Carapace olive, having large ocelli in center but smaller ocelli or spots at sides; ocelli often interrupted; pale rim of carapace not four or five times wider posteriorly than laterally, separated from darker ground color of carapace by one dark marginal line; large females often having remnants of ocelli at sides of carapace on mottled and blotched background; pattern on snout of pale, dark-bordered stripes that unite forming acute angle in front of eyes; well-defined dark markings in subocular and postlabial region; pattern contrasting [490] with ground color on side of head; postlabial stripe interrupted, diffuse; pale postocular stripe having blackish borders interrupted, not uniting with postlabial stripe; dorsal surface of soft parts of body having contrasting pattern, largest blackish marks on hind limbs; elongate tail of adult males having pale dorsolateral bands with well-defined lower blackish borders; underparts whitish, often having blackish marks, except in center of plastral area; dark marks on webbing of limbs, palms and soles; dark streaks often coincident with digits; small conical tubercles on anterior edge of carapace on adult males; conical or equilateral tubercles on anterior edge of carapace of large females; accessory knoblike tubercles in nuchal region and in middle of carapace posteriorly on large females.
Ontogenetic variation in PL/HW, mean PL/HW of specimens having plastral lengths 7.0 centimeters or less, 4.09, and exceeding 7.0 centimeters, 5.50; ontogenetic variation in CL/CW, mean CL/CW of specimens having plastral lengths 8.5 centimeters or less, 1.12, and exceeding 8.5 centimeters, 1.21; mean CL/PCW, 2.02; mean HW/SL, 1.30 (including subspecies hartwegi); mean CL/PL, 1.39.
Variation.—Variant individuals include: UMMZ 72512, an adult male, having some ocelli seven millimeters in diameter that are almost solid spots; UMMZ 89659 having postocular and postlabial stripes connected on right side of head; UMMZ 95615, 52948, 54402 having inner dark borders of pale stripes on snout represented by short dashes and dots (a ragged line connecting anterior margins of orbits on 54402); UMMZ 52948, 89659 having interrupted, black marginal lines on carapace with ends of some segments oriented inward and overlapping portion of adjacent segments; UMMZ 81699, female having plastral length of 19.0 centimeters, lacking conspicuous tubercles on anterior edge of carapace; UI 2403, CNHM 92204 having extensive dark mottling and marbling on throat and neck, undersurface of limbs and posterior portion of carapace.
Comparisons.—T. s. spinifer can be distinguished from all other subspecies of T. spinifer by the presence of large black ocelli (diameter 9-10 mm. on adult males, 2-3 mm. on hatchlings) in combination with only one dark marginal line. T. s. spinifer resembles asper in having ocelli or dots on the carapace but differs from asper in having only one dark marginal line and larger ocelli. T. s. spinifer differs from hartwegi only in the large size of the ocelli. T. s. spinifer resembles hartwegi and asper but differs from pallidus, guadalupensis and emoryi in having blackish spots and ocelli on the carapace and lacking whitish dots. T. s. spinifer resembles hartwegi, asper, and pallidus and differs from guadalupensis and emoryi in having conical or knoblike tubercles on the anterior edge of the carapace on large females.
T. s. spinifer differs from the subspecies asper, guadalupensis and emoryi in having a relatively narrower head, and from emoryi in having a relatively wider carapace. T. s. spinifer resembles hartwegi and asper but differs from the other subspecies in having the carapace widest at a plane approximately one-half way back on the carapace. The subspecies spinifer and hartwegi have longer snouts than pallidus, guadalupensis, and emoryi. T. s. spinifer differs from asper but resembles all the other subspecies in having a relatively longer plastron.
Remarks.—Lesueur's description of Trionyx spiniferus (1827:258-261, Pl. 6) seems to be based mostly, if not entirely, on a large female (length of carapace, 13 inches), which was "Le plus grand des individus observes …" [491] (op. cit.:258); an accompanying illustration depicting the dorsal surface of the bony carapace is unusual in lacking neurals (Pl. 6, E). Duméril and Bibron (1835:481) mentioned eight or nine additional specimens that Lesueur sent to the Museum of Natural History in Paris. Dr. Jean Guibé informed me under letter dated September 24, 1959, that a larger stuffed female, bearing catalog number 8808 is regarded as the holotype, and that there are seven additional specimens (1949, 4143, 8807, 8809-12) in the museum at Paris. All turtles were obtained by Lesueur from the Wabash River. To my knowledge no specimen that was available to Lesueur has been specifically designated as a type. Because the description seems to be based on one specimen, undoubtedly No. 8808, this specimen has been regarded as the holotype. However, Lesueur referred to several specimens and did not mention a type in the original description; consequently I prefer to regard No. 8808 as a lectotype.
Lesueur also described Trionyx ocellatus (op. cit.:261-263) as a variety of T. spiniferus having ocelli, or parts thereof, on the carapace and mentioned three specimens. The total number of specimens that were available to Lesueur is unknown. One young alcoholic specimen having ocelli is in the British Museum (Natural History) (Gray, 1855:69). The same letter from Dr. Guibé stated that a specimen in the Museum of Natural History, Paris, No. 6957, having a carapace 17 centimeters in length, conforms to the characters of ocellatus as mentioned by Lesueur, and was obtained from the Wabash River by Lesueur. Two of the specimens mentioned by Lesueur (loc. cit.) are stated to be females. No. 6957 is an adult male and clearly shows the juvenal pattern; it is regarded as the lectotype of T. ocellatus Lesueur, a name-combination, which is a synonym, based on a secondary sexual difference in pattern.
Rafinesque (1832:64) described a soft-shelled turtle from "the River Hudson between the falls of Hadley, Glen and Baker, and further up to the source" as Apalone hudsonica. The most outstanding characteristic was the presence of five claws on the digits of each limb. Rafinesque's recording of this characteristic was perhaps influenced by the illustration of a softshell in Bartram's Travels that showed each limb with five, clawed digits. Perhaps this was the basis for Boulenger (1889:245, footnote) regarding Apalone as "mythical." The large, yellowish, black-bordered spots, one behind and one in front of the eye presumably represent segments of the postocular stripe and the stripe on the snout; Rafinesque described the carapace as "entire … the margin is yellowish unspotted, then comes a circular black line …" and having "many round spots occulated and clouded by having a brown margin, with grey dots within." Except for five claws, the description is applicable to a softshell and referable to T. s. spinifer. To my knowledge, the only other records of the occurrence of soft-shelled turtles in the Hudson river drainage are those of Eights (in Bishop, 1923:120, Mohawk River at Cohoes), and DeKay (1842:7, Mohawk River and Hudson River near Albany); presumably these records are the basis for the comments of Holbrook (in Bishop, loc. cit.), and symbolized as an isolated locality by Conant (1958:318, map 35). The type locality of Apalone hudsonica is herein restricted to the Hudson River, near Baker's Falls, Saratoga County, New York.
Gray (1844:48) proposed the name Tyrse argus for a specimen reported to have come from Sierra Leone, West Africa; later (1855:68), he referred the species to the genus Trionyx. After comparison with a specimen of T. spiniferus Lesueur, Gray (1864:89) was "doubtful whether there must not have been some [492] confusion about the habitat of the specimen [which formed the basis of the description of Tyrse argus], and whether it is not more probably a North American species." The same author (1869:222; 1870:109) listed Tyrse argus as a synonym of Callinia spinifera (= Trionyx spiniferus Lesueur).
Agassiz (op. cit.:406-07) described Aspidonectes nuchalis on the basis of three adults from the Cumberland River and a number of young from the headwaters of the Tennessee River. Boulenger (1889:245, footnote 2) suggested that the status of A. nuchalis required further investigation. The species was not generally recognized after the turn of the century. Barbour and Loveridge (1929:226) listed MCZ 1908 (one of the juveniles) and 1623-25 as cotypes. Stejneger (1944:52) showed that nuchalis was not distinguishable from T. s. spinifer, and (op. cit.:49) listed MCZ 1623-25 as cotypes. Schmidt (1953:110) restricted the type locality to the Cumberland River, near Nashville, Tennessee.