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SR Timber Rattlesnake 2001 Eng

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COSEWIC

Assessment and Status Report


on the

Timber Rattlesnake
Crotalus horridus
in Canada

EXTIRPATED
2001
COSEWIC status reports are working documents used in assigning the status of wildlife species
suspected of being at risk. This report may be cited as follows:

Please note: Persons wishing to cite data in the report should refer to the report (and cite the author(s));
persons wishing to cite the COSEWIC status will refer to the assessment (and cite COSEWIC). A
production note will be provided if additional information on the status report history is required.

COSEWIC. 2001. COSEWIC assessment and status report on the Timber Rattlesnake Crotalus horridus
in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. vi + 24 pp.
(www.sararegistry.gc.ca/status/status_e.cfm)

Smith, Kim. 2001. COSEWIC status report on the Timber Rattlesnake Crotalus horridus in Canada, in
COSEWIC assessment and status report on the Timber Rattlesnake Crotalus horridus in Canada.
Committee on the Status of Endangered Wildlife in Canada. Ottawa. 1-24 pp.

For additional copies contact:

COSEWIC Secretariat
c/o Canadian Wildlife Service
Environment Canada
Ottawa, ON
K1A 0H3

Tel.: (819) 997-4991 / (819) 953-3215


Fax: (819) 994-3684
E-mail: COSEWIC/COSEPAC@ec.gc.ca
http://www.cosewic.gc.ca

Également disponible en français sous le titre Ếvaluation et Rapport de situation du COSEPAC sur le crotale des bois (Crotalus
horridus) au Canada

Cover illustration/photo:
Timber Rattlesnake — ©Illustration by Marisa Bonofiglio, Woodbridge, Ontario.

©Her Majesty the Queen in Right of Canada, 2011.


Catalogue No. CW69-14/111-2002E-IN
ISBN 0-662-31880-3

Recycled paper
COSEWIC
Assessment Summary

Assessment Summary – May 2001

Common name
Timber Rattlesnake

Scientific name
Crotalus horridus

Status
Extirpated

Reason for designation


The Timber Rattlesnake once occupied much of the Niagara Escarpment and other regions of southern Ontario, but
has not been seen in the province since 1941 despite intensive searches and its easy identification.

Occurrence
Ontario

Status history
Designated Extirpated in May 2001. Assessment based on a new status report.

ii
COSEWIC
Executive Summary

Timber Rattlesnake
Crotalus horridus

Description

The Timber Rattlesnake is a large venomous rattlesnake, characterized by yellow


or brown colouring with dark brown or black V-shaped bands across the back. Adult
sizes range from approximately one to two metres. The head is triangular with a distinct
“neck”. As with all pit vipers, there is an opening visible between the eye and the nostril.
Males are much longer and heavier than the females. Timber Rattlesnakes can be
confused with Massasauga rattlesnakes and some non-venomous species, such as the
fox snake. All non-venomous snakes lack the facial pit of the rattlesnakes. The
Massasauga rattlesnake has nine large scales on its head, whereas the Timber
Rattlesnake has many small scales on its head.

Distribution

This species was last sighted in Canada in 1941. Historically, the snake ranged
throughout southern Ontario and southern Quebec. As there have been no recorded
observations of the Timber Rattlesnake in almost 60 years, it is assumed not to exist in
Canada.

Habitat

The ideal habitats for this rattlesnake are forested areas with rocky outcrops, dry
ridges and second growth coniferous or deciduous forest. The winter habitat is primarily
the den, which is located on a south-facing rocky outcrop. In summer, the preferred
habitat for males and non-pregnant females is a well-canopied forest with thick
vegetation, whereas pregnant females prefer a more open forest with many fallen logs
and a warmer climate. The estimated area needed for a viable population is 50 km2.

General Biology

The Timber Rattlesnake is a “sit and wait” predator. The preferred prey is rodents
and other small mammals; however, the snakes also eat carrion, reptiles, amphibians,
and birds, and their eggs. Female snakes usually do not eat during gestation.

iii
The snakes mate in late summer, with the birth of 5-13 snakelings occurring from
late August to mid-September. The females reproduce once about every three years.
Male snakes mature at a mean age of 4 years, while females mature at a mean age of 6
years, depending on the location of the population. They can live up to 25 years,
reaching their adult size by about 4-5 years.

Timber Rattlesnakes are heliothermic, meaning that they regulate their


temperature through daytime basking. Though they are mostly active by day, during a
hot summer they may also be active at night. The mean summer temperature of a
Timber Rattlesnake is 26.9˚C and the mean winter (hibernation) temperature is 10.0˚C.
In the northern part of their range, they hibernate from September to April (an average
of 7.4 months) in communal dens. They are very loyal to their den site and will return
year after year. They are seasonally migratory — from the den site to the summer
habitat and back again. Males migrate further than females, likely to search for mates.

The rattle is the most obvious behaviour of these snakes, apparently used when
the individual feels angry or threatened. In general, they are very mild-mannered and
will not strike unless provoked. There is only a single recorded incident in Canada of a
Timber Rattlesnake bite resulting in death -- a soldier in the Battle of Lundy’s Lane in
1814. Male snakes engage in “combat dances” with other males to determine
dominance.

Population Size and Trends

There are no known populations of the Timber Rattlesnake in Canada and the last
sighting was almost sixty years ago. It is therefore assumed that the size of the
Canadian population of Timber Rattlesnakes is zero.

Limiting Factors and Threats

Populations of Timber Rattlesnakes are limited in the northern parts of their range
by a small number of suitable nest sites. The snake is slow to mature, has few
snakelings in each litter and a low juvenile survival rate. Taken together, these factors
result in a small number of new individuals added to the population each year.
Historically, they have been subject to human exploitation, such as bounty hunting,
collection and sport hunting. Today, road accidents and human destruction of habitat
(either directly or indirectly), and deliberate killing of venomous snakes all contribute to
the Timber Rattlesnake’s population reduction.

iv
Existing Protection

Each of the US states in which this snake is found affords the species some
degree of protection, however, the level varies from state to state. The last state lifted
the bounty on Timber Rattlesnakes in 1971. In Ontario, the Natural Heritage Information
Center determined that the Timber Rattlesnake is extirpated, the chance of rediscovery
being very small. In addition, it is designated as endangered by both the Ontario
Ministry of Natural Resources’ Committee on the Status of Species at Risk in Ontario
and the Ontario Endangered Species Act.

v
COSEWIC MANDATE
The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) determines the national status
of wild species, subspecies, varieties, and nationally significant populations that are considered to be at risk in
Canada. Designations are made on all native species for the following taxonomic groups: mammals, birds,
reptiles, amphibians, fish, lepidopterans, molluscs, vascular plants, lichens, and mosses.

COSEWIC MEMBERSHIP
COSEWIC comprises representatives from each provincial and territorial government wildlife agency, four federal
agencies (Canadian Wildlife Service, Parks Canada Agency, Department of Fisheries and Oceans, and the Federal
Biosystematic Partnership), three nonjurisdictional members and the co-chairs of the species specialist groups.
The committee meets to consider status reports on candidate species.

DEFINITIONS
Species Any indigenous species, subspecies, variety, or geographically defined population
of wild fauna and flora.
Extinct (X) A species that no longer exists.
Extirpated (XT) A species no longer existing in the wild in Canada, but occurring elsewhere.
Endangered (E) A species facing imminent extirpation or extinction.
Threatened (T) A species likely to become endangered if limiting factors are not reversed.
Special Concern (SC)* A species of special concern because of characteristics that make it particularly
sensitive to human activities or natural events.
Not at Risk (NAR)** A species that has been evaluated and found to be not at risk.
Data Deficient (DD)*** A species for which there is insufficient scientific information to support status
designation.

* Formerly described as “Vulnerable” from 1990 to 1999, or “Rare” prior to 1990.


** Formerly described as “Not In Any Category”, or “No Designation Required.”
*** Formerly described as “Indeterminate” from 1994 to 1999 or “ISIBD” (insufficient scientific information
on which to base a designation) prior to 1994.

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) was created in 1977 as a result of
a recommendation at the Federal-Provincial Wildlife Conference held in 1976. It arose from the need for a single,
official, scientifically sound, national listing of wildlife species at risk. In 1978, COSEWIC designated its first species
and produced its first list of Canadian species at risk. Species designated at meetings of the full committee are
added to the list.

The Canadian Wildlife Service, Environment Canada, provides full administrative and financial support to the
COSEWIC Secretariat.

vi
COSEWIC Status Report
on the

Timber Rattlesnake
Crotalus horridus
in Canada

2001

i
TABLE OF CONTENTS

INTRODUCTION............................................................................................................. 3
Taxonomy.................................................................................................................... 4
History of the Timber Rattlesnake in Canada .............................................................. 4
DISTRIBUTION ............................................................................................................... 5
North America.............................................................................................................. 5
Canada ........................................................................................................................ 6
HABITAT ......................................................................................................................... 6
Habitat Requirements .................................................................................................. 6
Habitat Trends ............................................................................................................. 7
Habitat Protection ........................................................................................................ 7
GENERAL BIOLOGY ...................................................................................................... 8
Food Habits ................................................................................................................. 8
Reproduction ............................................................................................................... 8
Growth and Survivorship ............................................................................................. 9
Physiology ................................................................................................................. 10
Specializations for Defence ....................................................................................... 11
Hibernation ................................................................................................................ 11
Movement and Migration ........................................................................................... 12
Behaviour .................................................................................................................. 12
Vulnerability ............................................................................................................... 13
POPULATION SIZES AND TRENDS............................................................................ 14
Population Size.......................................................................................................... 14
Population Distribution, Persistence and Trends ....................................................... 14
LIMITING FACTORS AND THREATS .......................................................................... 15
SPECIAL SIGNIFICANCE OF THE SPECIES .............................................................. 15
EXISTING PROTECTION ............................................................................................. 16
EVALUATION AND PROPOSED STATUS................................................................... 16
TECHNICAL SUMMARY............................................................................................... 17
ACKNOWLEDGEMENTS ............................................................................................. 19
LITERATURE CITED .................................................................................................... 19
THE AUTHOR............................................................................................................... 24
AUTHORITIES CONSULTED ....................................................................................... 24
Canada ...................................................................................................................... 24
United States ............................................................................................................. 24

List of figures
Figure 1. Distribution of the Timber Rattlesnake (Crotalus horridus) in North America.
From Conant and Collins, 1991....................................................................... 3
Figure 2. Distribution of the Timber Rattlesnake (Crotalus horridus) in Ontario. From
Oldham and Weller, 2000................................................................................ 5

ii
INTRODUCTION

The Timber Rattlesnake, Crotalus horridus, is the only wide-ranging woodland


rattlesnake of the deciduous forest biome of eastern North America (Brown, 1993) (See
Fig. 1) The most common colour phases in the northern parts of its range are termed
‘yellow’ and ‘black’, because the dorsal pattern consists of dark brown or black,
V-shaped crossbands on a yellow, brown or black ground colour (Schmidt and Davis,
1941). Variations of these colour phases occur in the western and southern portions of
its range (see canebrake rattlesnake, below)(Conant and Collins, 1991). Striped
mutations and amelanistic partial albinos have also been recorded (Hudson and Carl,
1985; Dundee, 1994a). Although some authors believe that colour phases are sex
related (Ditmars, 1907), others have suggested that the variation is a thermoregulatory
adaptation, as black specimens occur with greatest frequency in mountainous regions
(Schaeffer, 1969). Colour phases of newborns are readily distinguished after the first
moult (Ibid.). Newborn Timber Rattlesnakes have yellow on the lateral and ventral
surfaces of their tails, but are not thought to use their tails as “lures”, as do other
juvenile snakes with similar colouration (Neill, 1960).

The head is roughly triangular and abruptly distinct from the neck (Anderson,
1965). The facial or loreal pit, which is present between the eye and the nostril on all pit
vipers, functions in detecting the body heat of endothermic prey (Schmidt and Davis,
1941). The pupil of the eye is always vertically elliptical in the pit vipers, a feature
associated with nocturnal habits (Ibid.).

Figure 1. Distribution of the Timber Rattlesnake (Crotalus horridus) in North America. From Conant and Collins,
1991.

3
The pit and the vertical nature of the pupil are characteristics that can be used to
distinguish the Timber Rattlesnake from similarly patterned non-venomous snakes,
such as the eastern fox snake (Elaphe gloydi)(Harding, 1997). To discriminate between
timber and Massasauga rattlesnakes (Sistrurus catenatus), the number and size of the
scales on the dorsal surface of the head can be informative. The Timber Rattlesnake
has many small scales, whereas the Massasauga has nine large scales (Ibid.)

Taxonomy

Until recently, southern populations of the Timber Rattlesnake were widely


recognized as a separate subspecies, the canebrake rattlesnake (C. h. atricaudatus)
(Schmidt and Davis, 1941; Anderson, 1965; Martof et al., 1980). Although the
taxonomic status of the Timber Rattlesnake remains somewhat ambiguous in the
eastern portion of its range (Brown and Ernst, 1986), it is generally accepted that the
subspecific status of the canebrake rattlesnake is not warranted (Behler and King,
1996) and that Timber Rattlesnakes are monotypic (Collins and Knight, 1980).

Other common names include American viper, bastard rattlesnake, black


rattlesnake, common (timber) rattlesnake, eastern rattlesnake, great yellow rattlesnake,
mountain rattlesnake, mountain timber rattler, North American (horrid) rattlesnake,
Northern banded rattlesnake, northern rattlesnake, pit viper, rock rattlesnake, velvet tail,
yellowish brown rattlesnake and yellow rattlesnake (Wright and Wright, 1957). The
specific Latin name horridus means “dreadful”, in reference to the venomous nature of
the Timber Rattlesnake (Collins and Knight, 1980).

History of the Timber Rattlesnake in Canada

The earliest record of Timber Rattlesnakes in what is now Ontario dates back to
September 1669, where there is a reference in the journal of Rene de Brehart Galinée,
who was attached to M. de La Salle’s party (Logier, 1939). The locality was near an
Indian village called Otinaoustettaoua, which is near present-day Waterdown, in Halton
County (Ibid.). Another early account of a Timber Rattlesnake was of one collected by
Captain G. Wilkinson near the end of Point Pelee in September 1818 and reported by
Patch (1919, cited in Logier, 1925). It was “an old specimen measuring 56 inches in
length … as this one was such an old specimen and as no others have been taken
there in recent years, Mr. Patch was of the opinion that it was the last of its race”.
Another possibility was that it merely swam or drifted across Lake Erie from a
neighbouring population on one of the U.S. islands or mainland (F. Cook, pers. comm.).
Relict populations of Timber Rattlesnakes were known to have persisted on South Bass
Island and on the Catawba peninsula of Ohio until at least 1951 (Langlois, 1951).
Historical records are widely scattered and anecdotal and do not provide a clear picture
of the species’ former abundance and distribution (Cook, 1999; see also Fig. 2).

4
Figure 2. Distribution of the Timber Rattlesnake (Crotalus horridus) in Ontario. From Oldham and Weller, 2000.

DISTRIBUTION

North America

At the time of European settlement, the Timber Rattlesnake was found in 30 states
and was extremely abundant in areas of suitable habitat (Casper and Hay, 2001). By
the early 1970’s, the Timber Rattlesnake had been nearly extirpated in all but the most
remote sections of the United States (Morris, 1974). The species is still found in
Alabama, Arkansas, Connecticut, District of Columbia, Delaware, Florida, Georgia,
Iowa, Illinois, Kansas, Kentucky, Louisiana, Massachusetts, Maryland, Minnesota,
Mississippi, Missouri, North Carolina, New Hampshire, New Jersey, New York, Ohio,
Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, Vermont,
Wisconsin and West Virginia (Casper and Hay, 2001; Harding, 1997). They have been
extirpated from the states of Maine and Rhode Island, and may be close to extirpation in
New Hampshire (Brown, 1993). Timber Rattlesnakes are one of only a handful of North
American rattlesnakes that are found east of the Mississippi River (Schmidt and Davis,
1941; Morris, 1974). See Figure 1 for the North American range.

5
Canada

The Timber Rattlesnake is no longer found anywhere in Canada. The last Timber
Rattlesnake sighting was in the Niagara Gorge, Ontario in 1941 (Ontario Herpetofaunal
Summary [OHS], unpubl. data; Cook, 1999). Historically, the Timber Rattlesnake has
been reported from the counties of Essex, Halton, Kent, Elgin, Bruce, Peel, Niagara,
Welland, Hamilton-Wentworth and the Manitoulin District in Ontario (Logier and Toner,
1961; OHS, unpubl. data). There have also been scattered reports of Timber
Rattlesnake sightings in extreme southern Quebec along the U.S. border. However,
none of these reports has ever been fully substantiated, and the adjacent populations in
New York State (which may have acted as a source population for Quebec) have been
extirpated (see Cook, 1999 and Mélançon, 1950 for more detailed summaries of
Quebec reports). See Figure 2 for the Timber Rattlesnake’s range in Ontario.

HABITAT

Habitat Requirements

Timber Rattlesnakes prefer areas not frequented by people (Ditmars, 1907;


Anderson, 1965), although few such sites still exist (DeGraaf and Rudis, 1983). Ideal
habitats are forested areas with rocky outcroppings, dry ridges and second growth
coniferous or deciduous forests (Ibid.). Favourite basking rocks are used year after
year, and may attract many snakes at one time (Harwig, 1966).

The most important habitat component of northern Timber Rattlesnakes is the


communal den within which hibernation takes place. A population is defined by its den,
which is critical to the survival of the entire population (Brown, 1993). Granitic
escarpments and ledges with accumulations of talus are common features of den sites
(Brown, 1991), the majority of hibernacula being found on south-facing slopes (Galligan
and Dunson, 1979). Other habitat components are the summer habitat, where snakes
move and forage, and transient habitat located in between summer habitat and the den
(Brown, 1993).

Specific summer habitat requirements differ according to sex and age class. Males
and non-gravid females utilize forest habitat with greater than 50% canopy closure, thick
surface vegetation and few fallen logs (Reinert and Zappalorti, 1988b). In contrast,
gravid females prefer less densely forested areas with 25% canopy closure, equal
proportions of vegetation and leaf litter on the forest floor, frequent fallen logs and
warmer climatic conditions (Ibid.). An area of approximately 50 km2 of suitable habitat is
required to sustain a population (Brown, 1993).

6
Habitat Trends

In a letter to Francis Cook dated 15 September 1963, Frank Darroch described the
changes to the habitat where he collected the last known Timber Rattlesnake in Ontario
in 1941(Cook, 1999): “The place where I found the snake has in the last ten years been
entirely destroyed as a habitat, by the new road put in for the new hydro power plant.”
Thus, the persistence of Timber Rattlesnakes in that area of the Niagara region seems
highly unlikely.

Optimal forest composition and management strategies for Timber Rattlesnake


habitat remain a subject of debate because canopy cover is a significant factor in the
temperature profile of a den site. According to some researchers, the natural
progression of forest succession may in fact be incompatible with the long-term survival
of Timber Rattlesnake dens if the forest cover results in too much shade (Brown, 1993).
Others doubt the negative impacts of increased shading, and believe that selective tree
removal as a management strategy may actually do more harm than good (Reinert,
pers. comm. to Brown, 1993).

Habitat Protection

One seemingly harmless, yet potentially significant example of habitat alteration is


the repositioning of basking and shelter rocks by people looking for Timber
Rattlesnakes (Brown, 1993). Continuous disturbance of this sort may eventually cause
snakes to abandon these sites altogether (Ibid.). It is preferable instead to check for
snakes under rocks using mirrors and flashlights in order to minimize potential
disturbance (Harwig, 1966).

The remote areas preferred by Timber Rattlesnakes are becoming increasingly


less ideal because of enhanced access to such areas via four-wheel-drive and off-road
vehicles (Galligan and Dunson, 1979; Brown, 1993). Protected public lands and
privately owned nature preserves have contributed significantly to the conservation of
Timber Rattlesnakes where they still exist (Ibid.), although some public lands may have
been managed with the goal of eliminating Timber Rattlesnakes because of fears that
their presence might deter visitors (Cook, 1999).

7
GENERAL BIOLOGY

Food Habits

The foraging behaviour of Timber Rattlesnakes was studied in detail in


Pennsylvania (Reinert et al., 1984). The predominant ambush position of this sit-and-
wait predator involves coiling adjacent to a fallen log with the head positioned
perpendicular to the long axis of the log. Small mammals are the favoured prey;
different species are consumed in proportion to their prevalence at the site. White
footed mice (Peromyscus leucopus) and red-backed voles (Clethrionomys gapperi)
comprised the bulk of the diet of one sample, at 65% and 20% respectively (Ibid.). An
eastern chipmunk (Tamias striatus), a cottontail rabbit (Sylvilagus floridanus) and an
unidentified bird were also taken, each representing 5% of total prey consumed in the
above study. Similarly in Virginia, the three leading foods were mice, chipmunks and
cottontail rabbits (Uhler et al., 1939). Census data of white-footed mice around a
Pennsylvania rattlesnake den revealed a density estimate of 61 mice/ha (Galligan and
Dunson, 1979). Other studies also support the contention that the Timber Rattlesnake
consumes small mammals almost exclusively (Schmidt and Davis, 1941). However,
carrion, reptiles, amphibians, birds and their eggs are also eaten (Uhler et al., 1939;
Myers, 1956; Anderson, 1965; Keenlyne, 1972). Ironically, the food habits of the Timber
Rattlesnake make it an economically valuable species which, were it not for its
venomous nature, might actually have been encouraged to set up residence in Ontario
(Logier, 1939; Martof et al., 1980).

Keenlyne (1972) studied the sexual differences of feeding habits of Timber


Rattlesnakes in Wisconsin. Gravid females did not contain any food items, while 30.3%
and 35.0% of males and non-gravid females, respectively, contained at least one food
item. During follicle development, females either ate more, or were more efficient at
converting food to stored visceral fat. The cessation of feeding was apparently induced
by the development of large offspring. Observations on gravid females in captivity also
indicated that the snakes did not feed during gestation (Odum, 1979).

Reproduction

Male Timber Rattlesnakes reach maturity at an average age of 5.3 years in


northeastern New York (Aldridge and Brown, 1995) and 4 years in South Carolina
(Gibbons, 1972). Mean age of first reproduction for females averaged 7.8 years in
northwestern Virginia (Martin, 1993), 6 years in South Carolina (Gibbons, 1972), four
years in Kansas (Fitch, 1985) and 9-10 years in northeastern New York (Brown, 1991).

8
In the northern parts of their range females reproduce on average every three
years (Brown, 1993; Martin, 1993), with 10-75% of females in a population reproducing
in any given year (Galligan and Dunson, 1979). Fat stores are extremely diminished in
post-partum females (Gibbons, 1972), with a mean mass difference of 306 g between
gravid and post-partum females in a New York population (Brown, 1991). This weight
loss represents 41% of female body weight (Galligan and Dunson, 1979). Foregoing
reproduction in some years is apparently necessary to allow females to gain weight and
store yolk protein in the developing eggs (Galligan and Dunson, 1979; Brown, 1981).
Foraging success and the ability of females to regain mass may be the main
determinant of the length between successive reproductive events (Brown, 1991).

Ovulation occurs from late May to early June, while maximum spermatogenesis is
reached in July, continuing through September (Aldridge and Brown, 1995; Martin,
1993). Mating takes place in late summer (Martin, 1993) and young are born from late
August to mid-September (Galligan and Dunson, 1979). Opinion varies as to whether
females give birth at den sites or at ‘maternity rocks’ some distance away (Galligan and
Dunson, 1979). Typical litter size varies from five to thirteen young, according to
geographic location (Edgren, 1948; Anderson, 1965; Galligan and Dunson, 1979;
Brown, 1993) and gestation period can vary as much as four to six weeks, depending
on weather (Martin, 1996). Parturition of a brood of ten took 4½ hours to complete, with
the birth of each snakeling taking between five and 25 minutes (Trapido, 1939). Sex
ratio in a New Jersey brood was 1:1 (Odum, 1979).

Timber Rattlesnakes produce stillborn young and abort infertile eggs at a


frequency of about 20% (W.S. Brown, pers. comm. to Fitch, 1985). Of a litter of twelve
born to a New York snake, one was stillborn and an infertile egg was also deposited
(Stewart et al., 1960). Similarly, in a New Jersey litter of 13 born in captivity, one
snakeling was born dead, and another had a birth defect that would prevent it from
eating (Odum, 1979).

Male Timber Rattlesnakes may use scent trailing to locate receptive females
(Reinert and Zappalorti, 1988b). During courtship, the male repeatedly strokes the neck
region of the female with his chin (Anderson, 1965).

Growth and Survivorship

Contrary to popular belief, rattlesnakes cannot be aged directly by counting the


number of segments in the rattle. This is true for two reasons; first, a segment is added
each time a snake sheds (not necessarily once a year) and second, segments are
continually being broken off – most wild-caught individuals have between five and nine
segments in their rattles (Schmidt and Davis, 1941). If the rattle becomes accidentally
lost, the end of the tail remains blunt, never pointed (Ibid.). Zoos have reportedly been
known to attach additional rattles on the end of broken ones to make them appear more
impressive (Ditmars, 1907; Schmidt and Davis, 1941).

9
Average shedding rate of Timber Rattlesnakes in northern New York was 1.44
sheds per year (Brown, 1988), compared to two sheds per year in Kansas (Fitch, 1985).
Sheds can be measured directly by painting the rattle and then counting unpainted
segments upon recapture (Brown, 1991). Most rattle growth occurs within the first four
sheddings, and the increase in diameter of successive segments is less than 5% after
the seventh ecdysis (Fitch, 1985).

Timber Rattlesnake life history is characterized by delayed maturity, low


reproductive potential and high longevity (Fitch, 1985). In a Kansas population, mortality
was estimated at 45% in first-year young, and 25% annually thereafter (Ibid.). The
dynamics of this population suggests a rapid turnover, with newly matured adults
comprising a high proportion of the total population. Other studies have also indicated
that juvenile mortality is probably high (Odum, 1979). The natural lifespan of the Timber
Rattlesnake in the northern part of its range is approximately 25 years (Brown, 1993).
A Louisiana Timber Rattlesnake lived 36 years, 7 months and 27 days in captivity,
reaching a total length of 1770 mm (Cavanaugh, 1994).

In Pennsylvania, newborns ranged from 220-280 mm snout-vent length (SVL)


(Galligan and Dunson, 1979). They increased in size to 400-550 mm SVL in their third
year, 640-740 mm in their fourth year and 760-900 mm SVL by their fifth summer (Ibid.).
Timber Rattlesnakes in South Carolina reached lengths of 650-750 mm SVL by the end
of their second summer (Gibbons, 1972). Growth rates and size of newborns appear to
be highly variable geographically, and Brown (1991) presents a table summarizing both
growth rates and minimum age of first reproduction of Timber Rattlesnakes throughout
their range. Males grow considerably longer and heavier than females, and there is no
apparent reproductive advantage for females obtaining a larger body size (Gibbons,
1972). Growth rate slows after maturity (Galligan and Dunson, 1979), and adult total
length ranges from 900-1890 mm (Conant and Collins, 1991).

Physiology

The Timber Rattlesnake is a heliothermic species, with the ability to regulate its
temperature by radiation absorption throughout its daytime activities (Odum, 1979).
Approximately half of a Timber Rattlesnake’s daily activity cycle is spent basking, in
order to maintain its preferred body temperature (Ibid.). Temperatures of free-living New
York Timber Rattlesnakes ranged from 12.5ºC to 33.3ºC from early June to early
August (Brown et al., 1982). Mean body temperature during this time was 26.9ºC (Ibid.).
During hibernation in the same population, mean body temperatures from September
through May ranged from 4.3ºC to 15.7ºC, with a mean value of 10.5ºC (Brown, 1982).

10
Specializations for Defence

Timber Rattlesnakes are venomous. The severity of a snakebite depends on the


amount of venom injected, the toxicity of the venom, and the depth and location of the
bite (Minton, 1953). Fatal bites often involve small children (Guidry, 1953), who are
more sensitive to venom due to their smaller size. Death from a Timber Rattlesnake bite
can occur in as little as 35 minutes (Hutchinson, 1929), particularly if the individual is
allergic to pit viper venom (Parrish and Thompson, 1958). Symptoms of Timber
Rattlesnake poisoning include swelling, pain, respiratory difficulty, weakness, giddiness,
haemorrhage, weak pulse or heart failure, nausea, vomiting, ecchymosis, heart pain,
gastric disturbance, paralysis and unconsciousness or stupor (Hutchinson, 1929).
Freshly decapitated rattlesnakes are still dangerous, as muscles can contract to inflict a
deadly bite (Ditmars, 1907). Recently milked rattlesnakes are also capable of producing
serious bites, and there is no evidence that previously exposed victims develop any sort
of immunity through antibody production (Parrish and Thompson, 1958).

Bites from rattlesnakes at the time of shedding and replacement of fangs indicate
that the injection apparatus may not be fully functional at this time, as some victims
bitten during this period did not exhibit any symptoms of poisoning (Hutchinson, 1929).
The first shedding of maxillary fangs takes place at a very early age, as evidenced by
the fact that newborns have been found with fangs in the functional position (Barton,
1950). Venom is clear and watery in newborns, becoming bright yellow and
concentrated as the snake matures (Johnson et al., 1968).

Hibernation

Hibernation in the northern parts of the Timber Rattlesnake’s range occurs in the
cracks of rocky ledges, usually facing south (Odum, 1979). In Ontario, hibernacula were
most often located along the Niagara escarpment, following the limestone outcropping
northward toward the Manitoulin District (Logier and Toner, 1961).

Timber Rattlesnakes often hibernate with copperheads (Agkistrodon spp.) and


other snakes in the United States, due to the scarcity of hibernacula (Galligan and
Dunson, 1979; DeGraaf and Rudis, 1983; Conant and Collins, 1991). They exhibit high
fidelity to hibernacula (Odum, 1979; DeGraaf and Rudis, 1983), and some snakes follow
specific routes to and from hibernacula each season (Brown et al., 1982). Hibernation is
almost always communal, with only scattered reports of individuals hibernating singly
(Neill, 1948; Odum, 1979). Timber Rattlesnakes near the northern extent of their range
typically hibernate for 7.4 months, with the active period extending from early May
through late September (Brown, 1991). Spring emergence is likely affected, but not
completely determined, by temperature (Galligan and Dunson, 1979). Additionally,
some snake hunters claim that Timber Rattlesnakes will not leave the den until they
have shed (Ibid.).

11
Laboratory and field studies suggest that newborn Timber Rattlesnakes are able to
follow the scent trails of adults to communal hibernacula (Brown and MacLean, 1983;
Reinert and Zappalorti, 1988a). This ability could be crucial to the survival of newborns
through their first winter, as they may be born a considerable distance away from any
suitable hibernaculum (Reinert and Zappalorti, 1988a).

Movement and Migration

Timber Rattlesnakes are seasonally migratory, from den to summer range and
back (Brown, 1993). A variety of habitats are traversed throughout the active season,
depending partly on individual age and reproductive state. Timber Rattlesnakes may
swim across lakes during their seasonal migrations (Neill, 1948), and small Timber
Rattlesnakes have been found as high as 9 m in trees (Saenz et al., 1996).

Timber Rattlesnakes may be active by day or night; nocturnal activity is especially


common during hot summer nights (Martof et al., 1980). They tend to disperse upslope
to high ridges removed from human settlement when they emerge from hibernation
(Brown, 1981), and move in a looping pattern during the active season that returns
them to the same hibernaculum (Reinert and Zappalorti, 1988b). Female Timber
Rattlesnakes may or may not return to hibernation sites to give birth to young (Galligan
and Dunson, 1979; DeGraaf and Rudis, 1983). Movements of gravid females are
generally confined to thermally optimal gestation sites, such as open outcrop knolls in
the vicinity of the den (Reinert and Zappalorti, 1988b; Brown, 1991).

Migration distances of males in general are greater than those of females because
they actively engage in seeking mates (Reinert and Zappalorti, 1988b). The mean
maximum migratory distance from the den was 4.07 km for males and 2.05 km for
females in northeastern New York (Brown, 1993). The maximum single migratory
movement away from a den was 7.2 km for a male in the same population.
Translocated males have significantly larger activity ranges, range lengths and mean
distances moved per day than native males (Rupert and Reinert, 1992). Timber
Rattlesnakes collected during organized round-ups are often released far away from
their point of capture, and this action most likely leads to the death of the snake,
because of its unfamiliarity with its surroundings (Brown, 1993). Many translocated
snakes immediately leave the area in which they are released (Galligan and Dunson,
1979).

Behaviour

The most distinctive behaviour of rattlesnakes in general is the use of the rattle
itself. Rattles are vibrated sideways at about 48 cycles per second (Schmidt and Davis,
1941). Timber Rattlesnakes generally begin to rattle when approached within 1-2 m
(Barbour, 1950). Rattlesnakes are deaf to the sound of their own rattles, and are
thought to use them when angry or threatened (Ditmars, 1907).

12
Timber Rattlesnakes are considered the most mild-mannered of any of the North
American rattlesnakes, and individuals usually do well in captivity (Ditmars, 1907;
Anderson, 1965; Morris, 1974). In general, they do not strike unless provoked,
preferring instead to remain concealed (Ditmars, 1907). The “combat dance” of male
Timber Rattlesnakes has been interpreted either as an expression of sexuality, or as a
competitive interaction for food or dominance (Sutherland, 1958). This is a shy and
secretive species and reports of Timber Rattlesnakes biting humans are rare (Ditmars,
1907; Brown, 1981). The only Canadian report of a possible Timber Rattlesnake bite,
and subsequent fatality, was that of a soldier bitten during the Battle of Lundy’s Lane
near Niagara Falls in 1814 (see Cook, 1999 for details).

Vulnerability

Clearing of land, killing by humans and commercial exploitation have all


contributed to the decline of the Timber Rattlesnake throughout its range (Weller, 1982;
DeGraaf and Rudis, 1983). Habitat destruction of Timber Rattlesnake sites includes
blasting and filling-in of dens with concrete, logging, mining, and gas wells (Brown,
1993). Because they are large and poisonous, Timber Rattlesnakes engender a large
amount of human hostility (Plourde et al., 1989). The introduction of pigs into the
countryside contributed to the demise of the Timber Rattlesnake; pigs are protected
from envenomation by their thick layer of fat that prevents the venom from entering
circulation, and thus are able to kill and eat rattlesnakes (Nash, 1908). The colonial
nature of Timber Rattlesnakes was a factor contributing to their extirpation from Canada
by humans (Plourde et al., 1989).

Timber Rattlesnakes have been the object of bounty hunting since as early as
1719 (Klauber, 1956, cited by Galligan and Dunson, 1979; Brown, 1981; Casper and
Hay, 1998). In Wisconsin, bounty records showed a 70-80% decline over a seven-year
period (Casper and Hay, 1998). Bounty records from one county in Minnesota declined
from 4,955 in 1980 to 191 in 1987 (Ibid.). A number of populations from New York
appear to have been extirpated, primarily because of over-hunting (Brown, 1981). In
addition to bounty hunting, rattlesnakes have also been harvested for their oil (Ibid.) and
for their rattles (Anderson, 1965). Because of their gregarious nature, hundreds were
sometimes killed in a raid on a single den (Schmidt and Davis, 1941). Clearly, the
potential impacts of human persecution are enormous.

As with most other reptiles, road-kill is an additional aspect of human-induced


mortality (Martin et al., 1992; Dundee, 1994b; Jensen et al., 1994). In a Timber
Rattlesnake population in New York, male road-kills outnumbered female road-kills
3.9:1.0, and human-caused mortality rates in general were much higher for males than
for females (13:1) (Aldridge and Brown, 1995). Conversely, in a study in Georgia, gravid
females constituted the majority of road-killed snakes (Neill, 1948).

13
POPULATION SIZES AND TRENDS

Population Size

All available evidence indicates that the population size of the Timber Rattlesnake
in Canada is zero. The species has not been sighted in Canada in almost 60 years.

Population Distribution, Persistence and Trends

Frank Darroch collected the last confirmed Timber Rattlesnake in the Niagara
Gorge on August 22, 1941 (Cook, 1999). Since this time, many researchers have
conducted searches (including Frank Darroch, E.B.S. Logier, Frank Ross, Craig
Campbell and James Kamstra), but no Timber Rattlesnakes have been found (Ibid.). It
is almost a certainty that the Timber Rattlesnake has been extirpated, as demonstrated
by the following quotes:

1881 Garnier: “rapidly becoming extinct [in Ontario]”


1908 Nash: “formerly common and generally distributed throughout the province
now nearly extinct”
1939 Logier: “that the early distribution of this snake in Ontario was more
extensive seems likely”
1982 Weller: “may very well have been extirpated in Ontario”
1984 Cook: “the last specimen taken in Ontario was from Niagara Glen in 1941”
1989 Johnson: “extirpated from Ontario”
1989 Plourde et al.: “extirpated”
1993 Brown: “probably extirpated [from Ontario]”
1999 Cook: “almost certainly extirpated in Canada”

Population declines have also been noted throughout the Timber Rattlesnake’s
range in the United States, particularly in New York, Pennsylvania, Virginia, Connecticut
and New Jersey (Brown, 1981, 1993; Reinert, 1990). Timber Rattlesnakes once
occurred throughout the Carolinas and Virginia, but they have been eliminated from
areas of extensive deforestation and human settlement (Martof et al., 1980). In
Missouri, the Timber Rattlesnake population is gradually being reduced, and the
species has been practically exterminated in some areas where it was formerly common
(Anderson, 1965). Minor range extensions have been reported relatively recently in
Virginia (Martin et al., 1992), Louisiana (Lutterschmidt, 1992; Dundee, 1994b) and
Florida (Jenson et al., 1994). Of the populations that remain, many have been reduced
to 15-20% of their numbers a few decades ago (Martin, 1983). The minimum viable
population size for Timber Rattlesnake recovery is estimated at 30 to 40 individuals with
an even age distribution and at least four or five mature females per den (Brown, 1993).

14
LIMITING FACTORS AND THREATS

Human exploitation, through bounty hunting, commercial collecting and sport


hunting, is the leading cause of Timber Rattlesnake decline throughout the species’
range (Galligan and Dunson, 1979; Brown, 1993). Over a few decades, a single Timber
Rattlesnake hunter is known to have collected 2900-5000 snakes from New York alone
(Stechert, 1982; Brown et al., 1994). Snake hunters report that it is not difficult to ‘hunt
out’ (i.e. extirpate) an entire den (Galligan and Dunson, 1979). Gravid females are
particularly susceptible to persecution due to their preference for more open habitats
and the predictability with which they occupy specific sites (Brown, 1993). Gravid
individuals comprised 84% of female Timber Rattlesnakes turned in during organized
snake hunts in Pennsylvania (Reinert, 1990 in Brown, 1993).

Several biological traits of the Timber Rattlesnake greatly reduce its ability to
recover from large-scale losses of adults in a population. These features include slow
maturation rates, low reproductive output, low juvenile survival and slow population
replacement rate (Harding, 1997). Because females do not mature until about eight
years old and reproduce on average only once every three years (Martin, 1993), most
females will give birth to no more than five broods throughout their lifetime, assuming a
lifespan of 22 years (Brown, 1991). These demographic characteristics also make the
Timber Rattlesnake a species in which there is little natural “harvestable” surplus of
adult individuals (Brown, 1993). One Utah rattlesnake population heavily hunted for only
one year had still not recovered 12 years later (Woodbury and Hansen, 1950 cited in
Galligan and Dunson, 1979).

SPECIAL SIGNIFICANCE OF THE SPECIES

Timber Rattlesnakes are large, oviviviparous, iteroparous, long-lived and slow to


mature, with a relatively long mean generation time (Brown, 1991). As such, they have
the capacity to contribute significantly to our knowledge of covariation in life history traits
(Ibid.).

Timber Rattlesnakes have been employed as study subjects to test a number of


novel techniques. Sadighi et al. (1995) used remotely triggered cameras to monitor the
occurrence of Timber Rattlesnakes in the wild. Additionally, the population genetics of
Timber Rattlesnakes have been analyzed using microsatellite markers (Bricker et al.,
1996; Villarreal et al., 1996; Bushar et al., 1998).

Timber Rattlesnakes in the Big Black Mountain area of Kentucky play a prominent
part in certain religious rites in the area (Barbour, 1950). A local pastor is said to have
captured a four-foot specimen in his bare hands after chanting the “magic words” (Ibid.).

15
EXISTING PROTECTION

In Ontario, the Timber Rattlesnake has been assigned a rank of ‘SX’ by the Natural
Heritage Information Centre as apparently extirpated from Ontario (Oldham, 1997).
The Ontario Ministry of Natural Resources’ Committee on the Status of Species at Risk
in Ontario (COSSARO) has designated the Timber Rattlesnake an endangered species
under the following criteria: “any native species that, on the basis of the best available
scientific evidence, is at risk of extinction or extirpation throughout all or a significant
proportion of its Ontario range if the limiting factors are not reversed” (Ibid.). The Timber
Rattlesnake was among the first snakes to be officially designated as an endangered
species under Ontario’s Endangered Species Act, 1973 (Weller, 1982).

The Timber Rattlesnake receives varying degrees of protection throughout its


range in the United States. It is an endangered species in Massachusetts, where it may
not be harassed, killed, collected or held in possession except under special permit
(Jackson and Mirick, 1996). Penalties can range as high as a $5,000 fine and/or
imprisonment for 180 days, in addition to a restitution payment of $2,000 per animal
killed (Ibid.). The practice of paying bounties on Timber Rattlesnakes in New York
ended in 1971 under the Fish and Wildlife Law (Brown, 1981). In 1971, Vermont
became the last New England state to remove the bounty on the Timber Rattlesnake
(DeGraaf and Rudis, 1983). According to Casper and Hay (2001), C. horridus is
designated as extirpated in Maine and Rhode Island, endangered in Connecticut,
Massachusetts, New Hampshire, New Jersey, Ohio, Vermont, and Virginia, threatened
in Illinois, Indiana, Minnesota, New York, and Texas, a protected species in Maryland
and Kansas, protected from take in Oklahoma and Pennsylvania and a protected
wild animal in Wisconsin

Since 1996, the Nature Conservancy has ranked the Timber Rattlesnake “G5”
(Oldham, 1997). However, in light of its range-wide decline, the U.S. Fish and Wildlife
Service is conducting a review of the Timber Rattlesnake for possible protection under
the federal Endangered Species Act (Casper and Hay, 2001). Although the Timber
Rattlesnake was proposed for listing under Appendix II of the CITES Convention in
1997, the proposal was not adopted because it was argued that international trade was
minimal, and that the species would benefit more by increasing protection in the United
States (Ibid.).

EVALUATION AND PROPOSED STATUS

The designation of status for the Timber Rattlesnake is quite straightforward: it has
not been reported in this country in almost 60 years. The Timber Rattlesnake has
already been designated as extirpated in Ontario by the Nature Conservancy (Oldham,
1997), and the vast majority of experts agree that the Timber Rattlesnake no longer
exists in Canada. Thus the proposed status for COSEWIC is extirpated.

16
TECHNICAL SUMMARY

Crotalus horridus
Timber Rattlesnake Crotale des Bois

Extent and Area information


• extent of occurrence (km²) 0
• specify trend (decline, stable, increasing, unknown) -
• are there extreme fluctuations in EO (> 1 order of magnitude)? -
• area of occupancy (km²) 0
• specify trend (decline, stable, increasing, unknown) -
• are there extreme fluctuations in area of occupancy (> 1 order -
magnitude)?
• number of extant locations 0
• specify trend in # locations (decline, stable, increasing, -
unknown)
• are there extreme fluctuations in # locations (>1 order of -
magnitude)?
• habitat trend: specify declining, stable, increasing or unknown unknown
trend in area, extent or quality of habitat
Population information
• generation time (average age of parents in the population) ~13 years
(indicate years, months, days, etc.)
• number of mature individuals (capable of reproduction) in the 0
Canadian population (or, specify a range of plausible values)
• total population trend: specify declining, stable, increasing or -
unknown trend in number of mature individuals
• if decline, % decline over the last/next 10 years or 3 -
generations, whichever is greater (or specify if for shorter time
period)
• are there extreme fluctuations in number of mature individuals -
(> 1 order of magnitude)?
• is the total population severely fragmented (most individuals found -
within small and relatively isolated (geographically or otherwise)
populations between which there is little exchange, i.e., < 1
successful migrant/year)?
• list each population and the number of mature individuals in -
each
• specify trend in number of populations (decline, stable, -
increasing, unknown)
• are there extreme fluctuations in number of populations (>1 -
order of magnitude)?
Threats (actual or imminent threats to populations or habitats) [add rows as needed]
- human destruction of nesting habitat through direct action, mining, logging, drilling gas wells.
- - decimation of population through hunting, road-kills, persecution because of discrimination against
venomous snakes.

17
Rescue Effect (immigration from an outside source)
• does species exist elsewhere (in Canada or outside)? yes
• status of the outside population(s)? declining
• is immigration known or possible? no
• would immigrants be adapted to survive here? yes
• is there sufficient habitat for immigrants here? unknown
Quantitative Analysis yes

18
ACKNOWLEDGEMENTS

This report could not have been completed without the assistance of a number of
people. Francis Cook graciously allowed me to include information contained in his
unpublished Timber Rattlesnake account in his upcoming book, “The Natural History of
Amphibians and Reptiles in Canada”. Mike Oldham provided me with pertinent
information from the Ontario Herpetofaunal Summary. Ron Brooks and Glenn Barrett
supplied advice and technical assistance. Funding provided by the Canadian Wildlife
Service, Environment Canada.

LITERATURE CITED

Aldridge, R.D. and W.S. Brown. 1995. Male reproductive cycle, age at maturity, and
cost of reproduction in the Timber Rattlesnake (Crotalus horridus). Journal of
Herpetology 29(3): 399-407.
Anderson, P. 1965. The reptiles of Missouri. University of Missouri Press, Columbia,
Missouri. 300 pp.
Barbour, R.W. 1950. The reptiles of Big Black Mountain, Harlan County, Kentucky.
Copeia 1950: 100-107.
Barton, A.J. 1950. Replacement fangs in newborn Timber Rattlesnakes. Copeia
1950: 235-236.
Behler, J.L. and F.W. King. 1996. National Audubon Society Field Guide to North
American Reptiles and Amphibians. Alfred A. Knopf, New York. 743 pp.
Bricker, J., L.M. Bushar, H.K. Reinert and L. Gelbert. 1996. Purification of high quality
DNA from shed skin. Herpetological Review 27(3): 133-134.
Brown, W.S. 1981. Conserving the Timber Rattlesnake. Conservationist 36: 27-29.
Brown, W.S. 1982. Overwintering body temperatures of Timber Rattlesnakes (Crotalus
horridus) in northeastern New York. Journal of Herpetology 16(2): 145-150.
Brown, W.S. 1988. Shedding rate and rattle growth in Timber Rattlesnakes. American
Zoologist 28(4): 195A.
Brown, W.S. 1991. Female reproductive ecology in a northern population of the Timber
Rattlesnake, Crotalus horridus. Herpetologica 47(1): 101-115.
Brown, W.S. 1993. Biology, status and management of the Timber Rattlesnake
(Crotalus horridus): a guide for conservation. SSAR Herpetological Circular No. 22.
pp. i-iv + 1-78.
Brown, C.W. and C.H. Ernst. 1986. A study of the variation in eastern Timber
Rattlesnakes, Crotalus horridus Linnae (Serpentes: Viperidae). Brimleyana
12: 57-74.
Brown, W.S., L. Jones and R. Stechert. 1994. A case in herpetological conservation:
notorious poacher convicted of illegal trafficking in Timber Rattlesnakes. Bulletin of
the Chicago Herpetological Society 29(4): 74-79.

19
Brown, W.S. and F.M. MacLean. 1983. Conspecific scent-trailing by newborn Timber
Rattlesnakes, Crotalus horridus. Herpetologica 39(4): 430-436.
Brown, W.S., D.W. Pyle, K.R. Greene and J.B. Friedlaender. 1982. Movements and
temperature relationships of Timber Rattlesnakes (Crotalus horridus) in
northeastern New York. Journal of Herpetology 16(2): 151-161.
Bushar, L.M., H.K. Reinert and L. Gelbert. 1998. Genetic variation and gene flow within
and between local populations of the Timber Rattlesnake, Crotalus horridus.
Copeia 1998(2): 411-422.
Casper, G. and R. Hay. 2001. Timber Rattlesnake web page.
http://www.mpm.edu/collect/vertzo/herp/timber/htm.
Cavanaugh, C.J. 1994. Crotalus horridus (Timber Rattlesnake) longevity. Herpetological
Review 25(2): 70.
Collins, J.T. and J.L. Knight. 1980. Crotalus horridus Linnaeus Timber Rattlesnake.
Catalogue of American Amphibians and Reptiles 253.1-253.2.
Conant, R. and J.T. Collins. 1991. Reptiles and amphibians: Eastern and Central North
America (3rd Edition). Houghton Mifflin Company, New York. xxx + 450 pp.
Cook, F.R. 1984. Introduction to Canadian Amphibians and Reptiles. National Museum
of Natural Sciences, National Museums of Canada, Ottawa, Ontario. 200 pp.
Cook, F.R. [1999]. Unpublished Timber Rattlesnake account in “The Natural History of
Amphibians and Reptiles in Canada”.
DeGraaf, R.M. and D.D. Rudis. 1983. Amphibians and reptiles of New England: habits
and natural history. University of Massachusetts Press, Amherst. 85 pp.
Ditmars, R.L. 1907. The reptile book. Doubleday, Page and Company. 472 pp.
Dundee, H.A. 1994a. Crotalus horridus (Timber Rattlesnake) coloration. Herpetological
Review 25(1): 28.
Dundee, H.A. 1994b. Crotalus horridus (Timber Rattlesnake). Herpetological Review
25(1): 33-34.
Edgren, R.A. Jr. 1948. Notes on a litter of young Timber Rattlesnakes. Copeia 1948:
132.
Fitch, H.S. 1985. Observations on rattle size and demography of prairie rattlesnakes
(Crotalus viridus) and Timber Rattlesnakes (Crotalus horridus) in Kansas.
Occasional papers of the Museum of Natural History, University of Kansas
118: 1-11.
Galligan, J.H. and W.A. Dunson. 1979. Biology and status of Timber Rattlesnake
(Crotalus horridus) populations in Pennsylvania. Biological Conservation 15: 13-58.
Garnier, J.H. 1881. List of the Reptilia of Ontario. Canadian Sportsman and Naturalist
1: 37-39. In Cook, 1999 (above).
Gibbons, J.W. 1972. Reproduction, growth, and sexual dimorphism in the canebrake
rattlesnake (Crotalus horridus atricaudatus). Copeia 1972: 222-226.
Guidry, E.V. 1953. Herpetological notes from southeastern Texas. Herpetologica
9: 49-56.

20
Harding, J.H. 1997. Amphibians and reptiles of the Great Lakes region. The University
of Michigan Press, Ann Arbor: MI. xvi + 378 pp.
Harwig, S.H. 1966. Rattlesnakes are where and when you find them. Proceedings of the
Ninth Annual Meeting of the Ohio Herpetological Society 5: 163.
Hudson, R. and G. Carl. 1985. Crotalus horridus (Timber Rattlesnake) coloration.
Herpetological Review 16(1): 28-29.
Hutchinson, R.H. 1929. On the incidence of snake-bite poisoning in the United States
and the results of newer methods of treatment. Bulletin of the Antivenin Institute of
America 3: 43-57.
Jackson, S. and P. Mirick. 1996. Timber Rattlesnake in Massachusetts, web site
http://www.umass.edu/ umext/snake/timber.html.
Jensen, J.B., B.W. Mansell, and P.E. Moler 1994. Crotalus horridus (Timber
Rattlesnake). Herpetological Review 25(4): 166.
Johnson, B. 1989. Familiar amphibians and reptiles of Ontario. Natural Heritage/
Natural History Inc., Toronto. 168 pp.
Johnson, B.D., J. Hoppe, R. Rogers and H.L. Stahnke. 1968. Characteristics of venom
from the rattlesnake Crotalus horridus atricaudatus. Journal of Herpetology
2: 107-112.
Keenlyne, K.D. 1972. Sexual differences in feeding habits of Crotalus horridus horridus.
Journal of Herpetology 6: 234-237.
Klauber, L.M. 1956. Rattlesnakes, Vols. 1 and 2. University of California Press,
Berkeley. 1476 pp. In Galligan and Dunson, 1979 (above).
Langlois, T.H. 1951. Timber Rattlesnakes on the Lake Erie islands. Ohio Conservation
Bulletin 15: 14.
Logier, E.B.S. 1925. Notes on the herpetology of Point Pelee, Ontario. The Canadian
Field-Naturalist 29(5): 91-95.
Logier, E.B.S. 1939. The reptiles of Ontario. Royal Ontario Museum of Zoology,
Handbook No. 4. University of Toronto Press, Toronto, Ontario. 63 pp.
Logier, E.B.S. and G.C. Toner. 1961. Check list of the amphibians and reptiles of
Canada and Alaska (2nd Edition). Contributions of the Royal Ontario Museum Life
Sciences 53: 1-92.
Lutterschmidt, W.I. 1992. Crotalus horridus (Timber Rattlesnake). Herpetological
Review 23(1): 26.
Martin, W.H. 1983. The Timber Rattlesnake in the northeast: its range, past and
present. Herpetological Bulletin of the New York Herpetological Society 17: 15-20.
Martin, W.H. 1993. Reproduction of the Timber Rattlesnake (Crotalus horridus) in the
Appalachian Mountains. Journal of Herpetology 27(2): 133-143.
Martin, W.H. 1996. Crotalus horridus (Timber Rattlesnake) reproductive phenology.
Herpetological Review 27(3): 144-145.
Martin, W.H., J.C. Mitchell, and R. Hoggard 1992. Crotalus horridus (Timber
Rattlesnake). Herpetological Review 23(3): 91.

21
Martof, B.S., W.M. Palmer, J.R. Bailey and J.R. Harrison. 1980. Amphibians and
reptiles of the Carolinas and Virginia. University of North Carolina Press, Chapel
Hill. 264 pp.
Mélançon, C. 1950. Inconnus et Méconnus: Amphibiens et Reptiles de la province de
Québec. La Société Zoologique de Québec.
Minton, S.A . Jr. 1953. Variation in venom samples from copperheads (Agkistrodon
contortrix) and Timber Rattlesnakes (Crotalus horridus). Copeia 1953: 212-215.
Morris, P.A. 1974. An introduction to the reptiles and amphibians of the United States.
Dover Publications, Inc., New York. 253 pp.
Myers, C.W. 1956. An unrecorded food item of the Timber Rattlesnake. Herpetologica
12: 326.
Nash, C.W. 1908. Batrachians and reptiles of Ontario. Pp. 11-15 In Manual of
Vertebrates of Ontario. Department of Education, Toronto, Ontario. L.K. Cameron,
Toronto, Ontario. 229 pp.
Neill, W.T. 1948. Hibernation of amphibians and reptiles in Richmond County, Georgia.
Herpetologica 4: 107-114.
Neill, W.T. 1960. The caudal lure of various juvenile snakes. Quarterly Journal of the
Florida Academy of Sciences pp. 173-200.
Odum, R.A. 1979. The distribution and status of the New Jersey Timber Rattlesnake
including an analysis of Pine Barrens populations. HERP Bulletin of the New York
Herpetological Society 15(1): 27-35.
Oldham, M.J. 1997. Natural Heritage Resources of Ontario: amphibians and reptiles.
Canadian Association of Herpetologists Bulletin 11(2): 3-9.
Oldham, M.J. and W.F. Weller. 2000. Ontario Herpetofaunal Atlas. Natural Heritage
Information Centre, Ontario Ministry of Natural Resources.
http://www.mnr.gov.on.ca/MNR/nhic/herps/ohs.html (updated 15-01-2001).
Ontario Herpetofaunal Summary. Unpublished data provided by Mike Oldham.
Parrish, H.M. and R.E. Thompson. 1958. Human evenomation from bites of recently
milked rattlesnakes: a report of three cases. Copeia 1958: 83-86.
Patch, C.L. 1919. A rattlesnake, melano garter snakes and other reptiles from Point Pelee,
Ontario. The Canadian Field-Naturalist 33(3): 60-61. In Logier, 1925 (above).
Plourde, S.A., E.L. Szepesi, J.L. Riley, M.J. Oldham and C. Campbell. 1989.
Distribution and status of the herpetofauna of central region, OMNR. Parks and
Recreational Areas Section, OMNR, Open File Ecological Report SR8903, Central
Region, Richmond Hill, Ontario. 30 pp.
Reinert, H.K. 1990. A profile and impact assessment of organized rattlesnake hunts in
Pennsylvania. Journal of the Pennsylvania Academy of Sciences 64: 136-144.
Reinert, H.K., D. Cundall and L. Bushar. 1984. Foraging behavior of the Timber
Rattlesnake, Crotalus horridus. Copeia 4: 976-981.
Reinert, H.K. and R.T. Zappalorti. 1988a. Field observation of the association of adult
and neonatal Timber Rattlesnakes, Crotalus horridus, with possible evidence for
conspecific trailing. Copeia 4: 1057-1059.

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Reinert, H.K. and R.T. Zappalorti. 1988b. Timber Rattlesnakes (Crotalus horridus) of the
Pine Barrens: their movement patterns and habitat preference. Copeia 1988(4):
964-978.
Rupert, R. Jr. and H.K. Reinert. 1992. A comparison of movement patterns: native vs.
translocated Timber Rattlesnakes (Crotalus horridus). Journal of the Pennsylvania
Academy of Science 65 (Suppl.): 198.
Sadighi, K., R.M. DeGraaf and W.R. Danielson. 1995. Experimental use of remotely-
triggered cameras to monitor occurrence of Timber Rattlesnakes (Crotalus
horridus). Herpetological Review 26(4): 189-190.
Saenz, D., S.J. Burgdorf, D.C. Rudolph and C.M. Duran. 1996. Crotalus horridus
(Timber Rattlesnake) climbing. Herpetological Review 27(3): 145.
Schaeffer, G.C. 1969. Sex independent ground colour in the Timber Rattlesnake,
Crotalus horridus horridus. Herpetologica 25: 65-66.
Schmidt, K.P. and D.D. Davis. 1941. Field book of snakes in United States and Canada.
G.P. Putnam’s Sons, New York. 365 pp.
Stechert, R. 1982. Historical depletion of Timber Rattlesnake colonies in New York
State. Herpetological Bulletin of the New York Herpetological Society 17 23-24.
Stewart, M.M., G.E. Larson and T.H. Matthews. 1960. Morphological variation in a litter
of Timber Rattlesnakes. Copeia 1960: 336-337.
Sutherland, I.D.W. 1958. The “combat dance” of the Timber Rattlesnake. Herpetologica
14: 23-24.
Trapido, H. 1939. Parturition in the Timber Rattlesnake, Crotalus horridus horridus
Linne. Copeia 4: 230.
Uhler, F.M., C. Cottam and T.E. Clarke. 1939. Food of snakes of the George
Washington National Forest, Virginia. Transactions of the 4th North American
wildlife conference. Pp. 605-622.
Villarreal, X., J. Bricker, H.K. Reinert, L. Gelbert and L.M. Bushar. 1996. Isolation and
characterization of microsatellite loci for use in population genetic analysis in the
Timber Rattlesnake, Crotalus horridus. The Journal of Heredity 87(2): 152-155.
Weller, W. 1982. Rare and endangered reptiles in Ontario (Part I). Canadian Amphibian
and Reptile Conservation Society Bulletin 20(2): 1-8.
Woodbury, A.M. and R.M. Hansen. 1950. A snake den in Tintic Mountains, Utah.
Herpetologica 6(3): 66-70. In Galligan and Dunson, 1979 (above).
Wright, A.H. and A.A. Wright. 1957. Handbook of snakes of the United States and
Canada, Vol. 2. Comstock Publishing Associates, Cornell University Press, Ithaca,
New York. 1105 pp.

Final edit by Ronald J. Brooks and Elinor J. Hughes.

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THE AUTHOR

Kim Smith completed her Honours B.Sc. at the University of Guelph in 1996, with a
specialization in wildlife biology. From 1996 to 1998, she conducted field work on a
variety of taxonomic groups, including marine invertebrates and sea ducks, marine
fishes, reptiles, waterfowl and mammals. She is currently working toward an M.Sc. on
wood turtles with Dr. Ron Brooks at the University of Guelph. Kim has also written
COSEWIC status reports on the queen snake (Regina septemvittata) and the northern
ribbon snake (Thamnophis sauritus septentrionalis).

AUTHORITIES CONSULTED

Canada

Francis Cook, Researcher Emeritus, Canadian Museum of Nature

United States

Howard Reinert, Department of Biology, The College of New Jersey

William S. Brown, Associate Professor Emeritus, Department of Biology

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