Chapter III: Part 3

Bartram failed to use a binomial name with his description of "the great soft shelled tortoise," which appeared in his _Travels_ (1791:177-179, Pl. 4 and unnumbered plate between pages 282 and 283) and two editions of a French translation (1799 and 1801, 1:307); see Harper (1940). Recently, Bartram's _Travels_ has been placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature, Opinion 447 (see Hemming, 1957). Bartram's description of a soft-shelled turtle has provided the basis for the proposal of at least three name-combinations. The first was _Testudo_ (_ferox?_) _verrucosa_ proposed in 1795 by Schoepff; it appeared simultaneously in _The Historia Testudinum_ and in a German translation, _Naturgeschichte der Schildkröten_ (see Mittleman, 1944:245). Stejneger (1944:26) listed the type locality as eastern Florida. Daudin (1801:74), also referring to Bartram's description in his _Voyage_ (French translation), proposed the name _Testudo bartrami_; Harper (_op. cit._:717) restricted the type locality of _T. bartrami_ from "Halfway pond," east Florida, to southwestern Putnam County between Palatka and Gainesville, Florida. Rafinesque (1832:64-65), relying on the authenticity of the illustrations in Bartram's _Travels_ that depict a soft-shelled turtle having five claws on each of the hind feet, tubercles on the sides of the head and neck, and ten scales in the middle of the carapace (presumably inaccuracies or a composite on the part of the artist), referred to Bartram's description as a new genus, _Mesodeca bartrami_, a name which Boulenger (1889:245, footnote) referred to as "mythical." Geoffroy (1809a:18-19) considered Bartram's description the basis for the recognition of a second species of _Chelys_ (binomial nomenclature not employed), and Duméril and Bibron (_loc. cit._) suggested that the description was based partly on a "Chelyde Matamata." The descriptive comments of Bartram are not clearly applicable to _Testudo ferox_ Schneider; _Trionyx ferox_, however, is the only species of soft-shelled turtle known to occur in the region of Bartram's observations (east Florida), and the type locality was restricted to Putnam County, Florida, by Harper. The name-combinations, _Testudo___ (_ferox?_) _verrucosa_ Schoepff, _Testudo bartrami_ Daudin, and _Mesodeca bartrami_ Rafinesque are junior synonyms of _Testudo ferox_ Schneider.

Schweigger (1812:285) referred _ferox_ to the genus _Trionyx_ following the description of that genus by Geoffroy in 1809. _Testudo ferox_ was listed as a synonym by Geoffroy in the description of _Trionyx georgicus_ (1809a:17); Duméril and Bibron (1835:432) mentioned that the specific characters of _georgicus_ were taken from Pennant. The name _Trionyx georgianus_ presumably appears for this taxon in Geoffroy's earlier-published synopsis (1809:367). _T. georgicus_ was listed as occurring in rivers of Georgia and the Carolinas; the type locality was restricted by Schmidt (_op. cit._:109) to the Savannah River, Georgia. The two specific names _georgicus_ and _georgianus_ are regarded as substitute names and junior synonyms of _T. ferox_.

Geoffroy (1809a:14-15) also described _Trionyx carinatus_, a name-combination that hitherto has been considered a synonym of _Trionyx ferox_. There is no indication from the description that _carinatus_ is applicable to _ferox_. Most comments pertain to a description of the bony carapace and plastron, which Geoffroy depicts in Plate 4. It is a young specimen judging from the small and isolated preneural; the seventh pair of pleurals is unusual in being fused (no middorsal suture), and the neurals seem large in proportion to the size of the pleurals. The anterior border of the carapace is described as having tubercles. Geoffroy listed _Testudo membranacea_ and _Testudo rostrata_ as synonyms of _carinatus_. Fitzinger (1835:127) listed _T. membranacea_, _T. rostrata_ and _T. carinatus_ as synonyms of _Trionyx javanicus_ (= _T. cartilagineus_), which was also described by Geoffroy (_op. cit._:15). Duméril and Bibron (_op. cit._:478, 482) considered _carinatus_ to be the young of _spinifer_ (_ferox_ as synonym). Gray (1844:48), however, referred _T. membranacea_ and _T. rostrata_ to the synonymy of _T. javanicus_, but considered _T. carinatus_ to be a synonym of _T. ferox_ (_op. cit._:50), an interpretation followed by all subsequent authors. _Trionyx carinatus_ is questionably listed as a synonym of _ferox_ by Stejneger (1944:27). Duméril and Bibron (_op. cit._:482) wrote that the young type of _carinatus_ is in the museum at Paris. Dr. Jean Guibé informs me in letter of September 24, 1959, that the type of Geoffroy's _T. carinatus_ cannot be found in the Natural History Museum at Paris. For the present, _T. carinatus_ is considered a _nomen dubium_. According to Stejneger (1944:27), _Trionyx brongniarti_ Schweigger is a substitute name for _T. carinatus_.

I am unable to add anything to Stejneger's (_op. cit._:32) account of _Trionyx harlani_; the mention of its occurrence in east Florida indicates that it is indistinguishable from _Testudo ferox_ Schneider.

_T. ferox_ was considered to be indistinguishable from Lesueur's _Trionyx spiniferus_ (described in 1827), until Agassiz (1857:401) pointed out the differences between the two species. However, Agassiz (_op. cit._:402, Pl. 6, Fig. 3) regarded juveniles of _T. spinifer asper_ as the young of _ferox_. Consequently, the geographic range of _ferox_, as envisioned by Agassiz, extended from Georgia and Florida west to Louisiana. Neill (1951:15) considered all American forms conspecific. Crenshaw and Hopkins (1955) and Schwartz (1956) demonstrated that _ferox_ is a distinct species.

Fitzinger (1843:30) designated the species _ferox_ as the type species of his genus _Platypeltis_ as follows: "Platypeltis. Fitz. Am[erica]. _Platypelt. ferox_. Fitz. Typus." If populations of soft-shelled turtles that are referable to _Testudo ferox_ Schneider are considered to comprise a distinct genus by future workers, _Platypeltis_ Fitzinger, 1835, is available as a generic name with _Testudo ferox_ Schneider, 1783, as the type species (by subsequent designation).

_Trionyx ferox_ in the northern part of its range is sympatric with _T. spinifer asper_. In the region of overlap, the two species are nearly always ecologically isolated; _ferox_ inhabits lentic waters, whereas _T. s. asper_ is partial to lotic waters (Crenshaw and Hopkins, _op. cit._:16). There is no evidence of intergradation or hybridization.

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_.

Carr (1940:107) recorded _ferox_ from Okaloosa County, Florida, in the western end of the panhandle, whereas Crenshaw and Hopkins (1955:16) list the known westward extent of range as Leon and Wakulla counties. AMNH 6933 from west of the Apalachicola drainage in Washington County, Florida, tends to substantiate Carr's record, which is not included on the distribution map.

_Specimens examined._--Total 144, as follows: FLORIDA: _Alachua_: UMMZ 64178, 100969; USNM 10545, 10704, "near" Gainesville; UMMZ 56599, Levy Lake. _Brevard_: AMNH 12878, Canaveral. _Broward_: UMMZ 109441, Hugh Taylor Birch State Park; USNM 109548, 22 mi. WNW, 6 mi. SSE Fort Lauderdale. _Collier_: USNM 86828, Tamiami Trail, "near" Birdon. _Dade_: AMNH 50936, UMMZ 10183, 110981, Miami; USNM 84079, 86942, 15 mi. from (west) Miami, Tamiami Trail; UMMZ 111371, 19 mi. W, 1.3 mi. S Miami; UI 28984, 35 mi. W. (Miami) Tamiami Trail; AMNH 69932-33, UMMZ 101582, 101584, 104024, 40-45 mi. W Miami, Tamiami Trail. _Glades_: UMMZ 100836, mouth of Kissimmee River. _Hendry_: UMMZ 106302, 10.2 mi. SE Devil's Garden; UMMZ 106303-04, 106321-22, 30 mi. S Clewiston, near Devil's Garden. _Hernando_: TU 13624, 0.5 mi. S Citrus Co. line on US Hwy. 19. _Highland_: AMNH 65537, 71618, Archbold Biol. Stat., Lake Placid; AMNH 65622, Hicoria. _Hillsborough_: TU 13960, Hillsborough River, _ca._ 20 mi. NE Tampa; USNM 51184, Tampa; USNM 71156, Plant City. _Indian River_: USNM 55316, Vero Beach; USNM 59318, Sebastian. _Lake_: UMMZ 36072, USNM 20189, 029210, 029339, 38123, Eustis; UMMZ 76754-56, Lake Griffin. _Lee_: UMMZ 102276, 14 mi. SE Punta Gorda. _Leon_: CNHM 33701, USNM 95767, Lake Iamonia; USNM 103736, Silver Lake. _Marion_: AMNH 8294-95, UMMZ 95613 (4), USNM 52476-83, 100902-04, Eureka; AMNH 63642, near Salt Springs. _Martin_: TNHC 1292, 8.4 mi. N Port Mayaca. _Okeechobee_: AMNH 57379-84, Lake Okeechobee; AMNH 5931-32, Kissimmee Prairie. _Orange_: USNM 51421, 56805, Orlando; KU 16528. _Osceola_: USNM 029448, 029450-64, 029467-68, 029470, 029474-75, Kissimmee. _Palm Beach_: UMMZ 54101, Palm Beach; USNM 73199, Delray Beach. _Pinellas_: USNM 51417-20, St. Petersburg. _Polk_: AMNH 25543, Lakeland; UMMZ 112380, 6.7 mi. S Lake Wales; USNM 60496, 60532, 60534, 61083-87, Auburndale. _Putnam_: USNM 4373, 7651, Palatka; USNM 26035, ponds "near" Welaka. _Sarasota_: USNM 61352, Lake Myakka. _Sumpter_: UMMZ 71791, Bushnell. _Volusia_: UMMZ 100673, Lake Helen. _Washington_: AMNH 6933, Washington. _County unknown_: AMNH 4758; USNM 8899, St. John's River: USNM 59727-28, Lake Okeechobee, "near" mouth Taylor's Creek; USNM 84080.

GEORGIA: _Baker_: SM 2083, USNM 029619, 38980-81, 70398, Mimsville. _Berrien_: USNM 62217, Banks Mill Pond. _Charlton_: AMNH 69934, Okefinokee Swamp, SW Billy's Island; UMMZ 90010, east edge Okefinokee Swamp; USNM 84603, Okefinokee Swamp, Chesser's Island. _Irwin_: USNM 56804. _Lowndes_: UMMZ 67706, 10 mi. S Valdosta. _McIntosh_: USNM 19621, Darien.

SOUTH CAROLINA: _Charleston_: USNM 9670, Charleston.

NO DATA: AMNH 22750; USNM 71608-09.

_Records in the literature._--FLORIDA: _Alachua_: 10 mi. ENE Gainesville (Schwartz, 1956:18). _Brevard_: Merritt Island (Neill, 1958:6). _Broward_: Fort Lauderdale (Schwartz, _op. cit._:19). _Charlotte_: (Carr, 1940:107). _Clay_: Green Cove Springs (Brimley, 1910:18); St. John's River (Crenshaw and Hopkins, 1955:21); Doctor's Inlet (Schwartz, _op. cit._:18). _Collier_: Royal Palm Hammock (Crenshaw and Hopkins, _op. cit._:20); 11.2 mi. E Monroe Station (Schwartz, _op. cit._: 19). _Columbia_: (Carr, _loc. cit._). _Dade_: Paradise Key (Schwartz, _loc. cit._); Homestead (eggs, Stejneger, 1944:43). _Duval_: 4-10 mi. S Jacksonville (Deckert, 1918:31). _Glades_: _ca._ 8 mi. SW Okeechobee State Park. _Lake_: Alexander Springs (Schwartz, _op. cit._:18). _Lee_: 18 mi. S Fort Myers (Conant, 1930:63); 6 mi. SE Fort Myers (Hamilton, 1947:209). _Levy_: Gulf Hammock (Schwartz, _loc. cit._); Brownson (Stejneger, _op. cit._:42). _Monroe_ and _Okaloosa_ (Carr, _loc. cit._). _Okeechobee_: 6 mi. E Kissimmee River; state hwy. 78 "near" Okeechobee-Glades co. line. _Palm Beach_: SW part of Lake Okeechobee, near Clewiston; Milton Island Cove (Schwartz, _loc. cit._). _Pasco_: mouth Pithlachascotee River (Neill, _op. cit._:26). _Pinellas_: Belleair (Brimley, _loc. cit._); Seminole (Conant, _loc. cit._); 5 mi. E Clearwater (Schwartz, _op. cit._:19); Gulf Port (Stejneger, _op. cit._:43). _Polk_: Lake Shipp, near Winter Haven (Telford, 1952:185). _Sarasota_: 15 mi. E Sarasota (Conant, _loc. cit._); Venice (Conant, _op. cit._:61). _Taylor_: "near" Foley. _Wakulla_: "near" Crawfordville (Crenshaw and Hopkins, _op. cit._:15).

GEORGIA: _Baker_: 5 mi. NW Newton, 5 mi. W Newton, 4 mi. N Newton. _Ben Hill_: 6 mi. E Fitzgerald (Crenshaw and Hopkins, 1955:15). _Bulloch_: 14 mi. SE Statesboro (Schwartz, 1956:19). _Decatur_: "near" Bainbridge (Crenshaw and Hopkins, _loc. cit._). _Emanuel_: "near" Midville. _Evans_: 8 mi. NE Manassas, Tattnall County. _Ware_: Laura Walker State Park (Schwartz, _loc. cit._). _Wilcox_: 3 mi. SE Forest Glen (Crenshaw and Hopkins, _op. cit._:19).

SOUTH CAROLINA: _Beaufort_: 7 mi. NE Gardens Corner (Schwartz, 1956:19). _Chatham_: Savannah River at Savannah (Schwartz, _op. cit._:8-9). _Colleton_: 5 mi. from Whitehall, Combahee River (Schwartz, _op. cit._:19).

=Trionyx spinifer= Lesueur

Spiny Softshell

_Range._--In Canada, southern Ontario and Quebec; in the United States, northwestern Vermont and western New York south to northern Florida, east to central Montana, eastern Wyoming and Colorado, and New Mexico; introduced into the Colorado River system of California, Nevada, Arizona and New Mexico; in México, the northern part of the states of Tamaulipas, Nuevo León, Coahuila, and eastern Chihuahua (see map, Fig. 19).

_Diagnosis._--Juvenal pattern uniform tan or brownish lacking markings, having whitish dots or spots, or having well-defined, blackish ocelli or spots; surface of carapace "sandpapery" in adult males; conical projections (in some subspecies) along anterior edge of carapace in large females; contrasting pattern of blackish marks on pale background (in some subspecies) on dorsal surface of limbs of adult males.

Opisthotic-exoccipital spur well-developed; epiplastral callosity, when present, not covering entire surface.

_Description._--Septal ridges present; external and proportional characteristics variable (see accounts of subspecies); range in length of plastron (cm.) of ten largest specimens of each sex (mean follows extremes), males, 13.8-16.0, 14.4; females, 26.0-31.0, 28.0.

Guide to subspecies:

1. _T. s. spinifer_
2. _T. s. hartwegi_
3. _T. s. asper_
4. _T. s. pallidus_
5. _T. s. guadalupensis_
6. _T. s. emoryi_
]

Greatest width of skull usually at level of squamosal (74%); foramen magnum rhomboidal; ventral surface of supraoccipital spine narrow proximally, usually having medial ridge; opisthotic-exoccipital spur well-developed (66%); distal part of opisthotic wing tapered, not visible in dorsal view; lateral condyle of articular surface of quadrate larger than medial articular surface, not tapered posteriorly; maxillaries in contact above premaxillaries (88%); usually a combination of seven neurals, seven pairs of pleurals and contact of seventh pair of pleurals (83%); angle of epiplastron approximately 90 degrees; callosities when present on epiplastron not covering entire surface; hyo-hypoplastral suture usually present.

_Comparisons._--_Trionyx spinifer_ can be distinguished from _T. ferox_ and _T. muticus_ by the presence of any one of the characters mentioned in the "Diagnosis." Both sexes and all sizes of _T. spinifer_ resemble _ferox_ but differ from _muticus_ in having septal ridges. Most individuals of _T. spinifer_ (except some large females) resemble _muticus_ but differ from _ferox_ and large females of _ater_ in having a pale outer rim that is separated from the ground color of the carapace by a distinct (_spinifer_) or dusky (_muticus_) dark line. Large females of the subspecies _spinifer_, _hartwegi_, _asper_ and _pallidus_ may have enlarged conical projections along the anterior edge of the carapace and, unless these projections are considerably worn, are readily distinguished from large females of _ferox_ (flattened, knoblike prominences), and _muticus_ and _ater_ (smooth surface, no prominences). Large females of the subspecies _guadalupensis_ and _emoryi_ resemble _muticus_ and _ater_, and to some extent _ferox_, in having low, scarcely elevated prominences along the anterior edge of the carapace. Some females of _emoryi_ resemble _ferox_ in that the plastron extends farther forward than the carapace.

_T. spinifer_ is intermediate in size between _ferox_ (larger) and _muticus_ (smaller); the maximum size of the plastron in adult males is approximately 16.0 centimeters (14.0, _muticus_; 26.0, _ferox_), and of females, 31.0 centimeters (21.5, _muticus_; 32.5, _ferox_). The head for all subspecies of _spinifer_ is proportionately narrower than in _ferox_ but wider than in _muticus_.

In the skull, _spinifer_ more closely resembles _ferox_ than _muticus_, but differs from both _ferox_ and _muticus_ in usually having a well-developed opisthotic-exoccipital spur. Skulls of _spinifer_ resemble those of _muticus_ but differ from those of _ferox_ in being widest at the level of the squamosal. Skulls of _spinifer_ resemble those of _ferox_ but differ from those of _muticus_ in having the 1) ventral surface of the supraoccipital spine narrow proximally, and usually having a medial ridge, 2) foramen magnum rhomboidal, 3) distal part of opisthotic wing tapered, 4) lateral condyle of articular surface of quadrate not tapered posteriorly, and larger than medial articular surface, and 5) maxillaries in contact above premaxillaries. _T. spinifer_ resembles _ferox_ but differs from _muticus_ in having the epiplastron bent at an approximate right angle. _T. spinifer_ differs from _ferox_ in having an epiplastral callosity, and from _muticus_ in that the callosity does not cover the entire surface of the epiplastron. The hyo-hypoplastral suture is present more often in _spinifer_ and _muticus_ than in _ferox_.

_Remarks._--Gray (1869:221) proposed the generic name _Callinia_ as a new name for _Aspidonectes_ as understood by Agassiz (1857:403). Gray referred _Trionyx spiciferus_ (= _spiniferus_) Lesueur to _Callinia_. Stejneger (1907:514) designated _Trionyx spiniferus_ Lesueur as the type species of _Callinia_. If _Trionyx spiniferus_ Lesueur is considered to be generically distinct from other soft-shelled turtles, _Callinia_ Gray, 1869, is available as a generic name with _Trionyx spiniferus_ Lesueur, 1827, as the type species by subsequent designation.

_Geographic variation._--_T. spinifer_ is the most variable and widespread species of the genus in North America. Size of ocelli on the carapace decreases from east to west on turtles inhabiting waterways of the Upper Mississippi River drainage. The most impressive gradient, geographically oriented from western Louisiana to southwestern Texas is seen in each of several features: decrease in size of tubercles on the anterior edge of the carapace, reduction in contrast of pattern on the dorsal surface of limbs and side of head, change in pattern on the dorsal surface of the snout, and increase in the size of white spots on the carapace. But the gradient in size of white spots is reversed in _T. s. emoryi_, which has small white spots on the carapace. Some of the characters at the western terminus of this geographical gradient are shared with _T. ater_ and _muticus_. Those subspecies comprising the _emoryi_ group also show proportional characters that correspond closely with those of _T. ferox_.

On the basis of tuberculation and pattern on carapace, side of head, dorsal surface of limbs and snout, _Trionyx spinifer_ may be divided into six subspecies.

=Trionyx spinifer spinifer= Lesueur

Eastern Spiny Softshell

Plates 33, 34, and 52

_Trionyx spiniferus_ Lesueur, Mém. Mus. Hist. Nat. Paris, 15:258,
pl. 6, December, 1827.

_T[rionyx] s[pinifer] spinifer_ Schwartz, Charleston Mus. Leaflet,
No. 26:11, May, 1956.

_Trionyx ocellatus_ Lesueur, Mém. Mus. Hist. Nat. Paris, 15:261,
December, 1827.

_Apalone hudsonica_ Rafinesque, Atlan. Jour., Friend Knowledge,
Philadelphia, 1(No. 2, Art. 12):64, Summer, 1832.

_Trionyx annulifer_ Wied-Neuwied, Riese Nord-Amerika, 1(pt. 3):140,
1838.

_Tyrse argus_ Gray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 48,
1844.

_Aspidonectes nuchalis_ Agassiz, Contr. Nat. Hist. United States,
1(pt. 2):406, 1857.

_?G[ymnopus] olivaceus_ Wied-Neuwied, Nova Acta Acad.
Leopold.-Carol., 32:55, pl. 5, 1865.

_Type._--Lectotype, Museum d'Histoire Naturelle, Paris, No. 8808; large stuffed female obtained by C. A. Lesueur from the Wabash River, New Harmony, Posey County, Indiana (Pl. 52).

_Range._--Northeastern United States and extreme southeastern Canada in tributaries flowing into the Mississippi River from the east, and the St. Lawrence River drainage; extreme southern Quebec and Ontario, Canada, east through southern Great Lakes region to Wisconsin, and south through New York, western Pennsylvania and Illinois to Tennessee and western Virginia (see map, Fig. 19).

_Diagnosis._--Juvenal pattern of large, thick-bordered black ocelli, often 9-10 millimeters in diameter in center of carapace on adult males, and 2-3 millimeters in diameter on hatchlings (mean OD/PL, Michigan, .066); only one dark marginal line separating pale rim of carapace from dorsal ground color.

_Description._--Plastral length of smallest hatchling, 2.7 centimeters (UMMZ 89950, INHS 3143); of largest male, 14.5 centimeters (UMMZ 72512); of largest female, 31.0 centimeters (UMMZ 40866).

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 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 ..." (_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 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.

Agassiz (_loc. cit._) mentioned that _nuchalis_ "differs strikingly from Asp. spinifer in the much more elongated form of the male, and in the great development of the marginal spines and of the tubercles upon the carapace, ... But the most prominent specific character consists in the marked depressions on either side of the blunt median keel, and also in the triangular dilation of that keel behind the front margin of the carapace." These characters seem to be of no taxonomic worth. I have seen three syntypes (MCZ 1623-25) that undoubtedly correspond to the three adult specimens mentioned by Agassiz. All are females, measuring 19.5, 22.0, and 19.0 centimeters, respectively, in plastral length, and lack a contrasting mottled pattern on the carapace; the juvenal pattern is obscured, except for blackish spots at the edge of the carapace on MCZ 1625, and parts of an ocellus on MCZ 1624. The dorsal surfaces of the limbs are boldly marked. MCZ 1623, showing the diagnostic feature mentioned by Agassiz, is photographed by Stejneger (_op. cit._:Pls. 14, 15), and may be regarded as the lectotype of _Aspidonectes nuchalis_ Agassiz. MCZ 1908 is one of the young syntypes mentioned by Agassiz, and is referable to _spinifer_. The juvenal pattern consists of spots and ocelli; the plastron measures 3.1 centimeters in length, and the carapace 4.2 centimeters.

Wied-Neuwied (1865:55-57, Pl. 5) described the species _?G_[_ymnopus_] _olivaceus_, but was uncertain whether his interpretation was based on a species, a variety or a secondary sexual difference. Wied-Neuwied mentioned that Lesueur had already named this soft-shelled turtle as _Trionyx ocellatus_, and agreed with Lesueur that those turtles having occulated spots on the carapace were distinguishable from _T. spiniferus_ and _T. muticus_. But because Duméril and Bibron in their _Erpétologie Général_ failed to recognize _T. ocellatus_, Wied-Neuwied felt obliged to bring it to the attention of his American colleagues and he renamed it. Wied-Neuwied also stated, in the context of a synonym, "Beschreibung einer Reise in Nord-America Bd. I., pag. 140." This comment presumably refers to his earlier description of _T. annulifer_ (1838:140); seemingly Wied-Neuwied considered _T. annulifer_ and _G. olivacea_ as conspecific, although there is no mention of _annulifer_ in the text proper. Stejneger (_op. cit._:49) designated the type locality of _T. annulifer_ as the Ohio River at Pittsburgh, Pennsylvania, and of _Gymnopus olivacea_ as New Harmony, Wabash River, Illinois (_lapsus_ for Indiana).

_Trionyx spiniferus_ was questionably considered distinct from _T. ferox_ by Lesueur who listed "Testudo ferox Gm. Tortue de Pennant?" and "Trionyx georgicus Geoffr.?" as synonyms. Subsequently, most authors considered _T. spiniferus_ synonymous with _T. ferox_ until Agassiz (1857) pointed out differences between the two species.

The average size of the ocelli on the carapace of the subspecies _spinifer_ decreases westward toward the Mississippi River; ocelli of different sizes occur on different individuals from the same state and presumably from the same population. For example, INHS 2281, plastron 9.9 centimeters in length, from Effingham County, Illinois, has some ocelli eight millimeters in diameter, whereas a larger male from the same locality, UI 1322, plastron 11.6 centimeters in length, has the largest ocelli only five millimeters in diameter. 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. Intergradation with _asper_ possibly occurs in that part of the Tennessee River in eastern Tennessee as exemplified by UMMZ 59198.

Published reports indicate that _T. s. spinifer_ is not abundant in some of the northeasterly parts of its geographic range. Adams and Clark (1958:10) wrote that few softshells at Long Point on the Canadian side of Lake Erie are "ever collected and the area's game keepers report ... (none) ... seen in recent years. They also tell of recurrent severe stormy winters in which the muddy bottom of the marshland was repeatedly churned up and frozen. Such climatic conditions could easily destroy a large part of the _Trionyx_ population overwintering in the mud bottom." Wright (1919:8) reported that softshells are "rarely seen" in bays on the New York side of Lake Ontario, and Babcock (1938:53) wrote that _spinifer_ "is not common in Lake Champlain."

_T. s. spinifer_ probably extended its geographic range into the Hudson River drainage of New York _via_ the Erie Canal (connected Buffalo and Albany) after its completion in the early 1800's (DeKay, 1842:7). Now, the New York Barge Canal (essentially the Erie Canal, but with minor changes in course and the addition of several spurs) provides an avenue for dispersal of _spinifer_ to the Hudson River drainage, Lake Ontario and intervening waterways in New York (Mertens, 1928:199). Netting (1944:86-87), however, suggested that _spinifer_ occupied Lake Champlain, the Finger Lakes, Mohawk River and upper Hudson in the late stages of the formation of the Great Lakes.

A publication not seen by me is that of Mansueti and Wallace (1960). Its title suggests that _Trionyx_ occurs in Maryland.

The unsuccessful introduction of _T. s. spinifer_ in the Delaware drainage in New Jersey has been discussed by Fowler (1907:213), who wrote that they were found as early as the late 1860's and were introduced when young presumably to stock aquaria. Records of occurrence include Cooper's Creek, Camden County (Stone, 1906:168); Woodbury, Gloucester County (Cope, 1894:889); and Paulins Kill at Hainesburg, Warren County (Johnson, 1894:889).

Surface (1908:122) believed that soft-shelled turtles "have doubtless been introduced into the eastern part of Pennsylvania through the canal from the Western and Central part of New York," and Roddy (_in_ Neill, 1951:21) suggested that the species may be found in the Susquehanna River. Babcock (1919:420) mentioned a young specimen of _spinifer_ in the collection of the Boston Society of Natural History that was obtained "in White River, Vermont," a tributary of the Connecticut River of the Atlantic Coast drainage; seemingly this record has not been accepted and the species is not established. To my knowledge, populations of _T. s. spinifer_ do not occur in rivers of the Atlantic Coast drainage, except probably the Hudson-Mohawk drainage.

Stockwell (1878:401) wrote that _spinifer_ was found "as high as Athabasca." Presumably Stockwell referred to Lake Athabaska in northern Alberta and Saskatchewan, Canada, a region where soft-shelled turtles are unknown; see also the comments by Stejneger (1944:52).

_Specimens examined._--Total 250 as follows: ALABAMA: _Morgan_: UMMZ 99578, "near" Decatur.

ILLINOIS: _Adams_: INHS 2150, Quincy. _Bond_: INHS 8345, Greenville. _Carroll_: CNHM 42116, Ordnance School Proving Ground. _Cass_: INHS 2151, Beardstown. _Champaign_: INHS 2273, 2311, 2413, 3142, "near" Seymour; INHS 4229, Champaign; INHS 6163, Sidney. _Christian_: INHS 1560, Pana. _Coles_: INHS 1968-69, 2 mi. W Charleston. _Cumberland_: INHS 2282, Greenup. _De Witt_: INHS 7674, Farmer City. _Effingham_: UI 1322, 2281, 19365, "near" Effingham. _Fulton_: INHS 5531, 2 mi. NE Bluff City, Schyler County; UI 23449, Liverpool; UI 24611, Spoon River, 18 mi. NW Canton. _Hancock_: USNM 53522, 59277, "near" Hamilton. _Iroquois_: INHS 6869-70, 2.5 mi. N Crescent City. _Jackson_: TU 1369 (12), Elkville. _Kane_: CNHM 42400, Aurora. _Kankakee_: CNHM 324, Momence. _Kendall_: UI 2411, Plano. _Logan_: INHS 7171-72, 6 mi. N Lincoln. _Madison_: USNM 60571. _Macoupin_: UI 2401-02, Beaver Dam Lake. _Mason_: CNHM 346, 470, INHS 1122, 1559, 5756-58, UI 42, 2404, Havana, Lake Chautauqua. _Mercer_: CNHM 3220, New Boston. _Morgan_: CNHM 2067 (2), 3290, 3303-04, 3306, INHS 2152, 2154, 5132-37, USNM 54747, Meredosia. _Moultrie_: INHS 8989, 2 mi. NW Lovington. _Peoria_: UI 2406-10, Peoria. _Pope_: INHS 5505, Lake Glendale. _Putnam_: UMMZ 81604-14, 5 mi. N Henry, Marshall County. _Schuyler_: UI 2405, "near" Ripley, Brown County. _Scott_: INHS 2149, 2153, Naples. _Union_: CNHM 18623, 6 mi. SW Jonesboro. _Vermilion_: INHS 3142, Muncie; INHS (1 untagged); UI 1970, 3209, Danville; UI 2403, 1.5 mi. E Oakwood; UI 16265, Kickapoo State Park. _Wabash_: USNM 12061, Mt. Carmel. _Winnebago_: INHS 7185, Kishwaukee Forest Preserve; INHS 7294, 1/2 mi. S Shirland. _County unknown_: USNM 7661.

INDIANA: _Bartholomew_: UMMZ 61060, 10 mi. W Columbus. _Carroll_: USNM 42905-06, Burlington. _Clark_: UMMZ 110599, 14-mile Creek, 3 mi. NW Charleston. _Decatur_: UMMZ 55416, 3 mi. S Westport. _Elkhart_: UMMZ 105598, Elkhart River, south of Goshen. _Gibson_: UMMZ 89744, Foot's Pond. _Johnson_: UMMZ 108062, 2 mi. S Trafalgar. _Knox_: USNM 22711, Vincennes. _Kosciusko_: AMNH 8379, UMMZ 84287 (5), Winona Lake; UMMZ 110235, Wawasee Lake. _Lake_: CNHM 11019, 11021-24, Crown Point. _Marion_: UMMZ 103393, Ravenswood; UMMZ 110236, 1 mi. N Lawrence. _Marshall_: CNHM 39299; USNM 33495, Yellow River north of Burr Oak; USNM 33496-501, 35404, 42583-84, Lake Maxinkuckee. _Wells_: UMMZ 63523, Wabash River, Bluffton. _County unknown_ (Lagrange or Marshall): USNM 50670, Twin Lakes.

KENTUCKY: _Casey_: UMMZ 112252, trib. of Green River, south of Yosemite. _Green_: UMMZ 116718, Little Barren River, 1.5 mi. E Monroe, Hart County. _Rockcastle_: UMMZ 98767, Rockcastle River, 5 mi. above Livingston.

MICHIGAN: _Allegan_: UMMZ 42112, Kalamazoo River. _Barry_: UMMZ 53874, Thornapple River, 3 mi. NW Hastings. _Bay_: UMMZ 74670. _Branch_: UMMZ 95615, 1 mi. S Kinderhook; UMMZ 70748, Hog Creek. _Calhoun_: UMMZ 89950 (3); UMMZ 79133, near Battle Creek. _Cass_: UMMZ 40866-67, 53005, Diamond Lake; UMMZ 40868, 52948, Long Lake. _Jackson_: UMMZ 72494. _Kalamazoo_: UMMZ 42130, 80534, Kalamazoo; UMMZ 90506, Gull Lake; UMMZ 92599, Kellogg Bird Sanctuary. _Lenawee_: UMMZ 72457, Devil's Lake; UMMZ 74662, Wolf Lake Park. _Livingston_: UMMZ 54401, 76190, Portage Lake. _Monroe_: UMMZ 44604-06, USNM 51213, "near" Monroe. _Newaygo_: UMMZ 63469. _Oakland_: UMMZ 64363, Hay's Creek; UMMZ 96539, Clinton River. _Ottawa_: UMMZ 81699. _St. Joseph_: UMMZ 38876, 38889, "near" White Pigeon; UMMZ 96537, Corey Lake. _Van Buren_: UMMZ 90003, Wolf River, west of Kalamazoo, Kalamazoo County. _Washtenaw_: SM 2035, 2038, 2105, UMMZ 39847, 96538, "near" Ann Arbor; UMMZ 35765, 35769, 74518 (2), Portage Lake; UMMZ 54402-03, Little Lake; UMMZ 89659, Huron River, Dexter; UMMZ 110583-85. _County unknown_ (Washtenaw or Livingston): UMMZ 54400, Huron River near Portage Lake.

MISSISSIPPI: _Adams_: MCZ 46615, UMMZ 76446, "near" Natchez; MCZ 46621, 46633, USNM 01084, 01086, Washington. _Coahoma_: AMNH 5289, 5285-86, Moon Lake. _Lafayette_: MCZ 37173, Oxford; USNM 7650, Abbeville? (reported from Abbeville, South Carolina by Pickens, 1927:113; see discussion by Stejneger, 1944:50, and my comments on page 509 beyond). _LeFlore_: USNM 73668-69, Greenwood. _Madison_: USNM 95192, Big Black River. _Washington_: USNM 115980, Deer Creek. _Yazoo_: UMMZ 86669, Panther Creek west of Yazoo City; UMMZ 83304, Yazoo City.

NEW YORK: _Monroe_: CNHM 92001-02, Genesee River, Rochester. _Wayne_: AMNH 69931, CNHM 92004, Sodus Bay.

OHIO: _Athens_: UMMZ 111793, east branch Shade Creek. _Franklin_: USNM 26290. _Lucas_: USNM 51214, Toledo. _Pike_: UMMZ 99309, Morgan's Fork, Sunfish Creek. _Warren_: AMNH 4763, Little Miami River, 3 mi. below Morrow. _County unknown_: USNM 21128-29, Cuyahoga River.

TENNESSEE: _Benton_: UMMZ 113036, Eagle Creek, 1/2 mi. E Holliday. _Bradley_: UMMZ 59197, west branch of Chestnee Creek, 7 mi. E Cleveland. _Claiborne_: USNM 86677, 5 mi. SE Cumberland Gap, Powell River. _Davidson_: MCZ 1623-25, Cumberland River near Nashville (restricted locality); USNM 7165-67, Nashville. _Decatur_: KU 3000, Perryville. _Hamilton_: USNM 131861, Chattanooga. _Monroe_: TU 16058, Little Tennessee River, 10 mi. N Madisonville. _Obion_: UMMZ 53199, USNM 102911, Reelfoot Lake. _Overton_: UMMZ 69561 (2), Wirmingham. _Sevier_: TU 16132, UMMZ 86735, USNM 86681-82, near Sevierville; UMMZ 86734, Walden Creek "near" Gatlinburg. _County unknown_: MCZ 1908, headwaters of Tennessee River.

VIRGINIA: _Smythe_: USNM 101386, Holston River, Seven Mile Ford.

WEST VIRGINIA: _McDowell_: USNM 33767, Dry Fork, Perryville (county questionable, perhaps Randolph County).

WISCONSIN: _Chippewa_: CNHM 8223, Lake Wissota, mouth of Yellow River, Anson Twp. _Polk_: UMMZ 72511-12, St. Croix River "near" Never's Dam. _County unknown_: CNHM 15971, Eau Claire River.

_Records in the literature._--ONTARIO: _Carleton_: Ottawa (questionable record). _Essex_: Point Pelee. _Haldimand_: Dunville. _Kent_: Lake St. Clair. _Norfolk_: Long Point. _Oxford_: Beachville. _Wentworth_: Hamilton Bay (Logier and Toner, 1955:51).

QUEBEC: _Iberville_: Richelieu River at Iberville (Logier and Toner, 1955:51).

ALABAMA: _Lawrence_: Courtland (Stejneger, 1944:53).

ILLINOIS: _Boone_: Belvidere. _Bureau_: Bureau. _Cass_: Chandlerville. _Clay_: Louisville (Cahn, 1937:189). _Cook_: Lake Michigan (Kennicott _in_ Stejneger, 1944:44); Evanston (Necker, 1939:10); Chicago (Schmidt and Necker, 1935:76). _Crawford_: Robinson. _Douglas_: northern part of county (P. W. Smith, 1947:39). _Fayette_: Vandalia. _Fulton_: Ellisville (Cahn, _loc. cit._). _Grundy_: Morris (Stille and Edgren, 1948:201). _Jackson_: Jacob (Cagle, 1942:158). _Jersey_: Grafton (Cahn, _loc. cit._). _Kane_: Batavia; Dundee Game Farm (Stille and Edgren, _loc. cit._). _Kankakee_: Kankakee River near Altort (Necker, _loc. cit._). _Lake_: Fox Lake. _LaSalle_: Streator (Cahn, _loc. cit._). _Lawrence_: (Hahn _in_ Stejneger, 1944:44). _Lee_: symbol on map (Cahn, _loc. cit._). _McHenry_: McHenry (Stille and Edgren, _loc. cit._). _Macon_: Decatur. _Macoupin_: Carlinville (Cahn, _loc. cit._). _Ogle_: Oregon (Garman _in_ Cahn, _loc. cit._). _Randolph_: Chester, Reily Lake. _Rock Island_: Barstow, Hillsdale, Rock Island (Cahn, _loc. cit._). _Saline_: Horseshoe Lake (Stein, 1954:312). _Stephenson_: Freeport (Cahn, _loc. cit._). _Union_: Bluff Lake (Garman _in_ Cahn, _loc. cit._). _Whiteside_: Sterling, symbol on map (Cahn, _loc. cit._). _Williamson_: Marion (Cagle, 1942:158). _Winnebago_: Rockton; symbol in western part of county (Cahn, _loc. cit._). _County unknown_: Fox River (Yarrow, 1882:29).

INDIANA: _Brown_: 1 mi. below Helmsburg (Myers, 1927:339). _Clay_: Eel River (Kirsch _in_ Stejneger, 1944:45). _Franklin_: (Hughes _in_ Stejneger, _loc. cit._). _Jasper_: Jasper-Pulaski Game Preserve (Swanson, 1939:690). _Jefferson_: Madison (Myers, _loc. cit._). _Marion_: Irvington (Stejneger, _op. cit._:55). _Marshall_: 2 mi. NW Culver (KKA). _Monroe_: Bloomington (McLain _in_ Stejneger, _op. cit._:45). _Newton_: Lake Village (Stille and Edgren, _loc. cit._). _Posey_: Wabash River at New Harmony (Lesueur, 1827:257). _Starke_: Grant (Stille and Edgren, _loc. cit._). _Steuben_: Fish Creek "near" Hamilton (Stejneger, _op. cit._:53). _County unknown_ (Knox or Starke): USNM 72387, Knox (Stejneger, _op. cit._:55); "White Water valley," east-central part of state (Butler, 1894:224). USNM 8359 (= _Trionyx spinifer asper_) has been erroneously recorded from Madison, Indiana, by Yarrow (1882:29) and Hay (1892:145); see discussion by Cahn (1937:200) and Stejneger, (_op. cit._:73, 75).

KENTUCKY: _Edmonson_: Green River, Mammoth Cave National Park (Hibbard, 1936:281). _Fleming_: Fox Creek (Welter and Carr, 1939:130). _Jefferson_: (Funkhouser, 1925:71). _Morgan_: (Stejneger, 1944:54). _County unknown_: Ohio and Pond rivers (Funkhouser, _loc. cit._).

MICHIGAN: _Berrien_: mouth of St. Joseph River at St. Joseph (Lagler, 1943:303). _Eaton_: Brookfield; Olivet (Clark _in_ Ruthven, Thompson and Thompson, 1912:133). _Genesee_: (Miles _in_ Ruthven, Thompson and Thompson, _loc. cit._). _Iosco_: (Lagler, 1943:283, symbol on map). _Kent_: (Lagler, _loc. cit._). _Montcalm_: (Clark _in_ Ruthven, Thompson and Thompson, _loc. cit._). _Muskegon_: Muskegon River "near" Muskegon (Lagler, _op. cit._:303). _Van Buren_: Reynolds Lake, 2.5 mi. E Lawrence (Edgren, 1942:180).

MISSISSIPPI: _De Soto_: Lake Cormorant (Stejneger, 1944:55). _Holmes_: Thornton (Cook, 1946:185). _Humphreys_: Belzoni (Stejneger, _loc. cit._). _Sunflower_: _Warren_: Vicksburg, Eagle Lake (Cook, _loc. cit._). _Washington_: Lake Washington (Smith and List, 1955:125); Greenville (Stejneger, _loc. cit._).

NEW YORK: _Albany_: Hudson River at Albany (DeKay, 1842:7); Mohawk River at Cohoes (Eights _in_ Bishop, 1923:120). _Cattaraugus_: Allegheny River and Red House Lake in Allegheny State Park (Eaton, 1945:115). _Chautauqua_: Lake Chautauqua (DeKay, _loc. cit._). _Monroe_: Braddocks Bay and Long Pond on Lake Ontario (Wright, 1919:8). _Saratoga_: Hudson River near Baker's Falls (restricted locality, Rafinesque, 1832:64). _County unknown_: Lake Cayuga; Mohawk River (DeKay, _loc. cit._).

OHIO (Conant, 1951:158-59, 264, except records from Allen, Geauga and Noble counties): _Allen_: Sugar Creek, 6 mi. N Lima (Adler and Dennis, 1960:27). _Ashland_: Long Lake, Lake Twp.; Black Fork, Sec. 27, Green Twp. _Athens_: Hocking River "near" Athens; "near" Fisher, Alexander Twp. _Auglaize_: Pusheta Creek, west of Wapakoneta. _Brown_: White Oak Creek, 1 mi. N Higginsport. _Butler_: Oxford. _Champaign_: Mad River, 4 mi. SW Urbana. _Coshocton_: Walhouding River, below dam. _Defiance_: Auglaize River, Shawnee Scout Camp, Defiance Twp. _Erie_: Huron; Sandusky. _Fairfield_: Buckeye Lake. _Franklin_: Alum Creek, Westerville; Columbus. _Geauga_: Chardon Twp. (Wood, 1959:8). _Greene_: Huffman Dam. _Hamilton_: Harrison; mouth of Miami River. _Hardin_: "near" Hepburn. _Henry_: Maumee River, east of Napoleon; Maumee River "near" Texas; Maumee River, 3 mi. W Texas. _Highland_: Little Brush Creek, 2 mi. N Sinking Spring. _Huron_: Huron River "near" Monroeville. _Jackson_: Canter's Cove, Jackson Twp.; Jackson Lake. _Knox_: Brinkhaven. _Lake_: east branch Chagrin River, Kirtland; Grand River, 4 mi. E Painesville. _Lawrence_: Pine Creek, Elizabeth Twp. _Logan_: Miami River, "near" Indian Lake. _Lorain_: Oberlin. _Lucas_: Lake Erie at Reno Beach, Jerusalem Twp.; Lake Erie, 1/2 mi. offshore from mouth of Crane Creek; Maumee River at Maumee; Swan Creek, W of Toledo; "near" Waterville; Swan Creek "near" Whitehorse. _Madison_: London. _Medina_: Hinckley Lake. _Meigs_: Shade River, below Darwin. _Miami_: Miami River, above Troy. _Monroe_: Cranenest Fork, Green Twp. _Montgomery_: Mad River, Dayton; Miami River, Dayton; Stillwater River, Dayton. _Morrow_: Kokosing River, Franklin Twp. _Noble_: Jct. Sharon Twp. 1 and St. Rt. 78. (Adler and Dennis, 1960:27). _Ottawa_: East Harbour, Catawba Island. _Pike_: Chenoweth Fork, Sunfish Twp.; Scioto River, Camp Creek Twp. _Ross_: Paint Creek near Bainbridge. _Vinton_: Lake Hope; Lake Alma. _Warren_: Fort Ancient. _Washington_: Dam No. 2, Muskingum River, "near" Marietta. _Williams_: 1 mi. S Blakesley; St. Joseph River "near" Blakesley; West Branch, St. Joseph River, Sec. 8, Bridgewater Twp.; Edgerton. _Wood_: Grand Rapids; Grassy Creek, Rossford; Haskins; Maumee River opposite Toledo.

PENNSYLVANIA: _Allegheny_: Monongahela River above McKeesport (Atkinson, 1901:154); Ohio River at Pittsburgh (Wied-Neuwied _in_ Stejneger, 1944:44, 49). _Armstrong_: (Swanson, 1952:165). _Clarion_: Clarion River "near" Clarion (Allen, 1955:228); Foxburg (= Foxbury?, Boulenger, 1889:260). _Crawford_: _Elk_: _Erie_: Edinboro Lake. _Forest_: (Swanson, _loc. cit._). _Indiana_: Plum Creek; Crooked Creek (Netting _in_ Stejneger, 1944:48). _McKean_: (Swanson, _loc. cit._). _Somerset_: Stoyestown (Surface, 1908:122). _Warren_: _Venango_: Allegheny River south of Franklin (Swanson, _loc. cit._).

TENNESSEE: _Chester_: South Fork, Forked Deer River just E Henderson (Endsley, 1954:40). _Clay_: Mill Creek, 3 mi. from Butler's Landing; Obey River above mouth of Wolf River at Lilydale; mouth of Wolf River (Shoup, Peyton and Gentry, 1941:75); Iron Creek "near" Willow Grove (Stejneger, 1944:56). _Fentress_: _Jackson_: (Gentry, 1941:332). _Lake_: Reelfoot Lake (Parker, 1948:29). _Obion_: Walnut Log (Parker, 1937:85); east shore of Reelfoot Lake, Samburg (Rhoads, 1895:386). _Overton_: Medlock Branch, tributary of West Fork Obey River north of Allred (Shoup, Peyton and Gentry, _loc. cit._). _Roane_: 2 mi. S Kingston (Stejneger, 1944:55).

VERMONT: _Chittenden_: Lake Champlain, mouth of Winooski River; "near" Burlington; Milton (= Minton) (Babcock, 1919:420). _Franklin_: Swanton (Stejneger, 1944:55).

WEST VIRGINIA: _Randolph_: Tygart River at Elkins (Green, 1937:116).

WISCONSIN: _Burnett_: _Crawford_: (Pope and Dickinson, 1928:83). _Dane_: Lake Wingra, Madison (Noland, 1951:54). _Grant_: (Pope and Dickinson, _loc. cit._). _Green Lake_: Berlin (AMNH 6840-41, listed in card file March 2, 1959). _Jefferson_: Lake Mills (Dickinson, 1950:75). _La Crosse_: West Salem (Pope, 1930:281). _Oneida_: _Pepin_: (Pope and Dickinson, _loc. cit._). _Racine_: Eagle Lake (Edgren, 1944:498); Burlington; Rochester (Stille and Edgren, 1948:201). _Sheboygan_: Sheboygan (KKA). _Trempealeau_: _Vernon_: "near" Viroqua (Pope, _loc. cit._). _Walworth_: Lake Beulah (Dickinson, _loc. cit._). _Washburn_: (Pope and Dickinson, _loc. cit._). _Waukesha_: Lac La Belle (Cahn, 1929:8). _Winnebago_: Wolfe River (Dickinson, _loc. cit._).

=Trionyx spinifer hartwegi= (Conant and Goin)

Western Spiny Softshell

Plates 35 and 36

_Amyda spinifera hartwegi_ Conant and Goin, Occas. Papers Mus.
Zool. Univ. Mich., No. 510:1, pl. 1, map 1, June 15, 1948.

_T[rionyx] s[pinifer] hartwegi_ Schwartz, Charleston Mus. Leaflet,
No. 26:11, May, 1956.

_Type._--Holotype, UMMZ 95365; alcoholic adult male; obtained at Wichita, Sedgwick County, Kansas, in May, 1945, by Robert Young.

_Range._--Central United States in tributaries flowing into the Mississippi River from the west, except the Red River drainage; eastern Montana, North Dakota, and southern Minnesota south to eastern Colorado, northern Oklahoma and Arkansas (see map, Fig. 19).

_Diagnosis._--Juvenal pattern of small ocelli, rarely as large as two millimeters in diameter, or usually solid black dots that are not much larger in center of carapace than at sides (mean OD/PL, Kansas, .022); only one dark marginal line separating pale rim of carapace from dorsal ground color.

_Description._--Plastral length of smallest hatchling, 2.8 centimeters (USNM 9928); of largest male, 13.1 centimeters (USNM 55687); of largest female, 25.5 centimeters (KU 2283).

Carapace olive, having small ocelli or black spots that are not much larger in the center of the carapace than at the sides; pale rim of carapace separated from darker ground color by one dark marginal line and not four or five times wider posteriorly than laterally; large females often having black dots at sides of carapace on mottled and blotched pattern; 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 with ground color on side of head; postlabial stripe broken, interrupted; pale postocular stripe having blackish borders interrupted, not joining with postlabial stripes; dorsal surface of soft parts of body having contrasting pattern, largest blackish marks on hind limbs; elongate tail of males having pale dorsolateral bands with well-defined, lower, blackish borders; patterns on soft parts of body usually obscured or absent on large females; 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; tubercles along anterior edge of carapace small and conical on adult males, and conical or knoblike on 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, 4.24, and exceeding 7.0 centimeters, 5.33; 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, 1.19; mean CL/PCW, 2.00; mean SL/HW, 1.30 (including subspecies _spinifer_); mean CL/PL, 1.38.

_Variation._--Variants include: CNHM 8949, UMMZ 72511 and TU 14591 having ocelli approximately 4 millimeters in diameter that are almost solid spots; KU 17728 having pale stripes on snout that lack black, inner borders; TTC 719 (female, plastral length 20.7 cm.), having distinct pattern on snout; USNM 14535, 17823, 55684, and 123446 (from different localities) having markings confined to margins of carapace (Stejneger, 1944:66, suggested that USNM 17823 probably came from Texas); UMMZ 92667 (female, plastral length 6.7 cm.) lacking pattern on carapace.

_Comparisons._--_T. s. hartwegi_ can be distinguished from all other subspecies of _T. spinifer_ by the presence of small dots and ocelli on the carapace that are all of approximately the same size in combination with only one dark marginal line. _T. s. hartwegi_ resembles _asper_ in having small blackish ocelli or dots on the carapace but differs from _asper_ in having only one dark marginal line. _T. s. hartwegi_ differs from _spinifer_ only in the small size of the ocelli. _T. s. hartwegi_ resembles _spinifer_ and _asper_, but differs from _pallidus_, _guadalupensis_ and _emoryi_ in having blackish spots and ocelli on the carapace and lacking small whitish spots. _T. s. hartwegi_ resembles _spinifer_, _asper_ and _pallidus_ but differs from _guadalupensis_ and _emoryi_ in having conical or knoblike tubercles on the anterior edge of the carapace on large females.

_T. s. hartwegi_ differs from the subspecies _asper_, _guadalupensis_ and _emoryi_ in having a narrower head, and from _emoryi_ in having a wider carapace. _T. s. hartwegi_ resembles _spinifer_ and _asper_ but differs from the other subspecies in having the carapace widest at a plane approximately one-half way back on the carapace. _T. s. hartwegi_ and _spinifer_ have longer snouts than do _pallidus_ and _guadalupensis_ or _emoryi_. _T. s. hartwegi_ differs from _asper_ but resembles the other subspecies in having a relatively longer plastron.

_Remarks._--The validity of _T. s. hartwegi_ has never been questioned. It intergrades with _spinifer_ over a broad area paralleling the Mississippi River. For convenience, specimens occurring west of the Mississippi River are referred to the subspecies _hartwegi_. Figure 8 shows much variation in size of ocelli on different individuals from the same state. For example, UMMZ 92667, plastral length 6.7 centimeters has a uniform pale brown carapace lacking any dark marks, whereas UMMZ 92652, plastral length 5.9 centimeters has some ocelli three millimeters in diameter on the carapace. Both are from Iowa. One specimen from Kansas, KU 1954 (Doniphan County, plastral length 11.8 cm.), has ocelli four millimeters in diameter, and USNM 7648 captured farther west at Fort Laramie, Wyoming, an adult male having a plastral length of 11.0 centimeters, has some ocelli five millimeters in diameter on the carapace. TTC 1090, an adult male from the panhandle of Texas has some ocelli so much as 5.5 millimeters in diameter. The size of the ocelli seemingly varies in the same local population.

Specimens of _T. spinifer_ in the lower Mississippi Valley are intergrades. Most individuals have small black dots on the carapace; some have small ocelli (TU 7216, 7501, 11912, 12123-24) interspersed with black dots (TU 5863), others have black dots confined to the edge of the carapace (TU 157, 4539, 7105), and still others have no pattern on the carapace (TU 7506, 13698.1, 10087.6). Two large males (TU 11580, 13025) have large ocelli (approximately five mm. in diameter) that have nearly black centers. In general, there is more dark pigmentation than farther north; some specimens have extensive pigmentation on the ventral surface of the carapace and soft parts of the body (TU 156, 5648). The dorsal surface of the limbs, especially the hind limbs, have a bold, black marbling and may be almost completely black (TU 5484, 5597). Many females, not exceeding plastral lengths of 7.0 centimeters, have a pale blotched pattern of lichenlike figures or have ill-defined black dots on the carapace (TU 10087, 13698.13, 13753.15).

Localities of specimens of _T. spinifer_ occurring in the Mississippi River drainage in Mississippi are arbitrarily listed under the account of the subspecies _spinifer_, whereas those in Louisiana (excluding _pallidus_) are listed under the account of _hartwegi_.

Neither Over (1943) nor Wheeler (1947:169) record _T. s. hartwegi_, respectively, from South Dakota or North Dakota; records from the Missouri River drainage in Montana suggest the occurrence of the species in that drainage in North and South Dakota.

_Specimens examined._--Total, 392 as follows: ARKANSAS: _Clay_: UMMZ 70735 (2), 7 mi. S St. Francis. _Crawford_: USNM 95352, Lee Creek, 7 mi. NW Natural Dam. _Drew_: CNHM 40785. _Lafayette_: KU 2225-29, 2944 (one of three specimens bearing last catalog number), 2963 (one of three specimens bearing this catalog number), 2964 (one of two specimens bearing this catalog number), Lewisville (see remarks under the account of the subspecies _pallidus_). _Lawrence_: CNHM 8949; CNHM 12598-600, 12602-04, TU 5855, UI 2413, Imboden; UI 2412, Black River at Powhatan. _Marion_: TU 14591 (6), White River at Cotter. _Prairie_: KU 1867, 1869, 1879, 1949-51, 2280-83, 2285-91 (2 specimens bear catalog number 2287), 2307, 2761-62, 2666, 2826, 2842, 3346-47, White River at DeValls Bluff. _Pulaski_: UMMZ 96540, Little Rock. _Saline_: USNM 17823, Saline River at Benton. _Searcy_: UMMZ 92755, Little Red River, 1.5 mi. SE Leslie. _Yell_: TU 14565, Petit Jean Creek, 10 mi. N Casa. _County unknown_: CNHM 28566-67, Ouachita River.

IOWA: _Allamakee_: UMMZ 72556-58, 92642-49, Mississippi River "near" Lansing. _Appanoose_: UMMZ 92667, Chariton River, 4.3 mi. N. Centerville. _Decatur_: UMMZ 92651, Grand River, 3.5 mi. WSW Decatur. _Dickinson_: UMMZ 55249, Milford; UMMZ 92655, Spirit Lake Twp. _Hamilton_: USNM 9928, Webster City. _Hardin_: UMMZ 92650, Eldora. _Louisa_: UMMZ 92654, Muscatine Slough, 12 mi. SW Muscatine, Muscatine County. _Muscatine_: INHS 7675, 5.5 mi. SE Muscatine; USNM 54730-32, Fairport. _Scott_: CNHM 433, Davenport; UMMZ 92656, Steamboat Slough, 2 mi. N Princeton. _Story_: UMMZ 92653, Squaw Creek at Ames. _Washington_: UMMZ 92652, English River, 2 mi. E Riverside.