobcats and coyotes to hab itat fragmentation and corridors in an urban environment. Biological Conservation 108:299-306. TRAUB, R.J., I.D. ROBERSTON, P.J. IRWIN, N. MENCKE AND A. THOMPSON. 2005. Canine gastrointestinal parasitic zoonos es in India. Trends in Parasitology 21:42-48. U.S. CENSUS BUREAU (USCB). 2004. http ://quickfacts.census.gov/qfd/states.

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39 WITTNICH, C. 1976. Entamoeba histolytica infection in a german shepherd dog. Canadian Veterinary Journal 17:259-263. WOODING, J.B., AND T.S. HARDISKY. 1990. Coyot e distribution in Florida. Florida Field Naturalist 18:12-14. WOODING, J.B., E.P. HILL, AND P.W. SUMNER. 1984. Coyote food habits in Mississippi and Alabama. Proceedings of the Annual Conference of the Southeast Association of Fish and W ildlife Agencies 38:182-188. ZAMAN, V. 1984. Atlas of Medical Parasitology. 2nd Edition. ADIS Health Science Press. Singapore, 170 pp. ZIERDT, C. 1991. Blastocystis hominis past and future. Clini cal Microbiology Reviews 4:61-79. ZIERDT, C., W. RUDE, AND B. BULL. 1967. Protozoan characteristics of Blastocystis hominis American Journal of Clinical Pathology 48:495-501.

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40 CHAPTER TWO: A COMPARATIVE ST UDY BETWEEN THE DIET OF COYOTES IN A PROTECTED AND URBAN HABITAT

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41 INTRODUCTION Diet is an important aspect of unde rstanding the ecology of the coyote ( Canis latrans ). Different factors affect coyote re production such as the amount of available food and the degree of human exploitation (Wi ndberg et al., 1997). As such, coyotes are very opportunistic and adaptable when it co mes to meeting thei r nutritional needs (Stratman and Pelton, 1997). A c oyote’s diet is reflective of the habitat it utilizes and varies across geographical expa nse. Previous research on the diet of coyotes has occurred throughout the United States (Wooding et al., 1984; Lee and Kennedy, 1986; Crossett and Elliott, 1991; Bartel and Knowlton, 2005). Most of the studies on the diet of coyotes have occurred in rural habitat (L ingle et al., 2005; Prugh, 2005; Azevedo et al., 2006). Relatively few studies have been conduc ted to determine the diet of coyotes in sub-urban habitats (MacCracken, 1982; Fedria ni et al., 2001) or ur ban habitats (Quinn, 1997; Grinder and Krausman, 2001). Even fewer studies have been conducted on coyotes in Florida, a state w ith rapidly changing habitats (i .e., drastic urbanization). Wagner and Hill (1994) conducted a study of the di et of coyotes in four different states (Florida, Alabama, Mississippi, and Arkansas), but only evaluated the effect of coyotes on wild turkey ( Meleagris gallopavo ). Stratman and Pleton (1997) and Thornton et al. (2004) also conducted studies on the diet of Florida coyot es but these studies were conducted on military facilities in northwest and south-central Florida, respectively. This

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42 current study compares the diet of Florida coyotes in the most de nsely populated county in Florida, Pinellas County (USCB, 2004). Coyotes are scavengers (Arjo et al., 2 002) and opportunistic omnivores (Blanton and Hill, 1989). As such, they have a wide spectrum of dietary items they consume. It is important to study these animal s throughout different geograph ical locations to document the great variety of food items consumed. Not only do coyote diet s vary across their geographical range, but also seasonally. Lee and Kennedy (1986) conducted a study in Tennessee and found seasonal variation in coyot e diet for rodents, insects, reptiles, amphibians, opossum ( Didelphis virginiana ), and persimmon ( Diospyros virginiana ). Wooding et al. (1984) found that livestock detec tion in coyote diet was highest in winter and spring in Mississippi and Alab ama, while white-tailed deer ( Odocoileus virginianus ) remains were more frequent during the summer and winter. Coyote diet varies across rura l, sub-urban, and urban grad ients as well. Fedriani et al. (2001) found that in the most urban area of their California study, anthropogenic foods comprised 25% of the coyote diet duri ng the dry season and 14% during the wet. In contrast, they found that in the rural areas, anthropoge nic foods accounted for 3% of their diet during the dry season and only trace amounts were detected during the wet season (Fedriani et al., 2001). Coyotes are relatively new to Florida a nd little research has been conducted to determine their diet in this region of th e country. In 1994, Wagner and Hill found wild turkey remains in only two scat samples from coyotes in Florida. Stratman and Pelton (1997) also conducted a diet study by collecti ng and analyzing the diet remains in scat samples and found that important diet items for coyotes in northwestern Florida were

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43 shrub/vine fruit (80%), beetles (55%), pers immon (27%), and white-t ailed deer (15%). Deer occurred most often (29%) during the fawning season. Wild hog ( Sus scrofa ; 13%) was only recovered during the spring. Thornt on et al. (2004) found that the majority of diet (via scat analysis) of c oyotes in south-central Florida consisted of white-tailed deer, wild hog, and domestic cow ( Bos taurus ). Contrary to Stratman and Pelton’s 1997 study, Thornton et al. (2004) recove red wild hog during every season. Also, coyotes observed in Stratman and Pelton’s (1997) study cons umed fruit more often than those in Thornton’s (2004) study (80% vs. 24.5%). The present study focuses on tw o aspects of coyote diet: di et diversity, defined as the different types of diet items consume d, and seasonal variation (wet season vs. dry season) in diet. The primary objective of this study was to investigate differences in diet between a protected and urban population of coyotes. More specifically, this study was designed in order to answer the following questions: Which habitat within this study (protected or urban) has highe r diet diversity? Does the co mposition of diet items differ between protected and urban habitats? Does habitat type (protected or urban) and/or season (wet or dry) affect whether or not coyotes c onsume anthropogenic waste (as determined by presence/absence of anthropogeni c waste in scat sample)? Does season (wet or dry) affect coyot e diet in the protected an d/or urban habitats? Diet diversity: Null hypothesis : Diet items consumed by coyotes in the protected habitat will not differ significantly from those diet items consumed in the urban habitats. Alternate hypothesis : Diet items consumed by coyot es in the protected habitat will differ significantly from those diet items consumed in the urban habitats.

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44 It is predicted that diet items consumed by coyotes in the protected habitat will differ significantly from those in the urban ha bitat. More specifi cally, coyotes in the protected habitat are ex pected to consume a wider variet y of diet items as opposed to those in the urban habitat, wh ich are expected to have a le ss varied diet, eating more of the same items consistently. This assumpti on is based on the protected habitat offering more variation in the types of diet items that coyotes could consume (i.e., more wildlife). Seasonal variation in diet: Null hypothesis : Coyote diet in protected and urban habitats will not differ significantly between seasons. Alternate hypothesis : Coyote diet in protected an d urban habitats will differ significantly between seasons. It is predicted that seasonal variation (wet season vs. dry season) will affect the diet of coyotes in both habita ts (protected and urban) due to changes in available diet items.

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45 STUDY SITES Pinellas County is the most densely popul ated county in Florida with over 1281 people per square kilometer (U SCB, 2004). Two types of ha bitat were compared during this study: protected and urban. Protected Habitat Brooker Creek Preserve (BCP; 27o54’ N, 82o41’W) was used for the protected habitat (Bean et al., 2005). BCP is an 8500 acr e wilderness area that is actively managed for natural resource protection. Located in the northeast corner of the county, the boundaries of BCP are shared with densely populated residential areas. The study site consists of extensive pine flatwoods and fr eshwater swamps. Fauna include white-tailed deer ( Odocoileus virginianus ), raccoons ( Procyon lotor ), armadillo ( Dasypus novemcinctus ), bobcat ( Lynx rufus ), otter ( Lutra canadensis ), wild turkey ( Meleagris gallopavo ), red-shouldered hawks ( Buteo lineatus ), wood storks ( Mycteria americana ), bobwhite quail ( Colinus virginianus ), and gopher tortoise ( Gopherus polyphemus ). Urban Habitat Different sites throughout Pinellas Count y were used for urban habitat. Sites were determined using GIS (ArcGIS v.8) to plot existing geospatial information including land-use categories, railroads, bi ke trails, and power lines throughout the

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46 county. Subsequently, a map revealing ru ral, sub-urban, and urban areas based on residential (land-use category) population density was genera ted. Sites were identified based on criteria expected to support urban coyot es. The sites were constrained such that they had land cover of urban sites similar to that of BCP, were traversed by power lines, bike trails, or inactive railroads, and were lo cated in urban areas. The wet season at both sites was defined as June 1 through Oct ober 31 and the dry season as November 1 through May 31 (Chen and Gerber, 1990).

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47 METHODS Field Methods Fresh coyote feces were collected over th e course of two years (May 2005 thru March 2007) from trails, power lines, and bike trails in the protected (n=49) and urban (n=71) habitats. Paths were traversed on f oot, bicycle, and by ATV three times a week during the course of this study. Each fecal sample was measured (length and diameter) and photographed in the field. Species origin of the fecal samples was determined by adjacent sign (tracks) and dimensions of feces. Upon confirmation of coyote scat, the sample was assigned a unique identification code and its longitude and latitude were recorded by use of a GPS unit. Finally, the sample was placed in its own paper bag. To avoid collecting bobcat feces, only those samples in excess of one inch in diameter were collected (Gompper et al., 2003). To avoi d collecting domestic dog feces, only samples which contained hair and bone fragments and/or which were accompanied by coyote tracks were collected (W ooding et al., 1984). Laboratory Methods Prey of the Florida coyote were iden tified to the lowest possible taxonomic category based on bone, teeth, nails, and hair that were recovered from each sample. After removing approximately 4 grams of th e sample for parasite examination, the remaining sample was oven dried at 60–80oC for at least 48h to kill any latent parasites

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48 (Wagner and Hill, 1994). After desiccation, eac h sample was individually placed in the top of a combination of wire mesh sieves (Stratman and Pelton, 1997) and washed thoroughly with a garden spray hose attached to a sink faucet. The remaining diet items were then transferred from each sieve onto pa per plates and allowed to dry overnight. After the contents on each paper plate (one for each sieve) were thoroughly dry, the remains were separated into the following categ ories: hair, bones, teeth, nails, feathers, reptile, insects, vegetative matter, berries, anthropogenic waste (i.e ., trash, rope, plastic wrappers) and unknown. Dorsal guard hair was then separated based on gross morphological characteristics (i.e. color, color bands, and color band locations) and slides were made for prey identification. Hairs were identified based on gross morphological characteristics and medullary configurat ions (Wilkins et al., 1982). To aid in identification, hair, bones, teeth, and nails we re compared with specimens housed at the Florida Museum of Natura l History in Gainesville (Thornton et al., 2004). Statistical Methods Diet items were recorded and the percen tage of coyote scat samples containing each item was determined. A Poisson regressi on, with the total number of diet items as the response variable, was ut ilized to determine the relation between diet items and habitat (H), season (S), and in teraction (H*S) because the va riable “diet items” is count data and follows a Poisson distribution. In or der to determine if presence or absence of anthropogenic waste in coyote feces was in re lation to habitat (H), season (S), or an interaction (H*S), a logistic regression was utilized because the diet item “Anthropogenic Waste” is a Bernoulli variable and follows a binomial distributi on. Thus, a logistic

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49 regression was used to examine what effect, if any, H, S, or H*S had on coyotes consuming anthropogenic waste. In both the Poisson regression and the logistic regression, the habitat (H) response was cont rolled for the effects of season (S).

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50 RESULTS Forty-nine fecal samples were collected from the protected habitat and were used to determine the diet of coyotes in this habi tat. Diet items recovered most often from coyotes in the protected hab itat were vegetative matter (96% ), Insecta (53%), Rodentia ( Sciurus carolinensis, S. niger shermanii, Sigmodon hispidus, and Geomys pinetis ) (45%), Cervidae ( Odocoileus virginianus ) (33%), berries (31%), and Lagomorpha ( Sylvilagus spp.) (29%) (Table 1). Among all diet items, Aves (12%), anthropogenic waste (8%), Testudines (4%), and Felidae (2%) were recovered the least in the protected habitat (Table 1 and Figure 1). In the urban habitat, 71 fecal samples were collected and were utilized to determine the diet of urban coyotes. The di et items recovered most often from urban coyote scat were berries (56 %), Lagomorpha (32%), vegetative matter (25%), Rodentia (18%), and anthropogenic waste (1 8%; Table 2). In the urban habitat, Aves (7%), Insecta (4%), Didelphidae ( Didelphis virginiana ) (3%), Testudines (1%), Felidae (1%), and Procyonidae (1%) were recovered the least (Table 2 and Figure 1). Analysis using a Poisson regression indicated that coyotes in the protected habitat had higher diet diversity than urban coyot es with a fitted regr ession of log(mean response) = 1.1386-0.6138H with a p-value = 0.0000. Thus, the diet of urban coyotes is less varied than that of coyotes in a protected habitat.

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51 The logistic regression showed a very weak relationship between habitat and anthropogenic waste consumed by coyotes. Th e most significant term in the model was the interaction between hab itat and season (H*S): logit( prob) = -2.0149+1.0594H*S with a p-value = 0.0821. Main effects of habita t was logit(prob) = -2.4204+0.9249H with a pvalue = 0.1265. In the protected habitat, ei ght percent (8%) of the samples contained anthropogenic waste (Table 3). In compar ison, eighteen percent (18%) of the samples collected from the urban habitat cont ained anthropogenic waste (Table 3). While the Poisson regression showed that neither season (S) nor interaction (H*S) could be used to determine any changes in th e number of diet items consumed by coyotes from either habitat, seasonal va riation did affect the types of diet items consumed. In the protected habitat, Insecta and berries were found more frequen tly in the samples collected during the wet season while Lagomorpha, R odentia, and Cervidae appeared more frequently during the dry season (Figure 2). It should also be not ed that the only time Felidae was present in any of the collected sa mples from the protec ted habitat was during the wet season (Figure 2). Additionall y, in the protected habitat, fawn Odocoileus virginianus were only consumed in the wet season while adult O. virginianus were only consumed in the dry season (Table 1 and Figure 3). In the urban habitat, Didelphidae, Pr ocyonidae, Felidae, Insecta, Aves, and Testudines were only recovered during the wet season (Figure 4). Berries and vegetative matter were recovered most often from th e urban habitat during the wet season while Lagomorpha and anthropogenic waste were re covered most often during the dry season (Figure 4). During the wet season, 68% of the fecal samples collected from the urban habitats contained berries, but this number declined to 22% during the dry season (Table

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52 2 and Figure 5). Conversely, during the wet season only 21% of th e samples collected from the urban habitat contained Sylvilagus spp., but during the dry season this number rose to 67% (Table 2 and Figure 5). When looking at seasonal variation among Florida coyotes (combining both protected and urban habitat data ), season (wet vs. dry) affect s consumption of certain diet items more than others. Overall, Sylvilagus spp., adult O. virginianus and vegetative matter were recovered more often from fecal samples of Florida coyotes during the dry season while berries and fawn O. virginianus were recovered more often during the wet season (Figure 6). Additionally, when combin ing protected and urba n habitat data to examine Florida coyote diet, vegetative ma tter (54%) and berries (46%) were recovered from more samples than any other diet item (Figure 7). The percentages of Florida coyote fecal samples containing Lagomorpha, Rodentia, and Insecta were 31%, 29%, and 24% respectively (Figure 7). Speci fically, of the Rodentia recovered, Sigmodon hispidus was recovered most often from Florida coyote scat (Table 3).

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53 Table 1. Diet items consumed by co yotes in protected habitat (n=49). % n%n % n Didelphidae000000Didelphis virginiana 000000Lagomorpha2263682914Sylvilagus spp. 2263682914Rodentia411150114522Sciurus carolinensis 410021 Sciurus niger shermanii 729284 Sigmodon hispidus 3083683316 Geomys pinetis 005121Cervidae3083683316Odocoileus virginianus ( fawn) 30800168 Odocoileus virginianus (adult) 00368168Procyonidae000000Procyon lotor 000000Felidae410021 Insecta63174195326 V egetative Matter9325100229647 Berries44121433115 Aves15492126 Testudines415142 Anthropogenic Waste729284 Diet Item: Wet Season (n=27) Dry Season (n=22)TOTAL (n=49)

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54 Table 2. Diet items consumed by co yotes in urban habitats (n=71). % n%n % n Didelphidae420032Didelphis virginiana 420032Lagomorpha211167123223Sylvilagus spp. 211167123223Rodentia21111121813Sciurus carolinensis 95112107 Sciurus niger shermanii 840064 Sigmodon hispidus 420032 Geomys pinetis 000000Cervidae000000Odocoileus virginianus ( fawn) 000000 Odocoileus virginianus (adult) 000000Procyonidae210011Procyon lotor 210011Felidae210011 Insecta630043 V egetative Matter28151732518 Berries68362245640 Aves950075 Testudines210011 Anthropogenic Waste1582851813 Diet Item: Wet Season (n=53) Dry Season (n=18)TOTAL (n=71)

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55 Table 3. Diet items consumed by Florida coyotes (protected and urban habitats combined).% n%n%n Didelphidae003222Didelphis virginiana 003222Lagomorpha291432233137Sylvilagus spp. 291432233137Rodentia452218132935Sciurus carolinensis 2110778 Sciurus niger shermanii 846478 Sigmodon hispidus 3316321518 Geomys pinetis 210011Cervidae3316001316Odocoileus virginianus 3316001316Procyonidae001111Procyon lotor 001111Felidae211122 Insecta5326432429 Vegetative Matter964725185465 Berries311556404655 Aves12675911 Testudines421133 Anthropogenic Waste8418131417 Combined (P+U) (n=120) Protected (n=49)Urban (n=71) Diet Item:

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56 0 10 20 30 40 50 60 70 80 90 100Vege ta tive Ma tt e r Insecta R od e ntia C erv id ae B err ie s Lagomorpha A ves An th ro po ge nic W a st eDiet ItemsPercentage of Samples Containin g Diet Item (%) Protected Habitat (n=49) Urban Habitat (n=71) Figure 1. Comparison of diet items consumed between coyotes in protected and urban habitats.

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57 0 10 20 30 40 50 60 70 80 90 100Lagomorpha R od enti a C e r vi d ae Fe l idae In s ect a V e ge t ativ e Matt e r Be r ri e s A ves Testudines An t hrop og eni c WasteDiet ItemsPercentage of Samples Containin g Diet Item (%) Wet Season (n=27) Dry Season (n=22) Figure 2. Seasonal variation in the percenta ge of coyote fecal samples collected in a protected habitat that co ntain each diet item.

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58 0 10 20 30 40 50 60 70 80 90 100S y lvilagus spp S. car ol in ensi s S. sh er m ani i S. hi spi dus G pi net i s O. virgi ni anus -F aw n O. virginianus -Adult F elidae I nsecta V eg eta t i ve M at te r B er ri es A ves Testu di nes Anthropogenic Wast eDiet Item% Samples Wet Dry Figure 3. Seasonal variation in the percenta ge of coyote fecal samples collected in a protected habitat that cont ain each diet item (repres ented by the lowest possible taxonomic level).

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59 0 10 20 30 40 50 60 70 80 90 100Did e lphi dae Lagom or pha Rode n tia Pr ocy o ni dae F el idae I nsect a Vegetative Matter Ber r i es Aves Te s tudines A nt hr opogeni c Wast eDiet ItemsPercentage of Samples Containin g Diet Item (%) Wet Season (n=53) Dry Season (n=18) Figure 4. Seasonal variation in the percentage of coyote fecal samples collected in urban habitats that contain each diet item.

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60 0 10 20 30 40 50 60 70 80 90 100D. virginiana Sy l vil a gu s s p p. S c aro l i n en si s S. s h erma n ii S. hispidus P l o t o r F e l i d ae Insecta Veg e t a t i ve Matte r B e r ri e s A v es Testu d ines Anthropogenic WasteDiet Item% Samples Wet Dry Figure 5. Seasonal variation in the percentage of coyote fecal samples collected in urban habitats that contain each diet item (represen ted by the lowest possible taxonomic level).

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61 0 10 20 30 40 50 60 70 80 90 100D. virg i niana Sy l v i la g us s p p. S. car ol i n en s i s S. sherma n ii S. hispidus G. p in e ti s O. virg i n ia n u s Fa w n O. virginian u s -Adu l t P. lotor F el i dae I ns e c t a Vegeta t ive Matte r Berries Aves T e s tu d in e s Anthropogenic WasteDiet ItemPercentage of Samples (%) Wet Dry Figure 6. Seasonal variation in the percenta ge of fecal samples collected from Florida coyotes (protected and urban habitats combined) that contain each diet item (represented by the lowest possible taxonomic level).

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62 54 46 31 29 24 14 13 9 3 22 1 0 10 20 30 40 50 60Vegetative Matter Berri e s Lagomorpha Rodentia Insec t a Anthropogenic Waste Cervidae Aves Testudines Didelphidae Fe l ida e ProcyonidaeDiet ItemsPercentage of Samples Containing Diet Item (%) Figure 7. Florida coyote diet (protected and urban ha bitats combined) (n=120).

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63 DISCUSSION Diet of Coyotes in Different Habitat Types Results indicate that my pred iction that the diet of coyot es in the protected habitat would differ significantly from that of coyotes in the urban habitat was correct. Coyotes in the protected habitat consumed a wider variet y of diet items (higher diet diversity) than the urban coyotes. This is lik ely attributed to the protecte d habitat offering more diverse wildlife species available for consumption. Of the diet items consumed in both habitat types, Sylvilagus spp. was the diet item that varied the least between urban (32%) and protected (29%) habitats while vegetative matter was the diet item that varied the most (96% protected, 25% urban). It should be noted that a limita tion to this study is that it only represents a snapshot in time. For example, the results do not indicate what percentage of coyote diet consists of each diet item, but rather what percentage of samples contains specific diet items. Deer were recovered in the protected hab itat from more samples than berries or Sylvilagus spp. Adult deer were rec overed in the protected ha bitat from as many fecal samples as fawns. It should be noted that the percentage of d eer found in coyote fecal samples from the protected environment is likel y an over-representation of deer mortality caused by coyotes (Stratman and Pelton, 1997). Coyotes are scavengers and opportunistic carnivores which allows them to obtain food without expending much

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64 energy (Arjo et al., 2002). Thus, coyotes may scavenge carrion or feed on one kill for multiple days (Stratman and Pelton, 1997). However, it is interesting that 8% of the coyote fecal samples collected in the protected habitat contained anthropogenic wa ste. The protected habitat is largely surrounded by dense urban housing. Thus, the question of how these coyotes obtained anthropogenic waste is of importance. Wh ile it is possible that anthropogenic waste could have been obtained from within the borde rs of the protected habitat, it seems more likely that the coyotes obtained it from the neighborhoods surr ounding the habitat. During the course of this study, coyote tracks were identified outsi de gates and fences lining the parameter of the protected habitat. Additionally, while obtaining scat samples in the protected habitat, areas were disc overed where coyotes had dug, and were actively utilizing as passageways, areas under the fe nce that surrounded th e study site. It appeared that these passageways were bei ng actively utilized as an egress from the protected habitat to the surr ounding neighborhoods and then la ter as an entryway back into the protected habitat. Further studies of the movements of coyotes could address this question. While the origin of the anthropogenic waste recovered from coyote fecal samples collected in the protected hab itat cannot be determined, it is assumed that some of it was obtained from the neighborhoods surrounding the study site. This assumption is supported by a comparable study (Fedriani et al., 2001) in which only 3% of the coyote samples contained anthropogenic waste, as oppos ed to 8% of the samples in the present study. Because coyote scat samples were co llected in the protected habitat over the course of this study and relatively few samp les contained anthropogeni c waste, it can be

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65 assumed that if the coyotes were leaving th e protected habitat it was probably for short durations of time. Additionally, the results of the present study indica te that the diet of coyotes in the protected habitat is reliant on and maintained by the biological diversity within the preserve rather than by human influence in the surrounding neighborhoods. Areas with higher human population densit ies would also have higher amounts of anthropogenic waste. Therefore, it is not surprising that anthropogenic waste was recovered over twice as often from coyote fecal samples collect ed in the urban habitat as opposed to those collected in the protected ha bitat. Eighteen percen t (18%) of the fecal samples collected in the urban habitat contai ned anthropogenic waste, as opposed to 8% in the protected habitat. It is presumed th at even fewer fecal sa mples collected in the protected habitat would contai n anthropogenic waste if coyotes remained in the protected habitat, as opposed to venturing out into the surrounding urban neighborhoods. While there are noticeable differences between the amount of anthropogenic waste consumed by coyotes in the protected an d urban habitats, the logistic regression showed a weak relationship be tween habitat and anthropogenic waste. It is suggested that the weak relationship shown by the logist ic regression may be due to coyotes in the protected habitat consuming anthropoge nic waste from the surrounding urban neighborhoods (urban habitat). Thus, rath er than a complete delineation between protected and urban habitats as main e ffects, the urban neig hborhoods surrounding the protected habitat may have cause d interference in the protected habitat data, resulting in a higher p-value. While the Poisson regression showed that season could not be used to determine any changes in the number of diet items c onsumed by coyotes from either types of

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66 habitat, seasonal variation did affect the types of diet items consumed. Diet items in the protected habitat most af fected by seasonal variati on were white-tailed deer, Sciurus carolinensis Geomys pinetis and Felidae. Fawn was only recovered from coyote fecal samples collected in the prot ected habitat during the wet season, while adult deer was only recovered during the dr y season. Additionally, S. carolinensis and Felidae were only recovered during the wet season from fecal samples collected in the protected habitat while G. pinetis was only recovered during the dry season. Seasonal changes were also recognized fo r other diet items recovered in coyote fecal samples collected in the protected habita t. For example, berries, Insecta, and Aves were recovered more often during the wet s eason. Berries were recovered from over three times as many samples collected in the protected habitat during the wet season (44%) as opposed to those collected during th e dry season (14%). Insecta was recovered from 63% of the samples collected in th e wet season, but only 41% of the samples collected during the dry season contained Insecta. Fifteen percent (15%) of the fecal samples collected in the wet season, as opposed to 9% of the samples collected in the dry season, contained Aves. Remains of Sylvilagus spp. were recovered more ofte n during the dry season from coyote fecal samples collected in the protected habitat. Thir ty-six percent (36%) of the fecal samples collected during the dry seas on from the protected habitat contained Sylvilagus spp., as opposed to only 22% of the sa mples collected in the wet season. While more coyote fecal samples collected in the dry season (9%), as opposed to the wet season (7%), from the protected habitat cont ained anthropogenic waste, the difference was negligible. Fedriani et al. (2001) found similar results for seasonal variation of

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67 anthropogenic waste recovered from coyote f ecal samples collected in rural areas of California. Fedriani et al. (2001) found that 3% of the fecal sample s collected in the dry season contained anthropogenic waste, but only trace amounts of anthropogenic waste were recovered during the wet season. Diet items in the urban habitat that were affected the most by seasonal variation were Didelphis virginiana Sciurus niger shermanii Sigmodon hispidus Procyon lotor Felidae, Insecta, Aves, and Testudines, all of which were only recovered during the wet season. No diet items were recovered during the dry season that were not also recovered during the wet season from fecal samples collected in the urban habitat. Therefore, in regards to the type of diet items consumed, the diet of urban coyotes changed the most during the wet season. Other diet items consumed in the urban habitat were also affected by seasonal variation. The percentage of coyote fecal samples collected during the wet season (68%) from the urban habitat that contained berries mo re than tripled when compared to that of the dry season (22%). Sylvilagus spp. was recovered from more than three times as many samples collected from the urban habitat dur ing the dry season (67%) as opposed to the wet season (21%). Additionally, half as many fecal samples collected during the dry season (11%) contained Rodentia as compared to those collected in the wet season (21%) from the urban habitat. Anthropogenic waste was also found in only half as many samples collected during the wet season (15%) as opposed to the dry season (28%) from the urban habitat. The results of the present study for seasonal variation of anthropoge nic waste in urban coyote fecal samples are very similar to those of Fedriani et al. ( 2001). Fedriani et al.

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68 (2001) conducted a study on c oyote diet in an urban area of California and found that during the wet season only 14% of the fecal samples contained anthropogenic waste, but during the dry season this number rose to 25%. Diet of Florida Coyotes When combining protected and urban habita t data to examine Florida coyote diet, most of the fecal samples contained vegeta tive matter (54%) and berries (46%). While Stratman and Pelton (1997) found that fruit accounted for 80% of the coyote diet in northwestern Florida, the present study rec overed berries from only 46% of the fecal samples collected. Rabbits were recovered from 31% of the fecal samples of Florida coyotes. Rabbits appeared in more samples of Florida coyotes ( 31%) than samples of coyotes in Kentucky (22%) (Crossett and Elliott, 1991). Rodentia and Insecta were recovered fr om 29% and 24%, respecitively, of the fecal samples collected from Florida coyotes Specifically, of the Rodentia consumed, Sigmodon hispidus was recovered most often (15%). Only seven percent (7%) of the fecal samples collected from coyotes in Florida contained S. niger shermanii (Sherman’s fox squirrel). Even so, it should be noted that the Sherman’s fox squirrel is ranked by Florida Fish and Wildlife Conservation Co mmission (FWC) as a Species of Special Concern (SSC) (FWC, 2004). Species are ranke d as SSC when there is future risk of extinction or, as is the case with the Sherman’ s fox squirrel, may already meet criteria for being classified as a threatened species, but conclusive data are lim ited or lacking (FWC, 2004).

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69 Felidae was only recovered from 1 (2%) coyote fecal sample collected in the protected habitat and 1 (1%) sample collected in the urban habitat. When combining samples from both habitats, Felidae was rec overed from only 2% of Florida coyote fecal samples. The results of the present study reve al dramatically less consumption of Felidae than other studies on coyote diet. Crossett and Elliott (1991) found remains of Felidae in 13% of the stomachs examined during necrops ies on coyotes in Kent ucky. It should be noted that it was not pos sible to differentiate, ba sed on gross morphological characteristics and medullary configurations, whether the remains in feces classified as Felidae were those of bobcat, domestic cat, or fe ral cat. Additionally, it is not possible to determine whether the remains of prey in coyote feces are the result from the coyote scavenging carrion or actually kil ling the prey. Therefore, it is likely that the one scat sample collected in the protected habitat that contained Felidae hair was actually a large (>1 inch diameter) bobcat scat. Genetic anal ysis of scat samples would need to be conducted to determine if samples were ind eed deposited by coyotes, or rather by bobcat or even domestic dogs. The results of the pr esent study indicate that cats are not a diet item frequently recovered from coyote scat in either habitat. These findings do not support the popular opinion among many of the general public that coyotes are a major threat to domestic cats. In regards to the importance of white-taile d deer for the diet of Florida coyotes, the results of the present st udy are similar to previous st udies conducted on the diet of Florida coyotes (Stratman and Pelton, 1997; T hornton et al., 2004). This study concluded that white-tailed deer was r ecovered from 13% of the f ecal samples collected from

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70 Florida coyotes. Similarly, Stratman a nd Pelton (1997) found that white-tailed deer accounted for 15% of the diet of coyo tes in northwestern Florida. Seasonal variation (wet vs. dry) did occur in the diet of Florida coyotes. Even so, some diet items were affected more than othe rs. For example, the remains of adult whitetailed deer were only recovered from fecal samples of Florida coyotes during the dry season. However, fawns were only recovere d during the wet season. These results are similar to those of other studies in which fawns were consumed more frequently (as carrion) during the same time of year (Cook et al., 1971; Salwasser, 1974; Berg and Chesness, 1978; Litvaitis and Shaw, 1980). It has previously been documented th at much of the large prey consumed by coyotes is indeed carrion (B erg and Chesness, 1978; Huge l, 1979; Weaver, 1979). Arjo et al. (2002) documented coyotes scavenging large prey carr ion, as opposed to capturing and killing large prey. Due to coyotes being sc avengers, it is difficult to determine if diet items recovered from a coyote’s feces were the result of the coyot e killing the prey or simply scavenging the carrion. Thus, the frequency of coyote predation on large mammals should not be predicted by the fr equency of large mammal remains found in coyote scat. Sylvilagus spp. and berries were other diet it ems of Florida coyotes that were dramatically affected by seasonal variation. When comparing the wet and dry seasons, Florida coyote fecal samples containing Sylvilagus spp. more than doubled in the dry season. Sylvilagus spp. was recovered from 50% of the fecal samples of Florida coyotes collected during the dry season, but only from 20% during the wet season. These results are similar to previous findings in which Sylvilagus spp. were reported as major diet

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71 items of coyotes during the winter months (Clark, 1972; Wagner and Stoddart, 1972). Similarly, when comparing the wet and dry seasons, the amount of Florida coyote fecal samples containing berries tripled in the we t season. In the wet season berries were recovered from 60% of the fecal samples, but less than 20% of the samples collected in the dry season contained berries. Conclusion Coyotes are relatively new inhabitants of Fl orida. Therefore, little is known about how Florida coyotes and other fl ora and fauna coexist. As opportunistic animals, coyotes are able to find available res ources in disturbed landscapes. Even so, they still rely on natural diet items (Riley et al ., 2003). While coyote diets ar e reflective of the habitats they inhabit, they also vary across geographical expanse. The present study documented that habitat and season interact to affect diet, further comp licating interpretation of coyote diet. Coyotes in the western United States ha ve been vilified as major predators (Mitchell et al., 2004), but this negative connot ation does not appear to be sufficient for coyotes in Florida, as evident by deer recove red from fecal samples of Florida coyotes. Over the course of the present study, deer was recovered from only 13% of all coyote fecal samples examined. Most reports (G ese and Grothe, 1995; Hugel and Rongstad, 1985; Ozoga and Harger, 1966) of coyotes capturi ng and killing large prey have occurred when there was sufficient amounts of snow on the ground, which hinders the prey’s ability to successfully escape pr edation (Arjo et al., 2002). Due to the climate in Florida, snow is not a factor. Deer was only rec overed from fecal samples collected in the

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72 protected habitat which had high deer densities. When deer densities are high, predation by coyotes has little effect on deer populations due to coyotes being more selective (i.e., primarily prey on weak indivi duals) (Patterson and Messier, 2 003). Due to the lack of snow in Florida and high deer densities in the protected habitat, it is presumed that coyotes mainly consumed carrion and/or dis eased or older individuals as opposed to killing healthy prey. Determining what diet items are consum ed by Florida coyotes and if seasonal variation occurs in the diet could support fu ture resource management plans. The results of the present study indicate that the prediction that the diet of coyotes in the protected habitat would differ significantly from that of coyotes in the urban habitat was correct. This prediction was based on the notion that in the urban habitat the amount of different diet items is limited while the protected ha bitat offers more variation (biological diversity) in the types of diet items that coyotes could consume (i.e., more wildlife). The present study found remains of the Sher man’s fox squirrel, which is a SSC, in only 8 coyote fecal samples. It is highly un likely that Florida coyotes pose a major threat to the continued existe nce of the Sherman’s fox squirrel. It is suggested that continued urbanization poses a greater threat to the future survival the squirrel rather than predation by coyotes. This suggestion is supported by the FWC. According to the FWC, the Sherman’s fox squirrel is a SSC because it “has a significant vulnerability to habitat modification, environmental a lteration, human disturbance, or human exploitation which, in the foreseeable future, may result in its becoming a threatened species unless appropriate protective or management tech niques are initiated or maintained” (FWC, 2004).

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73 While increased urbanization, habitat frag mentation, and complete habitat loss are inevitable, one of the most serious threats to biological diversity worldwide is the destruction of habitat (Wilcove et al., 1998) Documenting the diet of coyotes in protected and urban habitats is essential in order to deve lop a better understanding of the effects of habitat type on di et. These methods are especia lly critical due to the rapid disappearance of protected Florida habitat from urbanization.

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74 LITERATURE CITED ARJO, W.M., D.H. PLETSCHE R, AND R.R. REAM. 2002. Di etary overlap between wolves and coyotes in Northwestern M ontana. Journal of Mammalogy 83(3):754766. AZEVEDO, F.C., V. LESTER, W. GORSUCH, S. LARIVIERE, A.J. WIRSING, AND D.L. MURRAY. 2006. Dietary breadth and overlap among five sympatric prairie carnivores. Journal of Zoology 269:127-135. BARTEL, R.A. AND F.F. KNOWLTON. 2005. Func tional feeding responses of coyotes, Canis latrans to fluctuating prey abundance in the Curlew Valley, Utah, 1977 – 1993. Canadian Journal of Zoology 83:569-578. BEAN, D.L., E. ROJAS-FLORES, G.W. FOSTER, J.M. KINSELLA, AND D.J. FORRESTER. 2005. Parasitic helminth s of Eurasian collared-doves ( Streptopelia decaocto ) from Florida. Journal of Parasitology 91:184-187. BERG, W.E. AND R.A. CHESNE SS. 1978. Ecology of coyotes in northern Minnesota. Pages 229-247 in M. Bekoff, ed. Coyotes: biolo gy, behavior, and management. Academic Press, New York, New York.

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78 OZOGA, J.L. AND E.M. HARGER. 1966. Winter activities and feed ing habits of northern Michigan coyotes. Journa l of Wildlife Management 30:809-818. PATTERSON, B.R. AND F. MESSIER. 2003. Ag e and condition of deer killed by coyotes in Nova Scotia. Cana dian Journal of Zoology 81:1894-1898. PRUCH, L.R. 2005. Coyote prey selection and community stab ility during a decline in food supply. Oikos 110:253-264. QUINN, T. 1997. Coyote ( Canis latrans ) food habits in three ur ban habitat types of western Washington. Northwest Science 71:1-5. RILEY, S.D., R.M. SAUVAJOT, T.K. FULL ER, E.C. YORK, D.A. KAMRADT, C. BROMLLEY, AND R.K. WAYNE. 2003. Effe cts of urbanization and habitat fragmentation on bobcats and coyotes in Southern California. Conservation Biology 17(2):566-576. SALWASSER, H. 1974. Coyote scats as an indicat or of time of fawn mortality in the north Kings deer herd. Calif ornia Fish and Game 60:84-87. STRATMAN, M.R. AND M.R. PELTON. 1997. Food habits of coyotes in Northwestern Florida. Proceedings of the Annual Confer ence of the Southeastern Association of Fish and Wildlife Agencies 51:269-275.

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80 WILKINS, L., M. LANGWORTHY, C.D. RATHBUN, AND R. SULLIVAN. 1982. Identification of the dorsal guard hairs of some of Florida mammals. Technical Report prepared for the Florida Game and Fresh Water Fish Commission. University of Florida, Gain seville, Florida. 59 pp. WINDBERG, L.A., S.M. EBBERT, AND B.T. KELLY. 1997. Population characteristics of coyotes ( Canis latrans ) in the Northern Chihuahuan Desert of New Mexico. American Midland Naturalist 138:197-207. WOODING, J.B., E.P. HILL, AND P.W. SUMNER. 1984. Coyote food habits in Mississippi and Alabama. Proceedings of the Annual Conference of the Southeast Association of Fish and W ildlife Agencies 38:182-188. VOIGT, D.R. AND W.E. BE RG. 1987. Coyote. pp. 344-357 in M. Novak, J.A. Baker, M.E. Obbard and B. Malloch eds., Wild Furbearer Management and Conservation in North America. Ministry of Na tural Resources, Ontario, Canada.