Читать книгу North American Recent Soft-Shelled Turtles (Family Trionychidae) - Robert G. Webb - Страница 3

Type Species.—Trionyx aegyptiacus (= Testudo triunguis Forskål).

Diagnosis.—Cutaneous femoral valves absent; width of postorbital arch of skull less than diameter of orbit; pterygoids usually not contacting opisthotics; carapace lacking prenuchal bone and marginal ossifications; nuchal bone lacking conspicuous ventral ridges; posterior margin of nuchal overlying first pair of pleurals; lateral parts of nuchal bone overlying second pair of ribs; neurals seven or eight, rarely six or nine; pleurals seven or eight pairs, posterior one or two pairs sometimes in contact medially; distinct suture usually present between hyoplastra and hypoplastra; most laterad prong of posteromedial process of hypoplastra inserted between bifid anterolateral process of xiphiplastra.

Synonomy.—Geoffroy published a synopsis of the species he recognized (1809) prior to his formal description of the genus Trionyx (1809a). Schweigger, nevertheless, probably was the first person to recognize the soft-shelled turtles as a distinct group, and he proposed for it the name Amyda in an unpublished manuscript that he sent to Geoffroy. The latter author (1809a:15) relegated the name Amyda to the synonomy of Trionyx javanicus by means of the following entry: "Amyda javanica. Schweigger, dans un manuscript communique a l'Institut." Stejneger (1944:7) maintained that this publication of Schweigger's monotypic generic name clearly established its availability for the species congeneric with Amyda javanica (= Testudo cartilaginea Boddaert, 1770). Loveridge and Williams (1957:422) contend that this mere mention of the name Amyda neither constitutes the proposal of a new name nor validates it, and that the first valid usage of the name Amyda is that of Fitzinger (1835:120), who later (1843:30) designated the type species as Amyda subplana. The name Amyda cannot date from Oken (1816:348) as Volume 3 [Zoologie] of his Lehrbuch der Naturgeschichte published in 1815–1816 has been placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature with the Title No. 33; see Opinion 417 (Hemming, 1956).

There has been considerable debate as to whether Geoffroy did or did not designate a type species of the genus Trionyx (1809a). Although not specifically designated as the type species, Trionyx aegyptiacus (= Testudo triunguis Forskål) is considered by Smith (1930:2), Schmidt (1953:108, footnote), and Loveridge and Williams (1957:422) to have been sufficiently indicated by Geoffroy as the type species. But Stejneger (1944:6), H. M. [445] Smith (1947:122), Conant and Goin (1948:11), and Mertens and Wermuth (1955) maintained that Geoffroy did not adequately designate a type species, and that Fitzinger (1843:30) designated the type species as Trionyx granosus (= Lissemys punctata), a synonym of Geoffroy's species, coromandelicus.

If Fitzinger's designation of a type species is accepted, the name Trionyx is applicable to the forms herein referred to Lissemys, and Amyda to the American forms. If Geoffroy's designation is accepted, the American forms are referable to Trionyx, and Amyda is a synonym.

The preceding includes only those generic names (listed in chronological order) that have been applied to Recent American soft-shelled turtles. Generic synonyms of the genus Trionyx applicable to Old World species are listed by Stejneger (1907:514), Smith (1931:165), and Loveridge and Williams (1957:420–21).

Trionyx is the most widespread genus of the family; most of the species occur in southeastern Asia. All North American soft-shelled turtles belong to this genus.

For quick reference, all the specific and subspecific names proposed for soft-shelled turtles in North America are listed below in alphabetical order (left hand column) with their nomenclatural status as recognized in this paper. The synonyms are listed in the account of the appropriate species or subspecies, and are discussed under the subsection entitled "Remarks."

agassizi	Trionyx	spinifer asper
annulifer	Trionyx	spinifer spinifer
argus	Trionyx	spinifer spinifer
asper	Trionyx	spinifer asper
ater	Trionyx	ater
bartrami	Trionyx	ferox
emoryi	Trionyx	spinifer emoryi
calvatus	Trionyx	muticus calvatus
ferox	Trionyx	ferox
georgianus	Trionyx	ferox
georgicus	Trionyx	ferox
harlani	Trionyx	ferox
hartwegi	Trionyx	spinifer hartwegi
hudsonica	Trionyx	spinifer spinifer
mollis	Trionyx	ferox
microcephalus	Trionyx	muticus muticus
muticus	Trionyx	muticus muticus
nuchalis	Trionyx	spinifer spinifer
ocellatus	Trionyx	spinifer spinifer
olivaceus	Trionyx	spinifer spinifer
spiniferus	Trionyx	spinifer spinifer

Variation

Aside from qualitative variations and comparisons of patterns of pigmentation the following external measurements (to the nearest millimeter) were used.

Length of plastron: Maximal straight-line measurement (midventrally), from the anteriormost region of the ventral surface to the posterior end of the plastron; this measurement includes an anterior cartilaginous part.

Length of carapace: Maximal, straight-line measurement (middorsally), from the nuchal region to the posteriormost region of the free edge of the carapace.

Width of carapace: Maximal, straight-line measurement between the lateral margins of the carapace.

[446]

Plane of greatest width of carapace: Maximal, straight-line measurement from the posteriormost region of the free edge of the carapace to a point on the middorsal line at the level or plane of the greatest width of the carapace; this measurement and the last two, of course, include the fringing cartilaginous parts of the dorsal bony carapace.

Width of head: Maximal measurement between the lateral margins of the head.

Length of snout: Measurement from tip of snout to interorbital region of least breadth.

Diameter of ocellus: Maximal outside diameter of largest (not conspicuously ovoid or oblong) ocellus on carapace.

The following ratios were developed from the measurements. Reference to these ratios will be made by the abbreviations within the parentheses: length of carapace/length of plastron (CL/PL); length of carapace/width of carapace (CL/CW); length of carapace/plane of width of carapace (CL/PCW); length of plastron/width of head (PL/HW); width of head/length of snout (HW/SL); diameter of ocellus/length of plastron (OD/PL).

Secondary Sexual Variation

Size

In many species of turtles, females are larger than males; the difference in size between the sexes is probably most pronounced in aquatic emydids. The ten largest individuals of each sex were selected to indicate the relative difference in size between the sexes of the three American species of Trionyx (excluding ater, Table 2). Female soft-shelled turtles attain a larger size than males. T. ferox is the largest species; muticus is the smallest. The approximate maximal size of each sex and the difference in size between the sexes are more correctly expressed for spinifer and muticus than for ferox, because fewer specimens of ferox were examined; presumably the approximate maximal size of males and females of ferox is larger than is indicated in Table 2.

Table 2. Secondary Sexual Difference in Maximal Size of North American Species of the Genus Trionyx (excluding ater) Based on the Ten Largest Specimens of Each Sex of Each Species. The Extremes Precede the Mean (in parentheses).

Species	Plastral length (cm.)
ferox	males	17.0–26.0 (20.0)
	females	23.3–34.0 (27.9)
spinifer	males	13.8–16.0 (14.4)
	females	26.0–31.0 (28.0)
muticus	males	11.8–14.0 (12.3)
	females	17.7–21.5 (18.9)

Pattern

Secondary sexual differences in pattern are probably more pronounced in soft-shelled turtles than in other species of turtles, except perhaps for the well-known melanism and concomitant obliteration of pattern acquired by some adult males of the scripta section of the genus Pseudemys.

[447]

The difference in pattern between the sexes of American species varies with size of the individual and with the species and subspecies. The juvenal pattern of some individuals of T. spinifer asper differs according to sex. In the other species and subspecies, there are no secondary sexual differences in the juvenal pattern. That pattern in females of all species and subspecies is partly or entirely obscured by a mottled and blotched pattern as growth proceeds. This mottled and blotched pattern is present on females not yet sexually mature, and is of contrasting lichenlike figures, and in other individuals is less contrasting and a more uniform coloration. The largest males of T. spinifer retain a conspicuous juvenal pattern; in those of muticus the pattern may be well-defined or partly modified and obscured, whereas in large males of ferox the juvenal pattern is ill-defined or absent. No male normally acquires a contrasting mottled and blotched pattern on the carapace. The pattern on the carapace of many large individuals of ferox is not distinctive as to sex.

On the dorsal surface of the soft parts of the body there is a contrasting pattern in adult males and hatchlings of some forms, but in most large females the pattern is usually reduced to a near-uniform coloration; the pattern on adult males of ferox and muticus is not contrasting and resembles that on large females.

Coloration

Because most specimens examined were preserved, the detection of secondary sexual differences in coloration was difficult. 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.

Tuberculation

In all subspecies of spinifer the carapace of adult males is "sandpapery" owing to abundant, small, spiny tubercles distributed over its surface; all females lack spiny tubercles on the surface of the carapace.

Length of Tail

Elongation of the preanal region of the tail resulting in the extension of the cloacal opening beyond the posterior edge of the carapace occurs in males of several kinds of turtles, including Trionyx, at least in those from Louisiana, Texas, and Lake Texoma, Oklahoma (Webb, 1956:121). Probably this elongation is characteristic of males of all American softshells. Some females of spinifer and muticus that exceed the maximum size attained by males have the tip of the tail and cloacal opening extending a short distance beyond the posterior edge of the carapace. Some large females of ferox have more elongate tails than those of spinifer and muticus.

Width of Alveolar Surfaces of Jaws

Stejneger (1944:34–36, pl. 6) commented on a series of large skulls of ferox mostly from Kissimmee, Florida, some of which had conspicuously expanded alveolar surfaces. He suggested that the condition was confined to large males. A scattergram (Fig. 2) based on measurements obtained from 45 skulls of ferox shows widened alveolar surfaces of the upper jaws on some of the larger [448] skulls. Because the maximal size of adult males is unknown and the difference in size between the sexes of ferox is slight, such large skulls might represent either sex. The sex had been recorded for only three of the 45 skulls; none of the three exceeded 82 millimeters in basicranial length or had widened alveolar surfaces. Some of the larger skulls of approximately the same size differ markedly in width of the alveolar surfaces; this difference suggests that both sexes are included and that the sexes may be of approximately the same maximal size. On the other hand, the variation observed in skulls is possibly confined to one sex. To judge from what is known of the maximal sizes of the sexes of spinifer and muticus (see Table 2), skulls of ferox of more than 85 millimeters in basicranial length probably are of females. The largest alcoholic male (dissected) of ferox that I examined had a width of head of approximately 46.5 millimeters; that measurement corresponds to a basicranial length of 70 to 75 millimeters. The specimen of which measurements are depicted by the uppermost symbol in the scattergram (represented by KU 16528) was recorded as a female. Large females of T. s. asper from rivers emptying into the Atlantic Ocean have broadened alveolar surfaces.

Fig. 2. Basicranial length and greatest width of alveolar surface of upper jaw on 45 skulls of T. ferox. Some skulls (sex unknown) in which the basicranial length exceeds 85 mm. develop widened alveolar surfaces of the jaws.

Length of Claw

Secondary sexual differences in length of claw on the forelimb are pronounced in some kinds of turtles. Cahn (1937:178) stated that the female of Trionyx muticus usually has long claws on the hind feet, while the male has long claws on the forefeet, but I am unable to substantiate his statement. Measurements of length of the third claw on the hind limb taken in 41 males and 45 females of spinifer from Louisiana showed no secondary sexual difference.

Ontogenetic Variation

Pattern

In all species and subspecies the juvenal pattern is replaced in females as growth proceeds by a mottled and blotched pattern that is contrasting or of nearly uniform coloration. The blotched pattern (of lichenlike figures) is evident on the carapaces of most females that have plastra so long as 8.0 centimeters. The contrasting juvenal pattern on the dorsal surfaces of the soft parts of the body is correspondingly modified in females, but at a size larger than 8.0 centimeters. Size of ocelli (OD/PL) in T. s. spinifer and hartwegi seems to vary ontogenetically (see section on Geographic Variation).

Some hatchlings have blotched patterns (T. spinifer asper, TU 16689.2, plastral length, 3.5 cm.); the largest females examined that did not show any evidence of mottling were two asper having plastrons 7.6 and 8.0 centimeters in length. Variation in color and pattern probably is modified greatly by the environment (Heude in Stejneger, 1907:518, footnote d) and the physiological condition of the individual. Smith, Nixon and Minton (1949:92) reported that a female of T. s. hartwegi developed a striking melanistic pattern in captivity and they concluded that patterns of soft-shelled turtles may be produced not only by conventional chromatophores, but also by other depositions, both intra- and extracellular. TU 16170, taken from brackish water at Delacroix Island, St. Bernard Parish, Louisiana, is the only adult male I have seen that had a blotched pattern (orange-brown in life) on the carapace in addition to the juvenal pattern. One female of muticus, KU 48229, having a plastral length 14.5 centimeters, retained a well-defined juvenal pattern, and lacked a mottled and blotched pattern (see Pl. 46).

Tuberculation

Males of the subspecies of spinifer develop small, sharp tubercles on the dorsal surface of the carapace when sexually mature. As growth proceeds, the minute prominences along the anterior edge of the carapace on hatchlings of both sexes of spinifer change in shape to conical projections or low, flattened, scarcely-elevated prominences, depending on the subspecies (Fig. 8).

Large females of spinifer and ferox acquire enlarged, flattened knobs in the nuchal region and posteriorly in the center of the carapace.

Length of Tail

The preanal region of the tail rapidly elongates in males of all soft-shells when they are sexually mature.

Width of Alveolar Surfaces of Jaws

The alveolar surfaces of the jaws are conspicuously broadened in large adults of ferox, and females of that population of T. s. asper in the Atlantic Coast drainage.

Ratios

Width of head increases at a rate slightly slower than does the length of the plastron (PL/HW, Fig. 3). The change in proportions is most pronounced [450] at a plastral length of 7.5 to 8.0 centimeters. In general, the head is narrowest in muticus and widest in ferox. T. s. asper and emoryi seemingly have the widest heads among the subspecies of spinifer. Geographically width of head increases from spinifer and hartwegi through pallidus and guadalupensis to emoryi. T. ater terminates the cline; 12 specimens, ranging in plastral length from 9.6 to 18.4 centimeters, resemble ferox and asper in having wide heads (average PL/HW of 4.93).

Fig. 3. Ratio of length of plastron to width of head (PL/HW) in some American species and subspecies of the genus Trionyx. The size of each sample is given in parentheses following an indication of the range (< = less than, > = greater than) in length of plastron (in cm.) of each sample. The horizontal line indicates the observed variation; the vertical line, the mean; the white rectangle, four standard deviations; and the black rectangle, four standard errors of the mean. There is some ontogenetic variation in PL/HW. The head is narrowest in muticus and widest in ferox.

The carapace increases in width more slowly than it increases in length (CL/CW, Fig. 4). The change in proportions is most pronounced when the carapace is 8.0 to 8.5 centimeters in length. Ontogenetically muticus varies least and ferox most; large specimens of ferox have narrower carapaces than muticus of corresponding size. There is also an indication of a geographical gradient that parallels the cline mentioned above for PL/HW. There is a gradual decrease in width of carapace from pallidus through guadalupensis to emoryi. Of the subspecies of spinifer, emoryi has the narrowest carapace and [451] resembles ferox. In T. ater this cline is accentuated and terminates; 12 specimens, ranging in plastral length from 9.6 to 18.4 centimeters, resemble ferox and emoryi in having narrow carapaces (average CL/CW of 1.32).

Osteological Characters

Closure of the anterior, paravertebral fontanelles on the bony carapace, and size and number of plastral callosities are subject to ontogenetic variation (see sections entitled "Carapace" and "Plastron").

Fig. 4. Ratio of length of carapace to width of carapace (CL/CW) in some American species and subspecies of the genus Trionyx. Symbols as in Fig. 3. There is some ontogenetic variation in CL/CW (least in muticus). The carapace is narrowest in ferox and emoryi, and widest in muticus, pallidus and asper.

Fig. 5. Pattern on dorsal surface of snout of some American species and subspecies of the genus Trionyx. Note the gradual transition in pattern from that of hartwegi (b) and asper (c) to that of emoryi (h).

1 T. ferox (UMMZ 102276, × 1/3)

2 T. spinifer hartwegi (KU 46742, × 3/4)

3 T. spinifer asper (KU 50842, × 1)

4 T. spinifer pallidus (KU 2958, × 1/2)

5 T. spinifer pallidus (KU 2934, × 1/2)

6 T. spinifer pallidus (KU 2947, × 1/2)

7 T. spinifer guadalupensis (TU 10165, × 2/3)

8 T. spinifer emoryi (KU 48218, × 2/3)

9 T. muticus muticus (KU 48236, × 2/3)

Geographic Variation

Geographic variation occurs in Trionyx spinifer and T. muticus. The variant populations of spinifer are segregated into six subspecies, those of muticus into two. In the subspecies of spinifer there is both group variation and clinal variation.

Group Variation

The six subspecies of spinifer can be separated into two groups on the basis of the juvenal pattern. One group (subspecies spinifer, hartwegi and asper) has a pattern of dark spots or ocelli of various sizes on the carapace, whereas the other group (subspecies pallidus, guadalupensis and emoryi) has a pattern of small white dots or tubercles on the carapace. The two groups differ also in the manner in which the mottled and blotched pattern first appears on the carapace of females. Usually, contrasting lichenlike figures initially surround the dark spots or ocelli on the carapace in females of the spinifer group (less evident in pallidus), whereas females of the emoryi group usually lack a contrasting pattern early in ontogeny. In general, the two groups differ in the degree of pigmentation. The spinifer group has larger marks and more contrasting patterns on the head and limbs, and more extensive pigmentation on the ventral surface than members of the emoryi group. T. ater is more closely related to those subspecies of the emoryi group but differs in having the ventral surface heavily speckled with black and an over-all blackish, dorsal coloration; the underlying pattern of ater resembles that of emoryi.

Clinal Variation

Several characters are arranged in a geographical gradient or cline. Some characters are relatively uniform and represent a terminus in the spinifer group. Some characters change gradually and successively through the subspecies pallidus and guadalupensis, and terminate in emoryi and T. ater. Some characters of ater, in turn, show affinity with T. muticus and T. ferox.

Pattern on Snout

The pattern (Fig. 5) on the snout usually consists of pale, dark-bordered stripes that form an acute angle in front of the eyes in spinifer, hartwegi and asper, but the corresponding marks form a dark triangle the base line of which joins the anterior margins of the orbits in emoryi and usually in guadalupensis. In pallidus, the geographic range of which is between guadalupensis and hartwegi, there are different patterns that are in various degrees intermediate between those described immediately above for hartwegi and guadalupensis.

Pattern on Side of Head

The change in pattern (Fig. 6) and its contrast with the ground color on the side of the head parallels the sequence of changes in pattern on the snout. The pattern on the side of head contrasts with the ground color and consists of dark markings below the eye and on the neck, an indication of a postlabial stripe, and a pale, dark-bordered postocular stripe that may be variously interrupted (spinifer and hartwegi; asper usually has uninterrupted postocular and postlabial stripes that unite on the side of the head). The pattern is contrasting but variable in pallidus. T. s. emoryi and usually guadalupensis have fewer dark markings, sometimes none, and an interrupted postocular pale stripe that produces a pale blotch just behind the eye.

Fig. 6. Pattern on side of head of some American species and subspecies of the genus Trionyx. Note the gradual reduction in contrast of pattern and interruption of the postocular stripe from that of spinifer (b) to that of emoryi (f).

1 T. ferox (UMMZ 102276, × 1/3)

2 T. spinifer spinifer (UMMZ 54401, × 2/3)

3 T. spinifer asper (KU 50843, × 2/3)

4 T. spinifer pallidus (KU 50830, × 3/4)

5 T. spinifer guadalupensis (SM 659, × 2/3)

6 T. spinifer emoryi (KU 2922, × 3/4)

7 T. muticus muticus (KU 48228, × 2/3)

8 T. muticus calvatus (KU 47117, × 2/3)

Fig. 7. Pattern on the dorsal surface of the distal part of the right hind limb of some American species and subspecies of the genus Trionyx. Note the gradual reduction in contrast of pattern from that of hartwegi (a) to that of emoryi (d).

1 T. spinifer hartwegi (KU 15932, × 3/4)

2 T. spinifer pallidus (KU 40175, × 2/3)

3 T. spinifer guadalupensis (TU 10165, × 3/4)

4 T. spinifer emoryi (KU 3153, × 5/6)

5 T. muticus muticus (KU 48228, × 3/4)

6 T. ferox (UMMZ 102276, × 1/2)

Fig. 8. Shape of tubercles on anterior edge of carapace in some American species and subspecies of the genus Trionyx (× ¹/₂). Note the gradual reduction in size of tubercles from that of hartwegi (b) to that of muticus (h).

1 T. ferox (UMMZ 90010)

2 T. spinifer hartwegi (KU 3346)

3 T. spinifer pallidus (TU 13213)

4 T. spinifer guadalupensis (TU 10160)

5 T. spinifer emoryi (KU 2906)

6 T. ater (KU 46906)

7 T. muticus muticus (KU 48229)

8 T. muticus muticus (KU 48232)

Pattern on Dorsal Surface of Limbs

A corresponding sequence of change occurs in the size of dark markings on the dorsal surface of the limbs (Fig. 7). The hind limb usually has larger markings than the forelimb. The change is gradual from larger and darker markings (contrasting pattern) in hartwegi, spinifer and asper to smaller and paler markings (non-contrasting pattern) in emoryi.

Tuberculation

There is also a cline in tuberculation (Fig. 8) that parallels geographically the sequence of changes in patterns mentioned immediately above. The size of the tubercles along the anterior edge of the carapace changes in both sexes from those that are enlarged and equilateral or conical in shape in spinifer, hartwegi, asper and pallidus to those that are scarcely elevated in guadalupensis, emoryi and T. ater. Indeed, in the three kinds mentioned last, the tubercles are absent in some specimens. There seems to be a corresponding reduction in the size and number of small, sharp-tipped tubercles that cover the carapace in adult males; the carapace of T. ater is mostly smooth and has only a few small, whitish tubercles.

Fig. 9. Anteroposterior position of plane of greatest width of carapace (CL/PCW) in some American species and subspecies of the genus Trionyx. Symbols as in Fig. 3. The greatest width of carapace is midway between anterior and posterior ends in ferox, spinifer, hartwegi, asper and muticus, and farther posterior in the other subspecies of spinifer.

Ratios

The clinal tendencies in PL/HW (Fig. 3) and CL/CW (Fig. 4) that parallel those mentioned above for pattern and tuberculation have already been mentioned under the section "Ontogenetic Variation."

The ratio of CL/PCW (Fig. 9) was used in an effort to show further differences in the shape of the carapace, especially the plane on the carapace [459] where the greatest width occurs. Figure 9 shows the greatest width to be approximately midway between the anterior and posterior ends in the subspecies spinifer, hartwegi and asper, and in the species ferox and muticus (CL/PCW of 2.00). The greatest width of carapace is more posterior and at approximately the same plane in pallidus and guadalupensis, and farther posterior in emoryi. Calculated ratios for 12 specimens of T. ater average 2.15, a value that suggests closer affinity with pallidus, guadalupensis and emoryi than to the other species and subspecies.

Comparison of the relative lengths of snout (HW/SL, Fig. 10) in different populations of T. spinifer shows a character gradient. To facilitate a comparison utilizing large samples, the subspecies spinifer was combined with hartwegi, and pallidus with guadalupensis. The snout is longer in the subspecies spinifer and hartwegi than in emoryi; the length of the snout of emoryi resembles that of T. ferox. The snout is proportionately the longest in T. muticus. The average ratio of HW/SL for 12 individuals of T. ater is 1.37, and is nearer that of pallidus, guadalupensis, emoryi and ferox than that of muticus or the other subspecies of T. spinifer.

Fig. 10. Ratio of width of head to length of snout (HW/SL) in some American species and subspecies of the genus Trionyx. Symbols as in Fig. 3. Values for spinifer are combined with those of hartwegi, and those of pallidus with guadalupensis. The snout is proportionately the longest in muticus.

Size of the ocelli increases from west to east in populations of T. spinifer in the upper Mississippi River and Great Lakes drainages.

The ratio of OD/PL (Fig. 11) varies considerably but gradually increases from Kansas northeastward to Michigan. The minimal diameter of any ocellus recorded was one millimeter; solid dots on the carapace (hartwegi) were also recorded as one millimeter. 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. It is difficult to demonstrate [460] ontogenetic variation because specimens of corresponding size from the same general area may have ocelli of different sizes. The gradient in size of ocelli is also indicated by specimens from other states. I have the subjective impression that there is least variation in specimens from Michigan (Great Lakes-St. Lawrence River drainage), but this is not clearly shown by Figure 11.

Fig. 11. Ratio of diameter of ocellus to length of plastron (OD/PL) in T. spinifer from some states in the upper Mississippi River and Great Lakes drainages. Symbols as in Fig. 3. The size of the ocelli on the carapace gradually increases from Kansas northeastward to Michigan.

Character Analysis

Snout

The snout (Fig. 12) is tubate having terminal nostrils separated by a vertical septum. One of the principal characters distinguishing T. ferox and T. spinifer from T. muticus is a lateral, whitish ridge projecting from each side of the nasal septum (hereafter referred to as septal ridges but often referred to in the literature as a papilla). The shape of the end of the snout is truncate in T. ferox and T. spinifer, and the nostrils are larger than in T. muticus. In muticus the snout usually terminates somewhat obliquely, and the nostrils tend to be slightly inferior; also, the end of the snout is usually rounded and somewhat pointed, causing the nostrils to be visible in lateral view. Some T. muticus do not differ markedly from ferox or spinifer in shape of the end of the snout. Stejneger (1944:14) mentioned indication of a septal ridge that did not reach the opening of the nostril in muticus. I have slit the outer edge of the nostril on several specimens of muticus, and have not noticed an indication of a septal ridge.

Fig. 12. Shape of snout in T. spinifer (left, a-d, from KU 46907) and T. muticus (right, e-h, from KU 48236). Lateral views—a, e (× 1); anterior views—b, f (× 5); dorsal views—c, g (× 2.5); ventral views—d, h (× 2.5).

Tuberculation

Tubercles or obtuse prominences occur on the anterior edge of the carapace (Fig. 8) or on the dorsal surface of the carapace. Trionyx muticus lacks tubercles, although some individuals show shallow, widely spaced wrinkles that suggest prominences on the anterior edge of the carapace. Both sexes of T. ferox have prominences, resembling flattened hemispheres, on the anterior edge of the carapace and in the nuchal region. Large females of ferox have obtuse prominences in the center of the carapace posteriorly, some of which are often arranged in longitudinal rows. The surface of the carapace in both sexes of T. ferox has small closely-set, blunt tubercles arranged in rows that resemble longitudinal ridges (most evident in juveniles).

Large females of T. spinifer have obtuse prominences in the center of the carapace posteriorly, some of which in many specimens are arranged in longitudinal rows; I cannot discern any correlation of number or arrangement of prominences with size in spinifer or ferox. The carapace in adult males of spinifer bears small, sharp tubercles that make the surface feel like sandpaper. The tubercles on the anterior edge of the carapace in adults of both sexes vary from round to equilateral and conical to low and flattened (see comments on tuberculation under subsection entitled "Geographic Variation"). Some large females of the same subspecies have tubercles on the anterior edge of the carapace that may be conical (higher than wide) or equilateral. The difference in shape of the tubercles seems not to be correlated with size because one T. s. pallidus, 30.5 centimeters (TU 13212) has prominent but blunted and equilateral tubercles, whereas, another female of pallidus, 20.8 centimeters (TU 13210), from the same locality has higher, conical tubercles. The blunted, equilateral tubercles may be the result of environmental wear, or the difference in shape of tubercles may be due to individual variation.

Pattern on Carapace

Two features of the pattern on the carapace are of taxonomic worth: 1) the width and distinctness of the pale rim at the periphery of the carapace (marginal rim), if present, and 2) the kind of pattern on the carapace (juvenal pattern). 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; T. s. asper has more than one dark marginal line on the carapace. The marginal rim is ill-defined and blotched, or absent, in large females of all species of Trionyx.

The marginal rim is widest at the posterior end of the carapace and lacking in the nuchal area. The width of the pale marginal rim is very narrow, almost to the degree of being absent, in juveniles of T. ferox. T. s. emoryi has a pale, marginal rim that is four or five times wider posteriorly than it is laterally, whereas posteriorly the width of the rim in the other subspecies of T. spinifer and in the species T. muticus is only two or three times wider posteriorly than it is laterally.

The juvenal pattern commonly consists of whitish tubercles or dots (T. s. emoryi, T. s. guadalupensis, T. s. pallidus, T. ater), large black ocelli (T. s. spinifer), small black dots and ocelli (T. s. hartwegi, T. s. asper), large dusky spots or ocelli (T. m. calvatus), or small dusky dots or short streaks and dashes (T. m. muticus). Some hatchlings of pallidus and emoryi have a uniform pale brown or tan carapace; hatchlings of T. ferox have a distinctive pattern (Pl. 31). [463] Further comments and illustrations pertaining to kind of pattern on the carapace are offered under the accounts of species and subspecies.

Pattern on Dorsal Surface of Snout (Fig. 5)

T. ferox has pale stripes on a dark background that unite in front of the eyes; the dark ground color becomes paler with increasing size, but the stripes retain thick black borders. T. m. muticus has ill-defined, pale stripes that are evident just in front of the eyes and do not extend anteriorly to unite in front of the eyes, whereas T. m. calvatus lacks pale stripes on the snout. The kind of pattern on the dorsal surface of the snout that is characteristic for each of the subspecies of T. spinifer has been mentioned in the discussion of clinal variation.

Pattern on Side of Head (Fig. 6)

T. ferox has a pale broad, postocular stripe in contact with the orbit or not, and other pale marks on a dark background; the ground color becomes paler with increasing size, but the stripes and other marks retain thick black borders. T. m. muticus usually has an uninterrupted, dusky-bordered, postocular stripe, whereas T. m. calvatus (in adult males only) has pale postocular stripes with thick blackish borders. The pattern on the side of head that is characteristic for each subspecies of T. spinifer has been mentioned in the discussion of clinal variation.

Pattern on Dorsal Surface of Limbs (Fig. 7)

Young specimens of T. ferox have pale marks on a blackish background. As growth proceeds the distinctive contrasting pattern is obliterated and eventually is replaced by a uniform grayish coloration in large adults. The pattern on the limbs of T. muticus is not contrasting, and is almost a uniform grayish, consisting of fine, pale markings. The clinal variation in pattern and kind of pattern on the limbs of the subspecies of T. spinifer has been mentioned in the discussion of clinal variation. Dark markings tend to form streaks that are coincident with the digits, and larger markings occur on the hind limbs than on the forelimbs.

Marginal Ridge

The anterolateral edge of the carapace in T. ferox (both sexes and all sizes) is "folded over" into a ridge having a distinct inner margin (Pls. 1 and 2), which is hereafter referred to as the marginal ridge. Siebenrock (1924:184–85) referred to this ridge as a "Hautsäume" and mentioned its occurrence in Old World species of the genus Trionyx. The marginal ridge is not present in T. muticus, T. spinifer or T. ater.

Ratios