(DRAFT) - Taxonomy
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
TAXONOMY
NAME - TORTOISE, GOPHER
OTHER COMMON NAMES - TORTOISE, GOPHER; TURTLE and GOPHER; GOPHER;
ELEMENT CODE -
CATEGORY - Reptiles
PHYLUM AND SUBPHYLUM - CHORDATA,
CLASS AND SUBCLASS - REPTILIA,
ORDER AND SUBORDER - TESTUDINES,
FAMILY AND SUBFAMILY - TESTUDINIDAE,
GENUS AND SUBGENUS - GOPHERUS,
SPECIES AND SSP - POLYPHEMUS,
SCIENTIFIC NAME - GOPHERUS POLYPHEMUS
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Gopher Tortoise
Gopherus polyphemus (Daudin, 1802)
KINGDOM: Animal GROUP: Reptile
PHYLUM: Chordata CLASS: Reptilia
ORDER: Testudines FAMILY: Testudinidae
The gopher tortoise is a large (shell 15-37 cm long), dark brown
to grayish-black terrestrial turtle with elephantine hind feet,
shovel-like forefeet, and a gular projection beneath the head on the
yellowish, hingeless plastron or undershell (01).
Adults 200-400 mm carapace length with low, oval shell sides
tending to be somewhat parallel in large specimens, very flat top,
margins not greatly flared on lateroposterior border; gular
projections obtuse, short, slightly notched anteriorly at midline.
Forelimb ossicles moderately large, flattened, never fused;
antibrachial scales juxtaposed, rarely even slightly keeled. Front
foot flattened and wide: Auxiliary scale trapezoidal or triangular.
Head width 58 to 78 percent of hind foot width. Shell ground color
yellowish-white in hatchlings, darkening to brown to brownish-black
with age sometimes faintly marbled; younger individuals with lighter
areole. Margined scutes usually uniformly dark in adults, except in
Taxonomy - 1 (DRAFT) - Taxonomy
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
western range where often marbled with brownish-yellow. Plastron
usually immaculate yellow, except in western range where it may
possess darker rays or marbling.
Females generally smaller with less concave plastron, shorter
gular projection, and short blunt tail terminating at level of cloaca.
Males longer, more acute tail with terminal enlarged scale.
No subspecies of the Gopher tortoise are recognized and the
species may be separated from other turtles within its range by its
elongate gular projection, small elephantine hind limbs and
shovel-like forelimbs. The species was described in 1802 as Testudo
polyphemus by F.M. Daudin (10).
There were previously considered to be four extant species of
Gopherus; G. agassizi (Desert tortoise), G. berlandieri (Texas
tortoise), G. flavamarinatus (Bolson tortoise), and G. polyphemus
(gopher tortoise) (17). Wermuth and Mertens (12 in 17) considered all
extant Gopherus to belong to a single polytypic species (17).
However, each is considered a distinct species as indicated by
geographic discontinuity of the 4 taxa, behavior and morphological
differences (skull, shell and foot structure) (17).
Recently, the western species G. agassizi and G. berlandieri were
assigned to the genus Xerobates (13) leaving two extant species of the
genus Gopherus including the gopher tortoise. The following is a list
of scientific synonyms, other scientific names and specimens of the
gopher tortoise from Authenberg and Franz, 1978 (10) and Beyer 1900
(08):
Testudo polyphaemus Bartram, 1791:18 Nomen nudum (18);
Testudo polyphemus Daudin, 1802:256 No type specimen designated
(19);
Testudo depressa Guerin-Meneville, 1829:5 No type specimen
designated (20);
Testudo gopher Gray, 1844:4 Two syntypes presumably in the
British Museum (Natural History) (21);
Xerobates carolinus Agassiz, 1857:447 No type specimen
designated (22);
Xerobates polyphemus Beyer, 1897-1899:25-46 (08);
Gopherus carolinus Shaler, 1888:37 (23);
G. polyphemus Stejneger, 1893:161 First use of combination (24);
G. praecedens Hay, 1916:55 Holotype, Florida Geol. (25);
G. polyphemus polyphemus Mertens and Wermuth, 1955:371 (27).
Photographs can be found in Carr, 1952 (14) and Ernst and
Barbour, 1972 (01). Hatchlings are illustrated in Neill and Allen,
1957 (15) and Arata, 1958 (16), and skeletal material in Auffenberg,
1976 (11).
The Gopher tortoise is also commonly called "Gopher" and "Gopher
turtle."
Taxonomy - 2 (DRAFT) - Status
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
STATUS
Coded Status
Alabama; Federal Threatened
Alabama; State Listed
Louisiana; Federally Threatened
Louisiana; State Recognized
Mississippi; State Listed
T: Federal Threatened
Commercial
Commercial/consumption
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The gopher tortoise (Gopherus polyphemus) has been designated a
Threatened species pursuant to the Endangered Species Act of 1973
(50 CFR 17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540), as
amended. The species has this status wherever found west of Mobile
and Tombigbee Rivers in the States of Alabama, Mississippi, and
Louisiana.
This species is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild animal
(alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation, or foreign law; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
USFS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for management/recovery on Forest Service lands. The
Forest Service is responsible for integrating
management, protection, and conservation of federally
listed species into the Forest Planning process
(36 CFR 219.19 and 219.20).
Status - 1 (DRAFT) - Status
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Mississippi
DESIGNATED STATUS: Threatened
ADMINISTRATIVE AGENCY: Mississippi Dept. of Wildlife Conserv.
STATE STATUTE: Public Notice No. 2156, Eff. Sept. 1, 1981.
STATE: Alabama
STATUS: Game Animal (No open season)
ADMINISTRATIVE AGENCY: AL Dept. of Conserv. and Nat. Resources
STATE STATUTE: 1982-83 Annual Hunting Regs. pp. 6, 9.
STATE: Louisiana
STATUS: Recognized Threatened
ADMINISTRATIVE AGENCY: Louisiana Dept. of Wildl. and Fisheries
STATE STATUTE: LA. Stat. Annot. 56:1901 to 56:1907.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
Gopherus polyphemus is on Appendix II of CITES. The gopher
tortoise is also listed as vulnerable by the IUCN, 1982, and under
the common name Florida gopher tortoise in the 1986 IUCN Red List
of Threatened Animals.
ECONOMIC STATUSES:
"Gopher pulling" is not an uncommon practice among rural people
within the range of the western population. Some poor people, as well
as some not so poor, regard them as a significant source of dietary
protein. Occasionally, the species is sold or used as pets. Folklore
in the range of the gopher tortoise generally regards the meat of the
tortoise as having aphrodisiac properties.
82/12/30:47 FR 58454/ - Notice of Review
85/09/18:50 FR 37958/37967 - Notice of Review
86/07/08:51 FR 24723/24727 - Proposed Rule
87/07/07:52 FR 25376/25380 - Final Rule - threatened
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
SOCIETY OF AMERICAN FORESTRY TYPES
SAF TYPE STAGE CLOSURE
Longleaf Pine shrub--seedling
Longleaf Pine young tree
Longleaf Pine mature tree
Longleaf Pine Old Growth
Loblolly Pine-Shortleaf Pine shrub--seedling
Loblolly Pine-Shortleaf Pine young tree
Loblolly Pine-Shortleaf Pine mature tree
Loblolly Pine-Shortleaf Pine Old Growth
Shortleaf Pine-Oak shrub--seedling
Shortleaf Pine-Oak young tree
Shortleaf Pine-Oak mature tree
Shortleaf Pine-Oak Old Growth
LAND USE -
Residential
Transportation, communications, and Util
Mixed Urban or Built-up Land
Cropland and Pasture
Orchards, Groves, Vineyards, Nurseries,
Evergreen Forest Land
Mixed Forest Land
Sandy Areas other than Beaches
Transitional Areas
COMMENTS ON HABITAT ASSOCIATIONS -
Although the distribution of the western population of the
gopher tortoise falls within the longleaf pine belt, it occurs in a
great variety of habitat types, with the common factor in all occupied
habitats being the presence of sandy soils. These sandy soils need to
be about 1 m in depth to best accommodate burrowing. The gopher seems
fairly adaptable within the confines of the sandy soil criterion. The
following general habitat types all contain tortoises in varying
densities: old fields, planted pine, pine nurseries (Desoto Nat.
Forest), pine/oak, scrub oak, along the edges of pastures, corn fields,
clearcuts. in tung oil, peach and pecan orchards along highway
right-of-ways, in highway medians and along fences and hedgerows.
According to Spillers and Speake (03), edge habitat or ecotonal areas
appear important to tortoises, as evidenced by clustering of burrows
near edges. In this assessment, it was concluded that gopher tortoise
density was directly related to the amount of available edge.
The Spillers and Speake assessment (03), determined two
vegetation variables to be statistically correlated with tortoise
density. These variables were basal area and canopy closure. Optimum
values for these parameters based on tortoise density were:
0-2 square meters/ha basal area and approximately 10 percent canopy
closure. They found no definite trend in soil type relating to
tortoise density other than if there was no sandy soil available,
there would be few, if any tortoises.
Natural ecosystems in which gopher tortoises occur in good
densities are on droughty sand ridges normally dominated by longleaf
pine (Pinus palustris) stands with scattered scrub oaks and a
relatively open habitat conditions on the forest floor, maintained by
Habitat Associations - 1 fire. According to relocation experiences, and habitat studies by
Joan Diemer (06), the species requires some low ground cover and will
not normally exist in areas with an improved pasture look.
The traditional habitat of the western population of gopher
tortoises is natural xeric communities, mostly of the longleaf-scrub
oak type located on sand ridges. These natural plant associations are
fire dependent. The tortoise carrying capacity of these habitats is
proportional to the biomass of herbaceous foods such as broad-leaf
grasses (Poaceae) and legumes (Fabaceae) (04).
When these sandy sites are converted from longleaf stands to
slash or loblolly pine, the suppression of fire and consequent woody
understory development occurs along with a concomitant reduction in
fire-climax grasses and forbs, primary foods for the tortoise.
Accordingly, conversion of native longleaf stands often leads to a
decrease in tortoise numbers. As the sandy ridges are also high and
excessively drained, they have been and are preferred areas for road
building activities, which often leads to increasing tortoise
susceptibility to human predation.
Habitat Associations - 2 (DRAFT) - Food Habits
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
FOOD HABITS
TROPHIC LEVEL -
HERBIVORE
LIFESTAGE FOOD FOOD PART
General Arthropods
General Poaceae
General
General Forb Leaves/Stems
General Forb Flowers/Fruit/Seed
General Deciduous Shrubs-Flowers/Fruit/Seed
General Cactus-Flowers/Fruit/Seed
General Lianas-Flowers/Fruit/Seed
Food Habits - 1 (DRAFT) - Environment Associations
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Terrestrial Features: Burrows
G Availability of fence rows: roadside ditches and grassy
G
Environment Associations - 1 (DRAFT) - Life History
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
LIFE HISTORY
FOOD HABITS:
Conat (05) reports that gopher tortoises eat a mixed diet of
grasses, leaves, and wild fruits. Landers et al. (04) conducted a
food habit study of gopher tortoises in the longleaf-scrub oak
association of south Georgia, a habitat very similar to the natural
habitats of the western population. Landers et al. (04) concluded
that grasses and grass-like plants (certain species of Cyperaceae and
Asteraceae) were the principle foods. Carpet grasses (Axonopus
affinis), goose grass (Eleusine indica), side-seed grasses (Paspalum
ssp.), bahia (Paspalum notatum), crab grass (Digitaria sanguinalis),
nut grass (Cyperus esculentus), nutrush (Scleria ciliata), wiregrass
(Aristida stricta) were commonly utilized by adult tortoises in south
Georgia.
Forbs are apparently preferred by juvenile tortoises, with
utilization increasing as the growing season progresses. Legumes are
the most important forbs for tortoises (04). They eat the leaves and
stems. Important spring legumes are sensitive briar (Schrankia
microphylla), hoarypea (Tephrosia spp.), and rattlebox (Crotalaria
angulata). Summer legumes that are preferred by gopher tortoises
include milk-pea (Galactia spp.), butterfly pea (Clitoria marian),
pencil flower (Stylosanthes biflora) and partridge pea (Cassia
nictitans). Another important forb in the diet of gopher tortoises is
poor-Joe (Diodia teres). This plant pioneers in areas with past soil
disturbance.
Morning glory (Ipomoea pandurate) leaves are commonly eaten.
Dyschoriste (Dyschoriste oblongifolia) and braken fern (Pteridium
aquilinum) are major food items in the late spring and summer.
Fleshy fruits are commonly eaten in the summer and early fall.
The following is a list of those most commonly eaten in south Georgia
(04): blackberry (Rubus cuncifolius), sloe plum (Prunus umbellata),
blueberry (Vaccinium spp.), huckleberry (Gaylussacia spp.), prickly
pear (Opuntia spp.), maypop (Passiflora incarnata), muscadine (Vitis
rotundifolia) and hawthorne (Crataegus spp.).
HOME RANGE/TERRITORY:
Landers et al. (04) studied movement patterns and home ranges of
gopher tortoises in southwestern Georgia. This study produced
evidence that a dominance hierarchy based on body size exists in males
and is most often expressed in breeding colonies where densities are
high and social interaction frequent. Dominant (large) males display
agonistic behavior toward smaller males and defend their burrows
against them. The following description of the dominance hierarchy
between males is provided by Ernst and Barbour (01): "When
approaching an intruder, a highly motivated tortoise walks rapidly
with his neck fully extended; under normal circumstances the pace is
slower and the neck is only partially extended. When the tortoises
are about 25 feet apart, head-bobbing begins. At contact the vertical
bobs of both males become more lateral and finally lose any
recognizable pattern. Each sniffs the head and feet of the other.
The vertical bobs associated with these sniffing movements often are
interrupted by a lateral wiping motion across the surface of one of
Life History - 1 (DRAFT) - Life History
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
its forelegs. After 1 or 2 minutes of mutual sniffing, one of the
tortoises becomes more active than the other, and they demonstrate
dominance by their postures. The dominant tortoise supports its
weight on all four of its extended legs. He may walk around the
submissive one, stopping often to smell his rear legs. The submissive
tortoise holds his head low, with his neck partly extended. If the
dominant tortoise continues to investigate, the submissive one
positions himself at an angle of 45 degrees to the dominant one and
maneuvers to keep one side or the other presented to the front of the
dominant individual." Since females are significantly larger than
males, successful burrow defense on their part against males is not
uncommon, and is a part of territoriality.
There seems to be a skewed distribution of sub-adult males to
burrows and activity areas along the periphery of, or between
colonies, with large males utilizing burrows more near the center of
colonies (close to female burrows) and moving freely within colonies
to court several females. It is believed that gopher tortoises are
likely to emigrate from a colony prior to reaching maturity and that
females are non-receptive to nearly all courting displays of
subordinate males and that this may account for the larger breeding
movements of smaller males. The area (territory) where a defense
behavior may be found is near the center of the breeding colony and
nearby burrows in these core areas. Landers et al. (04) believes
territoriality is also exhibited by females in terms of burrow
defense.
According to Landers et al. (04) the mean feeding radius of
adults is generally less than 30 m from the burrow being used. The
feeding areas are non-linear, often resembling and elliptical shape.
Feeding ranges for the summer-fall are significantly larger than those
of spring. This is though to be due to a depletion of preferred foods
around the burrows by late summer. Other movements are related to
social interactions. April through early June is the approximate
breeding period. Home ranges of males are much larger than those of
females; most of the home range of males is used almost exclusively in
mate seeking and periodic relocation of preferred burrows rather than
feeding activity (04). Females do not search for other tortoises
during the breeding season. The mean home range size for telemetered
males in south Georgia was 0.45 ha and 0.22 ha for females (04).
PERIODICITY:
Gopher tortoises' movement is severely restricted during winter
months (Nov.-Feb.). As spring progresses the frequency of activity
increases linearly with gradual warming (04). The species is not
known to exhibit any crepuscular or nocturnal activity. During March
and April, outside burrow activity occurs mainly on warmer days with
most sightings being in the warmest part of the day (1600-1800 h). In
July and August, gopher tortoises studied by Landers et al. (04)
showed a bimodal feeding activity with peaks at mid-morning
(1000-1500 h) and mid-afternoon (1300-1500 h) and this was considered
to be related to avoidance of extreme temperatures. Following the
period of intense summer heat, activity resumed the unimodal pattern
of spring. Thus, the degree of activity throughout the year is
correlated with ambient temperature, with movement from the burrow
Life History - 2 (DRAFT) - Life History
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
being rare at cooler temperatures (<22 degrees Celsius), greatest at
28-31 degrees Celsius, and curtailed at temperatures exceeding 32
degrees Celsius (04).
MIGRATION PATTERNS:
The gopher tortoise is not migratory, but may evidence some
shifts in burrow location, or extension of activity ranges in response
to changing food supplies, colony description due to habitat
alteration, or in the instance of males during the breeding season,
activity ranges may be extended (04).
COVER/SHELTER REQUIREMENTS:
The burrow is the central feature in cover requirements for the
gopher tortoise and is dug in sandy soil. It is mostly dry but at its
deepest point is usually moist. Within the burrow the temperature and
humidity remain relatively constant. The burrow is usually straight
and unbranched and have been reported up to 47.5 feet long (01). At
the end of the burrow is a chamber large enough for the gopher
tortoise to turn around in. Natural ecosystems in which gopher
tortoises occur in good densities are on droughty sand ridges normally
dominated by longleaf pine (Pinus palustris) stands with scattered
scrub oaks and a relatively open habitat conditions on the forest
floor, maintained by fire. According to relocation experiences, and
habitat studies by Joan Diemer (06), the species requires some low
ground cover and will not normally exist in areas with an improved
pasture look
REPRODUCTIVE SITE REQUIREMENTS:
Courtship displays by the male which consists of circling, head
bobbing and biting occur in front of the female (01). The space where
these activities occur and also ultimately copulation, is often
associated with the female burrow but theoretically could occur at any
place a courting male and receptive female meet. Males are often seen
at the entrance to a female burrow doing the head bobbing routine.
Nests are dug in sand which is exposed to the full rays of the sun and
eggs are deposited about 6" below the surface (01). Landers et al.
(04) found nests to be mostly associated with burrow mounds, although
Ernst and Barbour (01) generalized that nesting most often occurred a
considerable distance from the burrow.
REPRODUCTIVE CHARACTERISTICS:
Landers et al. (04) considered 16-21 years to be required for
sexual maturity. This in males means a carapace length of 230-240 mm
and in females a length greater than 255 mm. The activity of
submaxillary glands is directly correlated with courtship and may be
considered an indication of sexual maturity. Females ultimately
control the actual period of mating through their response to male
courtship displays. In the south Georgia study (04) most copulation
occurred from late April through early June, although males continued
to show active submaxillary glands and display throughout the summer
and early fall.
Apparently there is but one breeding period per year and there is
no evidence of a female laying more than one clutch per year (04).
Life History - 3 (DRAFT) - Life History
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
The species seems promiscuous. Clutch size seems to increase with the
size of the female, the south Georgia tortoise (04) averaging 7 eggs
per clutch. The nesting period described by Landers et al. (04) was
mid-May through late June. South Georgia tortoises (04) averaged an
incubation period (laying to hatching) of 102 days.
PARENTAL CARE:
From the time eggs are deposited, no care is given to the nest,
and after hatching, the young receive no parental care (01). Growth
rates are probably similar to other members of the genus Gopherus,
with the first few years evidencing a rapid growth which rapidly
diminishes after 5 or 6 years (01). Females likely grow somewhat
faster than males.
POPULATION BIOLOGY:
Current limiting factors for the gopher tortoise include
conversion of habitat to urban areas, roads, croplands, and pasture
lands. Forest management which allows excessive shading of the forest
floor by sapling pine or because of fire suppression, which allows
excessive woody vegetation to invade the forest floor, cause a loss of
food supply. Human predation remains a threat to this species. The
gopher tortoise is a highly K-selected species. Assuming that all
mature females lay one clutch annually, the fecundity in the south
Georgia population (04) is 7 eggs per year. The annual birth rate
(number born alive) is only a small percentage of the fecundity due to
heavy nest predation. Landers et al. (04) found that the average
female would have a successful clutch only once in 10 years. Since an
average of 1.2 eggs per clutch did not hatch in the Georgia study, the
average hatch is reduced to 5.8, which translates to an effective rate
of reproduction of 5.8 hatchlings per mature female per 10 years.
Abercrombie (07) suggests that gopher tortoise populations may exhibit
reproductive pulses under the right conditions, and that these
occasional (very successful) pulses have a significant role in the
adaptive strategy. Landers et al. (04) estimated typical population
growth at 3-5 percent per year. Mortality in gopher tortoises is high
at the egg stage, often approaching a 90 percent nest loss. Mammals
such as foxes, raccoons, opossums and skunks are known nest predators.
Young hatchlings are also preyed upon by snakes and fire ants.
Abercrombie (07) reports a sex ratio of nearly 1:1 in hatchlings and
in others old enough to be sexed reliably by external characteristics.
In Abercrombie's (07) review of the literature to find population
model characteristics for Gopher polyphemus, sexual maturity was
assumed at 18 years. Longevity is estimated at 40-60 years in south
Georgia (04). Recovery of the western population seems to be
achievable and will be greatly dependent on the future land management
procedures of the U.S. Forest Service in south and southeast
Mississippi. Good adult densities are believed to be in the range of
one per acre (04).
SPECIES INTERRELATIONSHIPS:
Many animals find the burrow of a gopher tortoise a suitable
permanent home or a temporary shelter. Among vertebrates found in the
burrows are skunks, red foxes, raccoons, opossums, rats, rabbits,
Life History - 4 (DRAFT) - Life History
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
quail, burrowing owls, diamondback rattlesnakes, black racers, indigo
snakes, six-lined race-runners, gopher frogs, leopard frogs, and toads
(01). Arthropods also occur as commensals or obligates, including
some 32 species of spiders, ticks, and insects.
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 5 (DRAFT) - Management Practices
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Prescribed/controlled burning of habitat
Beneficial Selective Thinning
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Controlling/Restricting Herbicide Use
Beneficial Transplanting wild animals
Beneficial Restricting Poaching
Adverse Subsistence Hunting/Fishing/Trapping
Existing Subsistence Hunting/Fishing/Trapping
Adverse Collecting
Existing Collecting
Adverse Commercial Exploitation
Existing Commercial Exploitation
Adverse Food Supply Reduction
Existing Food Supply Reduction
Adverse Inherent Reproductive Characteristics
Existing Inherent Reproductive Characteristics
Adverse Predation
Existing Predation
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Highway/Railroads
Existing Highway/Railroads
Adverse
Existing
Adverse Suppressing wildfire
Existing Suppressing wildfire
Adverse Forest Alteration
Existing Forest Alteration
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
Factors which have been detrimental to the gopher tortoise are
not that dissimilar from present threats. Taking gophers for food or
sale remains a cultural practice in rural areas throughout the
gopher's range. The species, unlike most game species, is unable to
withstand significant taking in the short or long term aspects due to
an inherently low reproductive rate (estimated growth of 3-5 percent
per year) (04), and the lengthy period required for reaching sexual
maturity (assumed 18 years (04). Conversion of habitat to urban
areas, roads, croplands, and pasture lands has reduced the western
portion of the historic range by more than 80 percent and
fragmentation of their western range accentuates the impacts (02). In
addition to human predation, road mortality and taking the animals for
pets cumulatively affect gopher tortoise populations.
Forest management practices which result in an overly thick woody
understory, or on the other extreme, a shaded forest floor resulting
from densely spaced young pine, often times result in colony
Management Practices - 1 (DRAFT) - Management Practices
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
disruption and abandonment. Fairly open pine canopies combined with a
lack of fire often results in a thickly woody understory which is not
good gopher tortoise habitat.
The traditional habitat of the western population of gopher
tortoises is natural xeric communities, mostly of the longleaf-scrub
oak type located on sand ridges. These natural plant associations are
fire dependent. The tortoise carrying capacity of these habitats is
proportional to the biomass of herbaceous foods such as broad-leaf
grasses (Poaceae) and legumes (Fabaceae) (04).
When these sandy sites are converted from longleaf stands to
slash or loblolly pine, the suppression of fire and consequent woody
understory development occurs along with a concomitant reduction in
fire-climax grasses and forbs, primary foods for the tortoise.
Accordingly, conversion of native longleaf stands often leads to a
decrease in tortoise numbers. As the sandy ridges are also high and
excessively drained, they have been and are preferred areas for road
building activities, which often leads to increasing tortoise
susceptibility to human predation.
Although State laws in Alabama and Mississippi, and now the
Endangered Species Act, are in place to protect the gopher tortoise,
it is believed that illegal taking will continue to be a threat. In
Mississippi, over the next 30 years, cropland is expected to double,
and pasture land is expected to increase by 40 percent (02).
Additionally, within the tortoise's range these factors are expected
to continue and to adversely affect this species. On the other hand,
on the U.S. Forest Service lands in Mississippi, longleaf pine
management can be adjusted to be beneficial to the gopher tortoise.
UNAPPROVED PLAN:
At the current time, no recovery plan is available for the gopher
tortoise. The initiation date for writing the plan has not been set.
Probably the number one priority to aid in the recovery of the
western population of Gopherus polyphemus is "opening up" degraded
habitats either by prescribed burning or selective thinning of timber
stands to enhance herb growth; prescribed burning is the best means of
achieving that goal. As a close second to habitat enhancement, a law
enforcement strategy needs to be developed between the U.S. Fish and
Wildlife Service, the U.S. Forest Service, and the States of Alabama
and Mississippi which should stem the magnitude of taking.
Populations in marginal degraded and/or isolated habitats may
need periodic regular immigration of individuals from adjacent or
nearby "good" habitat (known as the "rescue affect") (09). In
addition, restricting or controlling development (habitat loss) and
herbicide use in colony areas (this could reduce food supply). Direct
effects of herbicide and pesticide use is uncertain.
Management Practices - 2 (DRAFT) - References
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Ernst, C.H., and R.W. Barbour. 1972. Turtles of the United
States. The Univ. Press of KY, Lexington. 347 pp.
02 U.S. Fish and Wildlife Service. 1986. Endangered and threatened
wildlife and plants; proposal to list the western population of the
gopher tortoise as threatened. Fed. Reg. 51(130):24723-24747.
03 Spillers, D.M., and D.W. Speaks. 1986. Status and distribution of
the Gopher tortoise (Gopherus polyphemus) in southern Alabama.
U.S. Fish and Wildl. Serv., 24 pp.
04 Landers, J.L., et al. 1981. The Gopher tortoise: Distribution
ecology and effects of forest management. Ga. Dept. of Nat. Res.
421 pp.
05 Conant, R. 1975. A field guide to reptiles and amphibians of
eastern and central North America, 2nd Ed. Houghton Mifflin Co.,
Boston. 429 pp.
06 Diemer, J. 1987. Pers. comm. Fla. Game and Freshwater Fish
Comm., Research Lab., Gainesville, FL.
07 Ambercrombie, C.L. A simulation model to the management of the
gopher tortoise (Gopherus polyphemus) and gopher tortoise habitat.
Unpubl. MS. 70 pp.
08 Beyer, G.E. 1900. Louisiana herpetology. Proc. Louisiana Soc.
Natur. 1897-1899:25-46.
09 U.S. Fish and Wildlife Service. 1987. Endangered and threatened
wildlife and plants; final rule for the western population of the
gopher tortoise as threatened. Fed. Reg. 52:25376-25380
10 Auffenberg, W. and R. Franz. 1978. Gopherus polyphemus.
Catalogue of American Amphibians and Reptiles. Soc. for the Study
of Amphib. and Rept. pp. 215.1-215.2.
11 Auffenberg, W. 1976. The genus Gopherus (Testudinidae): Part I
Osteology and relationships to extant species. Bull. Fla. State
Mus., Biol. Sci. 20(2):47-110.
12 Wermuth, H. and R. Mertens. 1961. Schildkroten, Krokodile,
Bruchenechsen. G. Fischer, Iena 422 pp.
13 Bour, R. and A. Dubois. 1984. Xerobates Agassiz, 1985, synonyme
plus ancien de Scaptochelys Bramble, 1982 (Reptilia, Chelonii,
Testudinidae). Extr. Bull. Mensuel Soc. Linneenne Lyon
53(1):30-32.
14 Carr, A.F., Jr. 1952. Handbook of turtles. The turtles of the
United States, Canada, and Baja California. Cornell Univ. Press,
Ithaca, NY. 542 p.
15 Neill, W.T. and E.R. Allen. 1957. The laminal spurs of the
juvenile gopher tortoise, Gopherus polyphemus (Daudin). Copeia
1957(4):307.
16 Arata, A.A. 1958. Notes on the eggs and young of Gopherus
polyphemus (Daudin). Quart. J. Fla. Acad. Sci. 21(3):274-280.
17 Auffenberg, W. and R. Franz. 1978. Gopherus. Catalogue of
American Amphibians and Reptiles. Soc. for the Study of Amphib.
and Rept. pp. 211.1-211.2.
18 Bartram, W. 1791. Travels through North and South Carolina,
Georgia, east and west Florida. Philadelphia. xxxiv + 522 pp.
References - 1 (DRAFT) - References
Species TORTOISE, GOPHER
Species Id ESIS154014
Date 14 MAR 96
19 Daudin, F.M. 1802. Historie naturelle, generale et particuliere
des reptiles, Vol. 2. F. Dufart, Paris. 432 pp.
20 Guerin-Meneville, M.F.E. 1838. Iconographie de Regne Animal de
G. Cuvier...Torre II. Reptiles J.B. Bailliere, Paris.
21 Gray, J.E. 1844. Catalogue of tortoises, crocodiles and
amphisbaenians in the collection of the British Museum. Edward
Newman, London. 80 pp.
22 Agassiz, L. 1857. Contributions to the natural history of the
United States of America, first monograph, vol. 1, part 2. North
American Testudinata. Little, Brown and Co., Boston. p. 233-452.
23 Shaler, N.S. 1888. Habits of the great southern tortoise. Pop.
Sci. Monthly 34(1):37-42.
24 Stejneger, L. 1893. Annotated list of the reptiles and
batrachians of the Death Valley expedition in 1891, with
description of new species. North Am. Fauna (7):159-474.
25 Hay, O.P. 1916. Descriptions of some Floridian fossil vertebrates
belonging mostly to the Pleistocene. 8th Ann. Rept., Florida State
Geol. Surv. p. 39-76.
26 Mertens, R. and H. Wermuth. 1955. Die rezenten Schildkroten,
Krokodile und Bruckenechsen. Zool. Journal Abt. Syst. 83:323-440.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Lohoefener, R., and Lohmeier, L. 1984. The status of Gopherus
polyphemus (Testudines, Testudinidae) west of the Tombigbee and
Mobile Rivers. U.S. Fish and Wildlife Serv. 126 pp.
02 Chandler, G. 1986. Pers. comm. Refuge Manager, Miss. Sandhill
Crane NWR, Gautier, MS.
References - 2