Robert G. Webb

“ But, different species of turtles prefer different kinds of aquatic habitats. Also, barriers occur in large drainage systems, such as the Mississippi, where, in general, the western tributaries are sluggish, turbid and shallow, and the eastern tributaries are fast-flowing, clear and deep. But in young, relatively small, river systems that do not traverse radically different physiographic regions, and that show no gross ecological differences, habitats or microhabitats that do exist probably are only partial barriers and seem not to prevent the dispersal of most kinds of aquatic turtles. ”

“ Baur (1891:637-38) thought that the family arose directly from the Amphichelydia, that the ancestors of the Trionychidae closely resembled Carettochelys in the structure of the carapace and plastron, and that a progressive reduction in ossification of those structures occurred. Nopcsa (1926:654) also wrote that the family originated from ancestors having a well-developed plastron; he maintained that the progressive reduction in ossification of the plastron was a specialization for aquatic life, and that the more primitive trionychids had the best developed bones and callosities. ”

“ The marginal rim is absent in females of T. ater, and only faintly evident in males. The marginal rim is obscured or absent (adult males and females) and is not separated from the ground color of the carapace by a dark marginal line in hatchlings of T. ferox. The carapace of T. muticus has a marginal rim that is usually separated from the ground color of the carapace by an ill-defined, dark marginal line; some individuals lack the marginal dark line. The subspecies of T. spinifer have a well-defined, dark, marginal line that separates the marginal rim from the ground color of the carapace ”

“ 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. ”

“ The smaller adult females probably mature sexually in their sixth year, but most probably do so when seven years old. Accordingly, some T. spinifer emoryi, which are sexually mature at a plastral length of 16.0 centimeters, are also sexually mature in their seventh year, whereas most spinifer (sexually mature at a plastral length of 18.0 to 20.0 cm., corresponding to a length of carapace of 25.2 to 28.0 cm. or about 10 to 11 inches) probably become sexually mature in their ninth year, and some when eight years old. ”

“ There is one difference in coloration between the sexes in the subspecies T. s. emoryi. Males from the Río Grande drainage, at least those from the Big Bend region of Texas, and southwestward in the Río Conchos into Chihuahua, México, are bright orange on the side of head (postlabial and postocular pale areas) ; an orange tinge also occurs in pale stripes on the snout, and pale orange blotches sometimes occur on the dorsal surfaces of limbs, especially the hind limbs. The coloration of these areas on females is pale yellow, lacking orange. ”

“ Surface of carapace smooth in adult males; large females lacking prominences posteriorly in center of carapace or in nuchal region; anterior edge of carapace smooth in both sexes, but occasionally having regularly spaced furrows or wrinkles (Fig. 8g) .
Variation.—Short dusky lines and streaks seem to be lacking from the juvenal pattern on the carapace more often in southern populations (Gulf Coast drainage of Texas) than in northern populations (Mississippi River drainage) . ”

“ Larger ratios are usually derived from measurements of larger individuals. Seemingly, there should be a clinal tendency in ontogenetic variation paralleling the size of ocelli and dependent on it; ontogenetic variation should be least in western populations in which the size of ocelli does not change appreciably with increasing size, and should be greatest in eastern populations in which the ocelli on adult males are larger than those on the carapace of young turtles. ”

“ The transition from the dark coloration of hatchlings to the paler coloration of adults is gradual and subject to individual variation. The loss of dark color ventrally occurs first on the plastral area, then the hind limbs, forelimbs, posterior part of carapace and last on the neck and throat. The soft parts of the body dorsally are gray or dark gray, and do not become so pale as the ventral surface. The smallest specimen that I have seen displaying the dark features of the hatchlings is a male, 7.7 centimeters (UMMZ 100673) ”

“ The seemingly greater restriction of muticus to running-water habitats suggests less vagility than in spinifer (Netting, 1944:86) .
Size and coloration are adaptations to habitat. Soft-shelled turtles of large size are best adapted to mesic, essentially continuous swampy or marshy habitats, whereas small size is an adaptation to less continuous, semi-isolated habitats. A turtle of the maximum size attained by ferox in the habitat of emoryi would, in a general way, probably be more conspicuous and exposed to its enemies, both in the aquatic environment and during overland excursions ”

“ In North America, soft-shelled turtles (genus Trionyx) occur in northern México, the eastern two-thirds of the United States, and extreme southeastern Canada. The genus fits the well-known Sino-American distributional pattern. In North America there are four species. Three (ferox, spinifer and muticus) are well-differentiated [591] and one (ater) is not well-differentiated from spinifer. Characters of taxonomic worth are provided by the following: size; proportions of snout, head and shell; pattern on carapace, snout, side of head, and limbs; tuberculation; sizes of parts of skull ”

“ For convenience, all softshells having locality data from states east of the Mississippi River are referred to spinifer, recognizing that intergradation occurs with hartwegi over a broad area paralleling the Mississippi River. The type locality of spinifer is in an area where most turtles do not have the larger ocelli (diameter of seven to ten mm. on adult males) ; however, some individuals from the Wabash River (UMMZ 63523, adult male, plastron 11.5 cm. in length, ocelli diameter seven mm.) agree with more "typical" spinifer to the east. ”

“ In polytypic species, the diagnosis of a subspecies is designed only to distinguish it from other subspecies of that species. The comments included under the subsection entitled "Description" pertain to individuals from an area where the taxon is most clearly differentiated. Because osteological characters are significant only at the specific level, they appear under the accounts of each species (excluding ater) . ”

“ Trionychid turtles may have traversed the Bering land bridge between Asia and North America in late Mesozoic times for they occur as fossils on the Atlantic Coast and in the Rocky Mountain-Great Plains region in Upper Cretaceous deposits. Shallow, inland seas may have afforded no barrier to the dispersal of softshells which presumably were tolerant of saline waters. The orogeny and volcanic action with subsequent erosion and sedimentation of the Rocky Mountain system, which was later accompanied by drier climates, tended to obliterate suitable habitats in the western United States ”

“ If this is true, corresponding patterns of distribution might indicate the same relative age of the population of turtles concerned. Generally, the genera of turtles that on morphological grounds are considered the oldest and most primitive (Macroclemys, Chelydra) show less differentiation into species and subspecies than those considered younger and more recently evolved (Graptemys, Pseudemys) . In the genus Graptemys, much differentiation occurs in the geologically, recently formed, Gulf Coast drainage systems of the southeastern United States. ”

“ From a distance I observed an individual of T. s. asper bury itself in shallow water on the Escambia River, Florida. Small individuals of muticus have been taken by hand along the shore of Lake Texoma. Along the Flint River near Bainbridge, Georgia, two hatchlings that were buried in sand in shallow water emerged at my approach and scurried a few inches, then buried themselves again. Larger turtles seem to be more wary. One that was disturbed, emerged from the sand and swam toward deep water.
In clear water, water-goggling may be effective in securing softshells. ”

“ Many characters of Trionyx ferox that are lacking in other North American forms are shared with some Asiatic softshells, such as the large size, longitudinal rows of tubercles that resemble ridges on the carapace, and the marginal ridge. It is thought that, of the living softshells in North America, ferox is more closely allied to Old World forms of the genus than to muticus or spinifer. ”

“ Remarks.—Intergradation of the subspecies pallidus and guadalupensis is of a clinal nature in which populations successively show a gradual resemblance to guadalupensis from western Louisiana and eastern Texas westward to central Texas. Because the sharpest break in this cline of characters occurs between the Colorado and Brazos River drainages, the turtles living in the Brazos River drainage and eastward are referred to pallidus, whereas those in the Colorado River drainage and westward are referred to guadalupensis. ”

“ Certain activities of softshells such as burying themselves in soft sand in shallow water or seeking crawfish and other food over a gravel-rock substrate or one that is débris-laden, are best carried on in different habitats. Repeated observations of turtles that are probably engaged in a specific activity in a restricted area may lead to erroneous general conclusions regarding the over-all preference for a specific habitat. ”

“ An arid environment in the central and southern United States and northern Mexico may have increased in area especially southward from Miocene times into the Pliocene (Dorf, 1959:189, 191) . The combination of physiographic changes and aridity, which modified the mesic, essentially continuous, aquatic habitats, may have isolated and aided in the differentiation of the ferox, muticus and spinifer stocks. ”

“ I doubt that a softshell can pursue and capture a healthy fish in natural waters. Recently dead fish are eaten and perhaps fish eggs, and senile and decrepit fishes. There is no evidence that soft-shelled turtles are active predators on any kind of fish. Of course in congested areas such as ponds of fish hatcheries, it is desirable to eliminate the turtles. ”