Title: Historic and present distribution of the American crocodile in Florida
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Title: Historic and present distribution of the American crocodile in Florida
Physical Description: Book
Creator: Kushlan, James A.
Mazzotti, Frank J.
Affiliation: University of Mississippi -- Department of Biology
Pennsylvania State University -- School of Forest Resources
Publisher: Society for the Study of Amphibians and Reptiles
Publication Date: 1989
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General Note: Drawn from Journal of Herpetology, Vol. 23, No. 1, pp. 1-7, 1989
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Journal of Herpetology, Vol. 23, No. 1, pp. 1-7, 1989
Copyright 1989 Society for the Study of Amphibians and Reptiles

Historic and Present Distribution of the
American Crocodile in Florida


JAMES A. KUSHLAN1 AND FRANK J. MAZZOTTI2,3

'Department of Biology, University of Mississippi, University, Mississippi 38677, USA
2School of Forest Resources, Pennsylvania State -.' .o .. University Park, Pennsylvania 16802, USA

ABSTRACT. The historic and recent distribution of the American crocodile (Crocodylus acutus) in Florida
is from Vero Beach and Tampa south to the lower Florida Keys. Its nesting distribution is southern Biscayne
Bay and northeastern Florida Bay. Both distributions reflect winter temperature. Nesting sites and non-
nesting habitat have been lost to development on Miami Beach and the upper Florida Keys, but this loss
has been compensated by the creation of artificial nesting sites on spoil banks along southern Biscayne
Bay and a westward addition to the nesting range in Florida Bay. Except for the shift in nesting away
from developed areas, the general distribution of the American crocodile in Florida is the same as that
historically documentable.


Habitat loss and concomitant decreases in dis-
tributional range are important factors in the
changing status of many crocodilian popula-
tions. The American crocodile (Crocodylus acu-
tus), although ranging widely in the neotropics,
is presently concentrated in a few population
centers in Jamaica, Hispaniola, Cuba, and south-
ern Florida. These populations are relatively
isolated from each other and have achieved a
degree of genetic distinctiveness (Menzies and
Kushlan, unpubl. obs.). It has long been thought
that the southern Florida population is at risk
of extirpation owing to excessive mortality and
habitat loss (Barbour, 1923; Ogden, 1978; Hines
et al., 1984), but such conclusions have been
based on limited information. Here we review
the historic distribution of the American croc-
odile in Florida, present data that document its
present distribution, and discuss factors that
might account for the observed distributional
patterns.

METHODS
Field work was concentrated at the southern
tip of Florida, USA. Surveys covered the entire
coastal zone, concentrating in Florida Bay from
Key Largo to Cape Sable. We observed croco-
diles on standardized surveys using power boat,
canoe, fixed-wing airplanes, and helicopter be-
tween July 1977 and September 1980. Infor-
mation on the intensity and effectiveness of our
methods is provided in Table 1. Efficiency av-
eraged one sighting every 3 hours during 456
survey hours. Night surveys were attempted by
helicopter without much success. Standardized


3Present Address: Department of Wildlife and
Range Sciences, University of Florida, 3245 College
Avenue, Davie, Florida 33314, USA.


night surveys using power boat and canoe were
conducted monthly over accessible habitat ad-
jacent to Florida Bay and along a west coast river
system. Sightings from boats were few because
of the small area covered, difficulty of access to
much of the habitat, and wariness of the ani-
mals. We also recorded the locations of all croc-
odiles sighted or captured by project biologists
or by National Park Service field personnel out-
side the survey. We do not include in this paper
multiple locations of hatchling, tagged, or ra-
dio-telemetered animals.
The most comprehensive and useful infor-
mation on crocodile distribution comes from
daytime helicopter and airplane surveys, which
were conducted monthly from July 1977 to July
1978 between Key Largo and Cape Sable. Fixed-
wing aircraft were flown at an air speed of 120
km/hr, and helicopters, at 80-100 km/hr, both
at altitudes of 50 to 80 m.
Nesting sites were found by observing, from
the air or on foot, signs of terrestrial activity,
followed by excavating each site for eggs.
Information on the historic occurrence of the
crocodile in Florida comes from the published
literature and unpublished information in the
files of Everglades National Park, and of Joseph
Moore (pers. comm.), who interviewed many
long-term residents of Florida Bay in the 1950s.

RESULTS

Historic ..e.-Rafinesque (1822) suspected
that a crocodile occurred in Florida, but a spec-
imen was not collected until 1869, in the Miami
River off Biscayne Bay (Wyman, 1870). From
that date through the 1960s, many scattered ob-
servations and second-hand reports of croco-
diles were published (Table 2), which together
provide a reasonable Pstimate nf t-hp hic-nrir





J. A. K. **, ..... AND F. J. MAZZUTTI


TABLE 1. Number of crocodile sightings per sur-
vey hour. Number of hours is in parentheses.

Breeding Non-breeding
Survey Overall season season
Boat 0.21 (289) 0.13 (132) 0.29 (157)
Helicopter 0.60 (133) 0.57 (81) 0.65 (52)
Fixed-wing 0.63 (34) 0.36 (11) 0.75 (23)
Mean (total) 0.35 (456) 0.30 (224) 0.42 (232)


distribution of the American crocodile in Flor-
ida (Fig. 1).
Crocodiles were credibly reported as far north
as Palm Beach on the Florida east coast. Most
reports centered on Biscayre Bay, well-visited
as the location of Miami, and the less accessible
northeastern Florida Bay. Willoughby (1913)
mentioned crocodiles on Cape Sable in north-
western Florida Bay, but no details were pro-
vided. Crocodiles definitely occurred in the up-
per Florida Keys as far south as Lower
Matecumbe Key (Carr, 1940). Allen and Neill
(1949) and Neill (1971) reported that the croc-
odile's range extended southward to Key West.
A published photograph of a crocodile report-
edly taken on a beach in Key West in 1935 is
the earliest concrete evidence for the American
crocodile in the lower Florida Keys. On the west
coast of Florida, crocodiles were observed pe-
riodically after 1940 at Naples, Sannibel, Os-
prey, and Pinellas near Tampa. LeBuff (1957b)
provided second-hand reports of west coast
sightings, including that of an animal killed in
Lee County and another captured near Sarasota.
The documentable historic nesting range of
the American crocodile in Florida is more re-
stricted than is the dispersion of individuals.
Crocodiles nested on islands (called keys) and
shores of Biscayne Bay and Florida Bay (Fig. 2).
A famous nesting site of long occupancy was
on Miami Beach (photo in Barbour, 1944). A
number of reports of nesting in Florida Bay are
available (National Park Service records; Moore,
pers. comm.). Recorded nesting there dates back
to at least 1914, in Alligator Bay. Other Florida
Bay locations used historically included Sam-
phire, End, and Club keys in 1930-1950, and
Cup-of-Whiskey Key in 1951. Black Betsy Key
was used for over 30 years. Before 1950, nesting
also took place in eastern Florida Bay, near the
main line of the upper Florida Keys. The record
of nesting on other islands is sporadic, as were
efforts by biologists to locate nest sites.
SI Distribution. -Our surveys demonstrat-
ed that crocodiles are well dispersed across the
coastal zone of extreme southern Florida from
Cape Sable to southern Biscayne Bay, including
Key Largo (Fig. 3). We found animals consis-
tently as far north as southern Biscayne Bay,
where they inhabit canals including those as-


TABLE 2. Historic
in Florida.


published reports of crocodiles


Location References
Florida east coast Henshall, 1884; Hornaday,
1891; Cope, 1898; Barbour,
1923; Pierce, 1970; Behler,
1978
Biscayne Bay, Wyman, 1870; Henshall, 1884;
Miami Beach Hornaday, 1891; Cory, 1896;
Barbour, 1923, 1944; Pierce,
1970; Monroe and Gilpin,
1980
South Biscayne Cory, 1896; Smith, 1896; Di-
Bay mock and Dimock, 1908;
Willoughby, 1913; Dickin-
son, 1953
Northeastern Dimock and Dimock, 1908; Di-
Florida Bay mock, 1918; Dickinson,
1953; Moore, 1953
Northwestern Willoughby, 1913
Florida Bay
Upper Florida Carr, 1940
Keys
Lower Florida Carr, 1940; Allen and Neill,
Keys 1949; Neill, 1971
Florida west coast, Willoughby, 1913; Barbour,
central Florida 1923 (but see Maynard,
1929); LeBuff, 1957a, b; Beh-
ler, 1978



sociated with the Turkey Point nuclear power
plant (see also Gaby et a!., 1985). The eastern-
most observations were on northern Key Largo,
where crocodiles occur primarily in old canals,
coves, and ponds in mangrove swamps (also P.
Moler, pers. comm.). Our observations were
concentrated in northeastern Florida Bay. Sixty-
nine percent of all sightings were in that area
during surveys that uniformly covered the en-
tire range of the crocodile in Florida Bay.
Crocodiles are periodically observed and re-
ported in Florida outside of Florida and Bis-
cayne bays (Ogden, 1978; Campbell, 1980; Ir-
vine et al., 1981; Alvarez, 1984; Hines et al.,
1984; D. Hubbard, pers. comm.; P. Moler, pers.
comm.). During the study period, crocodiles re-
sided in power plant canals in Fort Lauderdale,
and one was observed as far north as Vero Beach
on the east coast in 1974 (Behler, 1978). Along
the west coast, crocodiles were periodically re-
ported from Naples (recent records include 1973,
1975, 1976, 1980, and 1983). More northern ob-
servations have been made at Sannibel Island
and Tampa.
Information on the recent status of crocodiles
in the lower Florida Keys is scarce. Reports in
the 1960s and 1970s, especially by J. Watson (in
Ogden, 1978), were the last - sightings of
animals there. Jacobsen (1983) accepted only
three recent reports as hpinc rol;hio r Vno;..


















Tampa


' Vero -
S,B Beach


0 50 SO
______\ ^rBeach
km -\ al
Fort
Lauderdole
17 K I

18M' a mi
19
Key
West ------
20'''

FIG. 1. The overall distribution of the American crocodile in Florida (solid line) and average January air
isotherms (C; dashed lines).


sightings of crocodiles near Lake Okeechobee
are of animals formerly captive at a tourist at-
traction (C. Clemmons and J. Lang, pers. comm.).
The present nesting range of the American
crocodile is centered in northeastern Florida Bay.
Fig. 2 shows the location of nest sites discovered
in 1981 and 1982 (our data; Moler, pers. comm.,


for Key Largo; Gaby, pers. comm., for Turkey
Point). From 1970 to 1982, 74% of 141 clutches
of eggs found in southern Florida were asso-
ciated with Florida Bay. Most of the remaining
nests occurred at Turkey Point and on Key Lar-
go. Single nests have been found near Cape
Sable. As a result of these observations, the pres-


Miami
Beach


Florida
Bay


S0 20
km
FIG. 2. Historic nesting distribution of the American crocodile in Florida (dashed line) and present nesting
distribution (solid line). Each dot represents a nest site used in 1981 and 1982.





J. A. KUSHL- I AN)D P. J. M.AZZUTTI


Florida
Bay


FIG. 3. Locations of crocodiles observed in southern Florida in


ent nesting range of the American crocodile can
be delineated as southern Biscayne Bay, includ-
ing Turkey Point and Key Largo, and northern
Florida Bay to Cape Sable (Fig. 2).
Factors Affecting Distribution.--The American
crocodile is a tropical species, and it is likely
that climate limits the northern extent of its
range in Florida. Its overall range reflects win-
ter air isotherms (Fig. 1), corresponding gen-
erally to the 17C January isotherm. This simi-
larity suggests that winter temperatures may
ultimately be limiting the distribution of the
American crocodile in Florida. The only per-
manent population north of extreme southern
Florida is in the warm cooling canals of a Fort
Lauderdale power plant.
The nesting range coincides with the warm-
est winter location on the Florida mainland (Figs.
1, 2). The American crocodile appears to prefer
relatively deep estuarine habitats that are pro-
tected from wind and wave action (Kushlan and
Mazzotti, 1989), and these preferences appear
to account for the details of distribution within
the overall breeding range.

DISCUSSION
Historic Range.-The historic range of the
American crocodile has been a matter of some
disagreement (Moore, 1953; LeBuff, 1957b; Og-
den, 1978) and requires re-evaluation. The his-
torical presence of the American crocodile in
south Florida is well documented (Table 2).
Available reports make it clear that northeast-
ern Florida Bay, at the extreme tip of the Florida


20km
1977 to 1980. Each dot : a sighting.


peninsula, was the center of the crocodile's his-
toric distribution. Dimock (1918, p. 452) pro-
vided a precise description at the turn of the
century. He said crocodiles were "definitely
limited to the region at the extreme southern
end of the peninsula of Florida, a strip ten miles
long by three wide."
Individual crocodiles were reported sporad-
ically outside these limits. LeBuff (1957a) con-
cluded that crocodiles historically occurred
sparsely on the Florida west coast, a view dis-
puted by Moore (1953), who concluded that
there is no evidence that the American croco-
dile occurred there naturally. Ogden (1978) stat-
ed that such records were of escaped, released,
or storm-displaced animals. Certainly any of
these factors might contribute to displacement
of a crocodile, and it is likely that some of these
sightings were of previously captive animals.
Nevertheless, the record of observations is so
long and persistent as to suggest that the west
coast of Florida has been part of the overall
range since at least the 1940s. However, there
is no evidence that crocodiles lived there con-
tinuously during that period, and it is likely
that sightings were of transient individuals. A
similar explanation accounts for sightings in
the lower Florida Keys. Jacobsen (1983) con-
cluded that there is no evidence to suggest that
crocodiles ever occurred in substantial numbers
or permanently in the lower Florida Keys.
Reports from inland central Florida are es-
pecially tenuous. Barbour (1923) reported that
C. J. Maynard claimed to have killed a laree





CROCODILE POPULATION ECOLOGY


crocodile near Lake Harney in Volusia County,
central Florida, but Maynard (1929) later re-
tracted this record, admitting that he had mis-
identified an alligator. Interestingly, Maynard
based his retraction on a comparison made with
a crocodile that reportedly had been killed near
Lake Okeechobee by F. A. Ober. Willoughby
(1913) also reported a crocodile having been
killed near Lake Okeechobee, perhaps the same
one. Because of this report, we cannot rule out
the possibility that crocodiles have occurred in
the lake, although there is no evidence that they
lived there regularly.
The documentable historic nesting range in-
cludes northeastern Florida Bay and Biscayne
Bay as far north as Miami Beach. Evidence for
long-term nesting elsewhere in Florida seems
unconvincing at present. Pierce reported nest-
ing at Fort Worth (G. Voss in Ogden, 1978), but
supporting evidence is lacking.
Present Distribution.-Our systematic surveys
provided the first comprehensive data on the
distribution of this species in southern Florida.
Based on these data and other observations, the
present distribution over the core of its range
is well delineated (Figs. 2, 3).
Reports of crocodiles outside the breeding
range occur fairly regularly. As noted above,
this may be a natural result of the ability of
individual crocodiles to move long distances,
and/or the occurrence of previously captive an-
imals. In that records of introduced animals are
indistinguishable from those of native animals
and the presence of any crocodile indicates con-
ditions are appropriate for survival, distribu-
tion is appropriately evaluated using all re-
corded observations.
The ability of crocodiles to move long dis-
tances was well demonstrated in 1978 when a
crocodile made several long-distance move-
ments after being transported by wildlife offi-
cers. This animal, captured in Fort Lauderdale
(Grimm and Bubman, 1978) and transported to
Key Largo (T. Regan, pers. comm.), moved 100
km south before being recaptured on Big Pine
Key (W. Dunson, pers. comm.). Re-released on
Key Largo (J. Simon, pers. comm.), it was re-
captured 20 km south (P. Patty, pers. comm.),
and was released at the Fort Lauderdale power
plant (P. Rose and R. Wilcox, pers. comm.). Later
that year, after moving 10 km inland via canals,
it was recaptured and released near Naples on
the west coast (P. Moler, pers. comm.). It then
moved to Cape Sable, 150 km south (pers. obs.).
This individual undertook a succession of long-
distance movements and, with help from gov-
ernment agents, covered most of the known
range of the crocodile in Florida. Thus it is clear-
ly possible for crocodiles to periodically occur
well outside the nesting range.


Recent vs. Historic Distribution.-We find no
change in the overall distribution of the Amer-
ican crocodile in Florida. The Florida popula-
tion is, and apparently always was, restricted to
the southern part of the Florida peninsula, along
both coasts from Tampa and Vero Beach-Palm
Beach to the lower Florida Keys. Similarly, the
American crocodile is, and apparently always
was, most commonly reported from southern
Biscayne Bay to northeast :.. : Bay, its pres-
ent breeding range. The concurrence of this
distribution with winter temperatures suggests
that climate is the primary factor affecting croc-
odile distribution in southern Florida.
However, the nesting distribution of the
American crocodile has changed. Crocodiles no
longer nest on Miami Beach or frequent north-
ern Biscayne Bay. It is not known how impor-
tant Miami Beach was as a nesting habitat, but
it is likely suitable nest sites were limited along
the mangrove swamp-lined bay shore. Nesting
still occurs on both sides of Barnes and Card
sounds, the southern extensions of Biscayne Bay.
Nests there, at Turkey Point, and on Key Largo,
are on the spoil banks of canals dug through
mangrove swamps. It is probable that few suit-
able nest sites occurred naturally in these areas.
The current nesting activity in southern Bis-
cayne Bay in part compensates for the loss of
nesting habitat in northern Biscayne Bay and
Miami Beach.
Crocodiles also no longer nest on the Florida
Keys or on islands in nearby eastern Florida
Bay. We agree with Ogden (1978) that such a
loss must have been caused by the reduction in
suitable non-nesting habitat on the main line
of the Florida Keys. Our information from sur-
veys and telemetry conclusively shows that
crocodiles seldom occur in Florida Bay itself,
except for nesting, at the conclusion of which
they return to protected habitat (Kushlan and
Mazzotti, 1989). The off-season habitat for croc-
odiles nesting on islands in eastern Florida Bay
would have been the upper Florida Keys. Much
of the land area of the main line keys has been
developed, and conflicts between human resi-
dents and crocodiles, fatal to the crocodile, have
occurred in this area for over 40 years (R. P.
Allen, in litt.). The loss of habitat on the main
line of the Florida Keys, occupied for most of
the year, rather than conditions on or around
the Florida Bay nesting islands, undoubtedly
led to the decrease in the nesting range of croc-
odiles in eastern Florida Bay and the Florida
Keys.
In addition to islands, crocodiles nested his-
torically on the natural berm of coastal rivers
off Florida Bay. The same locations were used
by settlers and farmers as recently as the 1950s.
In that river nest sites are in thp vpar-rniinA





J. A. KUSHLAN AND F. J. MAZZOTTI


habitat of both adult and hatchling crocodiles
(Kushlan and Mazzotti, 1989), these seem to be
the most advantageous nest sites. In contrast,
nesting on offshore island beaches places hatch-
lings in some jeopardy. It is possible that use
of bay islands may have been the result of past
disturbance to the theoretically more suitable
nesting sites along coastal rivers. Nesting per-
sists on northeastern Florida Bay islands, and
there is no evidence that the number of these
sites has decreased. Actually documented use
of this area has increased in recent years, coin-
cident with an increase in efforts to find nests,
and undiscovered nesting sites undoubtedly ex-
ist westward in the bay and in the Cape Sable
area.
Thus the overall distribution of the American
crocodile in Florida remains generally undi-
minished from its historic extent, although the
nesting distribution has shifted away from de-
veloped areas. The provision of canal banks in
southern Biscayne Bay may have mitigated for
some of the loss of nesting habitat. The north-
ernmost population of the American crocodile
still occurs within its traditional range, limited,
it seems, principally by winter temperatures.

Acknowledgments. -We acknowledge with
pleasure the collaboration of our colleagues in
the study of the American crocodile in Florida,
Paul Moler, Ronald Gaby, William Dunson, and
Peter Lutz. We thank also those who assisted
in surveys or supplied data used in this paper:
Jim Abbott, Robert Austin, John Behler, Hank
Blatt, Barbara Bohnsack, Joanna Booser, Charlie
Brookfield, Dave Brown, Linda Campbell, Ann
Dunbar-Cooper, Ronald Gaby, Carol Hewes,
David Hubbard, Blair Irvine, Terri Jacobsen,
Richard Klukas, Lori Lagna, Jeffrey Lang, Alan
Litman, Mark McMahon, Don Mitchell, Paul
Moler, Joseph Moore, Amanda Muller, John
Ogden, Paige Patty, Tom Regan, William Rob-
ertson, Paul Rose, Robert Saddler, Mark Salz-
burg, Gene Saunders, Jeff Simon, Durban Tabb,
Fred Whitehead, Ron Wideman, and Ross Wil-
cox. Valuable suggestions and comments were
provided by I. Lehr Brisbin, William Dunson,
Ronald Gaby, Les Garrick, Ted Joanen, F. Wayne
King, William Magnusson, Harry Messel, and
Grahame Webb. We especially thank Lehr Bris-
bin for his assistance in preparing the manu-
script. The study was funded by the National
Park Service in cooperation with: the U.S. Fish
and Wildlife Service; Florida Game and Fresh
Water Fish Commission; Florida Power and
Light Co.; Connell Associates, Inc.; National
Science Foundation through a grant to W. Dun-
son, Pennsylvania State University; University
of Georgia Institute of Ecology; and Nova Uni-
versity Oceanographic Institute.


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Accepted: 12 November 1987.




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