2006 SOUTHEASTERN NATURALIST 5(3):443–452
Food Habits of Macrochelys temminckii (Alligator
Snapping Turtle) from Arkansas and Louisiana
Ruth M. Elsey
Abstract - Food habits of 109 Macrochelys temminckii (Alligator Snapping
Turtles) collected from Arkansas and Louisiana were studied by examination of
stomach and intestinal tract contents from harvested turtles. There was a positive
correlation between the turtle carcass mass and the gastrointestinal tract content
mass (r = 0.39106, p < 0.0001). The most commonly occurring prey item was
fish, followed by Procambarus clarkii (crawfish), molluscs, turtles, insects, and
Myocastor coypus (nutria). Other mammalian species occurred infrequently, as
did snakes, birds, and crabs. Several species (Dasypus novemcinctus [armadillo],
Didelphis virginiana [opossum], Sciurus sp. [squirrel], and Sus scrofa [hogs]) that
have not previously been reported as prey items for Alligator Snapping Turtles
were noted. Some prey items were recovered in intestinal tracts that were not
observed in stomachs, illustrating the importance of examination of the entire
gastrointestinal tract when evaluating food habits in this species. The results
suggest Alligator Snapping Turtles are opportunistic scavengers able to consume
a wide variety of prey species.
Introduction
Macrochelys temminckii Harlan (Alligator Snapping Turtle) is a large,
secretive freshwater turtle found in rivers of several southeastern states
(Ernst et al. 1994, Lovich 1993, Pritchard 1989). Limited population data
exist, but all range states have enacted laws to limit commercial take of this
species, valued for its meat (Buhlmann and Gibbons 1997, Levell 1997,
Roman et al. 1999) and thus possibly over-harvested. Recent work in
several states has been undertaken to conduct intensive surveys on distribution
and population status, including Arkansas (Wagner et al. 1996),
Florida (Moler 1996), Georgia (Jensen and Birkhead 2003), Louisiana
(Boundy 2003, Boundy and Kennedy 2006), and Oklahoma (Riedle et al.
2005). Alligator Snapping Turtles are difficult to study in the wild due to
their rarity, secretive nature, and tendency to remain dwelling on the bottom
in aquatic habitats with only infrequent basking (Zappalorti 1976). It
is thought that increased harvest in the last several decades has depleted
populations throughout the range (Roman et al. 1999). The United States
Fish and Wildlife Service (USFWS) listed the Alligator Snapping Turtle as
an Endangered Species Act Candidate in 1991 (USFWS 1991), although in
mid-1999, the USFWS concluded that M. temminckii did not warrant
Louisiana Department of Wildlife and Fisheries, Rockefeller Wildlife Refuge, 5476
Grand Chenier Highway, Grand Chenier, LA 70643; relsey@wlf.louisiana.gov.
444 Southeastern Naturalist Vol. 5, No. 3
listing under the Endangered Species Act (Boundy 2003). As of December
2004, when a ban on commercial take of M. temminckii was enacted in
Louisiana, this species cannot be harvested from the wild commercially
anywhere in its range (Boundy and Kennedy 2006). The species is now
listed as CITES Appendix III.
Numerous anecdotal reports exist on the food habits of this rare species,
but few detailed studies have been published on the diet of the Alligator
Snapping Turtle. It has been said that “there is astonishingly little scientific
information on the food of this species” (Pope 1939), and in particular “few
stomachs of wild juvenile Macroclemys have been examined” (Pritchard
1989). Dobie’s dissertation (1966) on the reproduction and growth of the
Alligator Snapping Turtle included a table listing food analyses of 32 of 231
M. temmincki from several states, although the majority were collected from
two sites in Louisiana (Manchac and Jonesville). Sloan et al. (1996) evaluated
stomach contents from 65 adult M. temminckii collected from March to
October 1986. Most specimens were harvested in Louisiana; a few were
from Arkansas and Mississippi. Harrel and Stringer (1997) evaluated the
fish species recorded as Alligator Snapping Turtle prey by identification of
otoliths collected from scats of 24 turtles live-captured in northeast Louisiana
and held in captivity.
Prior to the ban on commercial harvest of Alligator Snapping Turtles in
Louisiana, we had the opportunity to collect viscera from processed wildcaught
Alligator Snapping Turtles, including juveniles from several sites.
Any additional detailed information on this species might be useful to
wildlife biologists attempting to manage or enhance habitats to conserve the
Alligator Snapping Turtle.
Materials and Methods
Alligator Snapping Turtles were caught by commercial trappers in
southeastern Arkansas (n = 52: 19 males [M], 32 females [F], 1 unknown
sex [U]) and Louisiana in June, July, and August of 1993, and from
Louisiana in 1994. Eighteen samples were obtained from Louisiana in
1993; fourteen (6M, 5F, 3U) were categorized as being from north Louisiana,
six of these were caught in the Lake Providence area in East
Carroll Parish. Four M. temminckii were trapped near the Pearl River
(1M, 1F, 2U), which borders Mississippi. All samples from 1994 (n = 38:
10M, 15F, 13U) were trapped near Venice, LA in Plaquemines Parish
during May. A single sample of unknown sex was obtained in spring
1997 from the Atchafalaya River system near Butte LaRose in St. Martin
Parish. Body mass was obtained on 92 of the 109 specimens (average as
follows [kg ± SEM]: 19.72 ± 1.00 [SE Arkansas], 17.57 ± 2.30 [Venice],
15.28 ± 3.22 [North Louisiana], and 5.57 ± 1.72 [Pearl River]). No body
mass was available for the Atchafalaya River specimen.
2006 R.M. Elsey 445
All specimens (except the one Atchafalaya River specimen) were obtained
from commercial processors in Louisiana. Interviews with trappers
and processors revealed turtles were generally caught in baited hoop nets
or on baited hooks or trotlines. Often, a number of turtles were delivered
to a processor and held in a storage tank for several days prior to processing.
Viscera were collected and frozen for later analysis. Viscera were
later thawed, and the stomach and intestinal tracts were examined separately
for content analyses. Contents were rinsed over a sieve and items
(prey and non-food) identified, sorted, and weighed to the nearest 0.01
gram. If present, endohelminths were counted and preserved in 70% ethanol
for another study (West et al. 2000). Often, exact identification of
prey items was impossible due to extensive digestion having occurred,
but stomach/intestinal contents could be sorted into general categories
(turtle, mollusc, etc.). Miniscule stomach and intestinal contents were
categorized as unidentifiable.
Table 1. Alligator Snapping Turtle stomach and intestinal contents by site. Results are shown as the
percent frequency occurrence, and the sample size or number of occurrences is in parentheses.
Southeast Venice North Pearl Atchafalaya
Prey item Arkansas (52) (38) Louisiana (14) River (4) River (1)
Invertebrates
Crab 5.26 (2)
Crawfish 30.77 (16) 92.11 (35) 35.71 (5)
Mollusc 42.31 (22) 68.42 (26) 14.29 (2) 50.00 (2)
Insect 15.38 (8) 39.47 (15) 25.00 (1)
Vertebrates
Carp 40.38 (21) 28.57 (4)
Catfish 5.77 (3)
Gar 9.62 (5) 42.11 (16) 100.00 (1)
Fish, unidentified 75.00 (39) 86.84 (33) 85.71 (12) 50.00 (2) 100.00 (1)
Bird, unidentified 10.53 (4) 7.14 (1) 25.00 (1)
Snake 5.77 (3) 10.53 (4)
Turtle 42.31 (22) 15.79 (6) 28.57 (4) 25.00 (1)
Armadillo 1.92 (1)
Muskrat 1.92 (1)
Nutria 60.53 (23)
Opossum 1.92 (1)
Hog 3.95 (2)
Raccoon 1.92 (1)
Squirrel 1.92 (1)
Mammal, unidentified 11.54 (6) 2.63 (1) 25.00 (1)
Bones, unknown 13.46 (7) 18.42 (7) 14.29 (2)
Eggs/membranes 1.92 (1) 15.79 (6)
Other
Vegetation 100.00 (52) 100.00 (38) 92.85 (13) 100.00 (4) 100.00 (1)
Unidentifiable matter 92.31 (48) 100.00 (38) 100.00 (14) 100.00 (4) 100.00 (1)
Endohelminths 67.31 (35) 76.32 (29) 42.86 (6) 75.00 (3)
Non-food 15.38 (8) 36.84 (14) 35.71 (5) 25.00 (1)
446 Southeastern Naturalist Vol. 5, No. 3
Results
Fifteen of the 109 stomachs examined were empty; only one intestine
was empty. As expected, there was a positive correlation between the body
mass and the gastrointestinal tract content mass (Pearson correlation coefficient
r = 0.39106, p < 0.0001, n = 93 pairs). The percent frequency
occurrence (percentage of samples containing the prey item) for prey items
at each site is shown in Table 1. Table 2 shows the percent frequency
occurrence for all sites combined, as well as the average prey mass and
range for each prey item (stomach and intestinal contents combined).
Fish were the most common prey items at all sites (Tables 1 and 2) and
occurred in 79.82% of the gastrointestinal tracts examined. Some of the fish
species found were Cyprinus sp. (carp), Lepisosteus sp. (gar), and Ictalurus
sp. (catfish). Fish were almost certainly overrepresented, as fish (including
carp; B. Harrel, US Fish and Wildlife Service, Frankfort, KY, pers. comm.)
are often used as bait to trap turtles; however, fish are clearly an important
food item for this aquatic species.
Table 2. Alligator Snapping Turtle stomach and intestinal contents combined, for all sites
combined.
Percent Prey mass
frequency occurrence Average (g)
Prey type (of 109 samples) ± SEM Range (g)
Invertebrates
Crab 1.83 0.94 ± 0.59 0.35–1.53
Crawfish (Procambarus clarkii) 51.38 9.32 ± 1.82 0.01–59.32
Mollusc 47.71 3.53 ± 1.93 0.01–97.46
Insect 22.02 0.30 ± 0.10 0.01–1.78
Vertebrates
Carp (Cyprinus sp.) 22.94 110.95 ± 26.39 3.31–477.84
Catfish (Ictalurus sp.) 2.75 21.96 ± 20.36 0.18–62.60
Gar (Lepisosteus sp.) 20.18 68.09 ± 19.90 0.07–334.23
Fish, unidentified 79.82 26.47 ± 6.73 0.01–387.12
Snake 6.42 16.74 ± 6.40 1.84–50.62
Turtle 30.28 16.66 ± 6.58 0.11–184.82
Bird 5.50 3.51 ± 2.00 0.02–12.59
Armadillo (Dasypus novemcinctus) 0.92 123.62 (n = 1) 123.62 (n = 1)
Muskrat (Ondatra zibethicus) 0.92 133.80 (n = 1) 133.80 (n = 1)
Nutria (Myocastor coypus) 21.10 273.76 ± 41.73 18.15–647.00
Opossum (Didelphis virginiana) 0.92 44.33 (n = 1) 44.33 (n = 1)
Hog (Sus scrofa) 1.83 18.81 ± 16.85 2.01–35.61
Raccoon (Procyon lotor) 0.92 26.68 (n = 1) 26.68 (n = 1)
Squirrel (Sciurus sp.) 0.92 252.26 (n = 1) 252.26 (n = 1)
Mammal, unidentified 7.34 5.53 ± 2.78 0.01–20.30
Bones, unidentified 14.68 10.08 ± 7.67 0.08–123.96
Eggs/membranes 6.42 3.21 ± 0.89 1.24–8.12
Other
Vegetation 99.08 40.12 ± 8.27 0.01–662.07
Unidentified matter 96.33 9.49 ± 1.21 0.01–75.89
Non-food 25.69 2.71 ± 0.58 0.02–12.15
2006 R.M. Elsey 447
Procambarus sp. (Crawfish) were the next most commonly taken prey
category (51.38% occurrence; Table 2). This was due to the high occurrence
from the Venice samples (92.11%, Table 1), with only 30.77% and
35.71% of the SE Arkansas and North Louisiana samples, respectively,
having crawfish.
Molluscs and turtles were the third and fourth most frequent prey items
(Table 2) and occurred at all sites (Table 1), with the exception of the single
sample from the Atchafalaya River. One specimen from SE Arkansas contained
remains of at least six turtles, another contained remains of at least
three turtles, and several had consumed at least two turtles. In general, only
portions of the carapace or mandible remained, thus precluding further
identification; however, one Graptemys sp. was recovered, and two specimens
thought to be Trachemys scripta were noted.
Unidentified insect remains occurred in 22.02% of all gastrointestinal
tracts studied, but as expected, amounted to a very low amount of the
average prey mass (average = 0.30 ± 0.10 g, range = 0.01–1.78 g).
Myocastor coypus Kerr (nutria) was the sixth most frequently occurring
prey item (21.10% of all 109 samples; Table 1), but only occurred in the
samples from Venice (60.53% of 38 samples). Nutria was also highest in
terms of the average prey mass (273.76 ± 41.73 g, range 18.15–647.00 g)
and was the single heaviest prey item consumed (Table 2).
Several mammalian species were rarely taken, and only in the samples
from SE Arkansas. Mammals that were found once included: Dasypus
novemcinctus Linnaeus (armadillo), Ondatra zibethicus Link (muskrat), Didelphis
virginiana Linnaeus (opossum), Procyon lotor Storr (raccoon), and
Sciurus sp. Linnaeus (squirrel). Sus scrofa Linnaeus (hogs) were found in
two of the 52 specimens from SE Arkansas, and a few instances of unidentified
mammals occurred from SE Arkansas, Venice, and North Louisiana
(Table 1). Unidentified bones were often seen (Table 1 and 2) that were
probably mammalian, but could have been turtle remains and were thus
categorized separately.
Snakes occurred in 6.42% of all samples; four instances were seen in the
Venice samples (10.53%), and the other three were from SE Arkansas turtles
(5.77%). Due to their soft body parts and state of digestion, snakes generally
could not be more specifically identified, except two samples thought to be
Nerodia sp. (by the remaining scales) and one Regina rigida Say (Glossy
Crayfish Snake).
Birds were also taken infrequently (5.5% of all samples); four were
recovered from the Venice collections and one each from North Louisiana
and the Pearl River. In one sample from the Venice subset, the lobed
bird’s foot suggested the bird taken was a marsh wading bird, possibly a
gallinule or rail.
Crabs were rare findings, with only two instances from the Venice
samples, and accounted for only 0.94 ± 0.59 g prey mass.
448 Southeastern Naturalist Vol. 5, No. 3
A few cases of eggs/inner eggshell membranes were also noted; presumably
they were from turtles or birds, but due to digestion they could not be
classified further.
Vegetation was seen in 99.08% of samples examined, and may have been
purposefully ingested (acorns) or taken incidentally while foraging for other
prey. Various stems, sticks, leaves, seeds, roots, vines, and acorns were
recovered; one stomach contained 662.07 g of vegetative material.
Non-food items were seen in 25.69% of all samples. Usually, this consisted
of string and a fishing hook, which was the means by which the
specimen was caught. Rocks, pebbles, stones, and/or mineral fragments
were seen in one stomach and eight intestinal tracts. One stomach contained
about half of an artificial fishing bait (plastic worm). Two intestines contained
pieces of glass, one contained a piece of black thread, and another
contained a 4.18-g piece of aluminum foil.
Endohelminths were noted in 73 of the 109 samples examined; the
highest number recorded in a single turtle was 338 in the intestine of a turtle
from SE Arkansas. The percent frequency occurrence of endohelminths is
shown in Table 1. The actual numbers of endohelminths present by location
(stomach and intestinal contents combined) were 49.74 ± 11.54 (n = 35, SE
Arkansas), 23.07 ± 4.06 (n = 6, Venice); 20.00 ± 8.34 (n = 3, Pearl River),
and 9.66 ± 4.06 (n = 6, North Louisiana). Endohelminths were recovered in
far higher numbers from intestinal contents (35.21 ± 6.21 endohelminths per
intestine, n = 70) than stomachs (7.88 ± 2.52 endohelminths per stomach, n =
8) for all sites combined. Endohelminths identified were three species of
acanthocephalans and four nematode species; all except one were new
records for M. temminckii (West et al. 2000).
The average number of different prey item types (excluding unidentified
matter and non-prey items) observed in stomachs was 2.28 ± 0.12 (n = 90),
and 4.13 ± 0.17 for intestines (n = 108). Thus, M. temminckii in this study
consumed a variety of items, many of which would have been missed or
underrepresented if only stomachs were examined.
Discussion
In agreement with prior studies (Ernst et al. 1994, Harrel and Stringer
1997, Pritchard 1989, Sloan et al. 1996), my results suggest Alligator
Snapping Turtles are opportunistic scavengers, and fish is an important
component of their diet. Fish occurred in nearly 80% of the samples in our
study; as mentioned above, fish was used as an effective bait to catch the
turtles in many cases, and fish are therefore overrepresented as prey for
this species. Sloan et al. (1996) found fish in 56.9% of the adult M.
temminckii in their study; likewise, Dobie (1966) noted fish in 59.4% of
the 32 stomachs he examined.
2006 R.M. Elsey 449
Crawfish were seen more frequently in our study (51.38% overall, and in
92.11% of the samples from Venice) than other reports (26.2% frequency
occurrence in Sloan et al. study [1996] and 43.75% in Dobie [1966]).
Molluscs (freshwater mussels and snails) were the third most commonly
seen item in this study and occurred in 47.71% of all 109 samples. Dobie
(1966) found molluscs in 21.9% of 32 samples; Sloan et al. (1996) noted
only one snail recovered in their study. Other authors have noted Alligator
Snapping Turtles feed on molluscs (Dundee and Rossman 1989, Ernst et al.
1994, George 1987). Pritchard (1989) states that the jaws of large M.
temminckii show considerable adaptation to a molluscivorous diet, due to
their breadth, shortness, blunt crushing surface, and strength.
My findings of turtles as M. temminckii prey are in agreement with prior
studies; no new species of turtles were noted as prey. Although smaller turtles
are very frequently consumed by M. temminckii (B. Harrel, pers. comm.),
turtles may be somewhat overrepresented as a diet component, since each
one’s heavy dermal bone is probably very slowly digested as compared to soft
body parts of other species such as small fish or amphibians.
Surprisingly, insect parts were seen in 22% of the samples in this study,
but were not reported either by Dobie (1966) or Sloan et al. (1996). The
expected very low prey masses recovered (range 0.01–1.78 g) suggest insects
add little in terms of the overall nutrient intake of the Alligator
Snapping Turtle.
Nutria were the sixth most frequently occurring (21.1% overall, but
60.53% from Venice) prey in this study, but the highest in terms of average
prey mass. Sloan (1996) found 15.4% of their samples contained mammals
(raccoons, muskrat and rabbits), but none were nutria. Dobie (1966) recovered
mammal remains in 28.1% of his samples; only two of which were
identified as nutria.
Rare mammalian parts seen were armadillo, opossum, squirrel (one case
each) and hogs (two cases). To my knowledge these have not been reported
as prey items for M. temminckii. These uncommon mammals were all noted
in samples from SE Arkansas; Sloan et al. (1996) included a few samples
from Arkansas. Dobie (1966) used 231 M. temminckii from several states for
his reproductive study; the state of origin for the 32 samples selected for
food analyses was not specified. These mammalian species might have been
consumed as carrion; it seems less likely that a slow-moving M. temminckii
would catch some of these mammals as live prey.
Snakes were seen in 6.42% of all samples in this study; Dobie (1966)
noted snakes in at least 15.6% of the 32 samples he examined (one listing
of unidentified reptile), and none were recovered by Sloan et al. (1996).
Species noted by Dobie (1966) were Lampropeltis g. holbrookia and
Nerodia sp. The Glossy Crayfish Snake found in this study appears to be a
new prey-item record.
450 Southeastern Naturalist Vol. 5, No. 3
Birds were uncommonly seen (5.5%) in this study; similarly, Dobie
(1966) saw only two cases of bird remains, and Sloan et al. (1996) saw
birds in only three stomachs. In the present study, eggs/inner eggshell
membranes were seen in seven cases; at least one of these appeared to be
avian in origin.
Vegetation, which sometimes included acorns, was seen in 108 of the
109 gastrointestinal tracts examined. Of interest, Sloan et al. (1996) noted
acorns alone were the most abundant food item by weight (average =
129.7 g, range = 1–643 g) and volume in their study. Dobie (1966) also
noted plant material in all 32 samples he examined.
Endohelminths were found in 67% of the gastrointestinal tracts in this
study, a far higher frequency than the single case of leeches noted by Sloan
et al. (1996) in one stomach. Cahn (1937), as noted in Pope (1939), stated as
many as 250 specimens of the same species of nematode were taken from a
single large intestine of an Alligator Snapping Turtle. In this study, one
intestine from SE Arkansas contained 338 endohelminths.
The finding that M. temminckii stomachs contained an average of 2.28
different prey items and intestines contained 4.13 different prey items illustrates
that M. temminckii are adaptable to preying on a variety of species. Of
great importance, numerous prey items (some of which are new findings for
M. temminckii) would have been missed if only stomachs (and not intestines)
were evaluated.
The ability of Alligator Snapping Turtles to consume such a varied diet
of prey species which are readily available to them may be advantageous in
allowing it to adapt to a variety of aquatic habitats. Recent measures undertaken
by numerous southeastern states (prohibition of commercial harvest
and additional efforts to study population status and distribution), along with
habitat preservation and enhancement, may lead to maintenance or improvement
in Alligator Snapping Turtle populations.
Acknowledgments
Thanks are extended to Don Delesdernier, Harvey Kliebert, and Homer Wiley for
access to their seafood processing sheds and assistance with data collection. Several
Louisiana Department of Wildlife and Fisheries employees (especially Melvin
Bertrand and Donna Bowman) assisted with field carcass collections and lab dissections.
Mr. W. Parke Moore III is acknowledged for administrative support, and the
late Dr. Barry Moser conducted statistical analyses. Dr. Jeff Boundy reviewed an
earlier draft of the manuscript and provided several helpful references. I thank Karen
McCall for technical assistance in preparation of the manuscript.
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