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AMERICAN
BULLFROG
Rana
catesbeiana |
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| INTRODUCTION |
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The bullfrog is Rhode Island's largest anuran.
In contrast to other frogs in Rhode Island, bullfrogs spend most
of their life in the water where they hibernate, breed, and hunt.
During the winter months, they hibernate in the muddy substrate
of ponds and lakes. Once spring arrives, they emerge and spend
the warm months of the year lurking in shallow waters at the edge
of wetlands where they depend on aquatic vegetation for cover.
Bullfrogs usually linger in these vegetated shoals, although they
occasionally venture into deeper parts of lakes and ponds. Unlike
the similar green frog (Rana clamitans), bullfrogs also
primarily hunt from the water. They have a voracious appetite
and as adults, they are carnivorous and will eat anything that
moves. Thus, the size of adult's prey is limited only by the gape
of their mouth. In contrast, their tadpoles are largely vegetarian.
Due to their large size, bullfrogs are a favorite for those people
whom consider frog legs to be a delicacy. This has lead to their
introduction throughout North America and other continents, a
practice which has had devastating impacts on native fauna.
There is some evidence that some local populations of bullfrogs
may be declining (DeGraaf
and Yamasaki 2001), but for the most part their populations
appear to be stable in the region (Klemens
1993).
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| IDENTIFICATION |
- The only large, greenish frog
in Rhode Island without dorsal-lateral folds. The green
frog has prominent dorsal-lateral
folds (photo comparison).
- The bullfrog is the largest and
most aquatic North American anuran.
- Color is variable, but they typically
have a green or greenish brown dorsum and a whitish venter
with grayish mottling (Dickerson
1906; Conant and Collins
1991). Individuals can change color tone depending
on environment (Dickerson 1906; Albright
1999).
- Feet are strongly webbed.
- The vocal sac is under the throat
and extends backwards over the arm; therefore, when the
bullfrog croaks not only does the throat swell but also
the space between the ear and arm (Dickerson 1906). Both
males and females croak.
- Voice: Bullfrogs do not
sing in chorus. Dickerson (1906) described the bullfrog
mating call with 4 different phrases- "Be drowned,"
"Better go round," "Jug o' rum," and
"More rum"; all said in a deep hoarse voice.
Wiewandt (1969) described
three distinct calls; the "mating call"- 3 to
6
bass croaks, the 'gronk'- a single note of the mating
call, and the 'bonk'-
a "distinctly abrupt, forceful vocalization."
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Bullfrogs lack dorsal-lateral folds (green frogs have
prominent dorsal-lateral folds.)
On both the bullfrog and the green frog the tympanum (eardrum)
is larger than the eye.
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Sexual dimorphism
- Adult males have tympana
that are larger in diameter than the eyes, whereas the female's
tympana are approximately the same size as the eyes.
- Adult males have a bright yellow throat and chest. The female's
throat is white.
- Breeding males have strong, swollen thumbs (used to grasp
females during amplexus).
- Males tend to have a narrower head than females (Wright
and Wright 1949).
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| SIZE |
| AGE / SEX
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SVL (SNOUT VENT LENGTH) (cm)
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SAMPLE SIZE
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AVERAGE
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RANGE
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Std. Deviation
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ADULT FEMALE
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10.7
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8.0- 14.9
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1.6
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46
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ADULT MALE
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9.2
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7.4- 11.2
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1.5
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12
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JUVENILE
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5.7
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3.3- 8.2
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1.2
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95
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METAMORPH
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3.7
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2.5- 5.2
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0.9
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36
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AGE/ SEX
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MASS (g)
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SAMPLE SIZE
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AVERAGE
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RANGE
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Std. Deviation
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ADULT FEMALE
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115.8
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54.5- 200.0
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43.8
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46
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ADULT MALE
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76.5
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42.2- 124.4
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76.5
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12
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JUVENILE
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19.6
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5.7- 57.0
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11.8
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95
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METAMORPH
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5.0
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1.4- 9.9
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3.3
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36
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| RELATIVE ABUNDANCE |
Rhode Island
Bullfrogs are
widespread and locally common in our area (Raithel pers. comm.).
We found bullfrog tadpoles in 7 (23%) out of 30 permanent ponds
we sampled during the 2000 field season (Paton
and Egan 2001). |
Regional
Bullfrogs
are common throughout the region, but populations may be declining
in parts of New England (Degraaf and Yamasaki 2001). Klemens (1993)
classified bullfrogs as common in Connecticut; he characterized
them as widespread and flourishing in places where other frogs have
disappeared.{Relative
Abundance General} |
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| DISTRIBUTION |
Rhode Island
Bullfrogs are found throughout
mainland Rhode Island (Raithel pers. comm).
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Regional
Bullfrogs range
throughout the region except for northeastern Vermont, northern
New Hampshire, and northern Maine (Klemens 1993).
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General
Bullfrogs range
from Nova Scotia west to Wisconsin, south through the Great Plains
to eastern Colorado and Texas (Degraaf and Yamasaki 2001). They
occur all along the eastern seaboard from southern Florida to northern
Maine (Klemens 1993). They have been introduced throughout the western
United States, British Columbia and Mexico, as well as other countries
including Cuba, and Jamaica (Conant and Collins 1991).
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| HABITAT NON BREEDING |
| Bullfrogs overwinter
in permanent waters including marshes (Emlen
1968), lakes, and ponds (Willis
et al. 1956). They typically hibernate under mud, leaves, and
other debris (Wright 1914). |
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| HABITAT BREEDING |
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Bullfrogs are the most aquatic frog found in Rhode Island. They
may breed in the same water body that they overwinter or move
to a different wetland (Howard
1978). They are found in a wide array of freshwater
habitats, but primarily in permanent water (Klemens 1993). Klemens
(1993) collected bullfrogs in lakes, ponds, reservoirs, rivers,
stock ponds, marshes, woodland pools, floodplain swamps, bogs,
and a variety of manmade ponds and impoundments. Klemens (1993)
also found that juvenile bullfrogs move among waterbodies more
frequently than adults. We documented a similar pattern in Rhode
Island (P. Paton, unpubl. data).
Because bullfrogs tadpoles take more than 1 year to develop,
they require semi-permanent or permanent ponds for breeding (Degraaf
and Yamasaki 2001). Bullfrogs observed in vernal ponds are not
breeding, but feeding and rehydrating as they pass through the
landscape. It is more likely that these individuals are juveniles
(Klemens 1993), although we commonly have seen adult bullfrog
spend the summer in temporary vernal ponds that do not provide
suitable breeding habitat (P. Paton, unpubl. data).
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| HYDROLOGY |
Bullfrogs seem to prefer large
ponds and lakes where the shore is shaded by hydrophilic trees and
shrubs and the water has numerous aquatic plants for refuge (Dickerson
1906). They breed in semi-permanent or permanent ponds (Paton and
Egan 2001).
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| MOVEMENT CHRONOLOGY |
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In Rhode Island, adult bullfrogs first start immigrating to breeding
ponds at the end of March. Paton and Crouch (In press) started
capturing adults on average around 25 March, with peak immigration
by mid-April, and some individuals moving as late as mid-June.
Even though bullfrogs are present in wetlands as early as late
March, they do not initiate active choruses for at least a month-
until early May in Rhode Island (Crouch
1999). During the summer, movement among age cohorts and genders
of bullfrogs varies considerably. Raney
(1940), reported that some individuals move as little
as 100 feet (30 meters); whereas others may move up to 3,000 feet
(915 meters). Raney (1940) found no obvious correlation in these
movements.
Metamorph departure from breeding ponds is late compared to other
species. In southern Rhode Island, the first emigration movement
was detected in early August, and the latest movement witnessed
was at the end of October; the data showed that bullfrogs have
a very protracted emigration period (Paton
and Crouch In press).
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| REPRODUCTION |
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Bullfrogs are the last anuran to breed in Rhode Island, typically
beginning in late May and extending into July (Dickerson 1906).
Bullfrogs do not have a synchronized, rapid breeding period, such
as wood frog (Rana sylvatica) or spotted salamander (Ambystoma
maculatum). Rather, they have a prolonged breeding period
similar to green frogs and may breed twice in one season (Emlen
1968; Howard 1978). In Rhode Island, we have detected bullfrogs
calling at breeding ponds from early May through early August
(Crouch 1999); therefore, they also have a protracted calling
period compared to other species.
Having emerged from hibernation, bullfrogs either remain in the
pond where they overwintered, or move to another suitable breeding
site. For about a month, they remain silent (Albright 1999) waiting
for optimum conditions to begin courtship. In Maine, optimum conditions
are when air temperatures and water temperatures reach 20-22 °C
(68-72° F) (Albright 1999). The majority of mating rituals
(sustained choruses, male to male encounters, and copulation)
take place at night (Emlen
1976), though in the onset of the breeding season,
it is not uncommon to hear choruses during the day (Howard 1978).
In Michigan, Howard (1978) reported prolonged choruses starting
around 9:00 PM and continuing until dawn and copulation occurring
during the early morning hours. Interestingly, Howard (1978) found
that as the season progressed calling began later and later, frequently
not beginning until midnight. This may explain why Bridges
and Dorcas (2000)
only detected bullfrogs calling after midnight when
they monitored a pond with an automated data recording system
during June.
At breeding sites, males establish territories by spacing themselves
at regular intervals along the water's edge and remain in these
zones for up to 3 weeks (Emlen 1968). They are aggressive and
will engage in physical encounters with conspecific males (Emlen
1968; Wiewandt 1969). Females lured to the breeding site by the
calls of the males, initiate copulation by making contact with
a carefully selected male (Emlen 1968). Females generally deposit
eggs within the males territory. Howard (1978) observed 73 pairs
of bullfrogs in amplexus and only three pairs deposited their
eggs away from the original copulation site.
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| EGG MASS |
| Eggs are deposited on aquatic
vegetation in shallow water as a thin gelatinous mat similar to
the green frog. The 12,000 to 20,000 bicolored (black and white)
(Wright and Wright 1949) eggs float on the surface at first, but
typically sink to the bottom before they have fully developed (Kenney
and Burne 2000). The development of the thousands of eggs is
temperature dependent but hatching usually occurs within 5 to 20
days (Oliver 1955). |
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| LARVAE |
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The length of the larval period is unknown in Rhode Island.
Bullfrog larvae are known to overwinter for at least one
winter (Collins
1979), but more typically they require 2 to 3
years to metamorphose (Wright and Wright 1949; Degraaf
and Rudis 1983). At any given breeding
pond in Rhode Island, several sizes of bullfrogs tadpoles
can be observed, suggesting that it may take over 1 year
for bullfrog tadpoles to undergo metamorphosis in southern
New England (C. Heinz and S. Egan, pers. observ.).
Small tadpoles are easily confused with green frog tadpoles,
but as they grow they become distinctly different. Bullfrogs
typically have a green to greenish brown dorsum and a yellowish
venter. The transition between the dorsum and venter is
gradual, not abrupt as in the green frog. The back, tail
musculature, and dorsal fin of the bullfrog is peppered
with fine black spots. The green frog usually has a more
mottled appearance.
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Bullfrog tadpole
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| METAMORPHS |
| Newly metamorphosed bullfrogs
average approximately 3.8 cm (range 2.5-5.2) (Paton unpubl. data).
Size, however, is not necessarily a good way to assess the age of
a bullfrog, because the size at transformation is generally dependent
on the size attained as a tadpole (Dickerson 1906). Thus, larger
tadpoles result in bigger metamorphs. Because studies show that
bullfrogs can remain as tadpoles for 1 to 3 years, a wide range
of sizes are possible. Irregardless of size, recently metamorphosed
bullfrogs are easily distinguished from young green frogs by the
absence of dorsal-lateral folds. |
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| JUVENILES |
During warmer months, juveniles
tends to move across the landscape more frquently than adults. They
ulitize numerous types wetlands, including vernal ponds, for food
and cover.
Sexual maturity: Bullfrogs are not sexually mature until
4-5 years old (Albright 1999; Degraaf and Yamasaki 2001). |
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| FOOD |
Adults
Adult bullfrogs are aggressive
and will eat virtually anything that moves (Conant and Collins 1991).
Their diet regularly includes: snakes, fish, small turtles, mice,
young waterfowl, crayfish, diving beetles, dragonfly larvae, spiders
and other invertebrates (Albright 1999). Green
and Pauley (1987) reported finding bullfrogs with spring
peepers (Pseudacris c. crucifer) and American toads (Bufo
americanus) in amplexus in their stomachs. Stewart
and Sandison (1972) found frog remains in every bullfrog
stomachs they examined and noted that bullfrogs were frequently
observed chasing and consuming green frogs (Rana clamitans).
Bullfrogs are also cannibalistic, eating both their own tadpoles
and other adults (Albright 1999). |
Larvae
Tadpoles are largely
vegetarians, however they will feed on dead animal matter (Albright
1999).
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| PREDATION |
Adults
Adults are prey to a
large number of animals including wading birds, raccoons, minks,
snakes, and humans (Albright 1999).
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Larvae
Tadpoles are
prey to aquatic invertebrates such as predacious diving beetles
and dragonfly larvae, as well as, to adult bullfrogs (Albright 1999). |
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| CONSERVATION CONCERNS |
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Bullfrogs are common and presently secure throughout the state.
Their populations are increasing in some areas due to habitat
alternations (creation of permanent ponds and stream impoundments)
and degradation of wetlands because they are proficient at colonizing
newly created habitats (Klemens 1993).
Bullfrogs have been introduced to many areas, most notably the
western United States (Conant and Collins 1991). Once established
in a new area, bullfrogs can readily extirpate native anurans
(Klemens 1993; Conant and Collins 1991) and other species such
as garter snakes and small turtles (Conant and Collins 1991).
Bullfrogs are the primary source of frogs legs in the United
States; therefore, some areas they are considered economically
important (Klemens 1993; Albright 1999). To protect bullfrogs
from being overharvested, many states have listed bullfrogs as
a game species (Degraaf and Rudis 1983; Albright 1999). Harvesting
bullfrogs is regulated in Rhode Island.
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