HABITAT MANIPULATION AS A VIABLE STRATEGY FOR THE

CONSERVATION OF THE MASSASAUGA RATTLESNAKE IN NEW YORK

STATE

By
Kevin Thomas Shoemaker

A thesis submitted in partial fulfillment

of the requirements for the

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Master of Science Degree

State University of New York

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College of Environmental Science and Forestry
Syracuse, New York
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April 2007
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Approved:

Faculty of Environmental and Forest Biology

_________________________ __________________________
James P. Gibbs Dudley J. Raynal
Major Professor Chair, Examining Committee

_________________________ __________________________
Donald J. Leopold Dudley J. Raynal
Faculty Chairman Dean, Instruction and Graduate Studies
 UMI Number: 1442576

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UMI Microform 1442576

Copyright 2007 by ProQuest Information and Learning Company.
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 ACKNOWLEDGMENTS

I thank my advisor, James Gibbs, for providing key insights and guidance through all

stages of this research project and for inspiring me with his soft-spoken dedication to

wildlife conservation. I thank Alex Krofta for his innumerable hours of cheerful hard

work in the field and in the office, without which this project could not have been

completed. I would also like to acknowledge Bryan Windmiller, my first herpetology

mentor, whose passion for conservation is infectious – and apparently incurable.

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I am very grateful for the many people who volunteered their time to assist in
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this research project, including Helen Czech, Mike Fishman, Kari Gunson, Tom

Hennigan, Debra Joseph, John Koltz, Viorel Popescu, and Joel Strong. From the
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planning stages to the writing, this project has benefited from key comments and

suggestions from my steering committee – Glenn Johnson, J. Scott Turner, and Alvin R.
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Breisch. John Adamski of the Seneca Park Zoo in Rochester and Andrew Lentini of the

Metro Toronto Zoo provided me with essential training in handling venomous snakes.

Dr. Lianjun Zhang was instrumental in helping me to refine and improve my data

analyses. Chapter 1 benefited from suggestions for improvement given by Kent A.

Prior, Glenn Johnson, and Stephen J. Mullin. Chapter 2 was written in conjunction with

a course on writing for scientific publication (FOR 694). The instructor, Dr. Ruth Yanai,

as well as the other course participants, have been an enormous help in drafting this

manuscript. This research project was funded in part by a grant from the New York

State Biodiversity Research Institute (BRI).

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 For my first two years at ESF, I served as a GK-12 teaching fellow with the ESF

Science Corps; this fellowship not only paid the bills, but provided me with an

opportunity to grow both personally and professionally. I thank James Gibbs and

Dudley Raynal for encouraging me to apply for the GK-12 fellowship, and I thank the

wonderful high school teachers I worked with during my time as a teaching fellow:

Heather Weiland, Kathy Rizzo, and Myriam Ibarra.

I am grateful to everyone who took the time to respond to my requests for

information regarding the use of habitat manipulation in snake conservation –

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including John MacGregor, Boaz Shacham, Philip deMaynadier, Pat Gregory, Craig
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Kostrzewski, Geoffrey Sorrell, Juan Pleguezuelos, Stephen Friet, Scott Harp, Jennifer

Moore, Rebecca Christoffel, Rich Baker, Bob Zappalorti, Bruce Kingsbury, Wade
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Sherbrooke, Scott Gillingwater, Dave Roberts, Mark Howery, Dale Sparks, John Lamb,

Corey Anderson, Tom Anton, Josh Kapfer, Jaime Edwards, Rich Sadjak, Jeff Powers,
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Okla Thornton, and Kristin Stanford.

Thanks to all members of the Gibbs lab group from 2005-2007, for making work

both fun and productive. Thanks to all my ESF friends for making my time in Syracuse

so unforgettable. Finally – to my Mom, Dad and my entire extended family: your love

and support means the world to me.

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 TABLE OF CONTENTS

ACKNOWLEDGMENTS...........................................................................................................ii
TABLE OF CONTENTS............................................................................................................iv
LIST OF TABLES .......................................................................................................................vi
LIST OF FIGURES ................................................................................................................... vii
LIST OF APPENDICES.......................................................................................................... viii
ABSTRACT .................................................................................................................................ix
PREFACE ...................................................................................................................................... x
GENERAL INTRODUCTION.................................................................................................. 1
CHAPTER 1: HABITAT MANIPULATION AS A VIABLE STRATEGY IN SNAKE
CONSERVATION ...................................................................................................................... 4
INTRODUCTION..................................................................................................................... 4
Manipulation of Targeted Habitat Features......................................................................... 7
Basking sites .......................................................................................................................... 7

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Retreat sites ........................................................................................................................... 9
Hibernation sites................................................................................................................. 11
Foraging sites ...................................................................................................................... 14
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Nesting habitat.................................................................................................................... 14
Manipulation of habitat features as a viable conservation strategy ........................... 15
Manipulating the seral stage of natural communities ...................................................... 16
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Reversing vegetative succession ...................................................................................... 16
Promoting vegetative succession: herbivore exclusion ................................................ 21
Manipulation of vegetation communities as a viable conservation strategy ............ 22
Habitat manipulation from a landscape perspective........................................................ 22
Dispersal habitat................................................................................................................. 23
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Road-crossing structures................................................................................................... 24
Managing land-cover diversity ........................................................................................ 27
Managing patch size .......................................................................................................... 28
CONCLUSIONS ..................................................................................................................... 29
Moving towards evidence-based conservation ............................................................. 30
Developing an appropriate management plan .............................................................. 30
Monitoring management success..................................................................................... 32
Information dissemination................................................................................................ 33
SUMMARY.............................................................................................................................. 34
TABLES.................................................................................................................................... 36
FIGURES.................................................................................................................................. 44
CHAPTER 2: BASKING-SITE ENHANCEMENT AS A VIABLE CONSERVATION
STRATEGY FOR A THREATENED POPULATION OF MASSASAUGA
RATTLESNAKES IN NEW YORK STATE.......................................................................... 49
ABSTRACT.............................................................................................................................. 49
INTRODUCTION................................................................................................................... 50

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 METHODS .............................................................................................................................. 54
Study areas .......................................................................................................................... 54
Visual surveys for snakes.................................................................................................. 55
Measuring environmental temperature.......................................................................... 56
Characterizing Te at basking sites and random sites .................................................... 57
Transforming hourly Te data into metrics of thermal quality ..................................... 58
Measuring environmental characteristics at basking sites and random sites .......... 60
Analysis ............................................................................................................................... 60
RESULTS.................................................................................................................................. 63
Response of massasaugas to environmental temperature variation .......................... 63
Contrasting environmental characteristics between basking sites and random sites
............................................................................................................................................... 64
Contrasting basking sites at Cicero Swamp and Bergen Swamp................................ 65
Using microhabitat structure to simulate environmental temperature and crypsis at
Cicero Swamp ..................................................................................................................... 66

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Habitat management as an optimization problem........................................................ 67
DISCUSSION .......................................................................................................................... 68
Management implications................................................................................................. 70
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TABLES.................................................................................................................................... 74
FIGURES.................................................................................................................................. 80
LITERATURE CITED............................................................................................................... 91
APPENDICES .......................................................................................................................... 104
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APPENDIX 1: Datasheet for habitat characterization, 2006 field season..................... 104
APPENDIX II: Datasheet for rattlesnake observations, 2006......................................... 105
APPENDIX III: Summary of vital statistics of all massasaugas observed in 2006,
Cicero Swamp ....................................................................................................................... 106
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APPENDIX IV: Summary of vital statistics of all massasaugas observed in 2006,

Bergen Swamp ...................................................................................................................... 110
VITA .......................................................................................................................................... 111

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 LIST OF TABLES

CHAPTER 1

Table 1: Summary of relevant literature: reviewing the published documentation of

habitat manipulation in snake conservation.............................................pp.36-40

Table 2: Summary of anecdotal reports: reviewing the verbal and email responses to
our requests for information.......................................................................pp.41-43

CHAPTER 2

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Table 1. List of environmental variables measured in this study......... p.74

Table 2. Differences in environmental characteristics between basking sites and random

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sites..................................................................................................................p.75

Table 3. Comparison of candidate models (paired logistic regression) explaining

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environmental differences between basking sites and random sites at Cicero
Swamp.............................................................................................................p.76

Table 4. Contrasting basking sites at Bergen Swamp with basking sites at Cicero
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Swamp.............................................................................................................p.77

Table 5. Temperature models: building a model to simulate the effects of vegetation

management on average daytime Te in the burn area at Cicero
Swamp.............................................................................................................p.78

Table 6. Crypsis models: building a model to simulate the effects of vegetation

management on crypsis potential in the burn area at Cicero Swamp....p.79

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 LIST OF FIGURES

CHAPTER 1

Figure 1. Schematic representations of hypothetical temperature/crypsis tradeoffs in

three different habitats.............................................................................pp.44-45

Figure 2. Case study of the Australian broad-headed snake (Hoplocephalus

bungaroides)..................................................................................................p.46

Figure 3. Case study of the eastern massasauga rattlesnake (Sistrurus c.

catenatus)......................................................................................................p.47

Figure 4. Schematic: evidence-based wildlife management................p.48

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CHAPTER 2
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Figure 1. Maps of habitat areas surveyed at Cicero Swamp and Bergen
Swamp..........................................................................................................p.80
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Figure 2. The relationship between observed body temperature of basking snakes and
operative temperature at basking sites ................................................. p.81

Figure 3. Comparison of predicted and observed operative temperatures (Te) during a

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typical calibration period and validation period..................................p.82

Figure 4. The “model massasauga” used to estimate crypsis index for basking sites and
random sites...............................................................................................p.83

Figure 5. Contrasting average environmental temperature between Cicero Swamp and

Bergen Swamp for each major shared habitat category......................p.84

Figure 6a,b: Simulated average daytime Te and estimated crypsis at Cicero Swamp,
modeled as a function of canopy closure at ground level...............pp.85-86

Figure 7a-c. Basking site enhancement as an optimization problem: setting appropriate

management targets for massasauga rattlesnakes at Cicero
Swamp.....................................................................................................pp.87-90

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 LIST OF APPENDICES

APPENDIX I Datasheet: habitat characterization, 2006..................p.104

APPENDIX II Datasheet: snake observations, 2006..........................p.105

APPENDIX III Summary of vital statistics for all snakes observed at Cicero
Swamp, 2006................................................................pp.106-109

APPENDIX IV Summary of vital statistics for all snakes observed at Bergen

Swamp, 2006..................................................................p.110

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SHOEMAKER, KEVIN T. Habitat manipulation as a viable strategy for the conservation
of the massasauga rattlesnake in New York State.
Typed and bound thesis, 111 + x pages, 8 tables, 11 figures, 2007

ABSTRACT
Habitat manipulation is seemingly gaining acceptance in snake conservation. To
understand whether artificial hibernacula, prescribed fire and other such management
interventions have been effective in the recovery of snake populations, I reviewed the
available literature. Finding little evidence for management success, I caution against
the unquestioned use of habitat manipulation in snake conservation. To gain insight
into the effectiveness of basking-site enhancement as a management strategy for snakes,
I evaluated the selection of basking habitat by eastern massasauga rattlesnakes
(Sistrurus c. catenatus) at two locations in New York State – one of which is threatened
by woody plant succession. Although snakes at the heavily vegetated location – Cicero
Swamp – selected the warmest microhabitats available to them, the average

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temperature at basking sites was substantially lower than at an open-canopy reference
location – Bergen Swamp. Therefore, artificial enhancement of basking habitat will
likely improve the conservation status of the massasauga rattlesnake at Cicero Swamp.
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Key Words: snake conservation, habitat management, eastern massasauga rattlesnake
(Sistrurus c. catenatus), thermoregulation, operative temperature (Te), ecological
succession, crypsis, evidence-based conservation
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Author’s name in full: Kevin Thomas Shoemaker

Candidate for degree of: Master of Science Date: May 2007
Major Professor: James P. Gibbs, Ph.D.
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Department: Environmental and Forest Biology

State University of New York, College of Environmental Science and Forestry, Syracuse,
New York

Signature of Major Professor: _______________________________

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 PREFACE

This thesis document includes two manuscripts prepared for publication. Chapter 1 is a

review of the available literature on the use of habitat manipulation in snake

conservation, researched and written in collaboration with Drs. Glenn Johnson (SUNY-

Potsdam) and Kent A. Prior (Parks Canada). This manuscript, entitled “Habitat

manipulation as a viable strategy in snake conservation” has been conditionally

accepted for publication in a volume entitled “Snakes: Ecology and Conservation”,

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edited by Stephen J. Mullin and Richard A. Seigel. Chapter 2 is a research manuscript

evaluating the need for vegetation management to improve habitat for massasauga
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rattlesnakes at Cicero Swamp in New York. This manuscript was prepared for

publication as a contributed research paper in the journal “Ecological Applications”.

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My research was conducted in conjunction with a general survey of the

massasauga populations at Cicero Swamp and Bergen Swamp. These surveys were
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designed and conducted by Alex T. Krofta – a masters student at Clark University – and

myself. This research has been documented and submitted as a masters thesis (by Alex

Krofta) to Clark University. Alex and I intend to submit this thesis document (in

revised form) for publication in the Journal of Herpetology.

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 GENERAL INTRODUCTION

The eastern massasauga rattlesnake (Sistrurus c. catenatus Raf.) persists as two isolated

populations in the state of New York. One population is located in Cicero Swamp,

about 21 km northeast of the city of Syracuse, and the other population is located at

Bergen Swamp, between the cities of Rochester and Buffalo and 164 km to the west of

Cicero Swamp. Together, these populations represent the eastern edge of the

geographic range of the eastern massasauga rattlesnake and of the Sistrurus genus

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(Wright and Wright 1957). The preservation of these two rattlesnake populations is a

key conservation priority for the state of New York.

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A prairie-adapted subspecies, the massasauga rattlesnake is generally associated

with early-successional habitat (Johnson et al. 2000). Such habitat may enhance the
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ability of massasaugas and other reptiles to access solar radiation for effective basking

(Huey 1982). In addition, recently cleared areas may provide reptiles with enhanced
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foraging opportunities (Mushinsky 1987), improved nesting-site quality (Burger and

Zappalorti 1986, Shine et al. 2002), and may help reptiles meet other essential needs

such as dispersal (e.g., Brisson et al. 2003). For live-bearing species such as the

massasauga rattlesnake, access to open-canopy basking sites may be especially

important; several studies have suggested that body temperature regulation by gravid

females may be a key determinant of health and survival for both mother and offspring

(Reinert 1984, Peterson et al. 1993, O'Donnell and Arnold 2005). Previous research at

Cicero Swamp indirectly suggests that gravid female massasauga rattlesnakes rely on

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early-successional habitat for thermoregulation. Radio-tagged gravid female

massasaugas at Cicero Swamp tended to select open-canopy microhabitats with lower

stem density and shorter-stature shrubs than random points (Johnson 1995). Body

temperature records from temperature-sensitive radiotransmitters indicated that gravid

female massasaugas (n = 3) maintained body temperatures 5 ºC higher than non-gravid

snakes (n = 12).

Woody plant succession has been implicated as a key threat to massasauga

rattlesnakes across its range (Johnson et al. 2000). At Cicero Swamp, the primary

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massasauga habitat is a formerly open-canopy shrubland that has been slowly reverting
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to a closed-canopy state after an 1892 fire (Johnson and Breisch 1993). The loss of open-

canopy habitat at Cicero Swamp may be responsible for a decline in this population
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over the last several decades (Johnson and Breisch 1993). As woody plants have re-

colonized the swamp, basking sites may have become a critical limiting resource;
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however, this “thermal limitation” hypothesis has not yet been tested at Cicero Swamp

– and has rarely been tested for any reptile population (but see Webb et al. 2005).

Some wildlife professionals in New York State have called for aggressive habitat

management to reverse woody plant succession at Cicero Swamp (Johnson and Breisch

1993). Management action has been hampered by a lack of information on both the

status of the massasauga population at Cicero Swamp and the relative merits of

alternative management schemes. I initiated this research project to assess the case for

basking-site limitation at Cicero Swamp and to assist managers in setting informed

management goals.

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 The population of massasauga rattlesnakes at Bergen Swamp (located in New

York State between the cities of Rochester and Buffalo) has generally received less

attention than its counterpart at Cicero Swamp. I expanded my research project in 2006

to evaluate the status of this virtually unstudied population. The primary massasauga

habitat at Bergen swamp – an open-canopy sedge fen interspersed with clusters of short

(<0.5 m) shrubs – contrasts sharply with the heavily wooded shrubland habitat at

Cicero Swamp. As a result, basking habitat at Bergen Swamp is a useful benchmark

with which to compare basking habitat at Cicero Swamp. The results of my evaluation

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of the need for habitat management at Cicero Swamp (using Bergen Swamp as a
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benchmark) are presented in Chapter 2: “Basking site enhancement as a viable

conservation strategy for a threatened population of massasauga rattlesnakes”.

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Investigating the need for vegetation management at Cicero Swamp led me to

explore whether prescribed fire and other types of vegetation management have
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improved the conservation status of massasaugas at other locations. Finding little

documented evidence, I expanded my search to evaluate the use of vegetation

management for the conservation of other snake species. This literature review

expanded further when I was asked to co-author a chapter on habitat manipulation for

publication in an upcoming volume on snake conservation. The results of this broad

literature review are presented in Chapter 1: “Habitat manipulation as a viable strategy

in snake conservation”.

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 CHAPTER 1

HABITAT MANIPULATION AS A VIABLE STRATEGY IN SNAKE

CONSERVATION

INTRODUCTION

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Fifteen years ago, in a volume otherwise focused on snake ecology and behavior, R.A.

Seigel and J.T. Collins (1993) saw fit to include a chapter on snake conservation (Dodd
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1993: "Snake Conservation Strategies"). In his treatment of the topic, Dodd bemoaned
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the unquestioned acceptance of habitat manipulation practices such as conservation

corridors and road-crossing structures: “there is an urgent need to evaluate what are
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rapidly becoming accepted ... management techniques.”. Do conservation corridors

facilitate the movement of snakes through fragmented landscapes? How wide must we

build corridors to ensure that snakes will use them? Can we reduce overwinter

mortality by creating artificial hibernacula? Can artificial nesting or gestation habitat be

used to improve reproductive success? In an effort to answer such questions, we

review the available literature on habitat manipulation in snake conservation.

Habitat manipulation is often uncritically embraced as a practical management

“fix”. Its particular appeal may lie in the hope that habitat functions (e.g., the ability to

support viable snake populations) might simply be restored through direct

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manipulation of habitat remnants: no net loss, everybody wins – ecological costs and

benefits apparently optimized without harmful social or economic consequences.

However, the question remains: is habitat manipulation effective in improving the

conservation status of snakes, or would scarce financial resources be better spent

protecting existing habitat? Habitat manipulation projects may not function as

managers intend, and may in fact harm target species. Improperly designed habitat

alterations may present threats that snakes are evolutionarily unequipped to detect,

functioning as an “ecological trap”(Kolbe and Janzen 2002, Schlaepfer et al. 2002). Even

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habitat manipulation projects that are ecologically benign can potentially drain
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resources from more effective conservation strategies.

Our goals were to: 1) review the use of habitat manipulation in snake
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conservation, 2) evaluate the extent to which habitat manipulation has been successful

in achieving conservation goals, and 3) make recommendations regarding the use of

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habitat manipulation in future snake conservation endeavors. We hope that our results

will provide managers, planners, and field practitioners with a framework for making

informed habitat management decisions. Further, we hope to highlight knowledge

gaps and profitable applied research opportunities, the investigation of which should

lead to improved conservation practice.

We considered three broad categories of habitat manipulation: (1) manipulation

of targeted habitat features (e.g., basking sites, hibernacula), (2) manipulation of the

seral stage of natural communities (e.g., prescribed burning), and (3) manipulation of

ecological landscapes (e.g., increasing landscape connectivity). Published studies were

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identified by searching the ISI Web of Knowledge database, the online meta-search

engine, Google Scholar, and the bibliographies of relevant publications. To obtain

additional published materials, unpublished reports, and anecdotal reports,

information was broadly solicited by: 1) posting requests on relevant electronic mailing

lists (including the Partners in Amphibians and Reptile Conservation listserv), 2)

contacting state-employed herpetologists and non-game wildlife specialists (USA), and

3) contacting university scientists currently conducting snake conservation research in

the United States and abroad. Only studies that quantitatively evaluated the response

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of snakes to habitat manipulation were included in our review. Due to methodological
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variation and differences in types of data reported, meta-analysis of the rate of

management success or of the factors influencing management success (i.e., a systematic

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review sensu Gates 2002) was deemed impractical.

We were able to locate 34 published studies directly relevant to our three broad
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categories of habitat manipulation (Table 1). In addition, our general requests for

information yielded 14 relevant anecdotes and unpublished reports (Table 2). The

majority of studies (n = 22) investigated the response of snakes to large-scale vegetation

management (e.g., logging, prescribed fire). We also reviewed several studies of the use

of herbivore-exclusion fencing and road-crossing structures in snake conservation. The

response of snakes to manipulation of targeted habitat features (e.g., artificial

hibernacula, retreat sites and basking sites) was not well-documented in the literature.

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 Manipulation of Targeted Habitat Features

In general, snake habitat needs can be partitioned into five categories: basking, retreat,

hibernation and aestivation, feeding, and nesting.

Basking sites

Reptilian thermoregulation is so closely associated with physiological function (e.g.,

shedding, digestion, locomotion) that it may serve as a fundamental underlying driver

of habitat selection in many snake populations (Reinert 1993, Shine and Madsen 1996).

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Open-canopy basking habitat can be critically important for many snakes, especially

larger species (Stevenson 1985a). The need for open-canopy basking habitat is highly
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dependent on physiological state; basking behavior often accompanies digestion,

shedding, and gestation in snakes (Peterson et al. 1993, Reinert 1993).

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Thermoregulation is so important to gravid females of many ovoviviparous species that

they spend the majority of the gestation period within open-canopy “gestation” habitat
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(Reinert and Kodrich 1982, Brown 1993, Parker and Prior 1999). While at their chosen

gestation sites, gravid female snakes generally engage exclusively in basking behavior,

forfeiting opportunities for other essential behaviors such as feeding (Keenlyne and

Beer 1973, Seigel and Ford 1987, Seigel et al. 1987). Finally, many snakes bask

extensively upon emergence from hibernation. This behavior may be instrumental for

the completion of spermatogenesis in some species (Gregory 1982).

To facilitate thermoregulation, snakes tend to favor basking habitats that provide

low-cost access to a wide thermal gradient (Spellerberg 1975, 1988). For instance, black

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rat snakes have been shown to prefer forest/field ecotones, ostensibly because these

ecotones provide a wide gradient of thermal environments over a short distance

(Blouin-Demers and Weatherhead 2001a). Although canopy openness is generally a

prerequisite for basking habitat, snakes often prefer to bask in or near some form of

shaded cover (Nilson et al. 1999, Burger and Zappalorti 1988). Sparsely vegetated rocky

outcrops and talus slopes may afford a greater potential for both thermoregulation and

retreat than other available habitats, and are commonly used by many snake species

(e.g., Brown et al. 1982, Parker and Prior 1999).

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Declines and even extirpations of snake populations may be linked to loss of
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basking habitat. Anecdotal records suggest that an endangered population of

massasauga rattlesnakes in New York State has declined as basking habitat has reverted
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to a closed-canopy state after an 1892 fire (Johnson and Breisch 1993). One of us (KTS)

recently assessed the case for habitat manipulation (canopy removal) at this site. Using
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physical operative temperature models (sensu Peterson et al. 1993), this study showed

that basking massasaugas generally selected the warmest microhabitats available to

them. However, average temperatures at these basking sites were substantially

(approx. 3 ºC) lower than at sites selected by massasaugas at an open-canopy

benchmark location (K.T. Shoemaker, unpublished data). Habitat management to

improve basking habitat is therefore likely to prevent the further decline of this

endangered snake population.

Artificial basking habitat may consist of simple clear-cut patches or artificial tree-

fall gaps within a forested matrix (Schmidt and Lenz 2001, Gregory in press). Many

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reptiles are well-suited to take advantage of small canopy gaps (Vitt et al. 1998), and

may even follow tiny sun-flecks across a forest floor (Huey 1982). In addition, snakes

are not known to be territorial, and individuals of many species bask communally

(Gillingham 1987, Gregory et al. 1987). Efforts to create or improve basking habitat for

snakes should therefore focus more on the quality and strategic location of basking sites

than on the size of manipulated areas.

In some cases, vegetation removal may be insufficient to create optimal basking

habitat. As noted above, habitat heterogeneity can be important for effective

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thermoregulation; homogenization of basking habitat is probably undesirable in most
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cases. In addition, poorly-planned vegetation management may reduce habitat quality

for sedentary species that rely on crypsis as a predator-avoidance mechanism (e.g.,

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Graves 1989). Since vegetation management can improve microhabitat warmth at the

expense of crypsis potential, wildlife managers must be careful to set appropriate

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management targets for cryptic species. Quantifying the effect of vegetation removal

on crypsis potential (e.g., photographic analysis, visual estimation) as well as on

average environmental temperature may help in developing informed management

targets (Figure 1).

Retreat sites

Many snake species spend much of their activity periods within retreat sites (Huey et al.

1989, Whiting et al. 1997, Whitaker and Shine 2003, Pearson et al. 2005, Sherbrooke

2006). Retreat sites are generally used by snakes to hide from potential predators

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(Webb and Whiting 2005) and to gain shelter from extreme temperatures (Huey et al.

1989). Access to burrows and rock crevices can be important for thermoregulation

because these sites are often more thermally stable than surface habitat (Huey et al.

1989). In lieu of basking, some snakes access solar energy by selecting large flat rocks in

open, sunny places (colloquially termed "snake rocks") as retreat sites (Webb and Shine

2000). Snakes may also use retreat sites to protect themselves from desiccation (Clark

1970, Whiles and Grubaugh 1993), to lay eggs, and to forage (Whiles and Grubaugh

1993, Webb and Shine 2000, Russell et al. 2004). Because many snakes spend much of

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their time stationary within retreat sites, selecting a proper retreat site can improve
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fitness substantially (Webb and Shine 1998a, Kearney 2002, Pringle et al. 2003, Webb et

al. 2004).
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Artificial burrows have been successfully used to benefit the endangered pygmy

blue-tongue lizard (Tiliqua adelaidensis) in Australia (Milne and Bull 2000, Souter et al.
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2004). However, similar examples are difficult to find in the snake literature (Table 1).

Research in Australia suggests that artificial retreat sites (Webb and Shine 2000) and the

restoration of retreat-site habitat (Webb et al. 2005) may be important in the

conservation of the endangered broad-headed snake (Hoplocephalus bungaroides, Figure

2}. In one experiment, canopy cover was removed locally over several individual

overgrown rocks formerly occupied by broad-headed snakes. Open-canopy rocks

averaged about 10.3 ºC warmer than untreated rocks and were re-colonized by broad-

headed snakes and their primary prey species (the velvet gecko, Oedura lesueurii),

whereas untreated rocks remained unoccupied (Webb et al. 2005). In Texas, the

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endangered Concho watersnake (Nerodia harteri paucimaculata) makes extensive use of

retreat sites within rocky shoreline habitat. Because these microhabitats can be in short

supply during high water levels, the creation of an elevated rocky shoreline may

improve the conservation status of this snake (Whiting et al. 1997). Artificial rocky riffle

habitats have been constructed to benefit the Concho watersnake; apparently, snakes

have taken readily to these artificial riffles, and the riffles have been cited as a

management success (O. Thornton, pers. comm.).

In general, the placement of artificial rock piles may be an important means of

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improving retreat habitat for snakes, especially those adapted to talus slopes
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(Herrington 1988, Schmidt and Lenz 2001). Although not its primary purpose, riprap

(large stones and boulders used to stabilize waterways and prevent erosion) is
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reportedly used by snakes for basking and retreat in some locations (Herrington 1988,

Perry et al. 1996, Wylie et al. 2002). Strategically placed brush-piles may similarly serve
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to create valuable retreat habitat for snakes (Frier and Zappalorti 1983, Seymour and

King 2003).

Hibernation sites

Hibernacula are a special and particularly important type of retreat site for snakes in

temperate climates. Winter kill and other hibernation-related losses represent some of

the most important documented sources of mortality for snakes (Gregory 1982, Shine

and Mason 2004), suggesting that hibernation habitat may be a critically limiting

resource for many populations. Choosing a proper hibernaculum is exceptionally

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important for temperate snakes, as a poor choice is almost certainly fatal (Reinert 1993).

To be suitable, potential hibernacula must 1) provide protection from freezing

temperatures (Bailey 1949), 2) maintain relatively cool temperatures to reduce wasteful

metabolic expenditures (Goris 1971), 3) provide protection from desiccation, 4) provide

protection from predation (Burger et al. 1992), 5) provide access to an adequate supply

of oxygen (Gillingham and Carpenter 1978, Shine and Mason 2004) and 6) remain free

of molds and other pathogens (Goris 1971).

For populations that hibernate communally, hibernation-site improvement may

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be a relatively cheap, simple, and effective management strategy (Shine and Mason
IE
2004). For example, the red-sided garter snake (Thamnophis sirtalis parietalis) hibernates

communally by the thousands within limestone caverns in central Canada. Observing

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high overwinter mortality at some hibernation sites, researchers suggested that levee

banks could be erected to protect hibernacula against flooding, and that insulation of
PR

hibernacula may be used to protect snakes against freezing temperatures (Shine and

Mason 2004).

Many snake species make use of human-made structures such as building

foundations and sewer lines as hibernacula (Zappalorti and Reinert 1994, Seymour and

King 2003), raising the intriguing possibility that artificial hibernacula might be created

to benefit wild snakes. Needless to say, it can be very difficult to ensure that artificial

structures meet all the criteria of a suitable hibernaculum (listed above). In fact, several

attempts at creating artificial hibernacula have failed due to one or more critical

violations of these criteria (Bailey 1949, Goris 1971). Although failures documented in

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the literature generally involve captive snakes forced to use created hibernacula (see

Goris 1971), wild snakes may also be threatened by poorly-designed structures.

Artificial snake hibernacula may have promise as a management technique.

Man-made hibernacula effectively decreased overwinter mortality from nearly 100% to

approximately 10% at a commercial snake farm in Japan (Goris 1971). At this site,

hibernacula were created by filling a shallow hole with layers of gravel for drainage,

boulders to provide cavities large enough to house hibernating snakes, and packed soil

for insulation (Goris 1971). Another design used for captive snakes was a drained,

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concrete-lined hole partially stacked with concrete blocks and covered with soil (created
IE
primarily for a behavioral study). After a pump was installed to improve drainage at

the study site (high mortality was observed in the first year of the study due to
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flooding) overwinter mortality fell to around 15% (Gillingham and Carpenter 1978).

Artificial hibernacula similar to those described above were constructed for free-
PR

ranging northern pine snakes (Pituophis m. melanoleucus) in New Jersey. Many pine

snakes, as well as individuals of other species, have been documented to use these

structures (Zappalorti and Reinert 1994). The creation of artificial hibernacula was not

accompanied by rigorous monitoring of key population viability indicators (Zappalorti

and Reinert 1994), however, and the conservation success of these structures remains

unclear.

In general, there is little evidence demonstrating the effectiveness of artificial

hibernacula in snake conservation (Table 1). This is of concern because the construction

of artificial hibernacula is seemingly gaining acceptance as a management strategy for

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