COYOTE (Canis latrans) SPATIAL ECOLOGY AND INTERACTION WITH CATTLE (Bos
taurus) IN THE SUB-TROPICAL RANGELANDS OF FLORIDA
By
KE ZHANG
A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE
UNIVERSITY OF FLORIDA
2017
© 2017 Ke Zhang
3
ACKNOWLEDGMENTS
I would like to thank Dr. Stewart Breck and Dr. Michael Avery and the United States
Department of Agriculture, National Wildlife Research Center for providing funding to
undertake this study (#15-7449-1154-CA). A special thank you goes to Ralph Pfister of Adam’s
Ranch for his skill and guidance in how to trap coyotes efficiently, as well as Cary Lightsey,
Layne Lightsey, Jim Strickland, and Gene Lollis for allowing the work to occur on properties
within their cattle operations. I thank the MacArthur Agroecology Research Center of Archbold
Biological Station for logistical and data support. Thank you to Dr. Breck, Dr. Avery, Bethany
Wight, James McWhorter, Connor Crank, and Wes Anderson for their help capturing, sampling,
and tracking coyotes. I am very grateful to Connor Crank, Audrey Wilson, and Jane Anderson in
supporting me as I continued to learn English during my thesis writing process. I also thank my
committee members, Dr. Main and Dr. Basille, for their guidance during my proposal, and thesis
writing process. Above all, I am very grateful to Dr. Boughton for his patient guidance and
generous funding. This study was performed under IACUC #201408477 reviewed by the
University of Florida.
4
TABLE OF CONTENTS
page
ACKNOWLEDGMENTS ...............................................................................................................3
LIST OF TABLES ...........................................................................................................................6
LIST OF FIGURES .........................................................................................................................8
ABSTRACT ...................................................................................................................................10
CHAPTER
1 COYOTE HOME RANGE, MOVEMENT, AND ACTIVITY PATTERNS ..........................12
Introduction .............................................................................................................................12 Methods ..................................................................................................................................16
Study Sites .......................................................................................................................16 Capture, Restraint and Handling .....................................................................................17 Mortality Monitoring and Radio-telemetry .....................................................................18
Analysis Methods ............................................................................................................19
Results.....................................................................................................................................22 Capture and Retrieval ......................................................................................................22 Overall and Seasonal Home Ranges ................................................................................23
Movement Rates and Circadian Rhythm .........................................................................25 Effects of Temperature and Rainfall on Coyote Activity ................................................25
Coyote Interactions ..........................................................................................................26 Discussion ...............................................................................................................................28
Sizes of Sub-Tropical Coyotes ........................................................................................28
Home Range Sizes of Sub-Tropical Coyotes ..................................................................30 Movement Rates and Circadian Rhythm .........................................................................32
Effects of Temperature and Rainfall on Coyote Activity ................................................35
Individual Interactions .....................................................................................................36
2 COYOTE HABITAT SELECTION, INTERACTION WITH CATTLE ................................68
Introduction .............................................................................................................................68 Methods ..................................................................................................................................70
Land Cover Map Preparation ..........................................................................................70
Habitat Selection Analysis ..............................................................................................70 Cattle Movement Record .................................................................................................72
Coyote Habitat Selection of Pasture with Cattle/Calves .................................................73 Compare Calves Carcass Records with Coyotes Movements .........................................74
Results.....................................................................................................................................74 Simplified Land Cover Map ............................................................................................74
5
Overall Habitat Selection Result .....................................................................................74 Habitat Selection by Gender and Territoriality ...............................................................75 Cattle/Calves Pasture Selection Ratios ............................................................................75 Comparison of Calf Carcass Records with Coyotes Movements ....................................76
Discussion ...............................................................................................................................76 Coyote Habitat Selection .................................................................................................76 Cattle/Calves Pasture Selection Ratios, Carcass Visiting Behavior ................................78
3 CONCLUSION.......................................................................................................................101
LIST OF REFERENCES .............................................................................................................103
BIOGRAPHICAL SKETCH .......................................................................................................110
6
LIST OF TABLES
Table page
1-1 Coyote home range sizes in different studies ........................................................................38
1-2 Coyote capture records ..........................................................................................................40
1-3 Coyote overall home range summary ....................................................................................41
1-4 Coyote seasonal home range summary .................................................................................42
1-5 Coyote home range season, gender Mann-Whitney U test ...................................................43
1-6 Coyote core use area season, gender Mann-Whitney U test .................................................43
1-7 Hourly movement rate, time period of the day and season Linear Mixed-Effects Model
analysis ...............................................................................................................................43
1-8 Cumulative travel distance, time period of the day and season Linear Mixed-Effects
Model analysis ...................................................................................................................43
1-9 Hourly movement rate, time period of the day and season Tukey Honest Significant
Difference analysis.............................................................................................................44
1-10 Cumulative travel distance, time period of the day and season Tukey Honest
Significant Difference analysis ..........................................................................................45
1-11 Daily mean temperature and rainfall interactive effect on daily cumulative travel
distance Linear Mixed-Effect Model analysis ...................................................................46
1-12 Daily mean temperature and rainfall main effects on daily cumulative travel distance
Linear Mixed-Effect Model analysis .................................................................................46
1-13 Daily mean temperature, rainfall, and season interactive effect on daily cumulative
travel distance Linear Mixed-Effect Model analysis .........................................................46
1-14 Records of interactions of coyotes from two different home ranges ..................................46
7
2-1 Summary of reclassified land cover process .........................................................................82
2-2 Grazing records example from Bucks Island Ranch .............................................................84
2-3 Calculation selection ratio of pastures with cattle ................................................................85
2-4 Calculation selection ratio of pastures with cattle and calves ..............................................86
2-5 First-order level selection ratio Chi-Square Test ..................................................................87
2-6 First-order level selection ratio Chi-Square Test in winter ..................................................87
2-7 First-order level selection ratio Chi-Square Test in spring ...................................................87
2-8 Second-order level selection ratio Chi-Square Test .............................................................87
2-9 Second-order level selection ratio Chi-Square Test by territoriality ....................................87
2-10 Selection ratios of pasture with and without cattle..............................................................88
2-11 Selection ratios of pasture with and without cows calving .................................................88
2-12 Chi-Square Test of cattle/calve pasture selection ratio .......................................................88
2-13 Visits of coyote to pastures with incidental calf carcasses found by ranchers on Buck
Island Ranch.......................................................................................................................89
8
LIST OF FIGURES
Figure page
1-1 Research sites ........................................................................................................................47
1-2 Buck Island coyote home ranges ...........................................................................................48
1-3 Lightsey coyote home ranges ................................................................................................49
1-4 Blackbeard coyote home ranges ............................................................................................50
1-5 Circadian activity of coyote hourly movement in winter and spring ....................................51
1-6 Coyote hourly movement rates in different seasons and time periods ..................................52
1-7 Coyote hourly movement rates in different seasons and time periods Tukey Honest
Significant Difference ........................................................................................................53
1-8 Coyote cumulative travel distance in different seasons and time periods .............................54
1-9 Coyote cumulative travel distance in different seasons and time periods Tukey Honest
Significant Difference ........................................................................................................55
1-10 Predicted temperature and rainfall effect on daily travel distance ......................................56
1-11 Coyote daily travel distance by temperature and season .....................................................57
1-12 Instantaneous distance between female juvenile coyote F2 and F3 every 30mins .............58
1-13 Coyote F3 travel out of the home range on 3/13/2015 ........................................................59
1-14 Coyote F3 & M1 interaction on 3/10/2015 .........................................................................60
1-15 Coyote F2 travel out of the home range on 4/18/2015 ........................................................61
1-16 Instantaneous distance between female juvenile coyote F9 and F10 every 30mins ...........62
1-17 Coyote F8 & F9 & F10 interaction on 12/22/2014 .............................................................63
1-18 Coyote M13 & M15 interaction on 12/15/2014 ..................................................................64
1-19 Coyote M13 & M15 interaction on 4/5/2015 ......................................................................65
1-20 Coyote home ranges in southern states ...............................................................................66
1-21 Resident coyote home ranges in winter and spring season of Florida ................................67
9
2-1 Home range of two male coyotes and two female coyotes on the Buck Island Ranch
showing underlying simplified land cover categories .......................................................90
2-2 Home ranges of one male coyote and two female coyotes on the BlackBeard Ranch
showing underlying simplified land cover categories .......................................................91
2-3 Home ranges of two male coyotes on the Lightsey Ranch showing underlying
simplified land cover categories ........................................................................................92
2-4 First-order level resource availability and utilization proportions ........................................93
2-5 First-order level habitat selection ratios for whole study period ...........................................94
2-6 First-order level habitat selection ratios in winter .................................................................95
2-7 First-order level habitat selection ratios in spring .................................................................96
2-8 Second-order level habitat selection ratios by gender and season ........................................97
2-9 Second-order level habitat selection ratios by territoriality and season ................................98
2-10 Buck Island Ranch pasture boundary, carcass reported pastures, coyote home ranges
map .....................................................................................................................................99
2-11 Buck Island Ranch pasture boundary, carcass reported pasture, coyote home ranges,
and coyote location points on 11/12/2014 .......................................................................100
10
Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science
COYOTE (Canis latrans) SPATIAL ECOLOGY AND INTERACTION WITH CATTLE (Bos
taurus) IN THE SUB-TROPICAL RANGELANDS OF FLORIDA
By
Ke Zhang
May 2017
Chair: Raoul Boughton
Major: Wildlife Ecology and Conservation
Coyote are medium sized predatory, native American Canidae, that during the last one
hundred years has expanded its range extensively to include much of the North American
continent, including Florida. By the year 2000, coyote presence had been confirmed in all
counties of Florida (McCown & Scheick, 2007). The sub-tropical climate and intense cattle
husbandry in south-central Florida provide excellent environmental conditions for coyote to
survive, leading to changes in biological responses, such as increased phenotypic size and shifted
spatial behaviors. From November 2014 to May 2015, I retrieved extensive locational data from
9 coyotes spread over three research sites (Buck Island Ranch, BlackBeard Ranch, and Lightsey
Ranch). Using these spatial locations, I 1) tested for effects of season and gender on coyote home
range, 2) describe coyote circadian activity pattern and test for relationships between temperature
and rainfall with coyote movement, 3) infer interactions among coyote dyads using proximity
and describe the patterns of these occurrences, and 4) examined habitat selection of coyote
within rangelands of Florida, to understand if coyote have a preference for improved pasture
habitats, and if preference increases when pastures are stocked with cattle or cattle are calving.
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The average resident home range size of coyotes in my study area was 27.14 km2. Floridian
coyotes were most active during crepuscular and nocturnal periods of the day. In the spring,
coyote became more active, having higher hourly movements and longer cumulative travel
distance than in the winter. Increasing temperature and rainfall negatively correlated with daily
travel distance. Resident adult male coyotes in adjacent territories avoided encountering each
other during the period of the study, transient males infrequently encountered resident males. In
two cases females from the same group interacted frequently and are thought in one case to be
siblings and the other possibly a mother and daughter. At the population level, coyote preferred
improved pasture, forest, and scrub & shrub habitats. Where as they spent less time in wetland
areas, roads, open water, human communities and dry prairie. There was no evidence that coyote
preferred pastures with cattle or calves to any greater extent than the pastures without. Taken
together, coyote prefer improved pastures and although do occasionally take calves, their activity
patterns suggest there are many other factors that may drive their use of pasture other than the
presence of cattle and calves.
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CHAPTER 1
COYOTE HOME RANGE, MOVEMENT, AND ACTIVITY PATTERNS
Introduction
Coyotes originated in western North America, but are now commonly found throughout
the continent of North America. By the 1960s, coyotes had crossed the Mississippi river into the
southeastern United States (McCown & Scheick, 2007), entering northern Florida from Alabama
and Georgia. Populations were established in the panhandle and northern region of Florida by the
1970s (Layne, 1994), and the establishment of coyote populations in Florida was contributed to
by both natural expansion and release of coyotes used for hound hunts (Hill et al., 1987). A
survey conducted in 1983 documented coyotes in 18 Florida counties, primarily in the panhandle
(Brady & Campbell, 1983). The presence of coyote populations was confirmed in central Florida
in 1990 (Wooding & Hardisky, 1990). By the middle of 1990s coyotes were well established in
southern Florida (Maehr et al., 1996), and by 2000, coyotes were confirmed in 65 of Florida’s 67
counties (Main et al., 2000). Today, coyotes inhabit all counties of Florida (McCown & Scheick,
2007).
Coyotes can be described as resident or transient based on the spatial behavior patterns of
individuals. Residential coyotes are territorial, show aggressive behaviors toward intruders,
maintain consistent home range boundaries which manifest in consistent and repeated use of
specific areas over time, and usually have a smaller home range in general, while transients are
the alternative. Transients generally have large and disjointed home ranges, showing no fidelity
to any given area (Hinton et al., 2015). Transients may be coyotes that are unable to find a mate
and defend a territory. The delineation between resident and transient coyotes needs to be taken
into account when attempting to understand home range sizes and activity patterns of coyotes.
13
Coyotes show large variation in home range sizes across North America (Table 1-1).
Coyote home ranges have also been shown to vary by age, gender, season, territoriality, and land
cover types (Hotzman et al., 1992; Hinton et al., 2015; Gese et al., 1998; Laundre & Keller,
1984; Gese et al., 1990). One study of particular focus in Florida includes analyses of coyote
home ranges and activity patterns in an area dominated by natural communities with only about
3% pasture. Within these natural communities of Florida, annual coyote home range sizes
averaged 24.8 km2 (95% Confidence interval = 16.96 – 32.64 km2, n = 7, 95% fixed Kernel) and
did not differ between dry and wet seasons (Thornton et al., 2004). Coyotes preferred scrub
habitat within core home ranges (50% fixed Kernel) with the highest selection ratios among 8
land cover types in both the dry and wet seasons, but preference to scrub was only significant in
the wet season. The preference to scrub habitat could be explained by potential prey base and
high rodent densities in this type of environment (Franz et al., 1998). Coyotes avoided wet
habitats such as swamp and marsh within core home ranges, especially during the wet season
(Thornton et al., 2004).
Coyote are more active during crepuscular and nocturnal hours and are relatively inactive
during diurnal hours. Although coyote are active during crepuscular hours, the peak of activity
can be either dawn or dusk depending on seasonality, breeding condition, and anthropogenic
patterns of activity and influences (Arias-Del Razo et al., 2010; Andelt & Gipson, 1979; Way et
al., 2004). In Florida, coyote display a circadian rhythm of movement rates with nocturnal and
crepuscular periods higher than diurnal periods (Thornton et al., 2004). Sun exposure and severe
heat in the sub-tropical environment may be the factors that drive coyotes to avoid diurnal
activities. However, research on the relationship between day length, temperature, and coyote
activity in the sub-tropical environment like Florida is scarce.
14
Coyotes are omnivores, exhibiting broad diet capabilities and complicated interactions
with local food resources which, in turn, define activity and spatial use. For example, coyotes
prey on small mammals, reptiles, birds, insects, and larger species such as white-tailed deer
(Odocoileus virginianus) (Stout, 1982), mule deer (Odocoileus hemionus) (Truett, 1979), elk
(Cervus canadensis) (Robinson, 1952), and moose (Alces alces) (Benson & Patterson, 2013).
The distribution, seasonal availability, and density of these prey items are likely to influence
coyote behaviors, activity patterns, and spatial use (Bekoff & Wells, 1986). One particular study
from the Great Basin of northern Utah and southern Idaho, where jackrabbit (Lepus californicus)
is the major prey of coyote, indicated that coyote home ranges can be significantly larger when
prey is scarce compared to when prey is abundant, and there is a higher chance of increased
proportion of transients when food is lacking (Mills & Knowlton, 1991). In the western coastal
area of Florida, insects and berries were found most frequently in coyote fecal samples in a
wildland reserve area during the wet season, while rabbits, rodents, and deer were found most
frequently during the dry season. (Grigione et al., 2011). Similarly, in Avon Park located in
central south Florida coyote fecal samples contained vegetation material, white-tailed deer, feral
swine (Sus scrofa), cattle, rodents, and rabbits. There was 5% occurrence of cattle found across
samples, even though 97% of habitat coyotes used was native (Thornton et al., 2004). In 2014,
there were 1.7 million head of cattle and calves in Florida, with Okeechobee, Highlands,
Osceola, Polk, and Hardee counties being the top five producers, all of which distributed in
central south Florida (Florida Department of Agriculture and Consumer Services, 2014). The
large cattle and calves industry in these regions may provide abundant food resources through
modification of native habitats to improved pasture and a possible increase in biomass of prey,
15
reducing the necessity of larger movements and home ranges for coyotes to access food
resources.
Coyote range expansion has also led to increased interactions with other predators. For
example, coyotes may compete with lynx (Lynx canadensis) for highly important prey items
such as the snowshoe hare (Lepus americanus) (Kolbe et al., 2007). Coyote are highly
competitive with other species of similar size, and have been show to kill bobcats (Felis rufus)
(Gipson & Kamler, 2002), and rarely, lynx (O’Donoghue et al., 1997). From the measurement of
food habit overlap among carnivores in south Florida, there was a speculation that coyote were
more competitive with bobcats, black bears (Ursus americanus), and Florida panthers (Puma
concolor coryi) than local carnivores competed with each other (Maehr, 1996). A study of the
interactions between coyotes and bobcats in south-central Florida suggested that coyotes and
bobcats display similar habitat selection, activity patterns, and overlapping home ranges.
Segregation may occur between the two species at finer scales of core use areas (50% Fixed
Kernel), suggesting a level of territoriality and competition (Thornton et al., 2004), this may
reduce availability of resources to bobcat. Other effects caused by competition between coyotes
and other predators need to be further investigated but are beyond the scope of this study.
Studies of coyote ecology in Florida have addressed population expansion, home range,
habitat selection, diets, and parasites (Maehr et al., 1996; Thornton et al., 2004; Grigione et al.,
2001; Foster et al., 2003). Since the home range, activity pattern, and habitat selection analysis of
coyote in Florida have only been carried out in a natural environment (Thornton et al., 2004), it
is largely unknown how the dominant land use in south central Florida, cattle ranching, has
impacted the behavioral ecology of coyote. The altered land cover and abundant distribution of
cattle may have a variety of potential effects on coyotes. To understand how coyotes may behave
16
differently in this agricultural landscape, I undertook a study to describe home ranges, movement
rates, activity patterns, and spatial interactions among coyotes captured on ranches in south
central Florida. The hypotheses are 1) coyote have different home range sizes and circadian
activity patterns in different seasons of the year, 2) the temperature and rainfall have effect on
coyote daily cumulative travel distance, 3) there are frequent interaction among coyotes from the
same group. I specifically examined home range size across gender, age, season, and
territoriality; describe movement activity across time of day and test if there is a seasonal (winter
versus spring) effect upon circadian activity; investigate temperature and rainfall influence upon
movement behavior; and described interactions among coyotes when they occurred.
Methods
Study Sites
Coyotes were trapped in November and December 2014 on three different ranches
located in south central Florida (Figure 1-1). The ranches were located approximately 50 km
apart. The Lightsey Ranch is located 10 km east of the city of Lake Wales, south of Tiger Lake
(27°51’, -81°22’). The Buck Island Ranch is located 6 km east of the city of Lake Placid (27°09’,
-81°06’). The BlackBeard’s Ranch is north of Myakka River State Park (27°16’, -82°08’). The
density of human residents living within each site was very low (< 8%) except around the
Blackbeard’s Ranch. The human densities were slightly higher (10%) at the Blackbeard’s Ranch
site because the low-density residential areas occurred in the north and west sides of the ranch.
All research areas had similar topography and habitat types typical of south central Florida.
These habitats consisted of palmetto palm prairies, shrublands, open pine flatwoods, hammock
forest, improved pasture, and marsh wetlands. The climate of the area is subtropical, with a
winter season from November 2014 through March 2015 (Daily mean temperature range: 6.65 ~
17
26.4 °C) and spring season from March 2015 through May 2015 (Daily mean temperature range:
13.9 ~ 29.72 °C). Annual rainfall averaged 132 cm, of which 75% typically falls during the wet
season in the summer (Swain et al., 2007).
Capture, Restraint and Handling
At each study site, I searched for signs of coyote presence (i.e., footprints, scats, scrapes
and dens). Once potential areas of coyote activity were found, I set a trapping array of 1-4 soft
catch foothold traps (MB-550- Rubber Jaw 4 coil). These soft catch foot-hold traps had rubber
pads applied to the jaws to minimize foot injury. Each trap was dyed and waxed before
deployment to minimize human scent and maintain trap condition. I attached a trap tranquilizer
device (TTD) to each foothold. The TTD is a chewable pouch containing a tranquilizer, in this
case 600 mg propiopromazine hydrochloride (Sahr & Knowlton, 2000; Zemlicka et al., 1997), to
induce lethargy in the coyote and reduce injury when trying to escape from the trap. I used a
variety of scent lures including skunk scent, coyote urine, deer meat and fermented horse flesh
mingled with a sight lure such as a cow bone to attract coyotes to the traps. I set up traps during
the evening and checked traps regularly to avoid any coyotes suffering from heat or dehydration.
If a coyote was captured, I used a Y pole at the neck to pin the coyote to the ground. The
animal was then restrained by hobbling three feet, muzzling and applying a hood. The coyote
was then carefully removed from the foothold and processed in a shielded area on a cover sheet.
During the capture I monitored temperature and recorded morphometrics (length, weight, teeth
eruption and wear), gender, reproductive status, estimated age, and then collected samples for
other studies including blood, hair and scat. Body length (cm) and weight (kg) were reported as
mean, median, and 95% Confidence interval (95% CI). Mann-Whitney U test (Ruxton, 2006)
was used to evaluate the difference on body length and weight between genders. Refurbished
18
GPS LOTEK 3300 store-on-board collars with timed release drop-off mechanisms (set to 6
months) were fitted to each coyote. The GPS was set to record location every 30 minutes from
16:00 to 10:00 and hourly between 10:00 to 16:00 to conserve battery when coyotes are
suspected to move the least. The collars were recovered using collar specific Very High
Frequency (VHF) transmitters set to transmit continuously. I set collar mortality functions to go
off after 4 hours of stationary activity, at which point the pattern of VHF signal increased in bip
rate to signify stationary behavior of the collar (e.g., slipped, or dropped from animal, or animal
deceased).
Coyotes were released and their condition was observed until their revival from the effect
of tranquilizer, after which they were able to walk away on their own accord. I returned to the
release location after one hour to confirm that the coyote had left and was mobile using a VHF
radio receiver (Communication specialists model # R1000) with a Yagi antenna (Lotek centered
on 165mhz).
Mortality Monitoring and Radio-telemetry
I tracked coyotes weekly to check for mortality events and roughly recorded coyote
locations using triangulation, to make sure they had not left the general area. Drop-off
mechanisms were designed to go off six months after the deployment date, at which time I
located and retrieved the collars. For coyotes that exited the study area and were no longer
trackable via ground VHF antenna, I attempted to locate the deployed collars using aerial
surveys. Two flights were undertaken near the end of the 6 months on each property in either a
Cessna 150 or Cubcrafters Carbon Cub EX. A survey of 50 km2 radius was performed. All collar
data were then downloaded to a computer using the LOTEK interface (Download-link version 1)
and the software GPS host 3300. Data were cleaned before analyses with relocation points of
19
greater probability of low accuracy (hdop > 7.5) removed and relocations truncated to from
deployment date to date when collars were dropped or slipped from coyotes.
Analysis Methods
Estimation of Home Range
I defined November 2014 to the end of February 2015 as winter season, then beginning
of March 2015 to June 2015 as spring season. I estimated overall and seasonal home range areas
for each coyote using two methods: Minimum Convex Polygon (MCP) and fixed Kernel
Utilization Distribution (KUD). In both cases a 95% cut-off was used; in MCP the smallest area
containing 95% of fixes and in the KUD the 95% of kernels with highest probability of use,
converted to polygon areas. I also estimated core areas using 50% MCP and 50% KUD.
For the Kernel Utilization Distribution model, I used bivariate normal kernel, with
parameter h estimated by “reference bandwidth” that is equal to:
ℎ = 𝜎×𝑛16
when
𝜎 = 0.5×(𝜎𝑥 + 𝜎𝑦)
The 𝜎𝑥 and 𝜎𝑦 are the standard deviations of the horizontal and vertical coordinate of
the location points, n is the amount of the location points.
The analyses were performed in the adehabitatHR package (Calenge, 2006) using
Rstudio (version 0.99.893, Rstudio team, 2015). I reported the results of both, but only used
polygons derived from the fixed KUD method to undertake statistical analyses because it is more
specific and accurate in identifying actual likely use of a specific area compared with MCP
(Worton, 1989). The home range sizes were reported as mean, median, and 95% CI by groups. I
used Mann-Whitney U test (Ruxton, 2006) to evaluate the difference in the sizes of home ranges
20
between genders, ages, resident and transient, and in different seasons. I plotted the result of
home range analysis with average value and standard error, compared my findings with other
known home range.
Hourly Movement Rate and Circadian Activity
I used the adehabitatLT package in R (Calenge, 2006) to calculate the hourly movement
rate. I extracted the points recorded on the hour from the original data. The hourly movement
rate was defined by the distance between two consecutive location points, that should result in 24
hourly distance records. However, because of non-fixes or location points removed with low
accuracy, there were 3% of points absent, these points and the points next to them were not used
to calculate the hourly speed of movement. I calculated mean hourly movement rates of each
coyote, and then combined all coyotes together to estimate average hourly movement rates of the
population and plotted by hour to show circadian activity trends.
I obtained daily sunrise and sunset times of the research period from Astronomical
Application Department U.S website (http://aa.usno.navy.mil/). I divided 24 hours into four time
periods of the day: dawn (sunrise time ± one hour), dusk (sunset time ± one hour), diurnal
(sunrise time + one hour to sunset time – one hour), and nocturnal (the rest of hours). I calculated
the cumulative travel distance in each time period as the sum of all hourly movement rates in the
time period. Due to the change in day length, I defined November 2014 to the end of February
2015 as the winter short daily photoperiod, then beginning of March 2015 to June 2015 as spring
long daily photoperiod. I used boxplot to present the hourly movement rate and cumulative
distance travelled within each time period in winter and spring season separately.
All the hourly movement rates and cumulative travel distances were square-root
transformed before the statistical tests. I used Linear Mixed-Effects Model to test if hourly
21
movement rate and cumulative travel distance are effected by time of the day and season. I used
Tukey’s Honest Significant Difference (Yandell, 1997) after the Linear Mixed-Effects Model to
test the interactive effects among time period of the day and the season. Random effects from
coyote individuals and location sites were blocked in the analysis. Linear Mixed-Effects Model
analysis was carried out by nlme package in R (Pinheiro et al., 2017). Tukey’s Honest
Significant Difference analysis was carried out by multcomp package in R (Hothorn et al., 2008).
Effects of Temperature and Rainfall on Coyote Activity
To understand how weather and local climatic variables may influence coyote activity, I
obtained daily mean temperature and rainfall data from Florida State Weather Center website
using the closest weather station to each research site. The stations were Archbold Biological
Station (27.16670, -81.35000), 14 km away from the Buck Island site, MTN lake Station
(27.93330, -81.58330), 23 km away from the Lightsey site, and Myakka River SP Station
(27.23330, -82.30000), 5 km away from the Blackbeard’s Ranch site. I calculated daily
cumulative travel distance as the sum of the distances coyote moved in each hour of the day. All
the cumulative travel distances were square-root transformed, the rainfall record was converted
to the category of rain and no rain based on daily rainfall. I used Linear Mixed-Effects Model to
investigate the relationship between daily cumulative travel distance, temperature, presence of
rainfall, and the seasons. Random effects from coyote individuals and location sites were
blocked in the analysis. Linear Mixed-Effects Model analysis was carried out by nlme package
in R (Pinheiro et al., 2017).
Spatial Interactions
I described the spatial interactions of coyotes by calculating the overlapping area between
their home ranges, estimating instantaneous distance, and recording occasional visit behavior
22
into other coyote’s home ranges. I used the result from home range analysis to calculate the
overlapping areas between coyote home ranges. Pythagorean theorem was used to calculate the
instantaneous distance between every two coyotes. I plotted the instantaneous distances between
females who sharing same home ranges, to present their long-term interaction through the
research period. I defined the average coyote movement rate as 70 m/minute, and because there
was a one-minute variation in recording location on GPS, I considered two coyotes with an
instantaneous distance of < 140 m (the farthest distance two coyotes can be dispersed from each
other in one minute) as interacting at that moment. I then plotted the location points of these
coyotes with their home ranges on the maps to present their interactions and visiting behavior
toward each other’s home range.
Results
Capture and Retrieval
I captured and placed collars on 15 coyotes across three research sites from November
through December 2014 (Table 1-2). No coyotes were injured during the capture and all coyote
were left in the shade to revive after handling. All coyotes revived and left by themselves within
one hour after the handling.
Among the coyotes I captured, eight of them were female (two adults, eight juveniles),
and seven of them were male (all adults). Average male body length (mean = 99.57 cm, median
= 98.00 cm, 95% CI = 95.63 - 103.51 cm; n = 7) was similar (U = 13.5, p = 0.10) to average
female body length (mean = 92.38 cm, median = 88 cm, 95% CI = 86.87 – 97.89 cm; n = 8).
Average male weight (mean = 16.19 kg, median = 16.4 kg, 95% CI = 17.39 – 14.99 kg; n = 7)
was heavier (U = 6, p = 0.0009) than average female weight (mean = 12.72 kg, median = 12.1
kg, 95% CI = 11.39 – 14.05 kg; n = 8).
23
By the beginning of June 2015, I had successfully retrieved 11 collars from the field.
Four remained missing and were not found by aerial surveys. Among the 11 collars I retrieved,
three of the coyotes were killed by humans; one was hit by a vehicle on 11/16/2014 and one was
shot on ranch on 12/23/2014. The third was shot by a rancher over 21 km from the capture
location, but the collar still contained five months of data. Due to the short deployment period,
the first two coyote collars killed by humans were not used in analyses. The nine remaining
collars included 5 male adults, 1 female adult, and 3 female juveniles. Among these nine
coyotes, average male body length (mean = 98.6 cm, median = 98 cm, 95% CI = 93.78 – 103.42
cm; n=5) was similar (U = 6.5, p = 0.46) to average female body length (mean = 93.5 cm,
median = 93 cm, 95% CI = 86.11 – 100.89 cm; n=4). Average male weight (mean = 16.34 kg,
median = 16.4 kg, 95% CI = 14.95 – 17.73 kg; n=5) was similar to (U = 2, p = 0.06) that of the
females (mean = 12.62 kg, median = 12 kg, 95% CI = 10.78 – 14.46 kg; n=4).
Coyote location data began on 11/12/2014, the second day after the release of the first
captured coyote, and ended on 6/6/2015, the last day before collar drop off for the last coyote.
Although collars were all programed to drop off from coyotes after 6 months recording time,
some collars were collected before the scheduled day because of coyote’s death or collar-
slipping for unknown reason. The average number of location points recorded by each collar was
6737. Coyote M1 had the most location points out of any collar (7587), and coyote F9 had the
least location points (4667; Table 1-2).
Overall and Seasonal Home Ranges
Given the shape and size of their home ranges during the six-month study, I classified
seven coyotes (three males, four females) as residents, and two coyotes (both males) as transients
(Figure 1-2; Figure 1-3; Figure 1-4). The two male transients (M8 and M15) had large home
24
ranges of 921 km2 and 88 km2 under 95% KUD, respectively. Mean home range for all residents
was 27.14 km2 (median = 27 km2, 95% CI = 19.00 – 35.27 km2; n = 7) under 95% KUD, and
mean core home range (50% KUD) for all residents was 4.86 km2 (median = 4 km2, 95% CI =
2.08 – 7.64 km2; n = 7). The home ranges were not significantly different for male residents
(mean = 20.33 km2, median = 20 km2, 95% CI = 8.45 – 30.21 km2; n = 3) compared to female
residents (mean = 32.25 km2, median = 31 km2, 95% CI = 23.16 – 41.34 km2; n = 4) under 95%
KUD (U = 2, p = 0.23). Similarly, core areas were not significantly different for male residents
(mean = 4.33 km2, median = 2 km2, 95% CI = -1.26 – 9.92 km2; n = 3) compared to female
residents (mean = 5.25 ± 1.7 km2, median = 4.5 km2, 95% CI = 1.92 – 8.58 km2; n = 4) under
50% KUD (U = 4, p = 0.59; Table 1-3).
Under 95% KUD and 50% KUD, I found no significant differences in resident coyote
home range sizes among seasons (Table 1-4; Table 1-5; Table 1-6). In the winter, female
residents had an average home range size of 25.25 km2 (median = 25 km2, 95% CI = 16.80 -
33.70 km2; n = 4), while male residents had an average home range size of 26 km2 (median = 32
km2, 95% CI = 9.10 – 42.90 km2; n = 3). In the spring, female residents had an average home
range size of 40.75 km2 (median = 38.5 km2, 95% CI = 28.17 – 53.33 km2; n = 4) under 95%
KUD, while male residents had an average home range size of 17 km2 (median = 10 km2, 95%
CI = 3.28 – 30.72 km2; n = 3). Among all three female juvenile coyotes, the average home range
size increased from 22 km2 (median = 26 km2, 95% CI = 14.16 – 29.84 km2; n=3) to 42.67 km2
(median = 42 km2, 95% CI = 25.68 – 59.66 km2; n=3) from the winter to the spring, although the
difference is insignificant (U = 0, p = 0.08).
25
Movement Rates and Circadian Rhythm
Average coyote hourly movement rates began to increase at dusk and were maintained
through the night until dawn. After dawn, coyotes became inactive and traveled less during the
diurnal period. Descriptively, from November 2014 to early March 2015, hours of maximum
activity were 18:00-19:00, 21:00-22:00, while from early March 2015 to June 2015, hours of
maximum activity were 19:00-20:00 and 22:00-23:00. Coyote maximum average movement rate
in the spring was 1060 m/hour at 19:00, while the maximum average movement rate in the
winter was 900 m/hour at 19:00 (Figure 1-5).
Both time period of the day and season had effects on average hourly movement rate and
cumulative travel distance, there were also interactive effects among time period of the day and
the season (Table 1-7, Table 1-8). In both of the spring and winter, coyote had higher hourly
movement rates at dawn, dusk, and nocturnally, compared to diurnal rates (Figure 1-6). The
average hourly movement rates in each time period of the day in the spring were higher than the
corresponding hourly movement rates in the winter (Table 1-9, Figure 1-7). In both the spring
and winter, coyote traveled the longest distance during the night while distances were relatively
low during dawn, diurnal, and dusk (Figure 1-8). The travel distance in dawn, diurnal, and dusk
in the spring were longer than the corresponding distance in the winter, whereas the nocturnal
travel distances were similar in winter and spring season (Table 1-10, Figure 1-9).
Effects of Temperature and Rainfall on Coyote Activity
During the research period (November 2014 – May 2015) daily mean temperature varied
from 6.65 to 29.72 ℃, and daily mean rainfall varied from 0 to 6.05 cm, and averaged 0.26 cm.
There was no interactive effect of temperature and rainfall on distance travelled (Table 1-11).
However, both mean daily temperature and the presence of the rainfall significantly and
26
negatively impacted daily travel distance when the interaction was removed the model (Table 1-
12). The predictive equation model based on temperature and rainfall to estimate the distance
when there is no rain was:
D = [t×(−0.234) + 117.34]2
The predictive equation model to estimate the daily movement when there is rain should
be:
D = [t×(−0.234) + 114.38)2
Where D is the cumulative daily movement (m/day), t is the daily mean temperature (℃),
and it is squared to adjust back from square root distance. In this model as daily mean
temperature increases by 5 ℃, the daily distance travelled decrease about 270m/day, and when it
rains the daily distance travelled decreases a further 650m/day (Figure 1-10).
There was no interactive effect of temperature, season and rainfall (Table 1-13) and the
relationship of decreased distance travelled with increasing temperature was the same in both
seasons but coyotes travelled less overall in winter (Figure 1-11).
Coyote Interactions
Male adult residents at Buck Island (M1, M5) had exclusive home ranges (Figure 1-2).
There was a small shared area (2.5 km2) between the 95% KUD home ranges of these two male
adult residents, which was about 12.5% of M5 home range (20 km2) and 8% of M1 home range
(31 km2). There was no overlap between the core areas of M1 and M5.
Male transient coyote (M15) had large 95% KUD home range (88 km2) which covered
the home range of one male resident (Figure 1-3). Male transient coyote(M8) had huge 95%
KUD home range (921 km2) which covered the home ranges of the two female residents (Figure
27
1-4). However, the core areas of the transients did not overlap with the core areas of any male or
female residents that we had collared.
I only had data from one female adult resident (F9) who shared her home range (95%
KUD) almost completely with a juvenile (F10), except their core use areas (50% KUD) were not
overlapped with only 1.8 km2 shared, which was 18% of adult core area and 35% of juveniles
(Figure 1-4). In the case of two juvenile females they shared their 95% and 50% KUD almost
completely (Figure 1-2).
Collared coyotes presented a variety of interactions influenced by age, gender, and home
ranges (Table 1-14). At the Buck Island site, two female juveniles (F2 & F3) interacted
throughout the whole research period, and were likely from the same litter. From the beginning
of February 2015, the interactions became less frequent, and the average distance between the
two coyotes increased (Figure 1-12). From the beginning of February 2015, F3 began to
frequently leave her home range, travel into the home range of M1. From 2/21/2015 to
5/22/2015, there were 20 days that F3 traveled into the boundary area of M1 home range. Most
of the travel ended up by F3 returning to her home range on the same day. During 3/13/2015-
3/14/2015, F3 traveled cross the home range of M1 to the northeast part of the ranch, and
returned to her own home range on 3/15/2015 (Figure 1-13). Although F3 frequently traveled
into the home range of M1, there was only one time inferred interaction between these two
coyotes. On 3/10/2015, the closest distance between F3 and M1 was 124 m, which happened in
the boundary area between their home ranges around 1:30 am (Figure 1-14). Another female
juvenile (F2) rarely left her home range. During 4/17 - 4/18/2015, F2 left her home range, and
travelled several kilometers to the west, then to the north, and eventually entered the home range
of the male coyote M5 (Figure 1-15). However, she went back to her home range right after this
28
travel. There was no interaction between F2 and M5; the closest distance between them was 4
km.
At the Blackbeard’s site, a female adult and a female juvenile (F9 & F10) interacted
throughout the whole research period, without a shift in the frequency or average distance
between the two coyotes (Figure 1-16). These two females both encountered transient male adult
(M8) on 12/22/2014, female juvenile (F10) at 4:30 and female adult (F9) at 5:00. This happened
by F9 traveled into the home ranges of F10 and F9, then dispersed and had no further
interactions with these females (Figure 1-17).
At the Lightsey site, transient M15 had a large home range which covered more than 85%
of the home range of resident M13 (Figure 1-3). From 12/14-2014 to 4/12/2015, there were at
least 30 days that M15 visited the home range of M13. However, there were only two instances
of interactions between M15 and M13. On 12/15/2014, the interaction began at 08:00, and
interestingly the distance maintained between these coyotes was only several hundred meters
until 18:00 when they separated (Figure 1-18). On 4/5/2015, these two males again interacted for
hours and the closest distance between them was 95 meters at 5:00 before they completely
separated (Figure 1-19).
Discussion
Sizes of Sub-Tropical Coyotes
In the subtropical environment of central south Florida, the average male coyote weight
(all adults) was heavier than average female coyote weight (two adults, six juveniles). This
suggested difference was likely because of the juvenile females. Compared to a separate study in
central south Florida in mostly native habitat (Thornton et al., 2004) which reported male
weights of 14.2 kg (95% CI = 13.42 – 14.98 kg, n = 3, adults) and female weights as 13.0 kg
29
(95% CI = 12.41 – 13.59 kg, n = 4, four adults, one juvenile), the adult coyotes from my research
are approximately 2 kg heavier. This 15% greater coyote weight could be due to a number of
reasons, such as small sample size. The coyotes in my research were captured and weighed in
November and December of 2014, although not described exactly in Thornton et al., (2014) it is
suspected that coyote weight data was collected during April and May before VHF telemetry was
began. In sub-tropical Florida April and May is the end of the dry season and food resources may
be limited compared to later in the year when I captured coyote. There is evidence for changes in
male and female coyote weights increasing from summer to winter and decreasing from winter to
summer, with suggested reasons that coyote have increased body weight in the winter to
withstand cold temperature, deficiency of food, or to provide the additional energy for the
breeding in the spring (Poulle et al., 1995). Another, more intriguing hypothesis is that the
habitats and resources provided in the predominantly improved grazing landscape in my research
allowed coyotes to be in better condition and grow larger.
The body weights of coyote in Florida are heavier than in many regions of United State
of America. The heaviest coyotes were in northeastern region, average weights of coyote were
16.4 kg and 14.7 kg for male and female according to the summarized data from several studies
(Way, 2007). In California, average weights of coyote were 10.9 kg and 9.8 kg for male and
female, respectively (Hawthorne, 1971). In Texas, average weights of coyote were 12.6 kg (n =
46) and 10.5 kg (n = 38) for male and female, respectively (Young & Jackson, 1951). According
to Bergmann’s Rule, warm-blood vertebrates from cooler climates tend to be larger than same
species living in warm climates. Although, the northeastern coyotes have the heaviest weights
comparing with other regions in the United States of America, Longitude was significantly more
correlated in body weights than latitude. 62% variation of the male weights (p < 0.0001) and
30
59% variation of the female weights (p < 0.0001) can be explained by longitude, while only 13%
variation of the male weights (p = 0.043) and female (p = 0.044) could be explained by latitude
(Way, 2007). The small size of Californian and Texan coyotes may be because of dry, hot arid
climates with scarce of resources. The richer forests of the eastern US could be providing greater
and more regularly resources for coyote allowing for increased body size in the populations. The
larger body size of eastern coyote may also be contributed to increased hybridization with the
timber wolf (Canis lupus) and red wolf (Canis rufus), genetic selection to adapt to bigger prey,
and phenotypic increased in body size because of the greater food supply (Larivière & Crête,
1993; Thurber & Peterson, 1991). Although the coyote hybridization with wolf is still undefined
in Florida, the large population of white-tailed deer and year-round cattle industry in central
south Florida may provide a greater amount of food for coyote resulting in heavier weight of
coyotes in eastern US and Florida.
Home Range Sizes of Sub-Tropical Coyotes
The home range size of coyotes estimated under 95% fixed Kernel were relatively
consistent with the home range size estimated under 95% MCP, however, the core areas of
coyotes estimated under 50% fixed Kernel were not always consistent with the core areas
estimated under 50% MCP (Table 1-3). In general, Kernel estimate provides a less biased home
range estimation result than MCP (Swihart & Slade, 1997). However, when the sample size is
large enough, the difference between the results from different estimate methods is not obvious
(Boyle et al., 2009). In this research, there were thousands of location points for each coyote
(Table 1-2), the large sample (under 95%) diminished the bias and resulted in fairly consistent
home range sizes under both methods (Table 1-3). The average home range size for residents
(mean = 27.14 km2, median = 27 km2, 95% CI = 19.00 – 35.27 km2; n = 7) was consistent with
31
Thornton et al (2004) average yearly home range size (mean = 24.8 km2, 95% CI = 16.96 –
32.64 km2; n = 7), both carried out under 95% fixed KUD. The relative consistency between
studies and similarity among genders (from my study) suggest that a breeding pair of coyotes if
they are defending territory together in sub-tropical Florida require approximately 25 - 30 km2 on
average.
The resident coyotes in Florida have larger home range than the coyotes distributed in
southwestern states at the similar latitudes, such as California and Arizona, but have similar
home range size to the resident coyotes distributed in the southeastern states, such as South
Carolina (Table 1-1, Figure 1-20). The estimation method, sample size of the studies, length of
study, season of study, reproductive status, prey availability, and vegetation composition can all
affect the size and shape of coyote home range. Due to the complexity of these variables, there
are considerable difficulties and limited significance in comparing coyote home ranges from
different studies carried out in different environments, other than gross generalities. Therefore, I
have focused on comparison between gender, season, resident and transient within my study.
After removing transients, I found no significant differences in resident home range sizes
(95% or 50% KUD) between genders or seasons (Figure 1-21). However, in the spring, the
difference between female and male home range under 95% KUD was marginally significant (U
= 1, p = 0.11). The average home range size of females was tending to increase from the winter
to the spring (U = 1.5, p = 0.08) and the majority were juvenile (three out of the four). Two of
the female juveniles increased home ranges in the spring, inflating the overall average female
home range (Table 1-4). These two juveniles interacted extensively and were likely siblings.
Juvenile coyotes may have increased home range before dispersing (Harrison et al., 1991), and
the dispersal could be the result of the increased aggressiveness from other juveniles in the same
32
litter as they grow older (Bekoff, 1977). These two coyotes had increasing average distance
between each other from the winter to the spring, which may have been caused by increased
aggressiveness, as well as exploratory behavior as they begin to disperse from their natal home
range and search for available mates. In California, a study found that 1st year female coyotes
started to enter proestrus in November, and that 50% of the 1st year female coyotes were in
proestrus by February (Sacks, 2005). During the timeframe of my research it is likely that
juveniles were entering or in proestrus during and searching mates. I show in the interaction
analysis that one female juvenile frequently left her home ranges and traveled into one neighbor
male resident home range. (Figure 1-13). This female may also have travelled across other
coyote home ranges and had other interactions between her natal territory and the male resident
visited. This exploratory behavior of visiting nearby territories may explain why home range
sizes of female juveniles are possibly increasing during the spring season.
Movement Rates and Circadian Rhythm
In the sub-tropical environment of Florida, coyotes have high average hourly movement
rates during dawn, dusk, and nocturnally, compared to relatively low rates in the day (Table 1-7,
Figure 1-7). The dawn and the dusk periods by default are only 2 hours in length, while and the
daytime is about 10 hours, this resulted in the similar cumulative travel distances among dawn,
dusk, and day, showing that there are some periods during the day when considerable
movements are made. The sum distance travelled at night is approximately the same as sum
distances travelled during the dawn, day, and dusk (Table 1-8, Figure 1-8). The crepuscular and
nocturnal movement rates in this study area are consistent with another Florida study that
indicated coyote movement rates were 600 m/h crepuscularly and 630 m/h nocturnally but
diurnal movements from my study were only 100 m/h on average compared to 300 m/h
33
(Thornton et al., 2004). Coyotes present similar circadian activity patterns as reported from many
studies with higher rates at dusk, dawn, and night compared to the daytime (Kitchen et al., 2000;
Hidalgo-Mihart et al., 2009; Arias-Del Razo et al., 2011). The coyotes in Florida were relatively
less active during the day time. The avoidance of the diurnal movement could be explained by
very open landscape with a small amount of shelter, combined with exposure to the strong
tropical sun and high temperatures in Florida rangelands. Another reason could be coyote
avoided getting shot by local hunters. The lower diurnal activity may also be due to difference in
the land use, predominantly being agricultural and an increased amount of human activity
compared to natural lands, as coyote activity can be strongly affected by humans. Kitchen et al.
(2000) indicated that coyotes are less active during the day and more active at night but 8 years
later in the same area when human activity was less coyote had increased their daytime activity.
At my research sites there was recurrent horseback riding for cattle working, driving of trucks
and tractors, sod cutting, and orange picking activity by humans. Coyote may decrease activity
during the day to avoid encounters with humans that in Florida ranchlands are also known to
have hunted them when seen.
In most coyote circadian activity research, periods of interest were either divided into
biological seasons such as breeding, gestation, pup rearing, and juvenile independence (Servin et
al., 2003), or divided into climatic seasons such as spring, summer, fall, and winter (Gese et al.,
1988). In Florida, the biological seasons of coyotes are not well ascertained, so I divided my
research period into winter and spring. These two periods probably covered breeding/gestation
and the beginning of whelping and pup rearing in Florida, with breeding and gestation occurring
in the winter period and pup rearing more likely across spring (Main et al., 1999). The hourly
movement rates of coyotes were higher in all time categories dusk, nocturnal, dawn and diurnal
34
in the spring than the movement rates in the corresponding time category in the winter (Table 1-
9). As expected the cumulative travel distances in each time category show similar patterns to
hourly movement rates, in that in the spring coyote traveled longer distances in most time
categories than in the winter, except nocturnally when there was no significant difference (Table
1-10). The non-difference in nocturnal distance travelled can be explained by the changes in
night length with slower hourly movement rates in winter being compensated by length of night
compared to faster hourly movements in spring in a shorter night. It has been suggested that
coyote may spend less time in hunting and more time in resting in winter if carcasses and larger
prey are available, as small mammal, fruits, and invertebrates become less abundant (Bekoff and
Well, 1980; Gese et al., 1996). Reduced hourly activity in winter may also be the most efficient
way to save energy output and improve individual survival and condition during harsher
conditions (Shivik et al.,1997). In the cattle stocked rangelands of Florida, the majority of calves
are born from October through February and access to both carcasses and afterbirth may alter the
way coyote behave in winter. However, compared to many other regions of the US, Florida has
an extremely mild winter and thermal energy costs and reduction in food resources may not be as
taxing as other locations. The opinion that coyote reduce activity in winter in order to save
energy may be valid but further investigation should be carried out on coyote breeding cycles in
Florida. Although there are no adequate studies to describe the breeding seasons of coyote in
Florida, the changes in coyote activity behavior could be related to the reproductive status. It is
likely pro-estrus occurs sometime early in the year as daytime increases and in female coyotes
this period can last 60-90 days (Kennelly, 1972). In many canids there are heightened levels of
activity behavior preceding breeding during pro-estrus, including raised leg urination, ground
scratching, howling and general activity (Thomson, 1992, Bekoff 1979), and if this period
35
coincided mostly with spring months (March 2015 to May 2015), it would explain the increased
activity observed. Coyotes may have been patrolling home range more intensely and increasing
above mentioned behaviors. Understanding the breeding phenology of coyote in Florida will
enhance our knowledge of changes in activity patterns.
Effects of Temperature and Rainfall on Coyote Activity
The general relationship between temperature variation and coyote daily travel distance is
complex because behaviors such as foraging, breeding, and territory patrol can all affect the
distance coyote travel per day. One research study on the movement patterns of coyotes in
Washington found an insignificant correlation between daily mean temperature and daily
movement (Springer, 1982). In Florida I expected high temperatures to reduce activity, and after
controlling for the random effects of location and individual coyote, I show that daily cumulative
travel distance was significantly negatively correlated with daily mean temperature. This result
seems to be contradictory to the conclusions from my seasonal home range and activity analysis
which indicates that coyote have larger home ranges, higher hourly movement rates and longer
cumulative travel distance in the spring, especially when the temperature increases from the
winter to the spring. However, further investigation showed that in both winter and spring
periods activity decreases with temperature but activity is overall higher in spring (Figure 1-1).
I found that rain had an effect on coyote daily distance travelled and this may be because
they don’t like getting wet, as a wet pelt has a significantly reduced insulating quality and
decreases the total thermal resistance of the coat of the mammal (Webb & King, 1984). The
effect of decreasing thermal resistance is more significant when environmental temperature is
low, as has been shown in black-tailed deer (Odocoileus hemionus columbianus; Paker, 1987).
36
Once coyote get wet, the heat loss may increase and coyote may decide to find cover and in turn
reducing daily movement.
Individual Interactions
Resident coyotes defend their territory from intruders from other groups and transients
(Gese, 2001). Therefore, direct interactions among coyotes from different groups are infrequent.
During this study resident coyotes from different groups with exclusive home ranges rarely
encountered each other. However, there were numerous interactions between the female
residents from the same group. There were two pairs of female residents in this study. The two
female juveniles (F2 & F3) at Buck Island could be considered siblings because of their highly
overlapped home ranges, the long-term repeated interactions, and being of the same age. Before
February, these two juveniles were resting and moving together consistently. From February
onwards one female (F3) began to expand her area of use (Figure 1-13), although there was still
frequent interactions, the average distance between the female juveniles increased (Figure 1-12).
A possible reason for the expanded active area and farther average distance between the siblings
could be the pre-dispersal behavior of juveniles, a period of preparation before young coyotes
leave their natal territory. There is a strong tendency for individuals to avoid others and to show
increasing independence before the dispersal (Bekoff, 1977) and to show within territory family
aggression between adults and siblings (Harrison et al., 1991). It is unknown that either F3 was
forced by other sibling to expand her home range to avoid increasingly aggressive interactions or
it could be that F3 was a more dominant coyote and capable of leaving the natal territory.
Unfortunately, we do not have the breeder home range to define territorial boundaries the
juveniles may have been safer within. The reason for the increasing average distance between
these two coyotes could also be exploration in order to find available male coyotes. I
37
documented coyote F3 visiting M1 near her home range boundary and frequently traveled into
M1 home range. At the same time, there was also one time that F2 left her home range and
eventually traveled 7 km to the north into the home range of M5.
Another pair consisted one female juvenile (F10) and one female adult (F9). These two
coyotes also had highly overlapped home ranges and long-term interactions, however the core
area did not have large proportion overlapping. The distances between these two coyotes did not
show trend of increasing through the research period. One potential reason for the long term
interactions could be less pressure from other members in the group, for example no siblings or
the male resident died. It could also be that this juvenile was slow to develop and stayed within
female adult home range as long as possible.
The two transient coyotes had large home ranges with overlap with the resident coyote
home ranges. Both these male transients were recorded interacting with other coyotes (Figure 1-
17, Figure 1-18, Figure 1-19). Usually, transient coyotes are dispersing juvenile and transient’s
movement can be restricted to the area between resident home ranges (Kamler & Gipson, 2000).
However, M15 was already a 2-year old adult when he was captured in 2014 winter. The reason
of the abnormal revisitings toward resident home range by transients can be that the residents
lacked of the ability to defend the territory. Another potential explanation can be the that there is
extraordinarily attractive food resource distributed in resident’s home range. Timm et al. (2004)
discuss coyote visiting areas near suburban-wildland interface. The home range of the male
resident (M13) was half surrounded by low density human community. The two times inferred
interaction between M13 and M15 both happened near human community by the lake. The
transient may have visited several times due to the attraction of the nearby human community
offering a potential food resource to the coyote.
38
Table 1-1. Coyote home range sizes in different studies.
Paper General Adult Juvenile Male Female Resident Transient Method Location Season
Boisjoly et al., 2010 121±7,
n=19
2689±252,
n=4
100%
MCP
Eastern
Quebec,
Canada
Annual
Bowen, 1982 13.7±1.35, n=18 95%
MCP
Alberta,
Canada Annual
Chamberlain et al., 2000 10.01±1,
n=16 24.68±7.48, n=10 95%
ADK Mississippi Spring
Gehrt, 2009 4.95±0.34,
n=84
26.8±2.95,
n=40
95%
MCP Illinois Annual
Gese, 1988 11.3±0.78,
n=56
106.5±6.93,
n=16
95%
MCP Colorado Annual
Grinder & Krausman, 2001 12.6±3.5,
n=13
105.2±37.9,
n=3
95%
MCP Arizona Annual
Grubbs & Krausman, 2009 22.9±4.2, n=9 95%
MCP Arizona Annual
Grubbs & Krausman, 2009 26.8±5.1, n=9 95%
FK Arizona Annual
Harrison & Gilbert,
1985 46.4±1.06, n=7 MCP* Maine
Summe
r
Harrison et al., 1991 11.2±2.9, n=7 MCP* Maine Winter
Jantz, 2001 10.61, n=14 95%
ADK Alabama Annual
Kamler et al.,
2003 12.5±0.4, n=7 95%
MCP Texas Annual
Kamler et al.,
2003 8.9±1.2, n=7 95%
MCP Texas Annual
39
Table 1-1. Continued
Paper General Adult Juvenile Male Female Resident Transient Method Location Season
Riley et al., 2003
4.69±1.
31,
n=28
4.18±1.06,
n=12
6.17±1.59,
n=22 2.84±0.66, n=18 95%
MCP California Annual
Schrecengost,
2007 30.5±8.6, n=18 20.61±5.4, n=16
95%
MCP South Carolina
Schrecengost,
2007 31.85±8.3, n=22 24.24±4.7, n=20 95% FK South Carolina
Thornton et
al., 2004 24.8±4, n=7 95% FK Florida Annual
This research 24±5.6
4, n=4
31.33±6.4
4, n=3
20.33±6.0
6, n=3
32.25±4.6
4, n=4
27.14±4.
15, n=7
504±416.
5, n=2 95% FK Florida
Winter
&
Spring
Note: MCP= Minimun convex polygon.
ADK=Adaptive kernel utilization.
FK= Fixed kernel utilization.
KUD= Kernel utilization distribution.
*= Proportion of points not descripted
n= Sample size
Unit: X ± SE km²
40
Table 1-2. Coyote capture records.
Location
Capture
date
Ag
e
Weight
(kg)
Length
(cm)
Gend
er Description Data start Data end
Fix
Num
1
Buck Island
Ranch
11/11/201
4 3+ 17.80 98 M
Collar dropped and
recovered
11/12/20
14
5/12/201
5 7,587
2
Buck Island
Ranch
11/11/201
4 <1 11.20 87 F
Collar dropped and
recovered
11/12/20
14
5/12/201
5 7,235
3
Buck Island
Ranch
11/22/201
4 <1 11.40 87 F
Collar dropped and
recovered
11/23/20
14
5/23/201
5 7,411
4
Buck Island
Ranch
11/23/201
4 <1 11.80 86 F Missing 0
5
Buck Island
Ranch
11/24/201
4 2.5 15.10 97 M
Collar dropped and
recovered
11/25/20
14
5/25/201
5 7,538
6
Blackbeard
Ranch 12/4/2014 1.5 16.10 105 F Missing 0
7
Blackbeard
Ranch 12/4/2014 <1 12.40 89 F
Coyote shot, excluded
from analyses
12/5/201
4
12/23/20
14 763
8
Blackbeard
Ranch 12/5/2014 1+ 14.40 91 M
Coyote shot, collar
recovered
12/6/201
5
5/11/201
5 6,391
9
Blackbeard
Ranch 12/5/2014 3+ 15.30 101 F
Collar slipped off, collar
recovered
12/6/201
5 4/1/2015 4,667
1
0
Blackbeard
Ranch 12/6/2014 <1 12.60 99 F
Collar dropped and
recovered
12/7/201
4 6/6/2015 7,326
1
1 Lightsey Ranch
11/11/201
4 1 14.20 98 M Missing 0
1
2 Lightsey Ranch
11/11/201
4 <1 11.00 85 F
Coyote hit by vehicle,
excluded from analyses
11/12/20
14
11/16/20
14 269
1
3 Lightsey Ranch
11/13/201
4 1+ 18.00 106 M
Collar dropped and
recovered
11/14/20
14
5/14/201
5 7,527
1
4
Blackbeard
Ranch
12/6/2014 5 17.40 106 M Missing 0
1
5 Lightsey Ranch
12/13/201
4 2+ 16.40 101 M
Coyote shot, collar
recovered
12/14/20
14
4/12/201
5 4,953
41
Table 1-3. Coyote overall home range summary.
Note: MCP= Minimum convex polygon.
KUD= Kernel utilization distribution.
Unit: km².
Coyote
ID Location Gender Age
MCP
95%
MCP
50%
KUD
95%
KUD
50% Data start Data end
Data
Fix
Number
1 BuckIsland Male Adult 29 14 31 10 11/12/2014 5/12/2015 7,587
2 BuckIsland Female Juvenile 26 3 23 2 11/12/2014 5/12/2015 7,235
3 BuckIsland Female Juvenile 72 4 44 4 11/23/2014 5/23/2015 7,411
5 BuckIsland Male Adult 15 5 20 1 11/25/2014 5/25/2015 7,538
8 Blackbeard Male Adult 963 523 921 125 12/6/2014 5/11/2015 6,391
9 Blackbeard Female Adult 32 11 35 10 12/6/2014 4/1/2015 4,667
10 Blackbeard Female Juvenile 30 6 27 5 12/7/2014 6/6/2015 7,326
13 Lightsey Male Adult 10 3 10 2 11/14/2014 5/14/2015 7,527
15 Lightsey Male Adult 92 21 88 13 12/14/2014 4/12/2015 4,953
42
Table 1-4. Coyote seasonal home range summary.
Coyote ID Gender Age KUD 95% Winter KUD 50% Winter KUD 95% Spring KUD 50% Spring
1 Male Adult 31.65 10.08 30.94 7.79
2 Female Juvenile 14.09 1.14 41.69 5.02
3 Female Juvenile 25.55 2.24 58.24 6.18
5 Male Adult 37.22 2.99 10.18 2.05
8 Male Adult 686.28 108.45 439.71 30.47
9 Female Adult 35.27 9.84 34.79 8.93
10 Female Juvenile 26.23 4.84 27.63 4.95
13 Male Adult 8.70 1.87 10.14 2.35
15 Male Adult 61.13 11.13 118.33 19.29
Note: KUD= Kernel utilization distribution.
Unit: km²
43
Table 1-5. Coyote home range season, gender Mann-Whitney U test.
Male 95% Female 95% U p-value
Winter 25.86 25.29 5 0.86
Spring 17.09 40.59 1 0.11
U 3 1.5
p-value 0.66 0.08
Note: KUD= Kernel utilization distribution.
Unit: km²
Table 1-6. Coyote core use area season, gender Mann-Whitney U test.
Male 50% Female 50% U p-value
Winter 4.51 4.98 7 0.86
Spring 4.07 6.27 3 0.37
U 3 5
p-value 0.64 0.46
Note: KUD= Kernel utilization distribution.
Unit: km²
Table 1-7. Hourly movement rate, time period of the day and season Linear Mixed-Effects
Model analysis. Value Std.Error DF t-value p-value
(Intercept) 21.62 0.95 5899 22.77 0.00
Winter-Spring -2.17 0.49 5899 -4.44 0.00
Diurnal-Dawn -10.89 0.52 5899 -20.91 0.00
Dusk-Dawn 5.26 0.52 5899 10.08 0.00
Nocturnal-Dawn 5.37 0.52 5899 10.32 0.00
Table 1-8. Cumulative travel distance, time period of the day and season Linear Mixed-Effects
Model analysis. Value Std.Error DF t-value p-value
(Intercept) 30.48 2.03 5899 15.03 0.00
Winter-Spring -2.99 0.99 5899 -3.01 0.00
Diurnal-Dawn 2.90 1.06 5899 2.73 0.01
Dusk-Dawn 7.51 1.06 5899 7.06 0.00
Nocturnal-Dawn 51.57 1.06 5899 48.65 0.00
44
Table 1-9. Hourly movement rate, time period of the day and season Tukey Honest Significant
Difference analysis.
Estimate Std. Error
z
value Pr(>|z|) Significant
Winter.Dawn - Spring.Dawn -2.17 0.49 -4.44 < 0.001 ***
Spring.Diurnal - Spring.Dawn -10.89 0.52 -20.91 < 0.001 ***
Winter.Diurnal - Spring.Dawn -12.66 0.49 -25.99 < 0.001 ***
Spring.Dusk - Spring.Dawn 5.26 0.52 10.08 < 0.001 ***
Winter.Dusk - Spring.Dawn 1.87 0.49 3.83 0.00 **
Spring.Nocturnal - Spring.Dawn 5.37 0.52 10.33 < 0.001 ***
Winter.Nocturnal - Spring.Dawn 2.77 0.49 5.69 < 0.001 ***
Spring.Diurnal - Winter.Dawn -8.72 0.49 -17.91 < 0.001 ***
Winter.Diurnal - Winter.Dawn -10.50 0.45 -23.42 < 0.001 ***
Spring.Dusk - Winter.Dawn 7.43 0.49 15.21 < 0.001 ***
Winter.Dusk - Winter.Dawn 4.03 0.45 8.99 < 0.001 ***
Spring.Nocturnal - Winter.Dawn 7.53 0.49 15.49 < 0.001 ***
Winter.Nocturnal - Winter.Dawn 4.94 0.45 11.01 < 0.001 ***
Winter.Diurnal - Spring.Diurnal -1.78 0.49 -3.64 0.01 **
Spring.Dusk - Spring.Diurnal 16.15 0.52 30.95 < 0.001 ***
Winter.Dusk - Spring.Diurnal 12.76 0.49 26.16 < 0.001 ***
Spring.Nocturnal - Spring.Diurnal 16.26 0.52 31.26 < 0.001 ***
Winter.Nocturnal - Spring.Diurnal 13.66 0.49 28.04 < 0.001 ***
Spring.Dusk - Winter.Diurnal 17.93 0.49 36.71 < 0.001 ***
Winter.Dusk - Winter.Diurnal 14.53 0.45 32.37 < 0.001 ***
Spring.Nocturnal - Winter.Diurnal 18.03 0.49 37.09 < 0.001 ***
Winter.Nocturnal - Winter.Diurnal 15.44 0.45 34.43 < 0.001 ***
Winter.Dusk - Spring.Dusk -3.39 0.49 -6.94 < 0.001 ***
Spring.Nocturnal - Spring.Dusk 0.11 0.52 0.20 1.00 #
Winter.Nocturnal - Spring.Dusk -2.49 0.49 -5.10 < 0.001 ***
Spring.Nocturnal - Winter.Dusk 3.50 0.49 7.19 < 0.001 ***
Winter.Nocturnal - Winter.Dusk 0.90 0.45 2.01 0.47 #
Winter.Nocturnal - Spring.Nocturnal -2.60 0.49 -5.34 < 0.001 ***
Note: ** means p<0.05 significant
***means p <0.005 significant
# means not significant
45
Table 1-10. Cumulative travel distance, time period of the day and season Tukey Honest
Significant Difference analysis.
Estimate Std. Error
z
value Pr(>|z|) Significant
Winter.Dawn - Spring.Dawn -2.99 0.99 -3.01 0.05 #.
Spring.Diurnal - Spring.Dawn 2.90 1.06 2.74 0.11 #
Winter.Diurnal - Spring.Dawn -4.63 0.99 -4.67 <0.001 ***
Spring.Dusk - Spring.Dawn 7.51 1.06 7.06 <0.001 ***
Winter.Dusk - Spring.Dawn 2.54 0.99 2.55 0.17 #
Spring.Nocturnal - Spring.Dawn 51.57 1.06 48.65 <0.001 ***
Winter.Nocturnal - Spring.Dawn 51.36 0.99 51.72 <0.001 ***
Spring.Diurnal - Winter.Dawn 5.90 0.99 5.94 <0.001 ***
Winter.Diurnal - Winter.Dawn -1.64 0.91 -1.80 0.62 #
Spring.Dusk - Winter.Dawn 10.51 1.00 10.56 <0.001 ***
Winter.Dusk - Winter.Dawn 5.53 0.92 6.04 <0.001 ***
Spring.Nocturnal - Winter.Dawn 54.56 0.99 55.05 <0.001 ***
Winter.Nocturnal - Winter.Dawn 54.35 0.91 59.47 <0.001 ***
Winter.Diurnal - Spring.Diurnal -7.54 0.99 -7.59 <0.001 ***
Spring.Dusk - Spring.Diurnal 4.61 1.06 4.33 <0.001 ***
Winter.Dusk - Spring.Diurnal -0.37 0.99 -0.37 1.00 #
Spring.Nocturnal - Spring.Diurnal 48.67 1.06 45.92 <0.001 ***
Winter.Nocturnal - Spring.Diurnal 48.46 0.99 48.80 <0.001 ***
Spring.Dusk - Winter.Diurnal 12.15 1.00 12.21 <0.001 ***
Winter.Dusk - Winter.Diurnal 7.17 0.92 7.84 <0.001 ***
Spring.Nocturnal - Winter.Diurnal 56.20 0.99 56.71 <0.001 ***
Winter.Nocturnal - Winter.Diurnal 55.99 0.91 61.27 <0.001 ***
Winter.Dusk - Spring.Dusk -4.98 1.00 -4.99 <0.001 ***
Spring.Nocturnal - Spring.Dusk 44.06 1.06 41.48 <0.001 ***
Winter.Nocturnal - Spring.Dusk 43.85 1.00 44.06 <0.001 ***
Spring.Nocturnal - Winter.Dusk 49.03 0.99 49.42 <0.001 ***
Winter.Nocturnal - Winter.Dusk 48.82 0.92 53.36 <0.001 ***
Winter.Nocturnal - Spring.Nocturnal -0.21 0.99 -0.21 1.00 #
Note: ***means p <0.005 significant
# means not significant
46
Table 1-11. Daily mean temperature and rainfall interactive effect on daily cumulative travel
distance Linear Mixed-Effect Model analysis. Value Std.Error DF t-value p-value
(Intercept) 120.29 7.20 1301 16.71 0.00
Temperature -0.12 0.08 1301 -1.52 0.13
Rain 3.75 12.29 1301 0.31 0.76
Temperature:Rain -0.10 0.17 1301 -0.55 0.58
Table 1-12. Daily mean temperature and rainfall main effects on daily cumulative travel
distance Linear Mixed-Effect Model analysis. Value Std.Error DF t-value p-value
(Intercept) 121.50 6.85 1302 17.73 0.00
Temperature -0.13 0.07 1302 -1.94 0.05
Rain -2.96 1.46 1302 -2.03 0.04
Table 1-13. Daily mean temperature, rainfall, and season interactive effect on daily cumulative
travel distance Linear Mixed-Effect Model analysis. Value Std.Error DF t-value p-value
(Intercept) 141.15 15.21 1297 9.28 0.00
Temperature -0.36 0.19 1297 -1.85 0.06
SeasonWinter 5.94 16.49 1297 0.36 0.72
Rain 7.28 48.42 1297 0.15 0.88
Temperature:SeasonWinter -0.24 0.23 1297 -1.01 0.31
Temperature:Rain -0.11 0.63 1297 -0.17 0.87
SeasonWinter:Rain 14.33 51.44 1297 0.28 0.78
Temperature:SeasonWinter:Rain -0.24 0.68 1297 -0.36 0.72
Table 1-14. Records of interactions of coyotes from two different home ranges.
Coyote ID Location Date Time Distance (m)
M1 & F3 BuckIsland 3/10/2015 01:30 124
M8 & F9 Blackbeard 12/22/2014 05:00 105
M8 & F10 Blackbeard 12/22/2014 04:30 137
M13 & M15 Lightsey 12/15/2014 08:00 127
M13 & M15 Lightsey 4/5/2015 05:00 95
47
Figure 1-1. Research sites.
48
Figure 1-2. Buck Island coyote home ranges.
49
Figure 1-3. Lightsey coyote home ranges.
50
Figure 1-4. Blackbeard coyote home ranges.
51
Figure 1-5. Circadian activity of coyote hourly movement in winter and spring.
0
200
400
600
800
1000
1200
1400
1600
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
mo
vem
ent
rate
(m
ean
+ 9
5%
co
nfi
den
ce i
nte
rval
m/h
r)
time (hr)
November 2014 - February 2015 March 2015 - June 2015
52
Figure 1-6. Coyote hourly movement rates in different seasons and time periods. Note: each box presents minimum, lower quartile,
median, upper quartile, maximum from bottom to top.
53
Figure 1-7. Coyote hourly movement rates in different seasons and time periods Tukey Honest Significant Difference. Note: mean
value and 95% confidence interval.
54
Figure 1-8. Coyote cumulative travel distance in different seasons and time periods. Note: each box presents minimum, lower quartile,
median, upper quartile, maximum from bottom to top.
55
Figure 1-9. Coyote cumulative travel distance in different seasons and time periods Tukey Honest Significant Difference. Note: mean
value and 95% confidence interval.
56
Figure 1-10. Predicted temperature and rainfall effect on daily travel distance.
10500
11000
11500
12000
12500
13000
13500
15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33
Dai
ly t
rave
l dis
tan
ce (
m/d
ay)
Temperature (℃)
Rain & Temperature effect on daily movement
no rain rain
57
Figure 1-11. Coyote daily travel distance by temperature and season.
58
Figure 1-12. Instantaneous distance between female juvenile coyote F2 and F3 every 30mins.
0
2000
4000
6000
8000
10000
12000
14000
16000
Dis
tance
bet
wee
n c
oyo
tes
(m)
Time (date)
Distance between F2 & F3
59
Figure 1-13. Coyote F3 travel out of the home range on 3/13/2015.
60
Figure 1-14. Coyote F3 & M1 interaction on 3/10/2015.
61
Figure 1-15. Coyote F2 travel out of the home range on 4/18/2015.
62
Figure 1-16. Instantaneous distance between female juvenile coyote F9 and F10 every 30mins.
0
1000
2000
3000
4000
5000
6000
7000
8000
Dis
tan
ce b
etw
een c
oyo
tes
(m)
Time (date)
Distance between F9 & F10
63
Figure 1-17. Coyote F8 & F9 & F10 interaction on 12/22/2014.
64
Figure 1-18. Coyote M13 & M15 interaction on 12/15/2014.
65
Figure 1-19. Coyote M13 & M15 interaction on 4/5/2015.
66
Figure 1-20. Coyote home ranges in southern states.
0
5
10
15
20
25
30
35
4095%
hom
e ra
nge
(Mea
n +
95%
CI
km
2)
Research site
Florida - Residents under 95% fixed
KUD (2014)
Florida - Residents under 95% fixed
KUD (2004)
California - Adults under 95% MCP
(2003)
Arizona - Residents under 95% MCP
(2001)
South Carolina - Residents under 95%
fixed KUD (2007)
South Carolina - Residents under 95%
fixed MCP (2007)
67
Figure 1-21. Resident coyote home ranges in winter and spring season of Florida.
0
10
20
30
40
50
6095%
hom
e ra
nge
(Mea
n +
95%
CI
km
2 )
Season * Gender
Spring Female
Spring Male
Winter Female
Winter Male
68
CHAPTER 2
COYOTE HABITAT SELECTION, INTERACTION WITH CATTLE
Introduction
Habitat use by coyotes can vary substantially. For example, coyotes have been reported
selecting open grassland in Kansas (Kamler & Gipson, 2000); however coyotes in Mississippi
avoided open habitats (Chamberlain et al., 2000). Diversity in habitat selection shown among
different research studies can be explained by the high behavioral plasticity of coyotes (Bekoff,
1977). Habitat use in coyotes has been shown to be influenced by season, gender, energy
requirements, habitat compositions, food supply, and physiographic characteristics (Bekoff &
Gese, 2003). These factors are not mutually exclusive, but among them changing food resources,
prey distribution and prey density are important factors as these factors will highly influence the
time coyotes spend in particular habitats.. For example, in southwestern Oklahoma, coyote select
savanna habitat because this type of habitat provides not only large prey population but also
great amount of vegetation food as fruits and seeds eaten by coyote (Livaitis et al., 1980).
Another example is that in central west Virginia that coyote select the area with recent timber
harvest, where contains great density of primary (white - tailed deer) and secondary (rodents)
prey of coyotes in this region (Crimmins et al., 2012). Other important factor has effect on
habitat use may be the selection of denning sites during the pup-rearing. Previous studies showed
that coyote preferred habitat with more cover when they were choosing area for dens (Hallet et
al., 1985).
Coyotes are an important predator of sheep, goat, and cattle. Coyotes were responsible
for the loss of 2.3% of the United States sheep population in 1999, and coyotes were blamed for
the deaths of 21,700 goats in Arizona, New Mexico, and Texas in the same year (Mitchell et al.,
2004). However, coyote predation on cattle was only a minor cause of loss to the cattle industry
69
in 2000, killing of less than 0.1% of the United States cattle population (Mitchell et al., 2004).
Most of the studies discussing the interaction between coyotes and livestock were carried out on
coyotes and sheep. One study particularly focused on the coyote predation on sheep indicated
that most of the coyote in the study area did not kill sheep (Sacks et al., 1999). Only the breeding
pairs who had territories overlapped sheep were the primary predators of sheep. At the same
time, the sheep carcass left by the breeding pairs might attract the non-breeding transients to
invade into territory of breeding pairs for scavenging (Sacks et al., 1999). If cattle are an
important food resource for coyotes on ranches in Florida, it could be assumed that coyotes
should spend relatively longer time periods in cattle pastures than the other areas. This may have
an effect on the overall habitat use of coyotes. Therefore, investigating the spatial relationship
between coyotes and cattle becomes an important part of coyote habitat selection analysis.
Research on coyote habitat use in Florida is insufficient. Thornton et al. (2004) indicated
that in south-central Florida, coyotes avoided cutthroat seepage slopes, swamp, and marsh, but
preferred scrub in the wet season. Coyotes only avoided swamp and did not show preference to
any of the other habitats in the dry season. In both the wet and dry seasons, coyotes did not show
any preference or avoidance of the pasture area. However, this research was carried out in an
area with only a small proportion of pasture (3%) compared to all other habitat types, and the
population size of cattle and access of cattle to different habitats was not described. The habitat
use of coyotes in agricultural areas of south-central Florida, especially in areas with large
proportion of pasture and high livestock numbers, is still unknown.
Coyote habitat selection may be affected by the hot weather in the late spring, the change
in land cover to large proportion of improved pasture, and the existence of abundant cattle
distributed on these rangelands in Florida. Due to these factors, I undertook a study to describe
70
coyote habitat selection and the interaction between coyotes and cattle. I specifically examined
coyote habitat selection across gender, age, and residents/transients. I investigated the interaction
between coyotes and cattle by estimating the amount of time coyotes spent in pastures with
cattle, and describe coyote pasture visits to known cattle carcasses.
Methods
Land Cover Map Preparation
I downloaded the Florida Cooperative Land Cover Map version 3.2 (October, 2016) from
the website of Florida Fish and Wildlife Conservation Commission
(http://myfwc.com/research/gis/applications/articles/Cooperative-Land-Cover). I then used
ArcMap (10.1) to redefine the existing map to a more simplified version and reduced number of
land cover categories. The original categories consisted of 56 land cover types (Table 2-1). There
were several reasons to simplify the classifications: 1) Difficulty of distinguishing several similar
land cover types from each other, 2) Incorrectly classified land cover types, 3) The detailed
classification scheme to 56 land cover types was not well represented across the study sites. I
redefined similar categories together according to the Florida land cover classification system
(Kawula, 2009). I also revised mistakes and changes in the original map by comparing with
satellite images obtained from Google Maps (Google Imagery, 2015). I then clipped the map to a
rectangle extent that bounded the 95% KUD home ranges of coyotes at each of the three research
sites, creating three new habitat maps to define habitat use in each area.
Habitat Selection Analysis
I divided the coyote movement data into two research periods as winter (November 2014
- February 2015) and spring (March 2015 - May 2015). I used Manly’s selection ratios with
Type I (first - order level) and Type II (second - order level) study design to infer habitat
71
selection (Manly et al. 2002). For both Type I and Type II, the land cover availabilities were
measured by the proportion of land cover type available within the research site boundaries.
Utilization of land cover types was measured by calculating the proportion of coyote locations
(GPS fixes) within each land cover type defined during simplification of Cooperative Land
Cover Map. For Type I design, I combined all coyote locations together to estimate population
level utilization. For Type II, I classified coyote locations as male-winter, male-spring, female-
winter, female-spring, resident-winter, resident-spring, transient-winter, and transient-spring to
detect if season, gender or territoriality have effects on coyote habitat selection. The selection
ratio of each land cover type was calculated by dividing the utilization of that land cover type by
availability of that land cover type. I used adeHabitatHS package (version 0.3.12) in Rstudio
(version 0.99.893) to conduct both Type I and Type II design analyses. A selection ratio of
greater than 1 infers a greater use of that land cover than availability, and that coyotes preferred
this type of land cover as habitat. A selection ratio of less than 1 suggests coyotes avoided this
type of land cover. If a selection ratio was close to 1, it meant coyotes did not prefer nor avoid
this type of land cover. For Type I design, to test the significant difference among selection
ratios, I used both Pearson’s Chi-Square Test and Log-Likelihood Chi-Square Test. Although
there was usually very small different between the results from two tests, I reported both but only
cared the result from Log-Likelihood Chi-Square Test because it had been adjusted (Manly et al.,
2002). For Type II design, First, a Chi-Square Test was used to test the null hypothesis that all
animals are using the resources in the same way (Khi2L1), no matter there was selection or not.
If the result was significant, it meant coyote individuals are using resources differently. Second,
a Chi-Square Test was used to test the null hypothesis that the utilizations are in proportions to
availabilities (Khi2L2). If the result was significant, it meant there was non-random selection.
72
Third, I used (Khi2L2 - Khi2L1) to test the null hypothesis that coyote were on average using
habitat in proportion to the availability (Manly et al., 2002).
Cattle Movement Record
The MacArthur Agroecology Research Center on Buck Island Ranch has maintained
cattle use and cattle reproductive stage by pasture in a database for the last 10 years. My study
was able to take advantage of this cattle “location” dataset to understand if coyote resource
selection was driven by local cattle presence and/or calving stage. I extracted cattle grazing
records (pasture location) inclusive of November 2014 to June 2015, the duration of my GPS
coyote dataset. The database consisted of pasture names, pastures areas, grazing periods, herd
ID, herd populations, calving information, and other basic information on grazing behavior on
Buck Island Ranch (Table 2-2).
I had four coyote location datasets distributed on this ranch, including two adult males
(M1 and M5) and two juvenile females (F2 and F3). I restricted the coyote location dataset by
those points that only fell within the Buck Island Pasture map using ArcMap 10.1 and a clip
function. I then spatially joined the pasture names to the coyote locations table and created a new
table which included coyote location points and the names of the pasture in which those points
fell in.
I created a subset of cattle grazing information that contained only pastures falling within
the coyote’s home range under 95% KUD. I then used a “merge” function in Rstudio to connect
coyote location data with cattle movement data by the same pasture names, and then used an “if
else” function to select the location points which matched a pasture that had cattle or calves on
the day and time of that coyote location.
73
Coyote Habitat Selection of Pasture with Cattle/Calves
I used a compositional analysis of habitat selection to test if coyote used pastures more
when cattle were present. I classified the pastures as with cattle, with cattle and calves, and
without cattle as three types of land cover. To calculate the selection ratio (𝛾𝑠) of cattle pasture
and calves pasture, I used the equation below:
𝛾𝑠 =𝜇𝑐𝛼𝑐
where 𝜇𝑐 is the utilization of cattle or calves pastures, and 𝛼𝑐 is the availability of
cattle or calves pasture.
To calculate the utilization (𝜇𝑐), I used the equation below:
𝜇𝑐 =𝑁𝑐
𝑁𝑎
where 𝑁𝑐 is the number of coyote location points fall into pasture with cattle or calves,
and 𝑁𝑎 is the number of all coyote location points within the research period.
To calculate the availability (𝛼𝑐), I used the equation below:
𝛼𝑐 =∑ 𝑆𝑖 ∗ 𝐷𝑖𝑛𝑖=1
𝑆𝑎 ∗ 𝐷𝑎
where 𝑆𝑖 is the area of the pasture with cattle or calves, 𝐷𝑖 is the number of the days
that cattle or calves stayed in the pasture, 𝑆𝑎 is the sum of the area of all the pastures within
coyote’s territory, and 𝐷𝑎 is the number of the days of the research period.
I calculated the selection ratios of the pasture without cattle or calves the same way. I
then used a Pearson’s Chi-squared test to infer if the selection ratios were significantly different
from each other (Table 2-3; Table 2-4).
74
Compare Calves Carcass Records with Coyotes Movements
I obtained an incomplete record of calves carcass from Gene Lollis, which contained
recorded calves deaths identified by ranchers. The record included the dates ranchers found the
carcass and the names of the pastures where the carcass was found. Due to the large number of
cattle and calves distributed across Buck Island ranch, not all of calves’ deaths could be recorded
by ranchers. From 11/12/2014 to 5/9/2015, 12 calf carcasses were recorded by ranchers. I plotted
coyote locations on the map and recorded whether or not coyotes visited the pasture in which a
calf death was recorded on the day that rancher the rancher found the calf carcass. If the coyote
spent more than one hour in the pasture at the same position, I considered this visit as a “stay”,
otherwise I considered the visit as a “pass by”.
Results
Simplified Land Cover Map
I reclassified the original 56 types of land cover into the following 10 general categories:
forest, scrub & shrub, dry prairie, improved pasture, wetland, water, agriculture, open area, road,
and human community (Table 2-1). For the Lightsey ranch site, I removed Lake Kissimmee and
Tiger Lake from the land cover map as these deep lakes are never traversable by coyotes
(Figure2-1, Figure2-2, Figure-2-3).
Overall Habitat Selection Result
At the first-order and second-order level, for all coyotes over the three sites, the
proportions of area of available resource were 25.5% wetland, 16.9% forest, 15.5% improved
pasture, 6% dry prairie, 2.3% water, 2.4% scrub & shrub, 8% human communities, 2.3% open
area, 19.5% agriculture, and 1.5% road (Figure 2-4).
At the first-order level, coyote preferred improved pasture, scrub & shrub, open area, and
forest, avoided wetland, road, dry prairie, water and human community (Table 2-5; Figure 2-5)).
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In the winter season, coyote preferred improved pasture, scrub & shrub, and open area, avoided
wetland, road, dry prairie, water, and human community (Table 2-6; Figure 2-6). In the spring
season, coyote preferred improved pasture, forest, agriculture, avoided open area, wetland, dry
prairie, road, water, and human community (Table 2-7; Figure 2-7).
Habitat Selection by Gender and Territoriality
At the second-order level, gender and season have effect on coyote habitat selection
(Table 2-8). Female coyotes preferred scrub & shrub and forest in both winter and spring
seasons, avoided open area and agriculture land. In addition, female coyote spent more time in
the scrub & shrub habitat in the winter than the spring, but spent more time in the forest habitat
in the spring than the winter. Male coyotes preferred open area and agriculture in both winter and
spring seasons, avoided scrub & shrub and forest. In addition, male coyote spent more time in
the open area in the winter than the spring, but spent more time in the agriculture land in the
spring than the winter. Both female and male preferred improved pasture, avoided wetland, road,
dry prairie, water, and human community (Figure 2-8).
At the second-order level, territoriality and season have effect on coyote habitat selection
(Table 2-9). Residents preferred scrub & shrub and open area in the winter, preferred forest and
agriculture in the spring. Transients preferred open area, forest and prairie, avoided agriculture
and scrub & shrub in the winter; preferred open area and agriculture, avoided scrub & shrub in
the spring. Both resident and transient preferred improved pasture, avoided road, water, and
human community (Figure 2-9).
Cattle/Calves Pasture Selection Ratios
Four coyotes were able to be included in the analyses, including two adult males (M1 and
M5) and two female juveniles (F2 and F3). Coyote M5 was a male adult coyote which occupied
76
territory that mostly consisted of orange groves, but on occasion visited pastures on Buck Island
Ranch. Coyote F2 and Coyote F3 were both female juveniles and at least half of their location
points were in pastures on Buck Island Ranch. Coyote M1, a male adult coyote, showed almost
all of his location points occurring in pastures on Buck Island Ranch. None of the four coyotes
showed preference toward pastures that were actively being grazed or had calving occurring
(Table 2-10; Table 2-11; Table 2-12).
Comparison of Calf Carcass Records with Coyotes Movements
Seven of the twelve calf carcass recorded had the pastures they were in visited by
coyotes.(Table 2-13). Only two of four coyotes visited pastures with a calf carcass, and of the
seven visits I classified five as staying for longer than one hours. The majority of the carcasses
found were in the home range of coyote M1 (Figure 2-10).
Discussion
Coyote Habitat Selection
Generally, coyote prefer open habitat (Kamler & Gipson, 2000). In central West Virginia
coyote preferred clear-cut area and avoided hardwood forest (Crimmins et al., 2012). In eastern
Québec, Canada, Coyote preferred clear-cuts of ages 5-20 years (Boisjoly et al., 2010). However,
in Jalisco, Mexico, coyote preferred not only open area, but also tropical semi-deciduous forest,
tropical deciduous forest, and secondary tropical forests (Hidalgo-Mihart et al., 2006). The
preference toward open area or forest can be influenced by many factors such as food resource,
human disturbance, environment temperature and possibly choice of denning sites. In my second
order analyses this preference to forest was shown to be driven by female coyotes and males did
not prefer forest in spring or winter. The heat and strong sun exposure in Florida can be a
potential explanation of preferring forest. Although coyote can tolerate heat in a dry
77
environment, they pant when the air temperature is higher than 34℃ and can be very heat
stressed sat 40℃ (Golightly et al., 1983). From March 2015 to May 2015, in the daytime, the
average maximum temperature was 31.64℃, it could be inferred that coyotes prefer forest land
cover with dense shade under the severe heat. The preference to forest can also be explained by
denning behavior for the pup raising. In Texas, in a research area consisted of native prairie,
farmland and tall permanent vegetation area, coyote preferred the habitat with tall permanent
vegetation for den sites, which provided the most cover for the natal den sites (Kamler et al.,
2005). Although both male and female take responsibilities during pup-raising, female are more
restricted to the den sites because of nursing responsibility (Harrison & Gilbert 1985).
Across my three ranch sites in south-central Florida with similar environments and land
use cover, at the first-order level, one other semi-open habitat coyotes preferred was scrub &
shrub environment, which accorded with Thornton et al. (2004) results from nearby more natural
site in southcentral Florida. In the Thornton study a higher selection ratio was observed for scrub
in the wet season than the dry season. In my research, the selection ratio was also positive for
scrub & shrub and but was higher in the winter than the spring but both these seasons are within
the dry season. After I investigated coyote locations by gender (second - order level), only the
female coyotes preferred the scrub & shrub habitat, while males avoided. Thornton et al. (2004)
indicated that the preference to the scrub habitats may be explained by a higher density of rodent
in this type of land cover (Franz et al., 1998), but this type of habitat is also in dry areas with
patches of thick cover such as palmetto (Seranora repens) that may also provide denning sites.
At Lightsey ranch at least two dens were known to be in scrublands with thick palmetto as cover
(Cary Lightsey personal communication).
78
In my research sites, there was large proportion (15.5%) land consisted of improved
pasture with intensive grazing behavior. At second-order level, coyotes across gender (male,
female), territoriality (resident, transient) and season (winter, spring) always showed preference
for improved pasture. Although there is less shelter for coyotes and frequent human activities
such as driving agricultural machinery, riding horse, and burning pasture occurring in this type of
land cover, the improved pasture is still the most preferred habitat by coyote. Coyote habitat
selection can be strongly influenced by the availability of the prey (Gosselink et al., 2003). Cattle
proportion was confirmed in coyote stomach (Gipson, 1974; Korschgen, 1957), and coyote have
accounted for the loss of calves directly in several occurrences (Dorrance, 1982; Fitch, 1948).
However, there is no evidence that the strong selection of improved pasture by coyote could be
related to the cattle and calves grazing in the pasture. Besides the predation, coyote also use
cattle as food resource by scavenging, especially in the winter season when the old, weak and ill
cattle die (Kamler et al., 2014).
Cattle/Calves Pasture Selection Ratios, Carcass Visiting Behavior
To address if coyotes were selecting improved pasture for aspects related directly to
cattle or calves, I tested if they spent more time when cattle or cattle with calves were in pastures
compare to pastures when the herds were absent. No coyotes presented high selection ratio for
cattle or calves pasture. This result seems to be contradictory to the conclusion that coyote
selected improved pasture because of the cattle resource in the pasture. However, the size of herd
was not taken into account in the statistical test. In addition, In addition, there was only a sample
size of four coyotes to use in combination with grazing records, and there is evidence to suggest
that predatory behavior among the coyotes differs depending on physical stature. According to
the research of effectiveness of removing breeding coyotes in reducing sheep predation (Blejwas
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et al. 2002), breeding pair of coyotes took most of the responsibilities of predation on sheep.
Some other researches indicated that most of the attacks to ungulate fawns caused by coyotes
involved coyote pairs or groups (Wells et al. 1982, MacConnell-Yount et al. 1978, Hamlin et al.
1979, Truett 1979). The reasons of coyote predation on ungulate were carried out by pairs or
groups could be increased energy demand by breeding coyotes (Blejwas et al. 2002). The
ungulates in my research area included white-tailed deer (Odocoileus viginianus), wild hog (Sus
scrofa), and also domestic cow (Bos taurus). For my research, M1 and M5, which had mutually
exclusive home range, could be speculated as having their mates. Unfortunately, there were no
evidence to confirm the existence of their mates. The only group members could be recognized
were two female juveniles, which had small body sizes, lacking of the abilities to participate in
ungulate hunting.
Another explanation can be coyotes were attracted by other food resource on improved
pasture. Improved pasture was usually designed to have larger vegetation biomass to meet the
particular requirements of using (Guo, 2007). It is not surprising if there were large population of
small mammals such as rodents and rabbits distributed in the improved pasture because the
vegetation can be their food. Thornton et al. (2004) indicated that in central Florida, rodent,
ungulates, and rabbit were the top three most common prey in coyote diet. All species that occur
and access improved pastures on grazing lands in Florida. Because of the number of other prey
items that could also be attracted to and be in high densities on cattle ranches in Florida, it is
difficult to identify the prey items that may be responsible for the increased selection of
improved pasture by coyotes. This type of data is beyond the scope of this study.
Among the four coyotes I captured, M1 had almost whole territory on pastures. This
coyote was also the largest male adult (17.8 kg on 11/11/2014), which gave him an increased
80
physiological ability to prey on cattle or calves. However, this coyote did not presented
preference toward pasture with cattle or calves. Nonetheless, this coyote was involved in all of
the visiting behaviors toward calves carcass pastures (Table 2-13). Unfortunately, there were
only 12 cattle carcass records through-out the research period which were incidentally found and
no consistent carcass searching was undertaken, and the data was insufficient to carry out any
statistical analysis. However, all of the pastures, where carcasses were found in, located within or
overlapped with the home range of M1 (Figure 2-10). The reason of the frequent carcass visiting
phenomenon by this coyote could be that this coyote patrolled its territory frequently through the
research period. For example, on 11/12/2014, M1 patrolled almost his whole home range within
one day, and his home range covered more than half of the pastures on this ranch (Figure 2-11).
The frequent patrol behavior expended the possibility of encountering calves carcass or even
weak calves to prey on, but there was no evidence coyote showing purposeful approaching
behavior toward cattle and calves.
Interestingly juvenile F3 also undertook a visit to a calf carcass pasture, which was on the
same day but at a different time to coyote M1. Although this coyote stayed in the south and
barely crossed the canal to visit the pastures in the north, this event happened to be during its
occasional visiting to the north area. As with all coyotes, both these coyotes who visited pastures
of fresh calf deaths did not show increased selection ratio for calving pastures. There are debates
on how coyote choose their prey. Some people indicate that coyote prey behavior accords with
optimal foraging theory, which means only the primary prey abundance should mostly affect
coyote diets (MacCracken & Hansen, 1987; Prugh, 2005). Some other people indicate that it is
not possible to easily determine if coyote choose prey as predicted by only an optimal foraging
theory or an opportunistic foraging theory (Boutin & Cluff, 1989). However, the data of the diets
81
of coyote in this research area is beyond the scope of my study. Nonetheless, my result would
suggest that constant targeted preying upon calves is not undertaken by coyotes and that they
may be searching widely for prey items and finding or killing calves is an opportunistic event.
82
Table 2-1. Summary of reclassified land cover process.
ID Area (pixel) Reclassified Original
1110 16477 Forest Upland Hardwood Forest
1120 61558 Forest Mesic Hammock
1150 834 Forest Xeric Hammock
1200 3422 Scrub & Shrub High Pine and Scrub
1210 45512 Scrub & Shrub Scrub
1300 3531 Dry Prairie Pine Flatwoods and Dry Prairie
1311 2191608 Forest Mesic Flatwoods
1312 77857 Forest Scrubby Flatwoods
1330 705016 Dry Prairie Dry Prairie
1340 8353 Dry Prairie Palmetto Prairie
1400 354254 Forest Mixed Hardwood-Coniferous
1500 277217 Scrub & Shrub Shrub and Brushland
1800 421 Human Community Cultural - Terrestrial
1821 1306543 Human Community Low Intensity Urban
1822 339543 Human Community High Intensity Urban
1830 483568 Open Area Rural
1840 339722 Road Transportation
1850 3814 Human Community Communication
1860 52966 Human Community Utilities
1870 30961 Human Community Extractive
1880 149 Open Area Bare Soil/Clear Cut
2100 21490 Water Freshwater Non-Forested Wetlands
2110 435730 Wetland Prairies and Bogs
2120 1362536 Wetland Marshes
2121 309635 Wetland Isolated Freshwater Marsh
2123 105582 Wetland Floodplain Marsh
2200 1495952 Wetland Freshwater Forested Wetlands
2210 2477 Forest Cypress/Tupelo
83
Table 2-1. Continued.
ID Area (pixel) Reclassified Original
2211 21195 Forest Cypress
2213 5642 Water Isolated Freshwater Swamp
2215 351 Water Floodplain Swamp
2220 3903 Wetland Other Coniferous Wetlands
2221 74600 Forest Wet Flatwoods
2230 1882 Forest Other Hardwood Wetlands
2231 8808 Wetland Baygall
2232 69295 Forest Hydric Hammock
2300 464 Water Non-vegetated Wetland
2400 430 Water Cultural-Palustrine
3000 8130 Water Lacustrine
3100 69432 Water Natural Lakes and Ponds
3200 217621 Water Cultural - Lacustrine
4000 1002 Water Riverine
4100 16528 Water Natural Rivers and Streams
4200 6889 Water Cultural - Riverine
5000 538 Water Estuarine
5240 1573 Wetland Salt Marsh
5250 35 Water Mangrove Swamp
7000 6705 Agriculture Exotic Plants
18331 2672051 Agriculture Cropland/Pasture
18332 343479 Agriculture Orchards/Groves
18333 38734 Agriculture Tree Plantations
18334 81007 Agriculture Vineyard and Nurseries
18335 53193 Agriculture Other Agriculture
22131 325 Water Dome Swamp
22132 23480 Water Basin Swamp
183313 1188240 Improved Pasture Improved Pasture
1833121 402478 Agriculture Sugarcane
84
Table 2-2. Grazing records example from Bucks Island Ranch.
Pasture Area
(km²) Herd ID
Date Into
Pasture
Date Out of
Pasture
Population
(head)
Period
(day)
South Marsh
East Lykes 3.62 Herd Z 7/3/2014 12/18/2014 142 168
W6 0.8 Herd Y 7/28/2014 12/18/2014 70 143
W3 0.77 Herd I 9/15/2014 11/13/2014 95 59
Tom's Yard 0.25 Herd K 9/15/2014 12/8/2014 35 84
Bull Field
North 2.32 D 10/2/2014 11/7/2014 338 36
Tropical East
South 2.39 G 10/6/2014 11/5/2014 60 30
800 Acres #1 1.58 A 10/7/2014 1/2/2015 60 87
W3 0.82 Herd N 10/13/2014 11/13/2014 106 31
S3 0.5 A 10/17/2014 11/3/2014 357 17
HeMarthria
North 1.33 M 10/17/2014 11/11/2014 499 25
W1 0.84 Herd Q 10/27/2014 12/18/2014 104 52
Ceiling #4 0.38 P 10/28/2014 12/1/2014 132 34
East Marsh
North 5.64 L 10/28/2014 2/19/2015 206 114
800 Acres #1 1.58 A 11/3/2014 1/2/2015 357 60
Griffith Park 2.29 O 11/4/2014 12/19/2014 137 45
Note: showing small proportion data of all the records, including name pasture, area of the pasture, identification of the herd, date
into and date out of pasture, size of herd (Populations) and the number of days in that pasture (period).
85
Table 2-3. Calculation selection ratio of pastures with cattle.
ID
𝑆𝑎 𝐷𝑎 𝑆𝑎 ∗ 𝐷𝑎 𝑆𝑖 ∗ 𝐷𝑖 𝛼𝑐 𝑁𝑐 𝑁𝑎 𝜇𝑐 𝛾𝑠
1 5968 182 1086176 444774 0.409486124 1993 7333 0.271785081 0.663722
2 2707 181 489967 220223 0.449464964 1741 3477 0.500719011 1.114033
3 6721 179 1203059 538621 0.447709547 1606 3828 0.41954023 0.937081
5 1646 177 291342 112274 0.385368399 28 66 0.424242424 1.100875
Note:
𝛾𝑠 is the selection ratio of cattle pasture and calves pasture.
𝜇𝑐 is the utilization of cattle or calve pastures, 𝛼𝑐 is the availability of cattle or calve pasture.
𝑁𝑐 is the number of coyote location points fall into pasture with cattle or calves
𝑁𝑎 is the number of all coyote location points within the research period.
𝛼𝑐 is the availability
𝑆𝑖 is the area of the pasture with cattle or calves, 𝐷𝑖 is the number of the days that cattle or calves stayed in the pasture
86
Table 2-4. Calculation selection ratio of pastures with cattle and calves.
Note:
𝛾𝑠 is the selection ratio of cattle pasture and calves pasture.
𝜇𝑐 is the utilization of cattle or calve pastures, 𝛼𝑐 is the availability of cattle or calve pasture.
𝑁𝑐 is the number of coyote location points fall into pasture with cattle or calves
𝑁𝑎 is the number of all coyote location points within the research period.
𝛼𝑐 is the availability
𝑆𝑖 is the area of the pasture with cattle or calves, 𝐷𝑖 is the number of the days that cattle or calves stayed in the pasture
ID
𝑆𝑎 𝐷𝑎 𝑆𝑎 ∗ 𝐷𝑎 𝑆𝑖 ∗ 𝐷𝑖 𝛼𝑐 𝑁𝑐 𝑁𝑎 𝜇𝑐 𝛾𝑠
1 5968 119 710192 75849 0.106800696 528 4792 0.110183639 1.031675
2 2707 102 276114 52045 0.188490986 66 1034 0.063829787 0.338636
3 6721 130 873730 120488 0.137900724 69 2303 0.029960921 0.217264
5 1646 109 179414 23306 0.129900677 1 52 0.019230769 0.148042
87
Table 2-5. First-order level selection ratio Chi-Square Test.
2 df p-value
Pearson 21726.2 9 <0.01
Log-Likelihood 22921.9 9 <0.01
Table 2-6. First-order level selection ratio Chi-Square Test in winter.
2 df p-value
Pearson 15747.2 9 <0.01
Log-Likelihood 15682.5 9 <0.01
Table 2-7. First-order level selection ratio Chi-Square Test in spring.
Table 2-8. Second-order level selection ratio Chi-Square Test
2 df p-value
Khi2L2MinusL1 22921.9 9 <0.01
Khi2L1 9530.67 27 <0.001
Khi2L2 32452.6 36 <0.01
Note: coyotes were divided into groups of Winter-Male, Winter-Female, Spring-Male, Spring-Female. Test was carried out among
groups.
Table 2-9. Second-order level selection ratio Chi-Square Test by territoriality.
2 df p-value
Khi2L2MinusL1 22921.9 9 <0.01
Khi2L1 4887.96 27 <0.001
Khi2L2 27809.9 36 <0.01
Note: coyotes were divided into groups of Winter-Resident, Winter-Transient, Spring-Resident, Spring-Transient. Test was carried out
among groups.
2 df p-value
Pearson 7157.95 9 <0.001
Log-Likelihood 8196.78 9 <0.001
88
Table 2-10. Selection ratios of pasture with and without cattle.
ID Cattle Pasture No Cattle Pasture
1 0.663722 1.233188
2 1.114033 0.906901
3 0.937081 1.051005
5 1.100875 0.936752
Table 2-11. Selection ratios of pasture with and without cows calving.
ID Calve Pasture No Calve Pasture
1 1.031675 0.996213
2 0.338636 1.153617
3 0.217264 1.125206
5 0.148042 1.127192
Table 2-12. Chi-Square Test of cattle/calve pasture selection ratio.
2 df p-value
Cattle Pasture Test 1.1574 6 0.9789
Calve Pasture Test 2.7466 6 0.8399
89
Table 2-13. Visits of coyote to pastures with incidental calf carcasses found by ranchers on Buck Island Ranch.
Event ID Date Pasture name Coyote ID Activity
1 11/12/2014 Ceiling #4 1 Stay 1.5 hours
2 2/9/2015 800 Acres Leftover 1 Stay 4.5 hours
3 3/9/2015 Tropical East North -- --
4 3/14/2015 North Kuhn 1,3 Stay 2 hours
5 3/16/2015 North Kuhn -- --
6 3/18/2015 North Kuhn 1 Passed by
7 3/26/2015 North Kuhn -- --
8 3/26/2015 Shopfield -- --
9 4/3/2015 South Kuhn -- --
10 4/4/2015 South Kuhn 1 Stay 1 hour
11 4/14/2015 South Kuhn 1 Passed by
12 5/9/2015 North Kuhn 1 Stay 1 hour
90
Figure 2-1. Home range of two male coyotes and two female coyotes on the Buck Island Ranch
showing underlying simplified land cover categories.
91
Figure 2-2. Home ranges of one male coyote and two female coyotes on the BlackBeard Ranch showing underlying simplified land
cover categories.
92
Figure 2-3. Home ranges of two male coyotes on the Lightsey Ranch showing underlying simplified land cover categories.
93
Figure 2-4. First-order level resource availability and utilization proportions.
94
Figure 2-5. First-order level habitat selection ratios for whole study period.
95
Figure 2-6. First-order level habitat selection ratios in winter.
96
Figure 2-7. First-order level habitat selection ratios in spring.
97
Figure 2-8. Second-order level habitat selection ratios by gender and season.
98
Figure 2-9. Second-order level habitat selection ratios by territoriality and season.
99
Figure 2-10. Buck Island Ranch pasture boundary, carcass reported pastures, coyote home ranges
map.
100
Figure 2-11. Buck Island Ranch pasture boundary, carcass reported pasture, coyote home ranges,
and coyote location points on 11/12/2014.
101
CHAPTER 3
CONCLUSION
The sub-tropical climate and agricultural improvement associated with the cattle industry
of south-central Florida impact the expression morphology, and the spatial-temporal behavior of
coyotes. Rangeland coyote in Florida have heavier body weights than coyotes in other states in
southern and western regions of the United States, which is consistent with the opinion that the
body size of coyote can be mostly explained by the longitude rather than the latitude (Way,
2007), but this of course is also related to increased resource abundance as coyote move from
southern and western arid areas to eastern grasslands and forests. The heavier body weight of
coyote in Florida may be a response to a greater food supply such as large population of white-
tailed deer and year round cattle husbandry practices.
The larger body size of coyote in Florida ensures their ability to maintain larger
territories. with the home range size of coyote in Florida being similar to others in the southeast
region of the United States, which are larger than the home range size of coyote in the western
regions of the United States. Juvenile female coyote tended to have a larger home range size in
the spring season than the winter. These increasing home ranges for juvenile female coyotes is
likely due to exploratory behavior as they ramp up their physiologically development to become
reproductively active and are in search of their own territory and mate.
Similar to the coyotes in other states, coyote in Florida are most active in the dawn, dusk,
and nocturnal periods of the day. Although, there is a negative correlation between temperature
and coyote movement, coyote still traveled faster and farther in most times of the day during the
spring season, compared to the winter season. It is not surprising that coyote may stay inactive
when the temperature is high. The relatively higher activity in the spring could be explained by
coyote conserving energy in the non-breeding season, but as spring occurs there is a general
102
increase in activity to find mates, defend territories, mate guard existing mates and possibly
increase food intake for gestations and lactation.
Generally, male coyote in Florida avoided encountering with other males, and seldom
leave their territories, except when they are transients. The two female coyote pairs we observed
from the same group maintained frequent interactions. However, the juvenile siblings began to
alienate each other starting in spring when they were more mature and also begin to travel farther
from their natal territory but this exploration was done independently of each other.
In central-south Florida, large proportions of the land cover consist of seasonal wetlands,
forest, and improved pasture with intense cattle grazing. Female coyote preferred scrub & shrub
area, with a shift to stay longer time in forest in the spring season. The male coyote also reduced
their activity in the open area in the spring. This can be the explained as coyote try to avoid
prolong exposure to the sun in the hot Florida spring season.
As a population all coyotes in this study presented a strong preference towards improved
pasture. The large population of cattle and calves in the improved pasture can be the incentive to
attract coyote to spend more time in this type of land cover. However, coyote did not show
preference toward the pasture with cattle or calves currently grazing, compared to the pasture
without cattle. The strength of these analyses is low as the ability to estimate the effect of cattle
and calves in pasture on coyote behavior with only four coyote collared on the one ranch with
these data was limited. Nevertheless, I did observe patterns of behavior in one male coyote that
had the most access to cattle pastures and often visited and stayed within pastures within his
territory when a new calf carcass was found. Further investigation will need to be carried out
with larger sample sizes of coyotes across more ranchers with accurate and detailed grazing
records coupled with consistent observation for dead cattle and calves.
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BIOGRAPHICAL SKETCH
Ke Zhang received his bachelor’s degree from Beijing Forestry University in China.
During his time in undergraduate, Ke majored in wildlife conservation and reserve management.
In the senior year of the undergraduate, he decided to come to the United States to continue his
study on ecology. He was accepted as a Master of Science degree student by the Wildlife
Ecology and Conservation Department at University of Florida. His research interests are spatial
ecology and evolutionary ecology. After graduating from University of Florida, Ke intends to
revise his thesis and have parts of his research published. He has the willing to apply for PhD
program and continue his study in the US.
