home range size and habitat use of the eastern indigo ...21424/datastream... · analysis between...

Home range size and habitat use of the Eastern Indigo Snake

(Drymarchon couperi) at a disturbed agricultural site in south Florida

A Thesis

Presented to

The Faculty of the College of Arts and Sciences

Florida Gulf Coast University

In Partial Fulfillment

of the Requirement for the Degree of

Master of Science

By S. Brent Jackson 2013

APPROVAL SHEET

This thesis is submitted in partial fulfillment of the requirements for the

degree of Master of Science

____________________________

Steven Brent Jackson

____________________________

Edwin M. Everham, III, Ph.D.

Committee Chair

____________________________

David W. Ceilley, M.S.

Committee Member

____________________________

John E. Herman, Ph.D.

Committee Member

The final copy of this thesis has been examined by the signatories, and we find that both the

content and the form meet acceptable presentation standards of scholarly work in the above

mentioned discipline.

III

Acknowledgements

My research would not have been possible without funding from the US Fish and

Wildlife Service and assistance and support from the Florida Fish and Wildlife Conservation

Commission as well as the Orriane Society. I would also like to thank my Committee: Dr.

Edwin M. Everham III, David W. Ceilley, and Dr. John E. Herman. Their advice, insight, and

support made this research a reality. I would also like to thank Dr. Billy Gunnels and Dr.

Marguerite Forest for their assistance and support in analyzing and presenting this research. My

gratitude also goes out to Colleen Clark for her countless hours in the field. Additionally, I

would like to thank the following individuals for their support and assistance in the field: Dr.

Jerry Jackson, Dr. Bette Jackson, Steve Mortallaro, Dana Dettmar, Jeff Talbott, Kory Ross, John

Ferlita, and Christian Lyon.

IV

Abstract

The Eastern Indigo Snake (Drymarchon couperi) is a species that is federally threatened

primarily because of habitat loss and fragmentation. Currently there is a paucity of data relating to

populations in the southern portion of this species range, which are believed to be different from

the northern populations due to climate and habitat factors. The objectives of this study were to

provide baseline data pertaining to home range size, habitat use, seasonal activity patterns, and

refugia use in disturbed habitats in south Florida using radio telemetry.

The field site for this study is the home to the future C-44 reservoir and stormwater

treatment area included in the Central and Southern Florida Project of the Comprehensive

Everglades Restoration Plan. It is an abandoned citrus grove intersected with canals, ditches, and

dirt roads located in western Martin County Florida.

A total of five snakes including four males and one female were sufficiently tracked for

analysis between the dates of January 2012 and March 2013. Total home range size varied from

9.71 - 65.78 ha. Several of the individuals tracked showed a preference for canal habitats

particularly during the winter months. All individuals tracked remained active all year long and

showed no significant difference in activity based on mean meters traveled per day when compared

between seasons. The two male snakes tracked for the longest period of time showed a significant

preference for artificial refugia in cooler temperatures and natural refugia in warmer temperatures.

D. couperi using the C-44 reservoir site demonstrate trends in home range size, habitat use,

seasonal activity patterns, and seasonal refugia preferences that differ from other populations of

this species. These differences highlight the need for conservation biologists to consider ecological

and behavioral differences across the range of a species, and within human-dominated landscapes,

when developing management plans. Understanding the role of disturbed habitats as possible

acceptable habitat for endangered and threatened species is integral to their continued survival.

V

Table of Contents

Page

Acknowledgements III

Abstract IV

Table of Contents V

List of Figures VII

List of Tables VIII

Chapter 1:

Introduction 1

Status 2

Taxonomy 3

Description 3

Range 4

Habitat 5

Refugia 6

Feeding 7

Reproduction 7

Home Range 9

Research Objectives 10

Chapter 2:

Methods 12

Study Site 12

Capture and Transmitter Implantation 15

Radio Tracking 16

Timeline 17

Analysis 17

Objective 1: Home Range 18

Objective 2: Habitat Preferences 19

Objective 3: Activity Patterns 20

Objective 4: Refugia Use 21

Chapter 3:

Results 22





VI

Page

Chapter 4:

Discussion 54





Conclusions 66

Literature Cited 70

Appendix 75

Appendix Table 1 Home Range Comparison Table 75

Appendix Table 2 Raw Data 76

Appendix Table 3 Key to Data 88

VII

List of Figures

Page

2.1 C-44 Study Area Map 12

2.2 C-44 Footprint Map 13

2.3 C-44 Weather with Seasonal Overlay 18

2.4 Canal Habitat Photo 19

2.5 Upland Habitat Photo 19

2.6 Great-Circle Distance Equation 21

2.7 Natural Refugia Photo 22

2.8 Artificial Refugia Photo 22

3.1 C-44 Indigo Snake Ranges 24

3.2 Monty Home Range 25

3.3 Vader Home Range 26

3.4 Dagwood Home Range 27

3.5 Paul Home Range 28

3.6 Nagini Home Range 29

3.7 Monty Winter Habitat Preferences 32

3.8 Monty Summer Habitat Preferences 33

3.9 Vader Winter Habitat Preferences 34

3.10 Vader Summer Habitat Preferences 35

3.11 Paul Winter Habitat Preferences 36

3.12 Paul Summer Habitat Preferences 37

3.13 Dagwood Winter Habitat Preferences 38

3.14 Nagini Winter Habitat Preferences 39

3.15 Nagini Summer Habitat Preferences 40

3.16 Nagini Seasonal Activity 42

3.17 Monty Seasonal Activity 43

3.18 Vader Seasonal Activity 44

3.19 Paul Seasonal Activity 45

3.20 Nagini Monthly Activity 46

3.21 Monty Monthly Activity 47

3.22 Vader Monthly Activity 48

3.23 Paul Monthly Activity 49

3.24 Monty Refugia Preferences 51

3.25 Vader Refugia Preferences 52

3.26 Nagini Refugia Preferences 53

VIII

List of Tables

Page

2.1 Plant Species List 14

3.1 Snake Home Ranges 23

3.2 Habitat Preferences 31

3.3 Average Daily Seasonal Movements 41

3.4 Refugia Type Versus Air Temperature 50

Appendix Table 1 Home Range Comparison Table 75

Appendix Table 2 Raw Data 76

Appendix Table 3 Key to Data 88

1

Chapter 1

Introduction

We currently face a global biodiversity loss that can be attributed primarily to

anthropogenic practices (Landweber and Dobson, 1999). The magnitude of this loss is so great

that many scientists are calling this event the “sixth extinction” (Frankham et al., 2002).

Scientists estimate that we are losing up to 27,000 species a year in tropical forests alone (Myers,

1993). Many scientists believe that secondary extinction events may occur as an indirect result of

a disruption in the food chain (energy flow) that occurs during mass extinction events (Bruno

and Cardinale, 2008). Therefore, it is important to minimize the current threats to Earth’s

biodiversity while concurrently exploring the ecological response of organisms to such threats.

One of the greatest threats to biodiversity is habitat loss (Gibbons et al., 2000). Areas where

habitat loss is the most extreme show high extinction rates and have been identified as

biodiversity “hot spots” (Myers et al., 2000).

Habitat loss is the main reason for decline in more than 85% of threatened species in the

United States and is the primary cause of decline in over 97% of herpetofaunal species in the

United States (Wilcove et al., 1998). Habitat loss, degradation, and fragmentation are also the

driving causes for herpetofaunal loss on a global scale (Gibbons et al., 2000; Hyslop, 2007).

Habitat loss may be of particular importance to reptiles because of their spatial

requirements. This is in part because of their furtive nature, comparatively large home range

sizes, low population densities, low fecundity and the lack of frequent congregation events

(Gibbons et al., 2000). Current evidence suggests that reptile declines constitute a worldwide

crisis (Gibbons et al., 2000).

2

One reptile whose range and population have been affected by habitat loss is the Eastern

Indigo Snake (Drymarchon couperi). In addition to facing habitat lose and fragmentation, this

large predator is challenged by relatively low fecundity, large home range requirements

(USFWS, 2008), association to another listed species, the Gopher Tortoise (Gopherus

polyphemus), for low temperature refugia in the northern part of its distribution (Hyslop, 2007),

and the negative stigma commonly applied to snakes by the general public (Knight, 2008).

D. couperi also represents a species with little known about its spatial and habitat

requirements at the southern end of its distribution, despite being federally listed (Hallam et al.,

1998; Hyslop, 2007). Knowledge of basic life history and home range requirements are

paramount to understanding population trends and the effect habitat loss has on populations and

ultimately to preserving this species.

Status:

D. couperi was listed as federally threatened under the Endangered Species Act on March

3, 1978. Its listing was attributed to habitat modification, collection for the pet trade, and the

gassing of G. polyphemus burrows during rattlesnake roundups (Speake et al., 1981b; USFWS,

1978; 1998). At the time of listing the species range was already believed to be limited to

Georgia and Florida (USFWS, 1978). Presently the species is still listed as threatened and the

species large home range size and wide distribution have complicated the collection of accurate

population data (Hallam et al., 1998; USFWS, 2008). Since the species listing the enforcement

of laws has curtailed detrimental activities such as collection for the pet trade and burrow gassing

(Hyslop, 2007; Lawler, 1977; USFWS, 1978). Despite the lack of reliable population data the

continued loss and degradation of habitat points to the continued decline of this species

(USFWS, 1998).

3

Taxonomy:

Holbrook first described the Eastern Indigo Snake in 1842 as Coluber corais. The type

locality for the first specimen was south of the Altamaha River in Georgia (Holbrook, 1842;

Hyslop, 2007). In 1853 Baird and Girard reassigned its genus to Georgia (Hyslop, 2007;

McCranie, 1980). In 1860 Cope reassigned the Eastern Indigo Snake yet again, this time to the

genus Spilotes (Hyslop, 2007; McCranie, 1980). In 1862 Cope declared the Eastern Indigo

Snake a subspecies of Spilotes corais (Hyslop, 2007; McCranie, 1980). Cope reassigned the

Eastern Indigo Snake a third time in 1900, this time to the genus Compsosoma (Hyslop, 2007;

McCranie, 1980). The current genus Drymarchon was assigned by Steineger and Barbour in

1917 (Hyslop, 2007; McCranie, 1980). Until recently Drymarchon was believed to be

monotypic through its range with multiple subspecies. The two subspecies found in the

southeastern United States included the Eastern Indigo Snake (Drymarchon corais couperi) and

the Texas Indigo Snake (Drymarchon corais erbennus) (Hyslop, 2007). In 1991 Collins

suggested that the Eastern Indigo Snake be raised to full species status because of geographic

separation and consistent differences in head scalation between it and the Texas Indigo Snake

(Collins, 1991). In 2001 the Society for the Study of Amphibians and Reptiles provisionally

accepted the raising of the Eastern Indigo Snake (Drymarchon couperi) to full species status

(Crother et al., 2001). Currently the genus Drymarchon contains five full species as well as five

subspecies (Wuster et al., 2001).

Description:

D. couperi is the longest snake in North America reaching lengths of up to 2.6 m

(Conant and Collins, 1998). Adult snakes range in size from 1.9 to 2.6 m in length

(Stevenson et al., 2009). They are uniformly black with a bluish sheen (Layne and Steiner,

4

1996). Additionally they have a variable reddish or cream coloration along their gular region

(Conant and Collins, 1998), though this coloring varies in extent and pigment. The scales of

this species are large and smooth with the exception of males who exhibit slight keeling on 1 to

5 middorsal scale rows (Layne and Steiner, 1996). D. couperi can be differentiated from the

Texas Indigo Snake (D. corais erebennus) by the lack of contact between the antepenultimate

supralabial scale and the temporal or postocular scales (Hyslop, 2007; Wright, 1957). Juvenile

D. couperi are similar in appearance to adults although some individuals display a blotched

dorsal pattern and more extensive ventral coloration (White and Garrott, 1990).

Range:

The genus Drymarchon is primarily a tropical genus that can be found from the

southeastern United States as far south as northern Argentina. The historic range of D. couperi

stretched from the coastal plains of Georgia, Alabama, and Mississippi, south to Florida (Carr,

1940; Diemer and Speake, 1983; Haltom, 1931; Moler, 1985a; USFWS, 2008). It is possible

that their range stretched as far north as southern South Carolina and as far west as southern

Louisiana (Hyslop, 2007; Smith, 1941). However, these additions to the species historical

range cannot be confirmed (USFWS, 1998). The current range of the species has been reduced

to Florida and the coastal plains of Georgia (Lawler, 1977; USFWS, 2008). The species is

now considered rare but it can still be found throughout peninsular Florida (Moler, 1985a;

USFWS, 2008). In the panhandle of Florida D. couperi is believed to persist in lower numbers

then the rest of Florida (USFWS, 2008). In south Florida, they are assumed to be concentrated

primarily in upland habitats. Their numbers are believed to be far less in wetland habitats such

as the Everglades (Duellman and Schwartz, 1958; Steiner et al., 1983).

5

Habitat:

The southeastern Coastal Plain is the herpetofaunal center of the United States

because it contains the highest diversity of reptile and amphibian species (Gibbons et al.,

1997; Trani, 2002). One of the reasons for this high diversity is due to unique habitats such

as longleaf pine forests (Landers et al., 1995). Longleaf pine forests represent areas of high

diversity supporting many organisms specifically adapted to the frequent disturbances such

as fire, and tropical storms (Landers et al., 1995). Many herpetofaunal species are endemic

to this habitat (Trani, 2002). Longleaf pine forests are home to 89 snake species including

three of the largest snake species in North America and seven species that are found nowhere

else (Trani, 2002). Of the original 30 million hectares of southeastern longleaf pine less than

1.2 million hectares of isolated fragments remain (Landers et al., 1995). This is less than

2% of its original extent. D. couperi remain important predators in this fragmented

ecosystem and the majority of research has focused on populations found within these

landscapes. However, this habitat does not represent the only habitat utilized by this species

(USFWS, 2008). Because of this populations that use different habitats are of particular

interest.

While D. couperi shows a preference for dry areas adjacent to water (Ernst and

Barbour, 1989), its range is large enough that climatic differences likely allow for the use of

different habitats depending on the snake’s geographic location. In Georgia D. couperi is

frequently found in sandhill habitats that also support populations of G. polyphemus (Speake et

al., 1978; Diemer and Speake, 1983). These sandhill communities are dominated by longleaf

pine-turkey oak forests (Hyslop, 2007; Wharton, 1977). These habitats are often used during

the winter months when burrow refugia is necessary in order to survive the cooler climate. In

6

summer months it is believed that snakes in the northern reaches of the geographic range move

away from the sandhills and occupy more hydric habitats such as river bottoms and wetlands

(Diemer and Speake, 1983; Lawler, 1977; Speake et al., 1981a).

Much of the species current range is located in Florida (Breininger, 2004). D. couperi

habitat use in Florida has historically been more varied than Georgia with snakes using habitats

such as mangrove swamps, wet prairies, xeric pinelands, hydric hammocks, citrus groves, and

scrub (Humphrey et al., 1992; Lawler, 1977). In the southernmost reaches of this species’

range tropical hardwood hammocks and pine uplands appear to be preferred, however

freshwater marshes, fallow fields, coastal prairie, mangrove swamps, and human impacted

habitats such as residential areas are also used to a lesser degree (Hyslop, 2007; Steiner et al.,

1983). These snakes are likely more habitat generalists than more northern individuals

because of climatic differences (USFWS, 2008). In south Florida D. couperi uses canal banks

where crab holes are commonly used as refugia in lieu of G. polyphemus burrows (Lawler,

1977; Speake et al., 1981a).

Refugia:

Throughout most of their range D. couperi require refugia for protection from extreme

temperatures, desiccation, and predators. They also serve as nesting sites (Holbrook, 1842;

Hyslop, 2007; Landers and Speake, 1980; Speake et al., 1981b; Speake et al., 1978). Studies

focused on water-loss in this species show that they are sensitive to desiccation and exposure to

direct sunlight with exposure times as short as ten minutes being fatal (Bogert et al., 1947; Ernst

and Barbour, 1989). The burrows created by G. polyphemus have been reported as particularly

important in the northern extent of the species range where they are heavily used in the winter

(USFWS, 2008). Additional refugia such as mammal burrows, stumps, logs, and debris piles are

7

also used for shelter (Lawler, 1977; Speake et al., 1978). Refugia use in the southern extent of

the species range has been considered less important because of the warmer climate (USFWS,

2008). D. couperi’s use of refugia in high temperatures has not been investigated.

Feeding:

D. couperi is characterized as a wide-ranging forager (Humphrey et al., 1992; Hyslop,

2007; Landers and Speake, 1980; Stevenson et al. 2010). It subdues prey items by chasing them

down and then immobilizing them with its powerful jaws (Keegan, 1944; Moulis, 1976;

Stevenson et al., 2010). D. couperi is not a constrictor (Stevenson et al., 2010). A study focused

on prey base of the Eastern Indigos determined that there were four major prey types utilized

throughout the snakes range. These prey items include anurans, G. polyphemus hatchlings,

snakes, and small mammals (Stevenson et al., 2010). In addition to hunting, D. couperi has been

documented scavenging carrion on multiple occasions (Stevenson et al., 2010). D. couperi is

capable of consuming large prey items such as Eastern Diamondback Rattlesnakes (Crotalis

adamanteus) as large as 1000 mm TL (Total Length) as well as adult Hispid Cotton Rats (Sigmodon

hispidus) (Stevenson et al., 2010). There is little known about the diet of hatchling and juvenile D.

couperi (Stevenson et al., 2010), but they are known to consume invertebrates as well as prey items

similar to adults (Layne and Steiner, 1996; Stevenson et al., 2010). The diverse diet and high vagility

of this species allow it to forage successfully in numerous habitats (Breininger et al., 2004; Hyslop,

2007; Speake et al., 1978; Stevenson et al., 2010).

Reproduction:

Gillingham and Chambers (1980) described courtship behavior. They broke the behavior

into three phases. During Phase I the male pursues the female and begins courtship. During Phase II

alignment and tail searching occur in preparation for intromission. During Phase III intromission

8

occurs. The entire courtship and copulation process can last over four hours (Gillingham and

Chambers, 1980). Information relating to reproduction in D. couperi is limited with most data

coming from captive populations (Hallam et al., 1998; Hyslop, 2007). Available reproductive

timelines for this species differ across their range and are therefore represented as broad periods of

time. The breeding season is considered to last from October through March (Groves, 1960;

Humphrey et al. 1992; Speake et al., 1978; Steiner et al., 1983). However, It is believed that it may

extend into April in Georgia (Hyslop, 2007; Moulis, 1976). D. couperi is oviparous and lays

clutches of four to 12 eggs (Speake et al., 1987; Steiner et al., 1983). The eggs range from 37 to 89 g

in mass (Speake et al., 1987; Steiner et al., 1983). The gestation period for this species can last for

130-140 days (Hyslop, 2007; Speake et al., 1987). Eggs hatch after approximately three months,

primarily between the months of August and September (Groves, 1960; Wright, 1957). Recorded

hatchling total length varied from 340–485 mm (Ernst and Barbour, 1989). Multiple studies

looking at sex ratios of hatchlings and juveniles show a 1:1 ratio of males to females (Moulis,

1976; Steiner et al., 1983). This ratio is reported as altered in adult snakes with males being more

numerous than females. A south Florida study reported an adult sex ratio of 1.54 males: 1 female

(Layne and Steiner, 1996). In a study using similar methods, Stevenson found an adult sex ratio of

2.1 males: 1 female in Georgia (Hyslop, 2007).

Sexual maturity is reached at 1500 mm total length (Layne and Steiner, 1996; Speake et al.,

1987). In a study of captive female snakes, sexual maturity was reached in three to four years

(Moulis, 1976). During the same study, Moulis noted that captive snakes commonly lay eggs

every year (Moulis, 1976). A two-year study of a wild population found that three of the five

females studied were gravid both years (Bolt, 1996). The maximum-recorded age of D. couperi in

captivity was 25 years 11 months (Snider and Bowler, 1992).

9

Home Range:

Understanding the effect of habitat loss on a species requires knowledge of the species

current use of the habitat (Hyslop, 2007). Unfortunately the secretive behavior, low densities,

and large home-range size of D. couperi make it difficult to obtain these data using traditional

field study techniques (Hyslop et al., 2009b; Parker and Plummer, 1987). Mark-recapture

studies are commonly used to determine population and home range size but are mostly

ineffective for D. couperi because of low recapture rates (Hyslop et al., 2009b; Parker and

Plummer, 1987). One field technique that has proven to be successful with D. couperi is radio

telemetry (Hyslop, 2007; Speake et al., 1978). Radio telemetry is time intensive and expensive

but ensures a greater likelihood of relocating individuals (Hyslop et al., 2009b). In addition

the transmitter implantation required for this type of study has been proven safe for D. couperi

(Hyslop et al., 2009b).

Previous studies looking at home range sizes of D. couperi have been carried out in

both southern Georgia and peninsular Florida with multiple studies carried out in southern

Georgia. Early studies primarily used relocated and captive bred specimens. These studies

found home range sizes ranging from 4.8 to >300 ha (Dodd and Barichivich, 2007; Moler,

1985b; Speake et al., 1978; Appendix Table 1). More recent studies on wild populations in

Georgia found home range sizes ranging from 35 to 1,530 ha, with male home ranges ranging

from 140 to 1,530 ha and female home ranges ranging from 35 to 354 ha (Hyslop, 2007).

These are the largest recorded home range size of any North American snake species (Hyslop,

2007). Other radio-telemetry studies conducted in Georgia show seasonal activity patterns

with smaller home range sizes during the winter months and home range expansion during the

summer months (Hyslop, 2007; Speake et al., 1978).

10

Radio telemetry studies conducted with D. couperi in peninsular Florida found home

range sizes ranging from 65 to 300 ha for males and 30 to 115 ha for females (Bolt, 2006;

Hyslop, 2007). A study conducted in northern Florida found home range sizes for males

ranging from 23 to 281 ha, (Hyslop, 2007; Moler, 1985b). A more recent study carried out in

central and east central Florida found home ranges between 12.8 – 538.4 ha (Breininger et al.,

2011). The southernmost telemetry study was carried out at Archbold Biological Station in

central Florida (Layne and Steiner, 1996). This study tracked 19 individuals and found the

average male home range to be 74.3 ha for males and 18.6 ha for females (Layne and Steiner,

1996). Home range sizes for southern populations of D. couperi remain a mystery.

Research Objectives:

To understand how habitat alteration will impact a species, it is important to first

understand the basic life history of the organism. Spatial requirements including home range

area, use of different habitat types, and seasonal changes in habitat use are key factors in

understanding the effect of habitat perturbations on the species

The goal of this study is to elucidate home range size, seasonal activity patterns,

differential habitat use, and refugia preferences of a population of D. couperi at an abandoned

agriculture site in southeastern Florida. The collection and analysis of this data is necessary to

provide guidance for the regional conservation management efforts of the species and provide

a baseline for future research. It will also serve to highlight possible intraspecific differences

that may relate to variation in climate and habitat differences throughout the species range.

1. Determine home range size of D. couperi in south Florida.

a. Describe total home range sizes of individuals

b. Examine seasonal (winter/summer) differences in home range size.

11

2. Determine habitat preferences of D. couperi in south Florida.

a. Describe habitat preferences of individuals

b. Examine seasonal (winter/summer) differences in habitat preferences.

3. Determine activity patterns of D. couperi in south Florida.

a. Describe overall activity patterns of individuals.

b. Examine seasonal (winter/summer) differences in activity patterns.

4. Describe refugia utilization of D. couperi in south Florida.

a. Describe refugia utilization by individuals.

b. Examine differences in refugia utilization in regards to air temperature.

12

Chapter 2

Methods

Study Site:

The field site for this study is an abandoned citrus grove area intersected with canals,

ditches, and dirt roads located in western Martin County Florida. The site will be home to the

future C-44 reservoir and storm water treatment area included in the Central and Southern

Florida Project of the Comprehensive Everglades Restoration Plan (USACOE, 2004). Prior to

agricultural use the site was likely similar to the surrounding Allapattah Flats Wildlife

Management Area, mixed slash pine and saw palmetto, with hardwood hammocks of oak and

sabal palm and patches of seasonal wetlands and marshes.

Figure 2.1 Map of study area

13

Figure 2.2 Aerial photo of C-44 footprint. Features in the southwest corner of the area are test

impoundments. Linear north/south features throughout the site are the ridges and swales

associated with the citrus orchard. Swales become seasonal wetlands.

The total C-44 project area is 4,856 ha and will contain a 4.6 m deep, 1,376 ha reservoir

(USFWS, 2006). Tree clearing and removal took place in 2006-2009. Removal of trees was

done using a backhoe and trees were piled and burned on site. Efforts were made to keep piles

away from canals and ditches were D. couperi had been previously sited (USFWS, 2006). The

citrus operation buildings are located within the footprint of the future reservoir and were

abandoned but still standing during the field portion of the study. In addition to buildings there

are abandoned equipment and pipes left over from the citrus operations during the course of the

study.

14

The study site for this research was limited to the 1,376 ha footprint of the future

reservoir. The reservoir footprint is dominated by native and non-native grasses and shrubs and

has limited canopy cover. Canals, ditches, and swales were typically covered with a mixture of

native and exotic aquatic and wetland vegetation throughout the site.

Table 2.1 List of most common plant species observed on the C-44 site.

Plant species list

Elephant Grass (Pennisetum purpureum)

Lantanas (Lantana sp.)

Smut Grass (Sporobolis indicus)

Beggar-Ticks (Bidens pilosa)

Common Ragweed (Ambrosia artemisiifolia)

Cogon Grass (Imperata cylindrica)

Dog Fennel (Eupatorium capillifolium)

Salt Bush (Baccharis halimifolia)

Brazilian Pepper (Schinus terebinthifolius)

Sabal Palm (Sabal palmetto)

Southern Wax Myrtle (Myrica cerifera)

Coastal Plain Willow (Salix caroliniana)

Bushy Bluestem (Andropogon glomeratus)

Yellowtop (Flaveria linearis)

Duck Potato (Sagittaria lancifolia)

American Crinum (Crinum americanum)

Pickerelweed (Pontederia cordata)

Torpedo Grass (Panicum repens)

Para Grass (Brachiaria mutica)

West Indian Marsh Grass (Hymenachne amplexicaulis)

Water Lettuce (Pistia stratiotes)

Water Hyacinth (Eichhornia crassipes)

Cattail (Typha sp.)

Fragrant Water Lily (Nymphaea odorata)

D. couperi had been documented within the study site and will be directly affected by the

construction and maintenance of the C-44 reservoir and surrounding storm water treatment areas

(USFWS, 2006). Biological surveys carried out in 2005 included one road kill and two live D.

15

couperi within the study area (USFWS, 2006). In addition, grove operators stated that D.

couperi were most often seen along roads and ditches on the property (USFWS, 2006).

The biological opinion for the site stated that many of the D. couperi sightings occurred

in the northern portion of the property bordering Allapattah Flats Wildlife Management Area

(USFWS, 2006). Population estimates for D. couperi were calculated using population estimates

from Archbold Biological Station, which assumes 2.6 snakes for every 100 ha (Layne and

Steiner, 1996; USFWS, 2006). All 4,856 ha of future reservoir and STA sites are classified as

D. couperi habitat. The use of a micro-irrigation system for the sites citrus operations provides

better snake habitat compared to a furrow-irrigated system, which requires additional flooding of

the land (USFWS, 2006). Due to the size of the study area, the US Fish and Wildlife Service

(USFWS), basing their estimate on D. couperi population density at Archbold Biological Station

(Highlands County, Florida), predicted that up to 126 adult D. couperi may be present at the C-

44 site. They also believed that the C-44 site represents less suitable habitat than that at

Archbold Biological Station. Thus they stated there are likely fewer than 126 adults at the C-44

site (USFWS, 2006).

Capture and Transmitter Implantation:

United States Fish and Wildlife Service permits allowed for the implantation of radio-

telemetry transmitters in up to five adult D. couperi. The transmitters used for this study were

Holohil Systems (Holohil Systems Ltd., Carp, Ontario) SI-2T internal transmitters. The

transmitters weigh 13 grams per transmitter, including batteries that are rated for an average of

two years of field use. Dr. Darryl Heard, (University of Florida, College of Veterinary

Medicine), supervised the implantation of the transmitters within the coelomic cavity of snakes

at the Small Animal Hospital at the University of Florida in Gainesville.

16

Snakes were hand captured opportunistically while conducting walking surveys in areas

where individuals were reportedly seen. A concurrent study being carried out by the USFWS

used coverboards as refugia to attract snakes and other herpetofauna to determine whether they

were present at the site. Areas around coverboards, abandoned buildings, and along canal banks

were frequently searched for snakes.

Captured individuals were examined to determine their sex and health. Sex was

preliminarily determined by looking for the presence of keels on middorsal scales, which are

characteristic of males (Layne and Steiner 1984; 1996; Stevenson et al., 2003). Any observation

related to health such as external damage and overall appearance of the skin were recorded at the

time of collection. All tracked snakes were sexed by examining genitalia at the time of

transmitter implantation.

For optimal post-surgery healing, individuals that appeared to be preparing to shed were

held until shedding occurred. Snakes were then transported to the University of Florida for

transmitter implantation. After surgery the snakes were held at Florida Gulf Coast University to

recuperate from the anesthesia and surgery. Snakes were typically held for three to five days

before being released back at the capture location. Typically snakes were released at the nearest

refugium to the site of capture. Implanted snakes were given a name in order to keep track of

individuals’ records to identify transmitter frequencies, and for ease of use in discussion.

Radio Tracking:

Individuals were tracked a minimum of twice a week and a minimum of once a day

during each tracking event. Tracking of individuals began immediately following release.

Snakes were tracked using a Communication Specialists (Communications Specialists, Inc.,

Orange, California) handheld yagi antenna and car-mounted roof antenna in conjunction with an

17

R-1000 handheld receiver. Once an individual being tracked was located, its location was

recorded using a handheld GPS unit. The observer recorded air temperature, atmospheric

conditions, and whether the snake was above ground or below ground. Information regarding

habitat and refugium type was also collected when applicable. Occasionally snakes were

recaptured to briefly examine their condition.

Timeline:

Our search for D. couperi at this site began in December 2011. As individuals were

captured they were implanted and released for tracking, so the duration of sampling varied

among the snakes. Data collection for this study was concluded at the end of one year.

Surviving snakes will continue to be tracked for an additional year after this study. At the

conclusion of the second year snakes will be recaptured and taken back to the University of

Florida Small Animal Hospital for transmitter removal. After recovery snakes will be released

back at the last site of capture.

Analysis:

Data were entered into Microsoft Excel for organization and were then imported to

ArcGIS 10.1 for spatial analysis and R package 3.0.1for statistical analysis. Seasons were

differentiated using ArcGIS 10.1 in conjunction with NOAA’s southeast Florida seasonal climate

categorization which used 42 years of temperature, dew point, and precipitation data to break the

year into two seasons: winter and summer (NOAA, 2009). The day summer started and winter

ended was May 21. The day that summer ended and winter started was October 17 (NOAA,

2009). These two dates provide the cutoff points for each season for this study (Figure 2.3).

18

Figure 2.3 Monthly average temperature and precipitation during study with lines delineating

seasonal breaks as determined by NOAA (2009). Monthly averages provided by Weather

Underground’s Indiantown station.

Objective 1: Home Range

Minimum bounding geometry polygons were created in ArcGIS in order to estimate

individual home ranges. Minimum bounding geometry (also commonly referred to as minimum

convex polygons) takes the sampled locations of each individual and creates the smallest

possible polygon using all included points. Kernel density estimations were also created using

ArcGIS to show high activity areas within home ranges. Kernel density estimations look at the

frequency at which an individual was sampled per square meter. This provided a visual

representation (color coded) for heavily used areas that would otherwise be represented only as

overlaying points. Representations of seasonal home ranges were similarly generated using

ArcGIS and NOAA designated seasons. This was accomplished using minimum bounding

19

geometry polygons that were comprised of points collected between the corresponding dates for

each season.

Objective 2: Habitat Preferences

The statistical package “R” was used to run a non-parametric binomial test to evaluate

habitat preferences between canal bank habitats (Figure 2.4) versus upland habitats (Figure 2.5)

in relation to seasonality. This test uses the proportion of each habitat type on the landscape as

the ‘expected’ frequency of habitat use and determines if the organism’s selection of the habitat

is non-random, i.e. shows a preference.

In order to differentiate habitats and determine their availability ArcGIS was used to select all

canals present within each individual’s seasonal ranges. The Florida Geographical Database Library

provided canal layers for Martin County. Selected canals were assigned an average width of 12

meters. All canals were assigned this width based on estimates from visual inspections at the field

site. In addition to the 12-meter wide canals an additional buffer was extended out 10 meters on

either side of the canal. The resulting canal and buffer area was used to represent canal habitat; their

areas were calculated within ArcGIS. The term “canal habitat” includes the actual water body,

Figure 2.4 Canal habitat Figure 2.5 Upland habitat

20

associated vegetation, canal banks, all structures in canals (drainage pipes, culverts, weirs, pump

stations), and the adjacent spoil uplands up to 10 meters on both sides of the canal, which often

includes agricultural access roads. The remaining habitat for each individual was classified as

upland habitat. The upland habitat is characterized as citrus grove fields with all citrus trees

removed. This “upland habitat” includes linear ridges, where citrus trees were planted and

adjacent swales that facilitated irrigation of the root zone. These seasonally flooded irrigation

swales are included in the generalized “upland habitat” category, and were not included as canals

or canal habitat because they were too narrow and shallow to be classified by the South Florida

Water Management District. These features were lumped in with upland habitat for this analysis.

The percentage of available canal habitat was then used in the binomial test. The statistical

significance of canal use for each season was examined in relation to the availability of canal habitat

for each snake during each season.

Objective 3: Activity Patterns

Activity patterns were calculated by using the great-circle distance equation to calculate the

minimum distance between data points (Nichols et al., 2000). This equation takes two points on the

surface of a sphere and determines the shortest distance between them.

Figure 2.6 Great-circle distance equation used to determine distance between sampling points

Once the distance between sampling points was determined it was divided by the number

of days between sampling events to determine daily movements. Average movement was

determined for individuals by calculating mean meters per day movement. When individuals

were located in the same location on successive tracking events an average daily movement of

21

zero was assumed. One sampling event was used per day and time of day in which sampling

occurred was not taken into account. For days where individuals were located more than once

during a single day the first location of the day was used. No weighting was used to account for

differences in days between sampling events. A non-parametric one-way permutation test was

then run in “R” using NOAA’s seasonal climate categories to test for differences in activity

patterns between seasons and among months.

Objective 4: Refugia Use

Refugia types were categorized as natural (Figure 2.7) or artificial (Figure 2.8). Natural

refugia included those made by mammals such as the burrows of Nine-banded Armadillos

(Dasypus novemcinctus) and Hispid Cotton Rats (Sigmodon hispidus), and any other naturally

occurring shelter. Artificial refugia included pipes, buildings, and waste left behind from citrus

operations. For this analysis, concrete piles and concrete waste were classified as artificial

refugia.

Air temperature was recorded during every sampling event and represents the air

temperature at the time sampling occurred. Air temperature data were collected using a digital

handheld thermometer. Locations where snakes were found in refugia were organized by type

and compared in relation to air temperature using “R” to run a one-way permutation test.

Figure 2.7 Natural refugia Figure 2.8 Artificial refugia

22

Chapter 3

Results:

A total of seven snakes including four males, two females and one juvenile of

undetermined sex were tracked between the dates of January 2012 and March 2013. Of these

snakes three were sampled at least 98 times; Monty was located a total of 107 times while Vader

and Nagini were both located 98 times (Table3.1). Two of the four remaining snakes were

successfully tracked enough times to include in analysis; Paul was located 35 times and

Dagwood 38 times (Table 1.1). The other two snakes were not included in the analysis due to

small sample size; Bette, a female, was located six times before being predated and Junior, a

juvenile, was successfully tracked 14 times before the external transmitter was shed.


After the field portion of this study was complete, polygons representing the home range

of each snake were established using minimum bounding geometry (Figure 3.1). Total home

range size ranged from 9.71 ha – 65.78 ha. Two males were tracked for every month of the year;

Monty had the largest total home range at 65.78 ha (Table 3.1 and Figure 3.2) and Vader had the

second largest total home range at 45.06 ha (Table 3.1 and Figure 3.3). The two other males

were tracked for less than 6 months out of the year; Dagwood had the third largest total home

range at 36.89 ha (Table 3.1 and Figure 3.4) and Paul had the fourth largest total home range at

23.52 ha (Table 3.1 and Figure 3.5). Nagini, the only female included in the analysis, was

tracked every month of the year; she had the smallest recorded total home range at 9.71 ha

(Table 3.1 and Figure 3.6).

Home ranges were broken into two seasons based on NOAA season categorization for

southeast Florida (NOAA, 2009). Winter home ranges included a maximum of 58.42 ha

23

(Monty; Table 3.1 and Figure 3.2) and a minimum of 1.96 ha (Nagini; Table 3.1 and Figure 3.6).

Summer home ranges included a maximum of 21.59 ha (Paul; Table 3.1 and Figure 3.5) and a

minimum of 9.71 ha (Nagini; Table 3.1 and Figure 3.6). Both Monty and Vader had larger

winter ranges than summer ranges, while Paul and Nagini displayed larger summer ranges than

winter ranges (Table 3.1).

Table 3.1 Snake home ranges for all snakes successfully tracked at least 35 times during the

course of the study. All sampling points were grouped to create home ranges represented by

Minimum Bounding Geometry polygons. Seasonal home ranges were calculated for individuals

with at least nine sampling events per season using the same technique.

Snake Points

Collected Sex

Winter Range

(ha)

Summer Range

(ha)

Total Home

Range (ha)

Monty 107 M 58.42 17.73 65.78

Vader 98 M 42.44 20.81 45.06

Dagwood 38 M 21.7 - 36.89

Paul 35 M 6.5 21.59 23.52

Nagini 98 F 1.96 9.71 9.71

24

Figure 3.1 Extent of the study area and minimum bounding geometry polygons using

ArcGIS 10.1, for all snakes tracked

25

Figure 3.2 Monty’s (male) total home range (top panel) as well as his winter and summer ranges

(bottom panels, black outlines represent total home range for comparison). All ranges were

created as minimum bounding geometry polygons with kernel density estimations highlighting

sampling occurrences per meter squared using ArcGIS 10.1.

26

Figure 3.3 Vader’s (male) total home range (top panel) as well as his winter and summer ranges




27

Figure 3.4 Dagwood’s (male) home range. This range was created as a minimum bounding

geometry polygon with kernel density estimations highlighting sampling occurrences per meter

squared using ArcGIS 10.1. No seasonal maps were created for Dagwood because he was not

tracked through the summer season.

28

Figure 3.5 Paul’s (male) total home range (top panel) as well as his winter and summer ranges




29

Figure 3.6 Nagini’s (female) total home range (top panel) as well as his winter and summer

ranges (bottom panels, black outlines represent total home range for comparison). All ranges

were created as minimum bounding geometry polygons with kernel density estimations

highlighting sampling occurrences per meter squared using ArcGIS 10.1.

30


Three male snakes and one female, (Monty, Vader, Paul and Nagini,) had enough sampling

points throughout the year to analyze seasonal habitat preferences for both seasons, while an

additional male (Dagwood) only had enough sampling points to look at habitat preferences during

the winter season (Table 3.2). The seasonal ranges for each snake were separated into two different

habitat types. Canal habitat consists of the canals and a ten-meter buffer along each side of the canal.

The remaining habitat within the seasonal range was classified as upland habitat. A one-way

permutation test was used to determine whether a significant preference for canal habitat exists for

each snake during each season based on the number of times canal habitat was used compared to the

percent of canal habitat within each snake’s seasonal home range.

Monty (Figures 3.7 and 3.8), Vader (Figures 3.9 and 3.10), and Paul (Figures 3.11 and 3.12)

showed a larger percentage of available canal habitat in their winter home ranges than in their

summer home ranges and all three were located more often in those canal habitats, demonstrating a

statistically significant preference for canal habitat, during the winter season (Table 3.2). Dagwood,

however, showed no significant canal habitat preference during the winter season (Table 3.2 and

Figure 3.13). Additionally, Vader was the only male to show a significant canal habitat preference

during the summer season (Table 3.2 and Figure 3.10). Interestingly, Nagini, the only female,

showed a conflicting trend from the males with a higher percent of available canal habitat during the

summer rather than winter season (Table 3.2). Also, except for Dagwood in the winter season,

Nagini’s seasonal home ranges contained the greatest percent of available canal habitat and she

showed a significant preference for those canal habitats during both seasons (Table 3.2 and Figures

3.14 and 3.15).

31

Table 3.2 Habitat use for all individuals sampled at least 35 times over the course of the study.

Canal habitat includes the canal itself as well as a ten-meter buffer along each side of the canal.

All other habitat was considered upland habitat. The percent of canal and upland habitat was

calculated based on the availability of each habitat type within each snake’s seasonal home ranges.

Seasons were based on NOAA seasonal data for southeast Florida (NOAA, 2009). The p-value

represents a one-way permutation test looking at preference for available canal habitat over

available upland habitat for each snake during each season. The color red denotes a significant p-

value.

Snake and Season %Canal

Habitat

Canal

Points

%Upland

Habitats

Upland

Points P-Value

Monty Winter 10.7 30 89.3 35 8.60E-14

Monty Summer 0.6 1 99.4 40 0.2187

Vader Winter 3.7 7 96.3 56 0.01278

Vader Summer 1.5 4 98.5 30 0.004596

Paul Winter 8.0 3 92.0 6 0.02979

Paul Summer 7.5 4 92.5 22 0.1501

Dagwood Winter 19.7 8 80.3 30 0.8369

Nagini Winter 18.5 8 81.5 29 0.0001461

Nagini Summer 34.2 13 65.8 3 2.07E-09

32

Figure 3.7 This figure displays Monty’s (male) winter habitat preferences. Habitats were broken

into two categories. Canal habitat includes the canal and a 10-meter buffer on either side of the

canal. Upland habitats represent all other available habitat within the winter home range.

During the winter season 10.7% of Monty’s winter home range was canal habitat and 89.3% was

upland habitat. The P-value represents a one-way permutation test looking at preference for

available canal habitat over available upland habitat during the winter season.

33

Figure 3.8 This figure displays Monty’s (male) summer habitat preferences. Habitats were

broken into two categories. Canal habitat includes the canal and a 10-meter buffer on either side

of the canal. Upland habitats represent all other available habitat within the winter home range.

During the summer season 0.6% of Monty’s winter home range was canal habitat and 99.4% was



34

Figure 3.9 This figure displays Vader’s (male) winter habitat preferences. Habitats were broken



During the winter season 3.7% of Vader’s winter home range was canal habitat and 96.3% was



35

Figure 3.10 This figure displays Vader’s (male) summer habitat preferences. Habitats were



During the winter season 1.5% of Vader’s summer home range was canal habitat and 98.5% was



36

Figure 3.11 This figure displays Paul’s (male) winter habitat preferences. Habitats were broken



During the winter season 8.0% of Paul’s winter home range was canal habitat and 92.0% was



37

Figure 3.12 This figure displays Paul’s (male) summer habitat preferences. Habitats were



During the winter season7.5% of Paul’s summer home range was canal habitat and 92.5% was



38

Figure 3.13 This figure displays Dagwood’s (male) winter habitat preferences. Habitats were



During the winter season 19.7% of Dagwood’s winter home range was canal habitat and 80.3%

was upland habitat. The P-value represents a one-way permutation test looking at preference for


39

Figure 3.14 This figure displays Nagini’s (female) winter habitat preferences. Habitats were



During the winter season 18.5% of Nagini’s winter home range was canal habitat and 81.5% was



40

Figure 3.15 This figure displays Nagini’s (female) summer habitat preferences. Habitats were



During the winter season 34.2% of Nagini’s summer home range was canal habitat and 65.8%

was upland habitat. The P-value represents a one-way permutation test looking at preference for


41


In order to determine seasonal activity patterns individuals had to be sampled at least nine

times during both winter and summer seasons. A total of four snakes were sampled enough times to

be included in this analysis (Table 3.3). Mean meters travelled per day was determined by taking the

distance travelled between tracking events, and dividing by the number of days between tracking

events, and then averaging movements for both seasons.

Average daily winter movements ranged from 88.05 m – 13.37 m. Average daily summer

movements ranged from 68.05 m – 41.79 m. Averages between the two seasons were tested to

determine whether movements were significantly different or not using a one-way permutation test.

Nagini, the female snake, showed a difference in activity between seasons that approached statistical

significance (Table 3.3 and Figure 3.16). However, the three male snakes, Monty (Figure 3.17), Vader

(Figure 3.18), and Paul (Figure 3.19) showed no significant difference (Table 3.3).

In addition to looking at activity differences between seasons, monthly activity patterns were

also tested for significance in Nagini, Monty, Vader, and Paul (Figures 3.20, 3.21, 3.22, and 3.23,

respectively). Breaking the activity patterns into a finer scale yielded similar results, with only Nagini

(Figure 3.20) showing a significant difference among monthly activity patterns.

Table 3.3 Activity patterns for all individuals that were located at least nine times in both the

summer and winter seasons. Average daily seasonal movements were determined by measuring

the distance between two consecutive sampling events and dividing that distance by the number

of days between events, and then averaging the daily movements for each season. Seasons were

based on NOAA seasonal data for southeast Florida (NOAA, 2009). The P-value was calculated

using a one-way permutation test.

Snake Winter

Points

Average Winter

Movements

(m/day)

Summer

Points

Average Summer

Movements

(m/day)

P-Value

Nagini 13 13.37 74 41.79 0.05761

Monty 50 52.67 39 50.71 0.8595

Vader 43 88.05 33 68.05 0.5499

Paul 9 26.99 23 64.12 0.2445

42

Figure 3.16 This figure displays Nagini’s (female) activity during the winter and summer seasons

based on mean meters traveled per day. Seasons were based on NOAA seasonal data for

southeast Florida (NOAA, 2009). The dark horizontal bars in this figure represent the median.

The boxes represent the 25th

to 75th

percentiles. The error bars represent the 5th

to 95th

percentiles. Circles represent individual data points outside the 5th

or 95th

percentiles. The P-

value was calculated using a one-way permutation test.

Summer Winter

050

100

150

200

25

0300

35

0Nagini

Seasons

Ave

rag

e M

inim

um

Dis

tance

Tra

vele

d P

er

Da

y in M

ete

rs (

+/-

95%

) p = 0.05921

43

Figure 3.17 This figure displays Monty’s (male) activity during the winter and summer seasons




to 75th


to 95th


or 95th

percentiles. The P-


Summer Winter

050

100

150

Monty

Seasons

Ave

rag

e M

inim

um

Dis

tance

Tra

vele

d P

er

Da

y in M

ete

rs (

+/-

95%

) p = 0.8595

44

Figure 3.18 This figure displays Vader’s (male) activity during the winter and summer seasons




to 75th


to 95th


or 95th

percentiles. The P-


Summer Winter

020

040

060

08

00

Vader

Seasons

Ave

rag

e M

inim

um

Dis

tance

Tra

vele

d P

er

Da

y in M

ete

rs (

+/-

95%

) p = 0.5499

45

Figure 3.19 This figure displays Paul’s (male) activity during the winter and summer seasons




to 75th


to 95th


or 95th

percentiles. The P-


Summer Winter

01

00

20

03

00

40

0

Paul

Seasons

Ave

rag

e M

inim

um

Dis

tance

Tra

vele

d P

er

Da

y in M

ete

rs (

+/-

95%

) p = 0.2445

46

Figure 3.20 This figure displays Nagini’s (female) monthly activity in mean meters traveled per

day. The error bars represent the 5th

to 95th

percentiles. The P-value was calculated using a one-

way permutation test.

P-value =0.0012

47

Figure 3.21 This figure displays Monty’s (male) monthly activity in mean meters traveled per


to 95th



P-value =0.9784

48

Figure 3.22 This figure displays Vader’s (male) monthly activity in mean meters traveled per


to 95th



P-value =0.1663

49

Figure 3.23 This figure displays Paul’s (male) monthly activity in mean meters traveled per day.

The error bars represent the 5th

to 95th

percentiles. The P-value was calculated using a one-way

permutation test.

P-value =0.09911

50


Use of refugia was examined in relation to air temperature. Individuals had to be tracked

through every month of the year in order to be included in this analysis. Therefore, three

individuals were included in this analysis. Refugia categories were broken into artificial and

natural. Available artificial refugia used during the course of the study included buildings, septic

tanks, pipes, and rock piles left behind from citrus operations. Additional artificial refugia were

placed at the field site for a concurrent D. couperi study being carried out by the US Fish and

Wildlife Service and the Florida Fish and Wildlife Conservation Commission, which included

artificial burrows and coverboards. Natural refugia available at the site can be primarily

characterized as mammal burrows. Correlations between refugia type and air temperature were

analyzed using a one-way permutation test (Table 4.1).

Monty (Figure 4.1) and Vader (Figure 4.2) both showed a statistically significant

preference for artificial refugia in cooler temperatures and natural refugia in warmer

temperatures (Table 4.1). Nagini (Figure 4.3) showed no significant preference for refugia type

in relation to air temperature (Table 4.1).

Table 3.4 Relationship between refugia type and air temperature. The p-value was calculated

using a one-way permutation test. Natural refugia included mammal burrows. Artificial refugia

included man-made structures, pipes, rock piles, cover boards, and artificial burrows.

Snake

Artificial

Refugia

Points

Average Temp.

When found in

Artificial Refugia

(C)

Natural

Refugia

Points

Average Temp.

When found in

Natural Refugia

(C)

P-Value

Monty 21 26 48 28 0.0171

Vader 35 26 22 29 2.2E-16

Nagini 26 26 44 28 0.7085

51

Figure 3.24 This figure displays Monty’s (male) preference for refugia type in relation to air

temperature. Natural refugia included mammal burrows. Artificial refugia included man-made

structures, pipes, rock piles, cover boards, and artificial burrows. The dark horizontal bars in this

figure represent the median. The boxes represent the 25th

to 75th

percentiles. The error bars

represent the 5th

to 95th


or 95th

percentiles. The P-value was calculated using a one-way permutation test.

Artificial Natural

22

24

26

28

30

32

34

Monty

RefugiaClass

Air T

em

pera

ture

C

p = 0.0171

52

Figure 3.25 This figure displays Vader’s (male) preference for refugia type in relation to air




to 75th


represent the 5th

to 95th


or 95th


Artificial Natural

20

22

24

26

28

30

32

Vader

RefugiaClass

Air T

em

pera

ture

C

p = 2.2e-16

53

Figure 3.26 This figure displays Nagini’s (female) preference for refugia type in relation to air




to 75th


represent the 5th

to 95th


or 95th


Artificial Natural

20

25

30

35

Nagini

RefugiaClass

Air T

em

pera

ture

C

p = 0.7085

54

Chapter 4

Discussion

Through this study I focused on the home range size, habitat use, activity patterns, and

refugia preferences of D. couperi in a fallow citrus grove in southeast Florida. During the course

of this study, a total of six adult snakes, four males and two females, were implanted with

transmitters. Of the six snakes captured, only five snakes could be tracked over an extended

period of time. Of these, four were located at least nine times during each of the NOAA seasons

allowing comparisons to be made between seasons. Three of the four snakes tracked through

both seasons were tracked for at least 307 days. The individuals tracked this length of time

included two adult males and one adult female. The small sample size for this study was due in

part to U.S. Fish Wildlife Services permitting restrictions, which limits the ability of this study to

provide population-level conclusions. However, the findings from this study do provide trends

that can be further tested in future studies.


Individuals in this study were tracked twice a week for 83 to 365 days. Total Home

range sizes for the four males varied from 23.52 – 65.78 ha and the only female tracked had a

home range of 9.71 ha. These home ranges are on the smaller end of the ranges recorded from

previous studies (Appendix Table 1). A radio telemetry study carried out in close proximity to

C-44 at Archbold Biological Station found average home ranges for males to be 74.3 ha and 18.6

ha for females (Layne and Steiner, 1996). The maximum recorded home range for males was

199.2 ha and 48.6 ha for females (Layne and Steiner, 1996). Layne and Steiner (1996) report

that these recorded ranges are likely smaller than the actual home ranges based on the correlation

they found between larger ranges with increased sampling. A study carried out recently in east-

55

central Florida included 107 snakes tracked once a week for 224 to 1,113 days. Home ranges

ranged from 12.8 - 538.4 ha and were calculated using minimum convex polygons (Breininger et

al., 2011). A recent study in Georgia tracked 32 snakes two to three times a week for 89 to 711

days. Home ranges ranged from 35 – 1,530 ha and were also calculated using minimum convex

polygons (Hyslop, 2007). The home range of 1,530 ha likely represents the largest recorded

snake home range in North America (Hyslop, 2007). Both of these studies had larger sample

sizes and tracked snakes over longer periods of time. Tracking frequency for this study was in

between referenced studies.

The smaller home ranges in this study could be a result of habitat quality, differential use

of habitat, and/or climate differences. The habitat at C-44 may represent a higher quality habitat

in relation to the needs of D. couperi compared to more natural habitats. The habitats used at

this site can be looked at as compressed in that it appears all of D. couperi’s requirements can be

found within a small area. This may allow for reduced home range sizes due to the closer

proximity of all necessary resources. The movements of a species and the area they occupy are

typically representative of the arrangement of necessary resources such as food, water, mates,

refugia, and appropriate thermal conditions (Gibbons and Semlitsch, 1987; Hyslop et al., 2009a).

Observations of prey items including small mammals and other snake species were frequently

seen at the field site indicating that prey items were prevalent. In addition both natural and

artificial refugia appeared to be abundant throughout canal and upland habitats. Water was also

abundant throughout the year although seasonal shifts caused the availability of water to decline

in upland habitats during the winter. Additionally, the climate of southeast Florida is warmer

and has less seasonal variation than that of previously referenced studies and may permit the

56

snakes to be more habitat generalists because of reduced restrictions on thermoregulation. This

may negate the need for individuals to increase their home range to include multiple habitats.

Both of the males tracked for more than 300 days had larger winter ranges than summer

ranges. Out of the three males with enough tracking events to create seasonal polygons, Paul was

the only male snake with a smaller winter range than summer range. Paul was predated 33 days

into the winter sampling period after having been located only nine times during the winter

season. It is possible that the small number of tracking events for this snake may have only

included part of his total home range and therefore underestimated its total size. The only female

tracked through both seasons showed a similar pattern to Paul in that she too had a larger

summer range than winter range. Females are undergoing gestation during much of the winter

and may not expand their range during this time (Hyslop, 2007).

Layne and Steiner (1996) found similar patterns at the Archbold Biological station for a

single male and female tracked long enough to make comparisons between seasons. A radio-

tracked male had a home range of 73 ha between the months of January and February and a

smaller home range of 42 ha in June and July. A radio-tracked female had a home range of 0.9

ha from January to March and a larger home range of 15 ha from April to May (Layne and

Steiner, 1996). Both of these snakes were sampled a minimal number of times but they represent

the closest study site nearest to ours at C-44, where D. couperi radio- tracking has occurred.

They also follow the same trends seen at C-44 in that the male had a larger winter range and

smaller summer range and the female had larger summer range and smaller winter range.

In Georgia populations Hyslop (2007) did not see any differences between seasonal home

range sizes in relation to sex, other than a reduction in spatial scale for female ranges throughout

the year. Any biological factors limiting female home range size during the winter in Georgia

57

would likely not be evident compared to males’ ranges because of the constraints on both sexes

from temperature. For this study it is possible that differences in seasonal home ranges between

sexes may simply be a result of small sample size.

D. couperi in Georgia were found to have larger summer ranges and smaller winter

ranges (Hyslop, 2007). Similarly a single D. couperi in central north Florida (Putnam County)

tracked over 323 days was also found to have a larger summer range and a smaller winter range

(Dodd and Barichivich, 2007). Colder temperatures restricted the movements of snakes in

Georgia and north Florida during winter so snakes spent more time in burrows. During the

warmer months snakes became more active and expanded their ranges to include different

habitats such as river bottoms and wetlands (Hyslop, 2007). The movement to new habitats may

have contributed to the much larger home ranges reported in these studies.

The geographical location of C-44 in southeast Florida provides a warmer climate with

mild winters. Because of the mild winters, D. couperi in the current study were not as restricted

by thermoregulation needs during the winter. This could contribute to larger winter home ranges

than those identified by Hyslop (2007). Studies tracking trans-located individuals in Georgia

showed temperature and seasonality are the two factors that most directly affected activity and

therefore home range size (Speake et al., 1978).

The larger winter home ranges compared to summer home ranges in the current study

may have been related to a smaller prey base and a reduced water supply during the winter.

During the winter water dries up across the landscape and likely restricts prey items to canals

that hold water all year long. Because of this water is a possible factor impacting changes in

home ranges between summer and winter at C-44. Based on the home ranges recorded in this

study it appears that as water dries up across the landscape during the winter, the snakes’ home

58

ranges increase to include a greater percentage of canal bank habitats. Similarly organisms that

constitute prey for D. couperi such as Banded Water Snakes (Nerodia fasciata) are likely

concentrated around the remaining water during the driest part of the year. On two occasions

during this study two different males were observed feeding on N. fasciata along canal banks.

Additionally the habitat preference findings in this study further support the importance of water

by showing that individuals spend a significant amount of time in canal habitats during the

winter. During the summer months this trend is not as strong with only Nagini and Vader

demonstrating a significant preference for canal habitat in relation to its availability. Seasonal

shifts in home range varied slightly among individuals but general trends become more evident

when examining individuals located over 300 days and when considering possible differences in

trends between sexes. The results of this study suggest trends in home range size for D. couperi

including: reduced summer season ranges: increased winter season ranges: and reduced total

home range size for individuals in southeast Florida at a fallow citrus grove.


The general habitat at C-44 is characterized as abandoned fallow citrus groves intersected

with irrigation canals. Abandoned citrus groves were listed as habitats commonly used by

foraging D. couperi at Archbold Biological Station (Layne and Steiner, 1996). Individuals

tracked in this study exclusively used this habitat and were never found outside of the citrus

groves. For this study the habitats at C-44 were divided into canal bank habitats and upland

habitats. Canal bank habitats consist of the canal itself as well as the surrounding embankment.

Upland habitats include fallow citrus groves bisected by swales and ditches that hold ephemeral

water during the summer. The preferences for canal banks versus uplands were examined

between seasons. Hyslop found that the most influential factor for microhabitat shifts is

59

seasonality (Hyslop et al., 2009a). They focused their microhabitat analysis around refugia

openings where snakes were located. Microhabitat was defined in regards to vegetation and

structure. They found that season rather than sex, size, or site played the biggest role in

determining what microhabitat snakes would use. Microhabitat analyses were not carried out for

this study but habitat preferences were analyzed in regards to available habitat and the amount of

time spent in these habitats during the winter and summer.

Rainfall and temperature are the major factors for distinguishing seasons in Florida

(NOAA, 2009). These factors cause seasonal changes in habitat and impact the activity patterns

of the snakes. Defining seasons in south Florida is a challenge. For this study, I used the

seasonal categories determined by NOAA, with a fixed date for the start and end of summer and

winter. Although the weather during the study period, January 2012 through March 2013, seem

to track well with the NOAA-defined seasons (Figure 2.3), it is possible that this seasonal

designation did not capture the seasons as perceived by the snakes, or their habit use in response

to the seasons. Some noticeable changes in the habitats during the summer season are the influx

of water and the explosion of vegetation. This affects both habitats. In the canal habitats the

canal bank is reduced during the summer because of the higher water levels. In the upland

habitats, ditches and other depressions fill with water. In addition, the growth of vegetation

provides cover for snakes to easily move across previously open fields. Movement across open

areas can be dangerous due to the risk of desiccation (Bogert et al., 1947) and predation. During

the winter season these ephemeral wet areas dried up. The canal water levels also dropped

exposing the banks and the cooler and drier weather results in a loss off much of the grassy

vegetation. By tracking snakes throughout the year it was possible to identify those areas where

60

the snakes were most commonly found which provided important information on habitat use and

seasonal preferences.

Identified seasonal home ranges indicate that the canals were an important habitat for D.

couperi during part of the year. Canals have also been documented as favorable habitat for D.

couperi in south Florida (Lawler, 1977). Lawler pointed to canals as likely places to locate this

species in south Florida and hypothesized that land crab burrows may provide refugia when G.

polyphemus burrows are not available. Studies in Everglades National Park have provided

anecdotal reports of D. couperi along canal banks in addition to agricultural fields and novel

habitats including mangrove swamps (Steiner et al., 1983).

All individuals tracked with the exception of Dagwood showed a significant preference

for canal habitats during the winter season. Within Dagwood’s range there was a ditch

intersection that held water throughout the winter. Dagwood spent a great deal of time along this

ditch. Despite the size of the ditch and the fact that it held water all year long it was not

classified as a canal. It is possible that this habitat served the same function as canal habitat.

Another trend indicated by my data was that all male snakes had a winter range with a

higher percentage of canal habitats than their summer range. Nagini, the only female in the

analysis, did not show this trend. She made extensive use of canal habitat throughout the year

and actually had the highest recorded percentage of canal habitat for any snake during the

summer. This follows trends noted by Hyslop (2007) where females and some males continued

to use habitat within their winter ranges all year long while other males only returned to over

wintering habitats during the winter months. The canals provided a reliable source of water

during the dry winter months and likely also an expanded prey base. In addition, the dry,

61

sparsely vegetated upland habitats likely contained fewer prey items and no cover. The lack of

cover may increase the likelihood of predation or desiccation.

During the summer months, the habitat preference for the male snakes switched to the

upland areas. This switch is likely driven by increased levels of precipitation, which increased

the distribution of water and caused an explosion of vegetation in the upland areas. In addition

the numerous swales and ditches bisecting the upland areas fill with water during the summer, so

the habitat was not exclusively typical upland. These flooded areas provided habitat for D.

couperi prey items. This was coupled with a reduced availability of foraging areas along the

canal banks as the water levels rose.

The use of canal habitat year round for Nagini and the use of canals primarily during the

winter for all males provides an alternative or contributing factor for why males may have visited

canal habitats with greater frequency during the winter. It is possible that canal habitats

represent a preferred habitat for female snakes and males enter these habitats for breeding

purposes. This fits trends of northern snakes during breeding season where males often

congregate around refugia being used by females (Stevenson et al., 2009). Previously

unidentified trends in habitat preferences and seasonal shifts for D. couperi include the

preference of canal habitat during the winter season by males and the possible yearlong

preference for canal habitats by females. These trends may be unique to southeast Florida

populations found in fallow citrus field habitats, and may help define alternative habitat to

include those areas with permanent water bodies.


Average distance moved between tracking events was used as an indicator of activity

level and patterns in this study. Using this method limits our ability to look at movements over a

62

short period of time and is only meaningful when used to gain a general pattern of activity. In

order to further refine activity levels using this method additional weekly sampling would be

necessary. By decreasing the number of days between sampling events you could analyze

activity on a finer scale. For this study, activity levels between NOAA seasons were compared.

The use of the NOAA defined seasons for the breakdown for this analysis may again possibly be

limiting as it breaks up the entire years worth of movements into two chunks that may or may not

capture what snakes are actually doing. An alternate way of determining seasons may have led

to different results regarding seasonal differences in activity. Because of this, activity patterns

were also compared across months to determine if analysis on a finer scale yielded any patterns

not evident between seasons.

Comparing trends in activity across different studies is difficult, as no standard for

quantifying activity exist. Moulis (1976) reported April as being the height of the active season

for D. couperi in Georgia, based on a nine-year monitoring study looking at the frequency of

sightings through the years. Another Georgia study using telemetry to track translocated

individuals found peak activity between the months of August and November based on

observations and shifts in home range sizes (Speake et al., 1978). A more recent telemetry study

looking at frequency of daily movements using the proportion of tracking days that an individual

changed locations found that females make smaller movements than males and that both sexes

make the fewest movements during the winter and the most during the summer (Hyslop, 2007).

A telemetry study that was conducted relatively close to C-44 in south-central Florida found that

the percentage of times snakes were recorded as active rather than inactive was highest from

August to October and then second highest from May to July (Layne and Steiner, 1996). Long

term monitoring in Everglades National Park shows peak activity based on frequency of

63

sightings between November and March (Steiner et al., 1983). It is possible that these findings

were skewed by the increased number of people making observations during those months.

Sightings at C-44 were also highest during the winter. Most individuals were captured during

this time although that could have been partially an artifact of the much lower levels of

vegetation present in the winter, making it easier to see and capture the snakes.

After analyzing all of the individuals’ activity patterns between seasons and months, none

of them showed a significant difference in their mean daily movements except for Nagini. Layne

and Steiner (1996) found similar trends in that females move much less during winter and spring

and more during summer and fall. Nagini’s height of activity (November) would have fallen into

Layne and Steiners’ fall season, and her lowest activity (December) would fall within their

winter season (Layne and Steiner, 1996). This trend is very weakly supported but, due to the

proximity of C-44 to Archbold Biological Station, it is plausible that individuals at both sites

may share similar activity trends. Although the male individuals at C-44 did not show a

significant difference, a constant pattern of higher mean activity in winter months was present

for sufficiently tracked individuals. In addition, home range sizes, a possible indirect measure of

activity, increased during the winter season. Activity levels are likely strongly impacted by

weather with snakes seeking refuge during the hottest periods in the summer and the coolest

periods in the winter but maintaining high activity levels throughout the year. Seasons in south

Florida are also tied to rainfall, which controls the distribution of moisture across the landscape.

The distribution of water is likely tied to both overall prey abundance, and the concentration of

potential prey items.

With the exception of Paul who had a small sample size, all males had significantly larger

ranges in the winter, but did not significantly increase their level of activity, as measured by

64

mean distance moved per day. The expanded winter ranges and slight trend in raised activity

through winter may have been contributed to by a reduction in prey availability in the cooler and

drier winter months as well as the need to include a reliable source of water. The larger ranges

and marginally increased activity in sufficiently tracked individuals could also be partially

caused by the males searching for females with which to breed.

What we do know about seasonal changes in activity patterns for this group of snakes is

that both male and female snakes remain active all year long with peak activity months falling in

the winter season for both sexes. These trends differ from those expressed by snakes in Georgia

(Hyslop, 2007). This is likely due to the cooler climate in Georgia restricting activity during the

winter. In addition the reduction of vegetation, the preference for canal banks, and the slightly

raised activity levels make the winter season the most likely time to see D. couperi. This may

also be the most dangerous time of year for D. couperi. Three out of the four snakes that were

predated during this study were predated during the winter. The additional snake was likely

killed in an intra-species conflict while expanding his home range.


Refugia type and availability may also play a role in where individuals are found during

certain times of the year. In Georgia, Hyslop (2007) found that D. couperi show a preference for

G. polyphemus burrows during the winter. These burrows are typically found in sandhill

environments and may represent the main draw for D. couperi to these habitats. In a south-

central Florida telemetry study G. polyphemus burrows represented 62% of refugia used (Layne

and Steiner, 1996). Natural ground holes and burrows of unknown origin accounted for another

25% of the refugia used (J. Layne & Steiner, 1996). No preference existed for refugia type in

relation to sex, activity, or season (Layne and Steiner, 1996). Additionally Layne and Steiner

65

(1996) observed D. couperi using a house, barn and concrete culverts as refugia during dry

weather. For this study, records were kept on the type and location of refugia used and

comparisons made between refugia preference in the two NOAA seasons. Only snakes tracked

for at least 300 days were included in this analysis. While snakes remained active throughout the

year, refugia were needed for resting, protection from predators, and protection during extremely

cool and hot periods.

Snakes were frequently observed using refugia along canal banks during the winter

months. It was also interesting that many of the refugia used were structures or items that were

left over from the citrus operations. Items like pipes, concrete piles, and even a septic system

were used as refugia during the winter. Many of the available artificial refugia were located

along canal banks. The snakes may have showed a preference for artificial refugia during the

winter because of the insulation some of the artificial items may offer, or simply due to its

availability in the canal habitats that the snakes preferred during winter.

It is possible that Nagini’s sex plays a role in her refugia preferences. Female D. couperi

may show preferences for refugia use based on quality of refugia in relation to nest site locations.

A study looking at translocated females tracked in north Florida showed that gestating females

show a preference for inactive G. polyphemus burrows (Hyslop et al., 2009a). This preference is

possibly due to the risk of eggs in active burrows being destroyed by other burrow denizens.

Factors such as this could skew any correlation between refugia type and air temperature. Nagini

also had fewer winter tracking events because her signal could not be located on multiple

occasions. It is probable that she was in a burrow that was sufficiently deep or insulated to

completely block her signal. This likely skewed our analysis of burrow preferences for Nagini.

Overall we see a trend for male snakes using artificial refugia in cooler weather and natural

66

refugia in warmer weather. Nagini showed no significant preference for certain refugia types in

relation to air temperature but her results were complicated by our inability to track her on a

number of sampling days. Despite this, the average temperature when Nagini was observed

using artificial refugia was identical to the average temperatures recorded when the other snakes

used artificial refugia. Based on these preferences for refugia types in relation to air

temperatures, it is plausible that there may be differences in how much insulation a particular

type of refugia may offer. Additional snakes need to be tracked and burrow temperatures

measured to determine whether this is the case. We now know that D. couperi at C-44 are not

dependent on G. polyphemus burrows during any part of the year and that they will use human

artifacts as refugia.

Conclusions

D. couperi using the C-44 reservoir site demonstrate trends in home range size, habitat

use, seasonal activity patterns, and seasonal refugia preferences that differ from other

populations of this species. Driving factors for these differences seem to be linked primarily to

differences in hydro period and climate. Some of the objectives of this study highlight

differences between the individuals tracked during this study and northern populations sampled

in previous studies.

One notable trend is the difference in sizes between seasonal ranges. The largest

seasonal range for males tracked for a full year at C-44 was winter. Winter represents the

smallest seasonal range for northern populations of D. couperi (Hyslop, 2007). Another

difference is habitat preferences. The individuals tracked during this study were tracked over a

highly disturbed landscape. The available habitat is much different than that of more northern

populations, and of the often-assumed need for pristine upland habitat for this species.

67

Availability of water appeared to be a significant factor in changes in home ranges observed over

the course of the study. Seasonal activity patterns of the snakes in the current study also differed

from those in other studies. Males tracked in this study showed no significant difference in

activity across seasons. This differs from northern populations and is likely caused by the cooler

temperatures and the resulting decrease in winter activity in northern D. couperi (Hyslop, 2007).

A final difference is the use of refugia. The C-44 site is unusual in that there are no G.

polyphemus present on site. The burrows of these tortoises were important refugia for northern

populations during the winter (Hyslop, 2007). The individuals tracked for this study relied

heavily on artificial refugia during the winter although natural refugia were available.

Future studies aimed at elucidating basic life history of D. couperi in unique habitats

should focus on prey base, a site-specific reproductive timeline, role of refugia, and causes of

mortality in novel environments. Understanding and quantifying prey base is important in

understanding the basic life history of an organism. The availability, type, and location of prey

likely play a role in the activity and home range size of a predator. Site-specific reproductive

timelines are also important in understanding the activity patterns and movements of an

organism, particularly for D. couperi who appears to shows varied reproductive timelines

throughout its range. Understanding the role certain refugia play is also important for D.

couperi. The availability of refugia may be a limiting factor for more northern snakes,

particularly if G. polyphemus burrows are not present. The final recommendation is to quantify

dangers to this species in novel sites. During the course of this study four of the snakes being

tracked were lost to mortality. Two of these snakes appear to have been predated by feral hogs

(Sus scrofa). Determining if S. scrofa is a significant predator of D. couperi is important in

developing land practices aimed at the continued survival of this species. These objectives cover

68

important basic life history traits that would be seminal in the development of conservation

efforts and land practices that cater to D. couperi in disturbed landscapes.

Some important findings from this study that impact management of the Indigo Snake

are its ability to populate and thrive in highly disturbed habitats and to make use of artificial

burrows. The smaller home ranges of the C-44 snakes may indicate that the habitat was quite

suitable and that snakes were not forced to travel long distances for adequate food, water, and

shelter. The small ranges may also have been partially a result of a relatively high density of

snakes. From a survey and monitoring perspective, the snakes were not always easy to find.

Sixty seven percent of the time we were tracking individuals we never actually saw the snake,

despite receiving a strong radio signal indicating the snake was likely within a few feet of us.

Additionally we were unable to locate individuals that were being tracked 11% of the time even

when we knew the snakes were somewhere in the area. This difficulty in finding the snake, and

the recognition that D. couperi can thrive in some heavily disturbed habitats, should result in

very conservative conclusions regarding the absence of snakes from a potential development site

when the survey includes only limited visual inspection.

Findings also demonstrated important differences between this population and other

populations, especially more northern populations. These differences highlight the need for

conservation biologists to consider ecological and behavioral differences across the range of a

species when developing management plans. The fact that this study documented a population

of D. couperi apparently thriving on an extremely impacted site, should not guide us toward

believing the snake is no longer in danger from habitat loss. While the C-44 site appears to be

good habitat; this tells us nothing about other human impacted landscapes.

69

However, in a world were landscape alteration is common practice, it is important to

understand that altered habitat can substitute for more natural habitats for many organisms.

Understanding the role of disturbed habitats as acceptable habitat for endangered and threatened

species and species of special concern is integral to their continued survival. Learning how to

effectively improve these atypical habitats for use by keystone species and others, allows us to

conserve natural flora and fauna in habitats that are becoming more common as human

populations increase, hopefully positively affecting the trend in global biodiversity loss.

70

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75

Appendix

Appendix Table 1 Home range comparison table

This table shows the results of D. couperi home range studies using radio-telemetry. All home range sizes were calculated using

100% minimum convex polygons or minimum bounding geometry.

Study Location

Average

Male Home

Range Size

Male

Sample

Size

Average

Female

Home Range

Size

Female

Sample

Size

Tracking

Duration

(Days)

Sampling

Frequency

Study

Authors

C-44 (Martin County) 42.8 4 9.71 1 83-365 Twice a Week Current

Study

Central Florida (Highlands

County) 74.3 12 18.6 7 8-197 Daily

Layne and

Steiner, 1996

Southeastern Florida

(Brevard County) 118 31 41 18 Unknown Unknown Bolt, 2006

North Florida (Levy

County) 141 4 - - Unknown Unknown Moler 1985

Central Florida

(Highlands, Polk

Counties)/ Southeastern

Florida (Brevard County)

201.7 23 75.6 21 224-1,113 Once a week Breininger et

al., 2011

Southeastern Georgia (Fort

Stewart Military

Reservation)

538 24 126 14 89- 711 Three times a

week Hyslop, 2007

76

Appendix Table 2 Raw Data

Snake_ID Date Time Temp_C Cloud N DD.DDDDD W DD.DDDDD Habitat Refugia Above

Ground Burrow

Vader(Capture) 1/19/12 12:00 17 1 27.09267 -80.44952 3 X 1 0

Vader(Release) 2/16/12 14:30 27 2 27.09270 -80.44960 1 4 2 2

Vader 1 2/17/12 10:52 24 2 27.09977 -80.45052 1 X 1 0

Vader 2 2/17/12 14:15 24 2 27.10070 -80.45115 1 X 1 0

Vader 3 2/21/12 11:30 24 2 27.09920 -80.45277 2 X 1 0

Vader 4 2/21/12 15:30 24 2 27.10103 -80.45330 2 X 1 0

Vader 5 2/23/12 11:53 26 1 27.09938 -80.45197 2 X 1 0

Vader 6 2/23/12 15:46 26 1 27.09652 -80.45232 3 X 1 0

Vader 7 2/24/12 12:34 29 2 27.09533 -80.45252 1 4 2 2

Vader 8 2/24/12 15:17 29 2 27.09267 -80.44990 1 X 1 0

Vader 9 2/28/12 12:17 23 3 27.09267 -80.44988 1 5 2 2

Vader 10 2/28/12 14:14 23 3 27.09267 -80.44988 1 5 2 2

Vader 11 3/1/12 12:18 27 3 27.09267 -80.44988 1 5 2 2

Vader 12 3/1/12 14:06 27 3 27.09267 -80.44988 1 5 2 2

Vader 13 3/2/12 12:54 28 2 27.09273 -80.44968 1 4 2 2

Vader 14 3/2/12 14:48 28 2 27.09273 -80.44968 1 4 2 2

Vader 15 3/6/12 11:27 24 2 27.09273 -80.44968 1 4 2 2

Vader 16 3/6/12 13:17 24 3 27.09273 -80.44968 1 4 2 2

Vader 17 3/8/12 11:50 24 3 27.09273 -80.44968 1 4 2 2

Vader 18 3/8/12 15:00 24 3 27.09273 -80.44968 1 4 2 2

Vader 19 3/9/12 12:52 27 3 27.09273 -80.44968 1 4 2 2

Vader 20 3/9/12 14:58 27 3 27.09273 -80.44968 1 4 2 2

Vader 21 3/13/12 12:33 25 3 27.09273 -80.44968 1 4 2 2

Vader 22 3/13/12 14:02 25 3 27.09273 -80.44968 1 4 2 2

Vader 23 3/15/12 12:22 27 3 27.09273 -80.44968 1 4 2 2

Vader 24 3/15/12 13:30 27 3 27.09273 -80.44968 1 4 2 2

Vader 25 3/20/12 13:10 25 2 27.09273 -80.44968 1 4 2 2

Vader 26 3/20/12 14:41 25 2 27.09273 -80.44968 1 4 2 2

Vader 27 3/22/12 13:01 27 2 27.09260 -80.44985 1 5 2 2

Vader 28 3/22/12 14:00 27 2 27.09260 -80.44985 1 5 2 2

Vader 29 3/28/12 11:30 26 2 27.09800 -80.45253 1 3 2 2

Vader 30 3/28/12 13:45 26 2 27.09740 -80.45322 1 X 1 0

Vader 31 3/30/12 12:55 27 2 27.09652 -80.45222 3 2 2 1

77


Ground Burrow

Vader 32 4/3/12 12:39 29 1 27.09922 -80.45048 1 2 2 1

Vader 33 4/6/12 13:04 29 2 27.10113 -80.45373 1 X X X

Vader 34 4/6/12 14:22 29 2 27.10113 -80.45373 1 X X X

Vader 35 4/10/12 13:31 27 2 27.09918 -80.45050 1 2 2 1

Vader 36 4/13/12 13:39 27 2 27.09648 -80.45222 3 2 2 1

Vader 37 4/13/12 15:07 27 2 27.09648 -80.45222 3 2 2 1

Vader 38 4/17/12 13:18 27 3 27.09587 -80.45267 4 4 2 X

Vader 39 4/19/12 13:42 28 2 27.09587 -80.45267 4 4 2 X

Vader 40 4/24/12 11:45 19 1 27.09587 -80.45255 4 4 2 X

Vader 41 4/26/12 13:08 27 2 27.09587 -80.45255 4 4 2 X

Vader 42 5/1/12 15:00 27 3 27.09835 -80.45373 2 X X X

Vader 43 5/4/12 12:20 30 2 27.09950 -80.45602 1 2 2 1

Vader 44 5/7/12 12:25 33 2 27.09953 -80.45575 1 2 2 1

Vader 45 5/11/12 8:05 26 2 27.09885 -80.45372 1 2 2 1

Vader 46 5/11/12 11:25 26 2 27.09888 -80.45375 1 X 1 0

Vader 47 5/15/12 12:27 29 2 27.10055 -80.45620 1 2 2 1

Vader 48 5/17/12 15:58 28 3 27.09937 -80.45512 1 X 1 0

Vader 49 5/22/12 11:39 29 2 27.09923 -80.45050 1 2 2 1

Vader 50 5/25/12 13:04 30 2 27.09920 -80.45050 1 2 2 1

Vader 51 5/29/12 14:01 30 3 27.09920 -80.45050 1 2 2 1

Vader 52 6/1/12 10:12 25 3 27.10035 -80.45598 2 2 2 1

Vader 53 6/5/12 13:19 30 3 27.10080 -80.45312 1 X 1 X

Vader 54 6/8/12 13:16 27 3 27.09918 -80.45292 1 2 2 1

Vader 55 6/12/12 12:56 28 1 27.10065 -80.45617 1 2 2 1

Vader 56 6/15/12 X 29 2 X X X X X X

Vader 57 6/19/12 X 29 3 X X X X X X

Vader 58 6/21/12 22:30 27 2 27.09590 -80.45258 4 4 2 2

Vader 59 6/21/12 1:32 26 2 27.09590 -80.45258 4 4 2 2

Vader 60 6/21/12 5:07 26 2 27.09590 -80.45258 4 4 2 2

Vader 61 6/26/12 12:55 31 3 27.09592 -80.45262 4 4 2 2

Vader 62 6/28/12 11:33 28 1 27.09973 -80.45375 1 2 2 1

Vader 63 7/3/12 14:14 32 3 27.10082 -80.45397 2 2 2 1

Vader 64 7/6/12 13:02 36 3 27.10097 -80.45370 1 X X X

Vader 65 7/10/12 13:26 32 2 27.09992 -80.45337 1 2 2 1

78


Ground Burrow

Vader 66 7/13/12 13:05 30 2 27.10038 -80.45598 2 2 2 1

Vader 67 7/16/12 12:35 26 3 27.09627 -80.45252 4 4 2 2

Vader 68 7/26/12 11:40 30 2 27.09974 -80.45375 1 2 2 1

Vader 69 7/30/12 19:00 29 3 X X X X X X

Vader 70 7/31/12 9:05 27 1 27.10019 -80.45645 1 X 1 0

Vader 71 8/3/12 8:50 29 2 X X X X X X

Vader 72 8/11/12 8:02 27 3 27.10008 -80.45340 1 X 1 0

Vader 73 8/14/12 9:50 31 2 X X X X X X

Vader 74 8/17/12 X 32 2 X X X X X X

Vader 75 8/21/12 X 29 2 X X X X X X

Vader 76 8/24/12 9:04 29 2 27.09631 -80.45245 3 4 2 2

Vader 77 8/28/12 13:00 30 2 27.10099 -80.45363 X X X X

Vader 78 8/31/12 10:20 31 2 27.09909 -80.45173 2 X 1 X

Vader 79 9/4/12 12:26 29 2 27.10104 -80.45139 1 2 2 1

Vader 80 9/7/12 10:50 30 2 27.09909 -80.45194 2 X 1 0

Vader 81 9/11/12 10:15 29 2 27.09652 -80.45247 3 X 1 0

Vader 82 9/14/12 11:40 29 4 27.09652 -80.45247 3 4 2 2

Vader 83 9/18/12 9:35 26 2 27.09650 -80.45245 3 4 2 2

Vader 84 9/21/12 12:21 30 3 27.09725 -80.45342 1 X 1 0

Vader 85 9/25/12 12:45 28 3 27.10002 -80.45236 1 X 1 0

Vader 86 9/28/12 11:42 30 2 27.10036 -80.45615 2 2 2 1

Vader 87 10/2/12 8:30 24 3 27.09899 -80.44998 1 X 1 0

Vader 88 10/5/12 12:00 31 2 27.09984 -80.45222 1 X 1 0

Vader 89 10/9/12 12:40 30 2 27.09653 -80.45237 3 X X 0

Vader 90 10/13/12 11:15 29 2 X X X X X X

Vader 91 10/16/12 14:49 29 2 27.09591 -80.45270 4 4 2 2

Vader 92 10/19/12 11:25 33 1 27.09651 -80.45235 3 X 1 X

Vader 93 10/24/12 10:15 30 2 27.09970 -80.45448 1 X 1 0

Vader 94 10/30/12 16:50 24 1 X X X X X X

Vader 95 11/2/12 10:30 24 1 27.09585 -80.45226 4 4 2 2

Vader 96 11/6/12 13:30 27 2 27.09800 -80.45083 1 X 1 0

Vader 97 11/9/12 15:50 25 2 X X X X X X

Vader 98 11/11/12 12:50 27 2 27.09611 -80.45266 4 4 2 2

Vader 99 11/16/12 14:30 24 3 27.09643 -80.45264 3 X 1 0

79


Ground Burrow

Vader 100 11/20/12 14:49 23 2 27.09306 -80.44984 1 X 1 0

Vader 101 11/24/12 18:15 21 2 X X X X X X

Vader 102 11/27/12 14:15 25 3 27.09612 -80.45272 4 4 2 2

Vader 103 11/30/12 11:40 24 2 27.09610 -80.45239 4 4 2 2

Vader 104 12/2/12 14:40 22 3 X X X X X X

Vader 105 12/5/12 12:00 24 3 27.10178 -80.45070 1 X 1 0

Vader 106 12/11/12 11:41 22 4 27.09240 -80.44976 1 X 1 0

Vader 107 12/14/12 16:05 24 2 27.09566 -80.45255 1 X 1 0

Vader 108(Dead) 12/18/12 13:00 32 1 27.09507 -80.45444 2 X 1 0

Monty(Capture) 1/10/12 11:00 18 2 27.09263 -80.44990 1 X 1 0

Monty(Release) 2/7/12 11:55 27 3 27.09263 -80.44990 1 X 1 0

Monty 1 2/7/12 13:30 27 3 27.09263 -80.44990 1 5 2 2

Monty 2 2/9/12 12:40 24 3 27.09322 -80.44770 1 X 1 0

Monty 3 2/9/12 14:47 24 3 27.09345 -80.44847 1 X 1 0

Monty 4 2/14/12 13:20 24 2 27.09043 -80.44953 3 3 2 2

Monty 5 2/16/12 X 27 2 X X X X 2 X

Monty 6 2/17/12 X 24 2 X X X X 2 X

Monty 7 2/21/12 X 24 2 X X X X 2 X

Monty 8 2/23/12 X 26 1 X X X X 2 X

Monty 9 2/24/12 13:10 29 2 27.09058 -80.44935 3 3 2 2

Monty 10 2/24/12 15:30 29 2 27.09058 -80.44935 3 3 2 2

Monty 11 2/28/12 12:37 23 3 27.09057 -80.44935 3 3 2 2

Monty 12 2/28/12 14:07 23 3 27.09057 -80.44935 3 3 2 2

Monty 13 3/1/12 11:04 26 3 27.09057 -80.44935 3 3 2 2

Monty 14 3/1/12 13:40 26 3 27.09057 -80.44935 3 3 2 2

Monty 15 3/2/12 12:42 28 2 27.09057 -80.44935 3 3 2 2

Monty 16 3/2/12 14:40 28 2 27.09057 -80.44935 3 3 2 2

Monty 17 3/6/12 11:15 24 2 27.09057 -80.44935 3 3 2 2

Monty 18 3/6/12 12:47 24 3 27.09057 -80.44935 3 3 2 2

Monty 19 3/8/12 11:10 24 3 27.09057 -80.44935 3 3 2 2

Monty 20 3/9/12 12:40 27 3 27.09043 -80.44932 3 3 2 2

Monty 21 3/13/12 12:22 25 3 27.09555 -80.44928 3 X 1 0

Monty 22 3/13/12 13:45 25 3 27.09613 -80.44932 3 X 1 0

Monty 23 3/15/12 12:30 26 3 27.09383 -80.44933 3 2 2 1

80


Ground Burrow

Monty 24 3/15/12 1:49 26 3 27.09383 -80.44933 3 2 2 1

Monty 25 3/20/12 12:40 27 2 27.09743 -80.44943 3 X 1 0

Monty 26 3/22/12 12:50 27 2 27.09707 -80.44930 3 3 2 2

Monty 27 3/22/12 13:54 27 2 27.09707 -80.44930 3 3 2 2

Monty 28 3/28/12 13:16 26 2 27.09707 -80.44930 3 3 2 2

Monty 29 3/28/12 14:00 26 2 27.09707 -80.44930 3 3 2 2

Monty 30 3/30/12 11:20 24 2 27.09707 -80.44930 3 3 2 2

Monty 31 4/3/12 12:03 29 1 X X X X X X

Monty 32 4/6/12 12:30 29 2 X X X X X X

Monty 33 4/10/12 12:59 27 2 27.09707 -80.44930 3 3 2 2

Monty 34 4/13/12 X 27 2 X X X X X X

Monty 35 4/17/12 12:28 27 3 27.09585 -80.44907 1 X 1 0

Monty 36 4/17/12 14:49 27 3 27.09588 -80.44933 3 X 1 0

Monty 37 4/19/12 12:45 28 3 27.09707 -80.44930 X X X X

Monty 38 4/24/12 15:47 22 1 27.09707 -80.44930 3 3 2 2

Monty 39 4/26/12 X 27 2 X X X X X X

Monty 40 5/1/12 X 27 3 X X X X X X

Monty 41 5/4/12 X 30 2 X X X X X X

Monty 42 5/7/12 X 33 2 X X X X X X

Monty 43 5/11/12 7:15 24 2 27.09465 -80.44905 1 X 1 0

Monty 44 5/11/12 11:00 24 2 27.09470 -80.44912 1 2 2 1

Monty 45 5/15/12 13:11 29 2 27.09170 -80.44898 1 X 1 0

Monty 46 5/17/12 15:37 28 3 27.08947 -80.44877 1 X 1 0

Monty 47 5/22/12 12:23 29 3 27.08998 -80.44722 1 2 2 1

Monty 48 5/25/12 12:46 30 2 27.08998 -80.44722 1 2 2 1

Monty 49 5/29/12 13:39 30 3 27.08960 -80.44713 1 2 2 1

Monty 50 6/1/12 9:51 24 3 27.09067 -80.44713 1 2 2 1

Monty 51 6/5/12 12:58 30 3 27.09067 -80.44713 1 2 2 1

Monty 52 6/8/12 12:51 28 3 27.09067 -80.44713 1 2 2 1

Monty 53 6/12/12 12:18 29 1 27.08948 -80.44642 1 2 2 1

Monty 54 6/15/12 12:21 29 2 27.08958 -80.44888 1 2 2 1

Monty 55 6/19/12 12:05 29 3 27.08975 -80.44687 1 2 2 1

Monty 56 6/21/12 11:06 26 2 27.08975 -80.44687 1 2 2 1

Monty 57 6/26/12 12:05 29 3 27.09010 -80.44672 1 X 1 X

81


Ground Burrow

Monty 58 6/28/12 12:24 28 1 27.09343 -80.44673 1 2 2 1

Monty 59 7/3/12 13:03 32 2 27.09402 -80.44835 1 2 2 1

Monty 60 7/6/12 12:30 33 3 27.09402 -80.44835 1 2 2 1

Monty 61 7/10/12 12:17 33 3 27.08958 -80.44662 1 2 2 1

Monty 62 7/13/12 12:31 30 2 27.08907 -80.44877 3 2 2 1

Monty 63 7/16/12 9:54 27 3 27.08985 -80.44852 1 X 1 0

Monty 64 7/26/12 10:27 29 2 27.09464 -80.44669 3 2 2 1

Monty 65 7/26/12 14:22 27 1 27.09466 -80.44670 1 2 2 1

Monty 66 7/30/12 18:09 30 2 27.09462 -80.44669 1 2 2 1

Monty 67 7/31/12 8:20 24 1 27.09462 -80.44669 1 2 2 1

Monty 68 8/3/12 9:18 29 2 27.09513 -80.44589 2 X 1 0

Monty 69 8/11/12 X 27 3 X X X X X X

Monty 70 8/14/12 9:35 31 1 27.09375 -80.44535 1 X 1 0

Monty 71 8/17/12 10:51 32 2 27.09142 -80.44671 1 2 2 1

Monty 72 8/21/12 11:47 29 2 27.08939 -80.44676 2 X 1 0

Monty 73 8/24/12 9:31 29 2 27.09333 -80.44632 2 X 1 0

Monty 74 8/28/12 12:00 29 2 27.09246 -80.44717 1 2 2 1

Monty 75 8/31/12 10:42 31 2 27.09246 -80.44717 1 2 2 1

Monty 76 9/4/12 13:10 29 3 27.09464 -80.44669 1 2 2 1

Monty 77 9/7/12 11:15 30 2 27.09419 -80.44794 1 2 2 1

Monty 78 9/11/12 9:40 25 1 27.09069 -80.44889 1 2 2 1

Monty 79 9/14/12 10:38 28 2 27.08976 -80.44634 1 X 1 0

Monty 80 9/18/12 8:30 25 1 27.09070 -80.44888 1 2 2 1

Monty 81 9/18/12 10:45 25 1 27.09042 -80.44874 1 X 1 0

Monty 82 9/21/12 13:00 30 3 X X X X X X

Monty 83 9/25/12 13:25 29 2 27.09243 -80.44715 1 2 2 1

Monty 84 9/28/12 12:36 31 2 27.09243 -80.44715 1 2 2 1

Monty 85 10/2/12 8:50 24 3 27.09029 -80.44866 1 X 1 0

Monty 86 10/5/12 12:19 31 2 27.09368 -80.44903 1 X 1 0

Monty 87 10/9/12 13:05 29 2 27.09335 -80.44900 1 X 1 0

Monty 88 10/13/12 10:35 29 2 27.09438 -80.44677 1 X 1 0

Monty 89 10/16/12 15:17 29 2 27.09214 -80.44894 1 X 1 0

Monty 90 10/19/12 9:15 26 1 27.09419 -80.44795 1 2 2 1

Monty 91 10/24/12 10:40 35 2 27.09419 -80.44796 1 2 2 1

82


Ground Burrow

Monty 92 10/24/12 11:50 35 2 27.09419 -80.44796 1 2 2 1

Monty 93 10/30/12 16:50 24 1 X X X X X X

Monty 94 11/2/12 11:30 26 1 27.08920 -80.44618 3 X 1 0

Monty 95 11/6/12 14:00 27 2 27.09003 -80.44386 1 2 2 1

Monty 96 11/9/12 13:50 25 2 27.08684 -80.44764 1 2 2 1

Monty 97 11/11/12 12:15 26 2 27.08712 -80.44761 1 2 2 1

Monty 98 11/16/12 15:00 21 3 27.09349 -80.44743 2 X 1 0

Monty 99 11/20/12 14:31 23 2 27.09104 -80.45184 1 X 1 0

Monty 100 11/24/12 18:43 21 2 27.09182 -80.44985 1 2 2 1

Monty 101 11/27/12 12:30 28 3 27.09182 -80.44982 1 2 2 1

Monty 102 11/30/12 10:45 24 3 27.09180 -80.44984 1 2 2 1

Monty 103 12/2/12 13:45 22 3 27.09179 -80.44984 1 2 2 1

Monty 104 12/5/12 12:30 25 2 27.09181 -80.44982 1 2 2 1

Monty 105 12/11/12 11:22 27 4 27.08955 -80.45299 1 X 1 0

Monty 106 12/14/12 15:26 24 2 27.08923 -80.45039 3 2 2 1

Monty 107 12/18/12 13:35 32 1 27.08925 -80.44650 3 X 1 0

Monty 108 12/21/12 11:40 18 1 27.08734 -80.44597 1 X 1 0

Monty 109 12/25/12 15:45 27 3 27.08809 -80.44893 1 X 1 0

Monty 110 12/28/12 12:30 23 1 27.08677 -80.44734 1 X 1 0

Monty 111 12/31/12 14:15 26 2 27.08789 -80.45084 1 X 1 0

Monty 112 1/5/13 12:26 27 3 27.08808 -80.44937 3 2 2 1

Monty 113 1/7/13 15:20 24 4 27.08687 -80.44636 1 X 1 0

Monty 114 1/9/13 13:30 29 2 27.08691 -80.44634 1 2 2 1

Monty 115 1/14/13 13:10 29 2 27.08767 -80.45098 1 2 2 1

Monty 116 1/18/13 14:20 23 2 27.09182 -80.44984 1 2 2 1

Monty 117 1/23/13 12:45 25 2 27.09184 -80.44982 1 2 2 1

Monty 118 1/25/13 12:35 21 1 27.09182 -80.44983 1 2 2 1

Monty 119 1/29/13 10:15 22 2 27.09201 -80.44632 1 X 1 0

Nagini(Capture) 2/23/12 10:35 26 1 27.11065 -80.44915 3 X 1 X

Nagini(Release) 3/30/12 11:07 24 2 27.11065 -80.44915 3 3 2 2

Nagini 1 3/30/12 13:15 24 2 27.11065 -80.44915 3 3 2 2

Nagini 2 4/3/12 12:20 29 1 27.11097 -80.44838 3 2 2 1

Nagini 3 4/6/12 12:40 29 2 27.11097 -80.44838 3 2 2 1

Nagini 4 4/6/12 14:13 29 2 27.11097 -80.44838 3 2 2 1

83


Ground Burrow

Nagini 5 4/10/12 13:15 27 2 27.11103 -80.44915 3 4 2 2

Nagini 6 4/13/12 14:01 27 2 27.11102 -80.44902 3 4 2 2

Nagini 7 4/17/12 13:44 27 3 27.11097 -80.44848 3 2 2 1

Nagini 8 4/19/12 14:06 28 3 27.11097 -80.44848 3 2 2 1

Nagini 9 4/24/12 10:57 16 1 27.11097 -80.44848 3 2 2 1

Nagini 10 4/26/12 12:53 27 1 27.11097 -80.44848 3 2 2 1

Nagini 11 5/1/12 12:28 27 3 27.11335 -80.44837 1 X 1 X

Nagini 12 5/4/12 12:50 30 2 27.11315 -80.44892 X 2 2 1

Nagini 13 5/4/12 14:10 30 2 27.11198 -80.44873 1 X 1 X

Nagini 14 5/7/12 X 33 2 X X X X X X

Nagini 15 5/11/12 X 26 2 X X X X X X

Nagini 16 5/15/12 12:45 29 2 27.11260 -80.44897 1 X 1 X

Nagini 17 5/17/12 16:11 28 3 27.11247 -80.44915 3 X 1 X

Nagini 18 5/22/12 14:07 29 2 27.11343 -80.44840 2 2 2 1

Nagini 19 5/25/12 13:42 30 2 27.11442 -80.44750 1 X X 1

Nagini 20 5/29/12 14:15 30 3 27.11152 -80.44830 X X X X

Nagini 21 6/1/12 10:36 25 3 27.11045 -80.44890 2 2 2 1

Nagini 22 6/5/12 13:30 30 3 27.11343 -80.44842 2 2 2 1

Nagini 23 6/8/12 X 28 3 X X X X X X

Nagini 24 6/12/12 X 29 1 X X X X X X

Nagini 25 6/15/12 X 29 2 X X X X X X

Nagini 26 6/19/12 13:11 29 3 27.11342 -80.44840 2 2 2 1

Nagini 27 6/21/12 21:16 27 3 27.11103 -80.44903 3 4 2 2

Nagini 28 6/21/12 0:42 26 3 27.11103 -80.44903 3 4 2 2

Nagini 29 6/21/12 4:20 26 3 27.11103 -80.44903 3 4 2 2

Nagini 30 6/26/12 13:27 31 3 27.11103 -80.44903 3 4 2 2

Nagini 31 6/28/12 10:33 27 1 27.11103 -80.44903 3 4 2 2

Nagini 32 7/3/12 13:56 32 3 27.11103 -80.44903 3 4 2 2

Nagini 33 7/6/12 13:38 33 3 27.11433 -80.44758 1 2 2 1

Nagini 34 7/10/12 14:27 33 2 27.11453 -80.44883 1 X 1 X

Nagini 35 7/13/12 14:18 30 2 27.11145 -80.44933 3 2 2 1

Nagini 36 7/16/12 10:57 27 3 27.11350 -80.44932 3 X 1 X

Nagini 37 7/26/12 X 30 2 X X X X X X

Nagini 38 7/30/12 19:20 27 2 X X X X X X

84


Ground Burrow

Nagini 39 7/31/12 8:00 23 1 27.11296 -80.44915 3 X 1 X

Nagini 40 7/31/12 9:35 23 1 27.11400 -80.44917 3 X 1 X

Nagini 41 8/3/12 8:31 27 2 27.11468 -80.44748 1 2 2 1

Nagini 42 8/11/12 X 27 3 X X X X X X

Nagini 43 8/14/12 10:10 32 2 27.11448 -80.44805 1 X 1 1

Nagini 44 8/17/12 9:31 29 2 27.11357 -80.44798 2 X 1 1

Nagini 45 8/21/12 10:32 29 2 27.11233 -80.44845 2 2 2 1

Nagini 46 8/24/12 10:47 34 2 27.11368 -80.44781 1 X 1 1

Nagini 47 8/28/12 11:17 29 3 27.11097 -80.44849 3 2 2 2

Nagini 48 8/31/12 10:00 28 2 27.11097 -80.44849 3 2 2 1

Nagini 49 9/4/12 11:51 29 2 27.10934 -80.44862 1 2 2 1

Nagini 50 9/7/12 10:19 29 2 27.11154 -80.44879 1 X 1 1

Nagini 51 9/11/12 8:45 25 1 27.11256 -80.44886 1 X 1 1

Nagini 52 9/11/12 11:00 25 1 27.11238 -80.44856 1 X 1 1

Nagini 53 9/14/12 9:47 28 2 27.11426 -80.44762 2 2 2 1

Nagini 54 9/18/12 7:50 24 1 27.11357 -80.44862 1 X 1 1

Nagini 55 9/18/12 11:00 24 1 27.11339 -80.44834 1 X 1 1

Nagini 56 9/21/12 11:35 31 3 27.11223 -80.44825 1 X X 1

Nagini 57 9/25/12 11:38 29 3 27.11101 -80.44902 3 4 2 2

Nagini 58 9/28/12 11:12 30 2 27.11101 -80.44902 3 4 2 2

Nagini 59 10/2/12 8:00 24 3 27.11103 -80.44906 3 X 1 2

Nagini 60 10/5/12 11:33 32 2 27.11246 -80.44836 1 X 1 1

Nagini 61 10/9/12 12:04 29 2 27.11277 -80.44877 1 X 1 1

Nagini 62 10/13/12 11:50 29 2 27.11425 -80.44764 1 X 1 1

Nagini 63 10/16/12 14:20 31 2 27.11203 -80.44855 1 X 1 1

Nagini 64 10/19/12 8:20 25 1 27.11060 -80.44917 3 3 2 2

Nagini 65 10/24/12 9:45 27 2 27.11060 -80.44918 3 3 2 2

Nagini 66 10/24/12 12:15 36 2 27.11060 -80.44918 3 3 2 2

Nagini 67 10/30/12 15:23 21 1 27.11098 -80.44846 3 2 2 2

Nagini 68 11/2/12 9:45 20 1 27.11104 -80.44806 3 X 1 2

Nagini 69 11/6/12 13:00 27 2 27.11423 -80.44758 2 X 1 2

Nagini 70 11/9/12 13:05 28 2 27.11334 -80.44830 1 X 1 1

Nagini 71 11/11/12 11:10 26 2 27.11192 -80.44895 3 X 1 1

Nagini 72 11/16/12 13:07 24 3 27.11637 -80.44865 1 X 1 1

85


Ground Burrow

Nagini 73 11/20/12 13:06 22 2 27.11400 -80.44842 1 X 1 1

Nagini 74 11/24/12 18:15 21 2 X X X X X X

Nagini 75 11/27/12 12:04 25 3 27.11201 -80.44897 3 X 1 1

Nagini 76 11/30/12 12:50 25 2 27.11681 -80.44882 1 X 1 1

Nagini 77 12/2/12 13:30 22 4 27.11075 -80.44917 3 X 1 2

Nagini 78 12/5/12 11:45 24 3 27.11062 -80.44916 3 3 2 2

Nagini 79 12/11/12 12:00 23 4 27.11062 -80.44917 3 3 2 2

Nagini 80 12/14/12 16:30 23 2 27.11062 -80.44917 3 3 2 2

Nagini 81 12/18/12 13:57 33 2 27.11085 -80.44887 2 X 1 1

Nagini 82 12/21/12 12:45 18 1 X X X X X X

Nagini 83 12/25/12 13:20 33 2 27.11294 -80.44914 3 X 1 2

Nagini 84 12/28/12 10:50 22 1 27.11283 -80.44916 3 X 1 2

Nagini 85 12/31/12 13:05 24 2 27.11289 -80.44916 3 X 1 2

Nagini 86 1/5/13 11:40 27 2 27.11288 -80.44918 3 8 2 2

Nagini 87 1/7/13 16:00 23 4 27.11293 -80.44933 3 3 2 1

Nagini 88 1/9/13 13:05 29 2 27.11205 -80.44915 3 3 2 1

Nagini 89 1/14/13 12:50 28 2 27.11060 -80.44915 3 3 2 2

Nagini 90 1/23/13 13:29 27 2 27.11062 -80.44914 3 3 2 2

Nagini 91 1/25/13 11:10 20 1 27.11064 -80.44916 3 3 2 2

Nagini 92 1/29/13 9:50 22 3 27.11096 -80.44847 3 2 2 2

Nagini 93 2/1/13 10:40 17 1 27.11151 -80.44780 1 2 2 1

Nagini 94 2/6/13 12:00 26 2 X X X X X X

Nagini 95 2/7/13 10:30 24 2 27.11151 -80.44843 1 3 1 2

Nagini 96 2/13/13 10:48 28 2 27.11135 -80.44845 2 X 1 X

Nagini 97 2/16/13 13:20 23 2 27.11463 -80.44839 2 X 1 X

Nagini 98 2/20/13 10:00 22 1 27.11147 -80.44847 2 X 1 X

Nagini 99 2/22/13 11:30 31 2 27.11152 -80.44842 1 X 1 X

Nagini 100 2/26/13 11:58 28 2 27.11151 -80.44843 2 X 1 X

Nagini 101 3/1/13 11:12 15 2 X X X X X X

Nagini 102 3/4/13 12:40 20 1 X X X X X X

Nagini 103 3/5/13 12:20 25 1 27.11148 -80.44842 2 2 2 1

Nagini 104 3/12/13 11:05 22 3 27.11212 -80.44803 1 X 1 X

Nagini 105 3/15/13 11:14 19 1 27.11213 -80.44779 1 X 1 X

Nagini 106 3/22/13 13:06 24 2 27.11098 -80.44870 3 8 2 2

86


Ground Burrow

Nagini 107 3/25/13 11:45 23 1 27.11101 -80.44904 3 5 1 2

Nagini 108 3/27/13 15:05 22 1 27.11110 -80.44850 3 X 1 X

Nagini 109 3/29/13 12:05 27 1 27.11099 -80.44850 3 X 1 X

Dagwood(Capture) 2/23/12 14:20 27 1 27.10642 -80.43307 3 6 2 2

Dagwood(Release) 3/1/12 12:00 27 3 27.10642 -80.43307 3 6 2 2

Dagwood 1 3/1/12 12:00 28 2 27.10642 -80.43307 3 6 2 2

Dagwood 2 3/2/2012 13:27 28 2 27.11023 -80.43333 1 2 2 1

Dagwood 3 3/2/12 15:50 24 2 27.11023 -80.43333 1 2 2 1

Dagwood 4 3/6/2012 12:20 24 3 27.10997 -80.43288 3 X X X

Dagwood 5 3/6/12 13:34 24 3 27.10997 -80.43288 3 X X X

Dagwood 6 3/8/12 12:19 24 3 27.10858 -80.43198 1 2 2 1

Dagwood 7 3/8/12 14:40 27 3 27.10737 -80.43210 2 2 2 1

Dagwood 8 3/9/12 13:30 27 3 27.10737 -80.43163 2 X X X

Dagwood 9 3/9/12 15:55 25 3 27.10713 -80.43170 2 X 1 X

Dagwood 10 3/13/12 13:10 25 3 27.10428 -80.43172 2 2 2 1

Dagwood 11 3/13/12 14:22 23 3 27.10428 -80.43172 2 2 2 1

Dagwood 12 3/15/12 12:57 23 3 27.10483 -80.43163 2 2 2 1

Dagwood 13 3/15/12 14:00 27 3 27.10483 -80.43163 2 2 2 1

Dagwood 14 3/20/12 13:53 27 2 27.10755 -80.43275 2 X 1 X

Dagwood 15 3/20/12 15:45 27 2 27.10755 -80.43275 3 X 1 X

Dagwood 16 3/22/12 13:19 27 2 27.10775 -80.43262 1 2 2 1

Dagwood 17 3/22/12 14:13 26 2 27.10750 -80.43230 2 X 1 X

Dagwood 18 3/28/12 12:45 27 2 27.10730 -80.43172 2 2 2 1

Dagwood 19 3/30/12 12:18 29 2 27.10735 -80.43175 2 2 2 1

Dagwood 20 4/3/12 13:12 30 1 27.10738 -80.43180 2 2 2 1

Dagwood 21 4/6/12 13:33 30 2 27.10738 -80.43180 3 2 2 1

Dagwood 22 4/6/12 14:37 27 2 27.10738 -80.43180 3 2 2 1

Dagwood 23 4/10/12 14:10 27 2 27.10545 -80.43272 2 2 2 1

Dagwood 24 4/13/12 14:20 27 2 27.10513 -80.43272 2 2 2 1

Dagwood 25 4/17/12 14:24 28 3 27.11452 -80.43267 2 X 1 X

Dagwood 26 4/19/12 14:31 20 2 27.11192 -80.43280 3 2 2 1

Dagwood 27 4/24/12 12:26 27 1 27.11075 -80.43152 1 X 1 X

Dagwood 28 4/26/12 13:30 27 2 27.11013 -80.43267 2 2 2 1

Dagwood 29 5/1/12 14:30 30 3 27.10210 -80.43443 1 X 1 X

87


Ground Burrow

Dagwood 30 5/4/12 12:30 33 2 27.11025 -80.43333 1 2 2 1

Dagwood 31 5/7/12 12:25 28 2 27.10678 -80.43383 1 2 2 1

Dagwood 32 5/11/12 8:40 28 2 27.10672 -80.43380 1 2 2 1

Dagwood 33 5/11/12 12:00 29 2 27.10672 -80.43380 1 2 2 1

Dagwood 34 5/15/12 13:24 28 2 27.10552 -80.43347 1 2 2 1

Dagwood 35 5/17/12 16:33 30 3 27.10497 -80.43353 2 2 2 1

Dagwood 36

(Dead) 5/22/12 13:20 86 3 27.10450 -80.43640 1 X 1 X

Paul(Capture) 6/28/12 13:15 29 2 27.07840 -80.44962 3 7 2 2

Paul(Release) 7/25/12 18:31 35 1 27.07841 -80.44963 3 7 2 2

Paul 1 7/25/12 1:46 27 1 27.07950 -80.44962 3 3 2 2

Paul 2 7/26/12 10:05 29 2 27.08199 -80.44936 3 X 1 X

Paul 3 7/30/12 18:50 28 2 27.07744 -80.44839 2 X 1 X

Paul 4 7/31/12 8:40 27 1 27.07710 -80.44839 2 X 1 X

Paul 5 8/3/12 X 29 2 X X X X X X

Paul 6 8/11/12 9:24 27 3 27.07682 -80.44820 2 2 2 1

Paul 7 8/14/12 9:12 32 1 27.07817 -80.44905 1 X 1 X

Paul 8 8/17/12 11:36 31 2 27.07588 -80.44898 1 X 1 X

Paul 9 8/21/12 12:23 32 2 27.07785 -80.44868 1 2 2 1

Paul 10 8/24/12 9:55 29 2 27.07700 -80.44821 1 2 2 1

Paul 11 8/28/12 12:32 29 2 27.07775 -80.44781 2 X 1 X

Paul 12 8/31/12 11:00 31 2 27.07776 -80.44781 2 X 1 X

Paul 13 9/4/12 X 29 4 X X X X X X

Paul 14 9/7/12 12:46 31 2 27.07920 -80.44865 1 X 1 X

Paul 15 9/11/12 9:00 25 1 27.07879 -80.44900 1 2 2 1

Paul 16 9/14/12 11:18 29 3 27.07599 -80.44873 2 X 1 X

Paul 17 9/18/12 9:00 25 1 27.07914 -80.45001 2 2 2 1

Paul 18 9/18/12 10:15 25 1 27.07914 -80.45001 2 2 2 1

Paul 19 9/21/12 1:27 29 4 27.07913 -80.45000 2 2 2 1

Paul 20 9/25/12 13:43 29 2 27.07913 -80.45000 2 2 2 1

Paul 21 9/28/12 13:03 31 2 27.07881 -80.44898 1 2 2 1

Paul 22 10/2/12 9:05 25 4 27.07792 -80.44807 2 X 1 X

Paul 23 10/5/12 12:47 31 2 27.07643 -80.44341 2 X 1 X

Paul 24 10/9/12 13:32 29 2 27.07695 -80.44813 2 X 1 X

88


Ground Burrow

Paul 25 10/13/12 10:15 28 2 27.07832 -80.44627 1 2 2 1

Paul 26 10/16/12 15:50 29 2 27.07830 -80.44628 2 2 2 1

Paul 27 10/19/12 10:30 36 1 27.07833 -80.44627 1 2 2 1

Paul 28 10/24/12 11:10 35 2 27.07880 -80.44899 1 2 2 1

Paul 29 10/30/12 16:24 25 1 27.07849 -80.44961 3 X 1 X

Paul 30 11/2/12 11:10 25 1 27.07810 -80.44961 3 X 1 X

Paul 31 11/6/12 14:45 26 3 27.07896 -80.45005 2 2 2 1

Paul 32 11/9/12 14:15 25 2 27.07935 -80.44973 1 2 2 1

Paul 33 11/11/12 12:30 27 1 27.07889 -80.44960 3 X 1 X

Paul 34 11/16/12 16:23 21 3 27.08036 -80.44748 2 X 1 X

Paul 35(Dead) 11/20/12 13:43 23 2 27.08076 -80.44747 2 X 1 X

Appendix Table 3 Key for Data

Cloud Cover Habitat Refugia Above Ground Burrow

1= Clear 1= Upland 1= Gopher tortoise 1= Above ground 0= NA

2= Partly cloudy 2= Wetland/ ditch 2= Mammal burrow 2= Below ground 1= Natural

3= Overcast 3= Canal/ bank 3= Pipe X= Unknown 2= Artificial

4= Rain 4= Forested 4= Rock pile - -

- 5= Other 5= Building structure - -

- - 6= Coverboard - -

- - 7= Junction box - -

- - 8= Other - -

- - X= Unknown - -

This table provides the meaning of numerals used to abbreviate raw data.

home range size and habitat use of the eastern indigo ...21424/datastream... · analysis between...

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