resembles Old World softshells more than it does any American species.
The northern part of the geographic range of _ferox_ overlaps that of
_T. s. asper_; there, the two species are ecologically isolated. _T.
spinifer_ is polytypic, has the largest geographic range, and is
composed of six subspecies, of which two are described as new
(_pallidus_ and _guadalupensis_). The subspecies are divisible into two
groups. One, the _spinifer_ group (_spinifer_, _hartwegi_ and _asper_)
is recognized by a juvenal pattern having black spots or ocelli; _asper_
is the most distinctive and shows little evidence of intergradation in
the lower Mississippi River drainage with the _spinifer-hartwegi_
complex, which, northward, is differentiated into two subspecies in
which there is an east-west cline in size of the ocelli on the carapace.
The _emoryi_ group (_pallidus_, _guadalupensis_, _emoryi_) is recognized
by a pattern of white spots; _emoryi_ is most distinctive. Each of
several characters behaves as a cline if traced from east to west
through the three subspecies. _T. s. pallidus_ intergrades with the
_spinifer-hartwegi_ complex in the lower Mississippi River drainage. _T.
s. emoryi_ is the most variable subspecies; in its most notable
population the males have orange coloration. _T. s. emoryi_ has been
introduced into the Colorado River drainage of Arizona. _T. ater_ most
closely resembles _T. s. emoryi_, but shows alliance with _T. muticus_
and _T. ferox_. _T. ater_ is confined to ponds of crystal-clear water in
central Coahuila, México. _T. muticus_ is completely sympatric with
_spinifer_, and is composed of two subspecies (_muticus_ and
_calvatus_). _T. m. calvatus_ shows no evidence of intergradation in the
lower Mississippi River drainage with _T. m. muticus_, corresponding
somewhat to the relationship of _T. s. asper_ with the intergradient
population of _T. spinifer_ in the Mississippi River.
Softshells have pharyngeal respiration and probably are incapacitated by
rotenone. _T. ferox_ and the subspecies of _spinifer_ occur in a wide
variety of fresh-water habitats; _muticus_ is more nearly restricted to
running water (especially in the northern parts of its range) than
_spinifer_, and may be less vagile than _spinifer_. _T. ferox_ is more
tolerant of marine and brackish waters than are _muticus_ or _spinifer_.
Small size and pallid coloration seem correlated with arid environments.
The largest species (_ferox_) and the smallest population of _spinifer_
(resembling _muticus_) both occur in the southernmost part of the range
of the genus. Diurnal habits include basking on shores or débris in
water, floating at the surface, procuring food, and burrowing in shallow
and deep water (no observations for _spinifer_ and _muticus_ in deep
water). Softshells are principally carnivorous; the food consists mostly
of crawfish and insects; there is evidence of cannibalism involving
predation on first- and second-year-old turtles. The capture of food is
triggered primarily by movement of prey; sight seems to be more
important than smell to _Trionyx_ in capturing food. There is no
indication of a food preference between species; enlarged crushing
surfaces of jaws in some _ferox_ and _asper_ may be an adaptation for
feeding on mollusks. Schools of fish are reported to follow softshells,
and presumably acquire food that is dislodged by the grubbing and
scurrying of the turtles on the bottom. Softshells are wary. They are
good swimmers, and travel rapidly on land. The depressed body is an
adaptation for burrowing and concealment. Permanent growths of algae do
not occur on the dorsal surface of softshells. There is evidence of some
nocturnal activity, and a general parallel in habits between trionychids
and chelydrids. Softshells sometimes move overland; they move little in
aquatic habitats. The normal annual period of activity of _spinifer_ in
latitudes 40° to 43° is approximately five months from April into
September, depending on the weather; they hibernate under a shallow
covering of mud in deep water. The southernmost populations may be
active throughout the year.
Males of _spinifer_ are sexually mature when the plastron is 9.0 to 10.0
centimeters in length (some when 8.0 long), whereas those of _muticus_
are sexually mature at 8.0 to 9.0 centimeters. In the mentioned size
range, the smaller adult males are probably in their fourth growing
season, and the larger males in their fifth. Most females of _spinifer_
are sexually mature at a plastral length of 18.0 to 20.0 centimeters and
are probably in their ninth year; the smaller individuals probably are
in their eighth. Females of _muticus_ are sexually mature when the
plastron is 14.0 to 16.0 centimeters long. Most of these are seven years
old but some are only six years old. Some large females contain immature
ovaries. The near-maximum length of carapace of _spinifer_ is 18 inches,
and such turtles are perhaps 60 years old; _ferox_ perhaps attains a
length of two feet.
_T. ferox_ deposits eggs from late March to mid-July, whereas northern
populations of _spinifer_ and _muticus_ usually deposit theirs from
mid-June to mid-July. Sandy sites are preferred for nests, although
movement to other sites occurs if the preferred sandy sites are
submerged or otherwise rendered unusable. _T. muticus_ limits its nest
sites to the open areas of sand bars and does not lay inland where it
must traverse vegetated areas, as does _spinifer_. Nests of _ferox_ and
_spinifer_ seem to differ from those of _muticus_ in being flask-shaped.
The seasonal reproductive potential is perhaps less in northern
populations (averaging 20 eggs per clutch and only one clutch per
season) than in southern populations (averaging about 10 eggs per
clutch, but three clutches per season). Larger females deposit more eggs
than smaller females. Eggs laid in northern latitudes are slightly
smaller than those laid farther south. In any latitude the incubation
period probably is at least 60 days. Hatchlings presumably leave nests
at dusk, nighttime or dawn, and may winter over in eggs or nests.
Man is a great enemy of softshells. Predation on eggs probably accounts
for most mortality. Physical conditions of the environment (overcrowding
of nest sites, inadequate hibernation sites) and probably some kinds of
parasitism contribute to mortality. Softshells are eaten locally and
sometimes appear in the market of large cities, but over most of their
range, there probably is no general demand and no special efforts are
made to capture them. Fish, mostly minnows, comprise a small proportion
of the diet. There is no evidence that softshells are active predators
on any kind of fish, but their known food habits suggest that they
compete with game fishes for food. Softshells are scavengers.
Fossil material was not studied in detail. The fossil softshells
indicate a more widespread, former distribution. Some osteological
characters and their variation in the living species are mentioned as an
aid to future workers concerned with an assay of fossil remains. Fossils
occur in marine, brackish and fresh-water deposits, and many are much
larger than the living species; the oldest American fossils are of Upper
Cretaceous age.
The interrelationships of the living species and subspecies suggest that
the species _spinifer_, _ater_, and _muticus_ are derivatives of a
_ferox_-like ancestor, and that they differentiated in North America;
most differentiation occurs in southwestern Texas and northern México
where characters of some populations indicate alliance with _ferox_. It
is hypothesized that aridity in the late Tertiary effected specific
differentiation by the modification and isolation of aquatic habitats.
Pluvial periods in the Pleistocene provided for confluence of aquatic
habitats and expansion of geographic ranges, and coupled with
physiographic changes, conceivably caused or enhanced some of the
subspecific variation.
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