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\begin{document}

\title{The Spatial Ecology of the Timber Rattlesnake (Crotalus horridus) in
southeastern Louisiana}



\author[1]{Timothy Borgardt}%
\author[2]{Kaleb Hill}%
\author[1]{Brian Crother}%
\affil[1]{Southeastern Louisiana University}%
\affil[2]{University of Northern Colorado}%


\vspace{-1em}



  
  \date{May 24, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
The Timber Rattlesnake (Crotalus horridus) has a large distribution
spanning much of the eastern United States. Because temperature, habitat
type, prey composition and abundance, and a variety of other factors may
dictate reptile behavior, populations of conspecific species may exhibit
behavioral differences across latitudinal and elevational gradients.
Using radio telemetry, we tracked 10 adult Timber Rattlesnakes (7 males,
3 females) from May 2016 to June 2017 in southeastern Louisiana to
examine the spatial ecology of male and non-gravid female snakes. Mean
annual and seasonal home ranges of non-gravid female Timber Rattlesnakes
were not statistically different from that of males. Mean seasonal home
range sizes and average distances travelled of both sexes was smallest
in winter, and had a general increasing trend beginning in spring with a
peak in fall. These increases seemed to coincide with the breeding
season, taking place from early July until the end of November.
Comparison of this study with other studies throughout its distribution
could have implications towards future management of conservation for
other southern populations of Timber Rattlesnakes.%
\end{abstract}%



\sloppy


\textbf{Introduction}

The advancement of radio-telemetry for wildlife studies has allowed
researchers to study the natural history of cryptic animals at a higher
resolution than ever before (Ujvari and Korsos 2000). Understanding the
spatial ecology of species and the temporal, biological, and ecological
factors controlling it is vital for implementing conservation strategies
for many species (Greene and Campbell 1992). This type of research is
especially important for temperate pitviper species, such as the Timber
Rattlesnake (\emph{Crotalus horridus} ), especially those that migrate
long distances annually, exhibit communal denning behaviors, and inhabit
areas near human activity centers. Anthropogenic activities causing
habitat fragmentation and structural changes that disrupt foraging and
den sites, migration routes, prey densities, and many other aspects of
species' life cycles and behaviors may be the greatest threat to
pitviper populations (Andrews and Gibbons 2008, Green and Campbell
1992). The Timber Rattlesnake is especially vulnerable to these types of
changes because of its long lifespan, low reproductive output, and
habitat requirements (Beaupre and Douglas 2009).

The movement patterns and home range use of the Timber Rattlesnake have
been extensively studied in central and northern portions of its range,
mostly above the 35\selectlanguage{ngerman}°N latitude (Brown 1982, Brown et al. 1982, Martin
1992, Reinert and Zappalorti 1988, Reinert and Rupert 1999, Sealy 2002,
Waldron et al. 2006b). In northern latitudes, populations of snakes
across multiple taxa migrate away from winter hibernacula during the
active season and return before the onset of the dormant season (Bauder
et al. 2016). This pattern is evident in populations of Timber
Rattlesnake, with males and non-gravid females generally migrating from
their hibernacula in spring and traveling in a looping pattern during
the active season that returns them to the same hibernacula before
temperatures drop below 15 \selectlanguage{english} in the fall or winter (Brown et al. 1982,
Reinert and Rupert 1989, Reinert and Zappalorti 1988).

Many snake species with large geographic ranges appear to exhibit more
pronounced migratory behavior at higher latitudes than lower latitudes
(Carfagno and Weatherhead 2008, Klug et al. 2011, Reed and Douglas 2002,
Rodriguez-Robles 2003). For example, a study on home range sizes of the
Grand Canyon Rattlesnake (\emph{Crotalus viridis abysus} ) found that a
population in Arizona had significantly smaller home range sizes than
northern population\emph{s} in Wyoming. The researchers attributed this
difference to higher prey abundance and suitable microhabitat for
hibernation and thermoregulation in the Arizona study area as compared
to the patchy distribution of prey and suitable microhabitat in Wyoming
(Reed and Douglas 2002). With a large distribution spanning much of the
eastern, northeastern, southeastern, and central United States (Powell
et al. 2016), it is possible that the Timber Rattlesnake exhibits
divergent spatial ecology patterns depending on geographic location.

Determining the extent, scale, and conditions by which movement patterns
and other aspects of a species' natural history differ among populations
is important because data from localized studies are often extrapolated
to represent other populations (Gomez and Gregor 2015). Using
extrapolated data to guide conservation and management practices can
create problems if such data do not represent the population being
managed. Because there are no published studies describing the spatial
ecology of the Timber Rattlesnake in Louisiana, any information
regarding home ranges and movement patterns must be extrapolated from
populations studied in other areas. Our study compares the annual and
seasonal movement patterns and home range sizes of males and non-gravid
females in a population of Timber Rattlesnakes in southeastern
Louisiana. Results from our study will provide insight into differences
or similarities that may exist in the spatial ecology of a population at
the extreme southern extent of the species' range.

\textbf{Materials and Methods}

\textbf{Field site description}

We conducted our study on four tracts of land within a 16
km\textsuperscript{2} area of Orleans Parish, southeastern Louisiana,
which we refer to as: ``Site B'', ``Site H'', ``Site M,'' and ``Site F''
(Fig. 2).

Site B encompasses a 118 ha stretch of second-growth bottomland hardwood
and degraded forest within the Bayou Sauvage National Wildlife Refuge.
This narrow strip of hardwood forest is situated on a natural levee,
causing an elevation gradient from -52 cm to 111 cm as one moves from
the marsh to the forest (Wall and Darwin 1999). It is bordered by a
four-lane highway to the south, a two-lane highway to the east, marsh
and open water to the north, and a gravel road paralleling a large bayou
to the west. The central and northeastern portion of the site is
populated by Chinese Tallow (\emph{Triadica sebifera)} , Live Oak
(\emph{Quercus virginiana)} , Bald Cypress (\emph{Taxodium distichum} ),
and a variety of other wetland tree species. Only a few areas in the
central part of the study site have an abundance of coarse woody debris.
Much of this debris consists of deadfall from Live Oak and Bald Cypress
trees that died following Hurricane Katrina in 2005 (Howard 2012). The
vegetation in the southwestern portion of Site B mostly consists of
emergent vegetation, such as Chinese Tallow, Elderberry (\emph{Sambucus
nigra)} , and Giant Reed (\emph{Arundo donax)} . Saw Palmetto
(\emph{Serenoa ripens} ), a few Live Oak, and Common Hackberry
(\emph{Celtis occidentalis} ) are also present. The central and
northeastern portions are dominated by Possumhaw holly (\emph{Ilex
decidua} ), especially at lower elevations, with patches of live oak at
higher elevations.

Site F is located on the east side of the canal bordering Site B. It is
privately-owned land dominated by weeds and vines, such as Blackberry
(\emph{Rubus sp.} ) and Common Cinquefoil (\emph{Potentilla simplex} ).
The invasive Chinese Tallow is also present. There are a few scattered
Live Oak, Bald Cypress, and Water Oak (\emph{Quercus nigra} ) present,
but most of the habitat at this site would best be described as an open
canopy, unmaintained field. At the time of our study, most of the land
was not being maintained, allowing emergent vegetation to become the
bulk of the plant biomass. This site is bordered by a four-lane highway
to the south, a gravel road and a bayou to the east, a small canal to
the west, and a major interstate to the north.

Site H is a privately-owned, 46 ha tract of land located approximately
2.5 km from Site B. Small transects of the property are managed for
hunting activities and New Orleans electric power infrastructure, but
most of the area is unmaintained forest. The forested area is dominated
by Chinese Tallow and is mostly a closed canopy habitat with large
amounts of deadfall. The site is bordered by a levee system to the
south, urban development to the east, a canal and gravel road to the
west, a larger canal to the northeast, and a gravel road and large body
of shallow water to the northwest. Because it is located far from the
natural levee built by the abandoned Mississippi River distributary, it
is an area of low elevation that is sometimes inundated with water
during times of high precipitation or when the water table is higher
than normal.

Site M is an 82 ha tract of privately-owned land located approximately 3
km from Site B. Only a small portion of the northwestern area of this
site is maintained. It has a few grass trails approximately 5 m in width
that are cut with a bush hog pulled by a large tractor biweekly. The
northern and northwestern boundaries are delineated by a canal
approximately 50 m wide and a moderately-used four-lane road. The
northeastern side of the property abruptly turns into a large, grass
field used for cement and rock storage, and further northeast is a large
canal and major interstate. The southern and southeastern areas are
contiguously forested, with the dominant vegetation consisting of
Chinese Tallow, Holly, and Carolina Ash (\emph{Fraxinus caroliniana} ).
All the forested areas have an abundance of coarse woody debris,
consisting of fallen trees and branches. This site has the highest
elevation and best-drained soils.

\textbf{Snake capture and transmitter implantation}

We found snakes for this study by scouting the study area in the winter
for possible hibernacula and suitable habitat. In March of 2016, eight
snakes (five males and three non-gravid females) were captured on top of
or near three separate hibernacula. These individuals were transported
to Southeastern Louisiana University where Wildlife Materials
International temperature-sensitive transmitters were implanted using
published methods approved by the Southeastern University IACUC (Reinert
and Cundall, 1982, IACUC Protocol \#0035). The first three anterior
rattles were painted a unique color of finger nail polish to facilitate
visual identification. After surgery, each snake was given approximately
one week to recover in a secure enclosure with a hide box and water
bowl. Upon recovery, individuals were released at their original capture
sites. While tracking already implanted snakes, we captured an
additional male snake and a non-gravid female snake in May of 2016.
These snakes were collected, implanted with transmitters, and released
using the methods described above. On June 7, 2016, one of the female
snakes died for unknown reasons. The transmitter from this female was
implanted into a male found within the study area and subsequently
released. Observations of this individual began on June 15, 2016, and
ended on June 4, 2017. In addition, the transmitter signal of one of the
male snakes in the study was lost on December 8, 2016. This individual
was not recovered.

Data collected from the deceased female were not used in any statistical
analyses. Data collected from the lost male were not used for the
statistical analyses of annual movements, but data collected during
monthly intervals with full data sets were used in the statistical
analyses of seasonal behavior. At the end of the study, all snakes with
working transmitters were recaptured, brought back to Southeastern
Louisiana University for transmitter removal and released at capture
sites.

\textbf{Telemetry and data collected}

We tracked snakes using a Wildlife Materials International, Inc.,
receiver with a three element Yagi antenna approximately once per week
for one year, April 2016-May 2017. After locating a snake, we recorded
date, time, and latitude/longitude using a Garmin Oregon 600 GPS unit.
For each observation, we recorded the behavior of the snake as foraging,
resting, moving, hidden, or breeding (if copulation was observed). We
also recorded a basic description of the location (vegetation type,
canopy closure, soil type, habitat type, etc.), weather conditions, and
any relevant behavioral observations.

\textbf{Movement and home range analyses}

We utilized ArcMap 10.2 (ESRI, 2011) to create maps of the study sites
with aerial photography images (1 meter resolution). We imported GPS
data from each snake as separate map layers, and generated home ranges
for each snake using the Home Range Tools (HRT) for ArcGIS 2.0.20
extension (Rodgers et al. 2015). We calculated home ranges using the
kernel density estimate application in HRT. The kernel method is a
non-parametric approach to estimating a Utilization Distribution (UD), a
probability distribution of areas utilized by an animal using the
locations it has previously used. It frees the UD from any parametric
assumptions, making better use of locational data (Worton 1989).

We calculated annual home range estimates for each snake that had nine
months or more data. Prior to analyses, we deleted duplicate GPS points
(if an individual was found at the same location at consecutive
recordings) to avoid skewing results. We generated a polygon
representing the 90\% isopleth and 50\% isopleth for each snake (Fig.
3). For this study, we defined the total home range of an individual by
the area encompassed within the 90\% isopleth, and defined the activity
core, or area most utilized by the individual, as the area encompassed
within the 50\% isopleth. We chose these values because most other
studies use these criteria, allowing for easy comparisons among studies.
We also generated seasonal home ranges for each snake using three-month
intervals representing seasons (December--February, winter; March--May,
spring; June--August, summer; September--November, fall). We only used
data from individuals that were tracked throughout each three-month
interval to calculate mean seasonal home range and activity core sizes.

We also used the HRT extension to generate the distance between
successive relocations for each snake. We used these results and
equation 1 to calculate the average distance moved between successive
relocations in each month. We then calculated the average for each three
month season.

\begin{equation}
\overset{\overline{}}{D}=\frac{\sum_{i=x_{1}}^{x_{n}}d_{i}}{\text{\ n}}\nonumber \\
 \end{equation}

Equation 1: Average distance moved between successive relocations.

\textsuperscript{1} \(\overset{\overline{}}{D}\) is the average distance moved
between successive relocations in a month

\textsuperscript{2} \(d\) is the distance moved between
each successive relocation

\textsuperscript{3} \(n\) is the total number of movements

\textbf{Statistical analyses of home range and movement data}

We used SYSTAT version 13 (Systat Software, San Jose, CA) to analyze
home range and movement data. For comparison of mean annual home range
and activity core sizes of males and non-gravid females we performed
two-sample t-tests

We used analyses of variance with sex, season, and the interaction
between sex and season as independent variables to compare the mean home
range sizes (dependent variable) of male and non-gravid female snakes
among seasons. We also compared the average distance moved between
successive relocations (dependent variable) using ANOVA with sex,
season, and the interaction between sex and season as independent
variables.

We used Kolmogorov-Smirnov one-sample test with the Lilliefors option on
residuals of each data set to test normality and Levene's test on
absolute values of residuals of each data set to test homogeneous
variance and to confirm assumptions of statistical analyses.

\textbf{Results}

\textbf{Annual home range}

We recorded a total of 383 observations for the ten snakes tracked from
May 2016 to June 2017. We found that male snakes had a mean annual home
range size of 26.02 \selectlanguage{ngerman}± 11.53 ha and a mean annual activity core size of
4.784 ± 1.54 ha (Table 2). We found that non-gravid female snakes had a
mean annual home range size of 30.89 ± 12.3 ha and a mean annual
activity core size of 6.90 ± 3.71 ha (Table 2).

Using two-sample t-tests, we found that there were no significant
differences between mean annual home range sizes (N=9;
t\textsubscript{8,0.025} = 0.638, p = 0.541) (Fig. 4) nor activity core
sizes (t\textsubscript{8,0.025} = 1.270; p = 0.240) (Fig. 5) of male and
non-gravid female snakes.

\textbf{Seasonal home range}

The mean home range and activity core sizes of male and non-gravid
female snakes were smallest in the winter (December-February) and
increased in size as the year progressed, with largest home range and
activity core sizes occurring in the fall (September-November) (Table
3)(Fig. 6).

Using ANOVA of the mean home range sizes of males and non-gravid females
among seasons, we found there were significant differences for the main
effects of sex (N=9; F\textsubscript{1,29,0.05} = 8.978, p = 0.006) and
season (F\textsubscript{3,29,0.05} = 28.739, p = 0.001), but no
significant difference for the interaction between sex and season
(F\textsubscript{3,29,0.05}=2.532, p = 0.077). According to \emph{a
posteriori} Bonferroni pairwise comparison of the effect of season on
mean home range size of both sexes of snakes in this population, the
mean fall home range size was significantly larger than all other
seasons (p = 0.001) (Table 4). Using pairwise comparison of the effect
of season and sex on mean home range sizes we found that male snakes had
significantly larger home ranges in fall compared to all other seasons.
We also found that female snakes had larger home range sizes in fall
compared to winter and spring (Table 5, Fig. 6).

We found that mean seasonal home range sizes of males and non-gravid
females were statistically equal and followed a similar pattern of
steady increase from winter to summer, but the home range size of male
snakes increased more sharply than that of female snakes from summer to
fall (Fig. 6). We also concluded that the mean fall home range size of
male snakes (25.30 ± 6.08 ha) was larger than the mean home range size
of female snakes (14.74 ± 2.48 ha), but they did not statistically
differ (p = 0.249) (Table 5).

\textbf{Movement}

We found that, annually, non-gravid females traveled an average of 54.36
± 34.06 m between successive relocations, and males traveled an average
of 62.73 ± 48.43 m (Table 6). Using an ANOVA comparing the effects of
sex, season, and the interaction of the effects of sex and season on the
average distances moved between successive relocations indicated a
significant difference for only the effect of season (N=9; Sex:
F\textsubscript{1,79,0.05}= 1.447, p= 0.233; season:
F\textsubscript{3,79,0.05}= 17.397, p\textless{} 0.001; interaction:
F\textsubscript{3,79,0.05}=0.516, p= 0.672). We concluded that snakes in
this population moved significantly longer distances in summer and fall
than winter and spring (Table 7, Fig. 7) using the \emph{a
posteriori}Bonferroni pairwise comparison of the effect of season on
average distance moved between successive relocations.

We also found that male and female snakes did not statistically differ
in their average distances moved during any season (Table 8). Both male
and female snakes shared the same annual trend, increasing distance
moved between successive relocations from winter to fall (Fig. 8).

\textbf{Discussion}

The spatial ecology of the Timber Rattlesnake has been well studied in
the northeastern and central parts of its distribution, but little is
known of southern populations. Because temperature, habitat type, and
prey composition and abundance dictate many aspects of reptile behavior,
populations of conspecific species may exhibit behavioral differences
across latitudinal and elevational gradients. Waldron et al. (2006a)
indicated that Timber Rattlesnakes within a population in South Carolina
had smaller mean annual home range sizes than conspecific populations in
Virginia and New Jersey, with the New Jersey population having the
largest home range. Furthermore, Mohr (2012) found Timber Rattlesnakes
within a population in northeastern South Carolina moved shorter
distances throughout the active season than populations previously
studied at higher latitudes. Our study shows the same pattern of
decreasing home range size from north to south.

Our analyses found that the annual home range sizes and average
distances moved between successive relocations of males in the current
study were statistically equal to those of non-gravid females (Figs. 6
and 7). Mohr (2012) also found that the average distances moved between
relocations did not differ among sex within a population in the South
Carolina mountains, but most other studies found male Timber
Rattlesnakes had larger home range sizes and moved further distances
than gravid and non-gravid female snakes (Reinert and Rupert 1999,
Reinert and Zappalorti 1988, Rudolph and Burgdorf 1997, Sealy 2002,
Waldron et al. 2006a, Waldron et al, 2006b). It is not clear why home
range sizes of snakes in our study's population are so small, but prey
density, habitat quality, climatic factors, body size, and reproductive
female density have all been shown to affect home range sizes
(Bjorneraas et al. 2011, Carrasco-Harris et al. 2020, Harestad and
Bunnel 1979, Lendrum et al. 2014).

Because sit-and-wait predators, like Timber Rattlesnakes\emph{,} will
remain concealed in a location until a prey item comes within a close
enough proximity to capture it, or until the probability of obtaining a
prey item decreases enough to warrant moving to a new location, they may
rely on high quality foraging habitats to gain energy (Schoener 1971).
Both prey density and suitable microhabitat availability may affect an
area's foraging quality (Beaupre 2008, Clark 2004a, Clark 2004b, Reinert
et al. 2011). Many populations of Timber Rattlesnakes travel long
distances to get from their winter den to summer foraging grounds. The
transient habitat in between the den and foraging grounds may be
suitable for thermoregulation, but not foraging (Brown 1992). Temperate
rattlesnakes, like many other ectotherms, must obtain enough energy from
food intake to satisfy energy requirements before temperatures become
too low for activity. An increase in the amount of time a snake spends
traveling between foraging sites may result in a net decrease in energy
at the end of the active season. If energy requirements are not met, the
snake may not be able to maintain its current lifestyle (Schoener 1971).
During this study, there was an absence of long movements away from
hibernacula in the spring. In fact, snakes were observed foraging within
2 m of known hibernacula. This likely indicates that prey availability
and foraging habitat quality are high enough that snakes in this
population have no need to travel long distances to foraging sites.
Individuals may only have to travel far enough that competition between
conspecifics is low. This could also play a role in the small home range
sizes of snakes in this population.

The study area is mostly a contiguously forested landscape, but there
are two major highways that run through it, bordering most of the study
sites (Fig. 3). There are also small patches of commercial and
residential development within the area. Roads may act as physical
barriers that prevent snakes from crossing them (Andrews and Gibbons
2005) and as genetic barriers that prevent neighboring populations from
exchanging genetic material (Clark et al. 2010). Species hesitant to
cross open spaces, such as large, heavy-bodied snakes, like the Timber
Rattlesnake, are especially susceptible to road barriers (Andrews and
Gibbons 2005).

Every home range within this study was bordered by a paved or gravel
road. Five of the snakes within this study crossed small gravel roads,
but none crossed paved roads throughout the entire study. This could
indicate either, the paved roads were acting as impervious barriers, or
the snakes did not travel across these roads because the habitat, prey
density, or some other variable across the road is of lower quality.
Site B, Site F, and the road adjacent to these sites are located on a
narrow natural levee formed by an abandoned distributary of the
Mississippi River, providing a small area of slightly higher elevation
than the surrounding areas. The land directly across the road is
forested, but abruptly turns to marsh and open water. The habitat across
the road adjacent to Site M has been clear cut or developed, and site H
is bordered by water and commercial development on all sides. These
habitats may not be suitable for Timber Rattlesnakes, so an attempt to
cross the road may not be profitable for snakes in this population.

In our study, we found that seasonal home ranges and movements of male
and female snakes were not statistically different from each other, but
males did have larger average home range sizes and movements than
females in September-November (Fall) (Fig. 6 and 8). These increases in
home range size and movement seemed to coincide with the breeding
season, which took place from early July until the end of November. An
increase in home range size and movements is common for male snakes
searching for receptive females in other populations of the Timber
Rattlesnake. Even though there was an increase in home range size for
males during the breeding season in southeastern Louisiana, it was not
as drastic as found in other populations (Anderson 2015, Lind and
Beaupre 2015, Reinert and Zappalorti 1988, Waldron et al. 2006b). Longer
term studies with larger sample sizes are needed to retest these
results.

Currently, there are many gaps in information regarding geographical
differences in behaviors and life history traits of Timber Rattlesnake
populations. Understanding the causes and effects of these differences
at a local level may help mitigate some of the fallacies of conservation
efforts that exist from extrapolating information from studies of
populations that are spatially distant. This baseline study of the
spatial ecology of a population of Timber Rattlesnakes in southeastern
Louisiana is another step towards a better understanding of this
species.

\textbf{Acknowledgements}

Much gratitude should be given to Dr. Brian Crother, Dr. Jeremiah Doody,
and Dr. Clifford Fontenot for all the guidance given during this
project. Also, thank you to the U.S. Department of Wildlife and
Fisheries in Lacombe, La, for all of their help. This study was
partially funded through the Edward. G. Schlieder Foundation.

\textbf{Disclosure Statement}

The authors of this manuscript have no conflicts of interest.

\textbf{Data Accessibility}

If editors decide that manuscript will be published, data will be
uploaded to one of the approved databases before publication.

\textbf{Protection of Human and Animal Subjects in Research}

All activities performed for this study were done so under IACUC
protocol \#0035 approved by the Southeastern Louisiana University IACUC
on 3/21/2016.

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Table 1. Individual study snake information.\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
Snake ID & Sex & Capture date & Study Site & Days tracked & SVL (cm) &
Total length (cm) & Weight (kg)\tabularnewline
350-1 & M & 4/2/2016 & Site B & 342 & 118 & 137 & 1.369\tabularnewline
350-2 & F & 4/2/16 & Site B & 342 & 126 & 135 & 2.323\tabularnewline
350-3 & M & 4/2/16 & Site B & 349 & 168 & 182 & 3.410\tabularnewline
350-4 & M & 4/10/16 & Site F & 348 & 132 & 144 & 1.708\tabularnewline
350-5 & M & 4/10/16 & Site F & 242 & 114 & 124 & 1.205\tabularnewline
350-7 & M & 4/5/16 & Site F & 351 & 138 & 149 & 1.498\tabularnewline
530-2 & F & 4/8/16 & Site H & 366 & 134 & 142 & 1.558\tabularnewline
530-3 & F & 4/10/16 & Site H & 58 & 120 & 128 & 0.944\tabularnewline
250-1 & F & 5/22/16 & Site M & 352 & 125 & 134 & 2.160\tabularnewline
250-2 & M & 6/15/16 & Site M & 355 & 141 & 150 & 2.205\tabularnewline
350-8 & M & 5/20/16 & Site B & 363 & 138 & 150 & 2.334\tabularnewline
\bottomrule
\end{longtable}

\par\null

Table 2. Average home range and activity core sizes of a population of
Timber rattlesnakes in southeastern Louisiana\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
Home range (ha) 90\% Isopleth & Activity core (ha) 50\% Isopleth & Home
range (ha) 90\% Isopleth & Activity core (ha) 50\%
Isopleth\tabularnewline
\midrule
\endhead
Males & Males & Non-gravid females & Non-gravid females\tabularnewline
26.0 \selectlanguage{ngerman}± 11.5 & 4.8 \selectlanguage{ngerman}± 1.5 & 30.9 \selectlanguage{ngerman}± 12.3 & 6.9 \selectlanguage{ngerman}± 3.7\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

\par\null

Table 3. Seasonal home range and activity center sizes of \emph{Crotalus
horridus} in southeastern Louisiana.\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
Season & Mean home range size of males (ha) (Minimum) \{Maximum\} 90\%
Isopleth & Mean home range size of females (ha) (Minimum) \{Maximum\}
90\% Isopleth & Mean activity center size of males (ha) (Minimum)
\{Maximum\} 50\% Isopleth & Mean activity center size of females (ha)
(Minimum) \{Maximum\} 50\% Isopleth\tabularnewline
\midrule
\endhead
December- February & 0.74 \selectlanguage{ngerman}± 1.08 (0.03)\{2.33\} & 1.40 \selectlanguage{ngerman}± 1.52
(0.32)\{2.47\} & 0.16 \selectlanguage{ngerman}± 0.22 (0.01)\{0.45\} & 0.45 \selectlanguage{ngerman}± 0.34
(0.21)\{0.70\}\tabularnewline
March- May & 8.70 \selectlanguage{ngerman}± 3.50 (3.40)\{12.81\} & 4.70 \selectlanguage{ngerman}± 2.89 (0.48)\{7.41\} &
2.03 \selectlanguage{ngerman}± 1.22 (0.60)\{3.27\} & 1.20\selectlanguage{ngerman}±0.60 (0.46)\{1.68\}\tabularnewline
June- August & 11.76 \selectlanguage{ngerman}± 4.41 (7.01)\{19.55\} & 9.07 \selectlanguage{ngerman}± 5.19
(4.87)\{17.22\} & 3.01 \selectlanguage{ngerman}± 1.14 (1.34)\{4.91\} & 2.25 \selectlanguage{ngerman}± 0.94
(1.35)\{3.22\}\tabularnewline
September- November & 25.30 \selectlanguage{ngerman}± 6.08 (21.83)\{34.36\} & 14.74 \selectlanguage{ngerman}± 2.48
(12.84)\{18.05\} & 6.45 \selectlanguage{ngerman}± 1.50 (4.93)\{8.52\} & 5.75 \selectlanguage{ngerman}± 3.11
(3.21)\{10.25\}\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

\par\null

Table 4: Results of Bonferroni pairwise comparison of the effect of
season on average home range size of male and non-gravid
female\emph{Crotalus horridus} .\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
& Dec--Feb & Mar--May & Jun--Aug\tabularnewline
\midrule
\endhead
Dec--Feb & X & X & X\tabularnewline
Mar--May & 0.031 & X & X\tabularnewline
Jun--Aug & \textless{}0.001 & *0.108 & X\tabularnewline
Sept--Nov & \textless{}0.001 & \textless{}0.001 &
\textless{}0.001\tabularnewline
\bottomrule
\end{longtable}

\textsuperscript{1} Numbers in columns represent p-values of Bonferroni
test

\textsuperscript{2} *=non-significant result

\par\null\par\null

Table 5: Results of Bonferroni pairwise comparison of seasonal home
range sizes of male and non-gravid female \emph{Crotalus horridus} in
southeastern Louisiana.\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
\begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Dec--Feb (F)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Dec--Feb (M)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Mar--May (F)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Mar--May (M)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (F)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (M)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
Sep-- Nov (F)\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Dec--Feb (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Dec--Feb (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.664\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Mar--May (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Mar--May (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.042\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.399\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.040\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.027\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.051\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sep-- Nov (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.004\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.006\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.352\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.634\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sep-- Nov (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.005\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.249\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\textsuperscript{1} Numbers in columns represent p-values of Bonferroni
test

\textsuperscript{2} *=non-significant result

\par\null

Table 6: Seasonal average distance traveled between successive
relocations by male and non-gravid female \emph{Crotalus horridus} in
southeastern Louisiana.\selectlanguage{english}
\begin{longtable}[]{@{}lll@{}}
\toprule
& Average distance traveled between relocation (m) Males & Average
distance traveled between relocation (m) Females\tabularnewline
\midrule
\endhead
Dec-- Feb & 18.56 \selectlanguage{ngerman}± 19.70 & 25.92 \selectlanguage{ngerman}± 27.72\tabularnewline
Mar-- May & 47.36 \selectlanguage{ngerman}± 33.11 & 30.43 \selectlanguage{ngerman}± 15.63\tabularnewline
Jun-- Aug & 95.24 \selectlanguage{ngerman}± 46.06 & 70.91 \selectlanguage{ngerman}± 44.25\tabularnewline
Sept-- Nov & 110.37 \selectlanguage{ngerman}± 69.88 & 100.87 \selectlanguage{ngerman}± 19.03\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

\par\null

Table 7: Results of Bonferroni pairwise comparison of the effect of
season on average distance moved between successive relocations
of\emph{Crotalus horridus} in southeastern Louisiana.\selectlanguage{english}
\begin{longtable}[]{@{}llll@{}}
\toprule
& Dec-- Feb & Mar-- May & Jun-- Aug\tabularnewline
\midrule
\endhead
Dec-- Feb & X & X & X\tabularnewline
Mar-- May & *1.000 & X & X\tabularnewline
Jun-- Aug & \textless{}0.001 & 0.002 & X\tabularnewline
Sept-- Nov & \textless{}0.001 & \textless{}0.001 & *0.441\tabularnewline
\bottomrule
\end{longtable}

\textsuperscript{1} Numbers in columns represent p-values of Bonferroni
test

\textsuperscript{2} *=non-significant result

\par\null

Table 8: Results of Bonferroni pairwise comparison of the effect of
season and sex on the average distances moved between successive
relocations of \emph{Crotalus horridus} in southeastern Louisiana.\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
\begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Dec-Feb (F)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Dec--Feb (M)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Mar--May (F)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Mar--May (M)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Jun-- Aug (F)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Jun-- Aug (M)\strut
\end{minipage} & \begin{minipage}[b]{0.10\columnwidth}\raggedright\strut
Sep-- Nov (F)\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Dec--Feb (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Dec--Feb (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Mar--May (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Mar--May (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.885\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.076\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.697\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Jun-- Aug (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.028\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.010\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.087\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sep-- Nov (F)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.027\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.013\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.096\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
X\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
Sep-- Nov (M)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.002\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textless{}0.001\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0.003\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*0.556\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
*1.000\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\textsuperscript{1} Numbers in columns represent p-values of Bonferroni
test

\textsuperscript{2} *=non-significant result

\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

Figure 1:~\emph{Crotalus horridus} in foraging position.

\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

Figure 2: Map of southeastern Louisiana study sites.

\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\par\null

Figure 3: Polygon representing home range area (90\% isopleth) and
activity core (50\% isopleth) of male \emph{Crotalus horridus} , 350-3.

\par\null\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

Figure 4: Average annual home range size (90\% Isopleth) of male and
non-gravid female \emph{Crotalus horridus} in southeastern Louisiana
(n=9)

\par\null\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

Figure 5: Average annual activity center size (50\% isopleth) of male
and non-gravid females from a population of \emph{Crotalus horridus} in
southeastern Louisiana (n=9)

\par\null\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

Figure 6: Average seasonal home range sizes of male and non-gravid
female \emph{Crotalus horridus} in southeastern Louisiana (n=9).

\par\null\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image7/image7}

\end{center}
\end{figure}

Figure 7: Seasonal average distance travelled between successive
relocations by a population of \emph{Crotalus horridus} in southeastern
Louisiana (n=9).

\par\null\par\null\par\null\par\null\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image8/image8}

\end{center}
\end{figure}

Figure 8: Monthly average distance travelled between successive
relocations males and non-gravid females from a population
of\emph{Crotalus horridus} in southeastern Louisiana (n=9).

\selectlanguage{english}
\FloatBarrier
\end{document}

