Chapter 4

Type.—Holotype, British Museum (Natural History) 1947.3.6.17; original number 53A, presumably that of Royal Society; stuffed adult female and skull; obtained from the Savannah River, Georgia, by Dr. Alexander Garden.Range.—Southern South Carolina, southeastern Georgia, and all of Florida except the Keys and perhaps the western end of the panhandle (see map,Fig. 18).

Range.—Southern South Carolina, southeastern Georgia, and all of Florida except the Keys and perhaps the western end of the panhandle (see map,Fig. 18).

Fig. 18.Map of southeastern United States showing geographic distribution ofTrionyx ferox.

Diagnosis.—Marginal ridge present; longitudinal rows of tubercles that resemble ridges on carapace of hatchlings; plastron often extending farther forward than carapace in adults; plastral area dark slate or gray in hatchlings; juvenal pattern of large slate or blackish blotches (often with pale centers) on a pale background; pale outer rim of carapace (absent on adults) narrow, not separated from ground color of carapace by distinct, dark line.Size large; head wide; carapace relatively long and narrow; snout short; greatest width of skull at level of quadratojugal; often no suture between hypoplastra and hyoplastra; callosities on epiplastron and preplastra usually lacking.Description.—Plastral length of smallest hatchling, 2.9 centimeters (UMMZ 95613), of largest male, 26.0 centimeters (AMNH 63642), of largest female, 34.0 centimeters (UMMZ 38123).Septal ridges present; over-all coloration of carapace and plastron, and soft parts of body of hatchlings slate or blackish; carapace having blackish, circular blotches, usually fused at margin, often with pale centers on buff background forming coarse reticulum; pale, narrow rim of carapace not separated from ground color by dark marginal line; pale rim, coincident with marginal ridge, absent from anteriormost nuchal region; longitudinal rows of tubercles on carapace resembling ridges; undersurface blackish, usually having posterior part of carapace pale with irregular blackish marks; blackish soft parts of body dorsally having large, pale markings, most consistent of which are postocular mark that may contact orbit, postlabial mark that curves around angle of jaws, inverted Y on top of snout, and one or two streaks on side of neck.Over-all coloration of adults grayish, paler than in hatchlings; carapace gray sometimes having slightly darker, large, irregular markings; mottled and blotched pattern on females not contrasting; sex of many large individuals not distinguishable on basis of pattern on carapace; pale rim of carapace obscure or absent; soft parts of body dorsally gray or brownish on large adults of both sexes, sometimes having slightly paler, large markings; small adult males usually having contrasting pattern on head; surface of carapace smooth (not "sandpaper") on adult males; undersurface whitish, throat often grayish; well-defined marginal ridge; anterior edge of carapace laterally to region of insertion of forelimbs studded with low, flattened tubercles resembling hemispheres, never conical; carapace usually having blunted tubercles, best developed anteriorly and posteriorly on midline, but sometimes linearly arranged, resembling ridges, especially at margins; anterolateral parts of plastron often extending farther forward than corresponding parts of carapace.Range in length (in cm.) of plastron of ten largest specimens of each sex (mean follows extremes), males, 17.0-26.0, 20.0; females 23.3-34.0, 27.9; ontogenetic variation in PL/HW, mean PL/HW of specimens having plastral lengths 6.5 centimeters or less, 3.52, and exceeding 6.5 centimeters, 4.87; ontogenetic variation in CL/CW, mean CL/CW of specimens having plastral lengths 8.0 centimeters or less, 1.18, and exceeding 8.0 centimeters, 1.30; mean CL/PCW, 2.01; mean HW/SL, 1.44; mean CL/PL, 1.26.Jaws of some skulls that exceed 75 millimeters in basicranial length having expanded alveolar surfaces; greatest width of skull usually at level of quadratojugal (72%); ventral surface of supraoccipital spine narrow proximally, usually having medial ridge; foramen magnum rhomboidal; opisthotic-exoccipital spur absent (82%), sometimes indicated by ridge (16%); distal part of opisthotic wing[481]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; usually a combination of seven neurals, seven pairs of pleurals, and contact of seventh pair of pleurals (56%), often eight pairs of pleurals (31%); angle of epiplastron forming approximate right angle; often no suture between hypoplastra and hyoplastra; callosities on preplastra and epiplastron usually lacking.Variation.—Crenshaw and Hopkins (1955:19) stated that in specimens from Lake Okeechobee and southward the carapace is wider relative to the width of the head, and Neill (1951:19) quoted Allen's observations thatferoxfrom southern Florida "average larger and darker than those collected farther north."Carr (1952:417) reported that the pale reticulum on the carapace is yellowish olive, the markings on head are yellow on an olive ground color, some markings more orange, and the plastron slate gray. Duellman and Schwartz (1958:271) mentioned that the carapace of hatchlings is edged in orange grading to yellow posteriorly and has a pattern of bluish-black blotches on a dull brown background, whereas the carapace is dull brown or blackish on adults. Neill (op. cit.:18) wrote "that the head stripes and the marginal ring of the 'carapace' are orange rather than yellow (yellow at the time of hatching, however)."The transition from the dark coloration of hatchlings to the paler coloration of adults is gradual and subject to individual variation. The loss of dark color ventrally occurs first on the plastral area, then the hind limbs, forelimbs, posterior part of carapace and last on the neck and throat. The soft parts of the body dorsally are gray or dark gray, and do not become so pale as the ventral surface. The smallest specimen that I have seen displaying the dark features of the hatchlings is a male, 7.7 centimeters (UMMZ 100673); a female, 9.5 centimeters (UMMZ 110987), is the smallest specimen having a whitish plastral area. The change from dark to pale coloration on the ventral surface occurs at a size of 8.0 to 9.0 centimeters. The largest specimens I have seen having indistinct, dusky blotches of the underside of the carapace are a female, 11.3 centimeters (UMMZ 100836), and a male, 16.0 centimeters (UMMZ 106322). A contrasting pattern on head and limbs, and a dark throat are still evident in a female 19.2 centimeters (UMMZ 106302).Comparisons.—Trionyx feroxcan be distinguished from all other species of the genus in North America by the presence of a marginal ridge, longitudinal ridges of tubercles on the carapace of juveniles (less evident in adults), and the unique juvenal pattern and coloration. The lack of a juvenal pattern and a smooth surface on the carapace (not gritty like sandpaper) distinguish adult males from those ofT. spinifer. Most adults of both sexes can be distinguished fromspiniferandmuticusby the extension of the plastron farther forward than the carapace (developed to a slight degree in some specimens ofT. s. emoryi). Both sexes of all ages can be distinguished frommuticusby the presence of knoblike tubercles on the anterior edge of the carapace, and septal ridges.T. feroxis the largest species in North America; the maximum size of the plastron in adult males is approximately 26.0 centimeters (16.0 inspinifer) and of adult females, 34.0 centimeters (31.0 inspinifer). The head is wider inferoxthan inmuticusand most subspecies ofspinifer(closely approached[482]byasper,guadalupensis,emoryiandT. ater). The carapace is narrower inferoxthan inmuticusand most subspecies ofspinifer(closely approached byemoryiandT. ater). The snout is shortest inferox, but almost as short inT. s. emoryiandT. ater.T. feroxhas proportionately the longest plastron in relation to length of carapace.Most skulls offeroxdiffer from those ofmuticusandspiniferin having the greatest width at the level of the quadratojugal (as do someT. s. asper; see account of that subspecies). In the skull,feroxresemblesspiniferbut differs frommuticusin 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. feroxresemblesmuticusbut differs from most individuals ofspiniferin lacking a well-developed opisthotic-exoccipital spur.T. feroxresemblesspiniferbut differs frommuticusin having the epiplastron bent at approximately a right angle;feroxdiffers from bothmuticusandspiniferin lacking a callosity on the epiplastron and probably in the more frequent fusion of the hyoplastra and hypoplastra.Remarks.—The early taxonomic history ofTrionyx feroxhas been discussed in detail by Stejneger (1944:27-32), who explained that Dr. Alexander Garden of Charleston, South Carolina, sent a description and specimen ofT. feroxto Thomas Pennant, and at the same time sent another specimen with drawings to a friend, John Ellis, in London. Pennant presented one of the specimens and drawings and the description to the Royal Society of London in 1771; the description was published in 1772 and included Garden's drawings. Because two specimens were involved the possibility exists that the description (text, drawings and type specimen) is a composite based on two specimens.I have not seen the type. Garden's original description (inPennant, 1772:268-271) leaves little doubt that the text subject is a large adult female offerox(see especially the statements, "fore part, [of carapace] just where it covers the head and neck, is studded full of large knobs, [and] The under, or belly plate, … is … extended forward two or three inches more than the back plate, …"). I am indebted to Mr. J. C. Battersby, British Museum (Natural History), Department of Zoology (Reptiles), for information concerning the type and for comparing it with the text description and three figures published by Pennant. The carapace of the type is approximately 16 inches long, 131/2inches wide, and has low, flattened, knoblike tubercles along the anterior edge. Some inaccuracies on the part of the artist (such as five claws on both feet on the right side ofFig. 3, and four claws on the left front foot ofFig. 2are evident), and slight changes in the proportions of the type would have occurred after death and preservation. It is the opinion of Mr. Battersby that the type, text description and three figures represent one specimen. Figures 1 and 2, dorsal and ventral views respectively, probably represent the same specimen from life; the neck is withdrawn and the tail tip is visible in dorsal view, but concealed beneath the posterior edge of the carapace in ventral view. Presumably the same specimen (probably drawn from dried and stuffed animal) is depicted inFigure 3(dorsal view); the neck is fully extended and a large part of the thick, pyramidal tail is visible in dorsal view. British Museum (Natural History) 1947.3.6.17 is considered a holotype. The three figures published[483]by Pennant have been duplicated by Schoepff (1795:Pl. 19) and Duméril and Bibron (1835:482). To my knowledge, the holotype was first specifically designated as the "(Type.)" ofT. feroxby Boulenger (1889:259). The skull of the holotype is figured by Stejneger (1944:Pl. 5).Garden did not list a specific locality for the two specimens that he sent to London, but did mention that the turtle was common in the Savannah and Altamaha rivers (of Georgia), and rivers in east Florida. Boulenger (loc. cit.) stated that the locality of the holotype was "Georgia." Baur (1893:220) restricted the type locality to the "Savannah river, Ga." Neill (1951:17), who believedT. feroxto be absent from the Savannah River, changed the type locality offeroxto east Florida. Schwartz (1956:8) reappraised the status of softshells in Georgia and Florida and reëstablished the Savannah River (at Savannah), Georgia, as the type locality ofT. ferox.Pennant failed to use binomial nomenclature when he published the type description of Garden. The first name-combination (Testudo ferox) was proposed by Schneider (1783:220).Lacépède (1788:137, Pl. 7) referred to Garden's description in Pennant only as "The Molle" but on a folded paper chart entitled "Table Méthodique des Quadrupèdes ovipares," which is inserted after an introduction of 17 pages, listedT. mollis; this name is again listed on another folded chart, entitled "Synopsis methodica Quadrupedum oviparorum," which is inserted between pages 618 and 619 under the genusTestudo. The illustration (Pl. 7) was taken from Pennant (Duméril and Bibron,loc. cit.). The type locality has been designated "(following Stejneger, 1944) as eastern Florida" by Schmidt (1953:108).Bartram failed to use a binomial name with his description of "the great soft shelled tortoise," which appeared in hisTravels(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'sTravelshas 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 wasTestudo(ferox?)verrucosaproposed in 1795 by Schoepff; it appeared simultaneously inThe Historia Testudinumand 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 hisVoyage(French translation), proposed the nameTestudo bartrami; Harper (op. cit.:717) restricted the type locality ofT. bartramifrom "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'sTravelsthat 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 ofChelys(binomial nomenclature not employed), and Duméril and Bibron (loc. cit.) suggested that the description was based partly on a "Chelyde Matamata."[484]The descriptive comments of Bartram are not clearly applicable toTestudo feroxSchneider;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?)verrucosaSchoepff,Testudo bartramiDaudin, andMesodeca bartramiRafinesque are junior synonyms ofTestudo feroxSchneider.Schweigger (1812:285) referredferoxto the genusTrionyxfollowing the description of that genus by Geoffroy in 1809.Testudo feroxwas listed as a synonym by Geoffroy in the description ofTrionyx georgicus(1809a:17); Duméril and Bibron (1835:432) mentioned that the specific characters ofgeorgicuswere taken from Pennant. The nameTrionyx georgianuspresumably appears for this taxon in Geoffroy's earlier-published synopsis (1809:367).T. georgicuswas 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 namesgeorgicusandgeorgianusare regarded as substitute names and junior synonyms ofT. ferox.Geoffroy (1809a:14-15) also describedTrionyx carinatus, a name-combination that hitherto has been considered a synonym ofTrionyx ferox. There is no indication from the description thatcarinatusis applicable toferox. 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 listedTestudo membranaceaandTestudo rostrataas synonyms ofcarinatus. Fitzinger (1835:127) listedT. membranacea,T. rostrataandT. carinatusas synonyms ofTrionyx javanicus(=T. cartilagineus), which was also described by Geoffroy (op. cit.:15). Duméril and Bibron (op. cit.:478, 482) consideredcarinatusto be the young ofspinifer(feroxas synonym). Gray (1844:48), however, referredT. membranaceaandT. rostratato the synonymy ofT. javanicus, but consideredT. carinatusto be a synonym ofT. ferox(op. cit.:50), an interpretation followed by all subsequent authors.Trionyx carinatusis questionably listed as a synonym offeroxby Stejneger (1944:27). Duméril and Bibron (op. cit.:482) wrote that the young type ofcarinatusis in the museum at Paris. Dr. Jean Guibé informs me in letter of September 24, 1959, that the type of Geoffroy'sT. carinatuscannot be found in the Natural History Museum at Paris. For the present,T. carinatusis considered anomen dubium. According to Stejneger (1944:27),Trionyx brongniartiSchweigger is a substitute name forT. carinatus.I am unable to add anything to Stejneger's (op. cit.:32) account ofTrionyx harlani; the mention of its occurrence in east Florida indicates that it is indistinguishable fromTestudo feroxSchneider.T. feroxwas considered to be indistinguishable from Lesueur'sTrionyx 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 ofT. spinifer asperas the young offerox. Consequently, the geographic range offerox, as envisioned by Agassiz, extended from Georgia and Florida west to Louisiana. Neill (1951:15) considered all[485]American forms conspecific. Crenshaw and Hopkins (1955) and Schwartz (1956) demonstrated thatferoxis a distinct species.Fitzinger (1843:30) designated the speciesferoxas the type species of his genusPlatypeltisas follows: "Platypeltis. Fitz. Am[erica].Platypelt. ferox. Fitz. Typus." If populations of soft-shelled turtles that are referable toTestudo feroxSchneider are considered to comprise a distinct genus by future workers,PlatypeltisFitzinger, 1835, is available as a generic name withTestudo feroxSchneider, 1783, as the type species (by subsequent designation).Trionyx feroxin the northern part of its range is sympatric withT. spinifer asper. In the region of overlap, the two species are nearly always ecologically isolated;feroxinhabits lentic waters, whereasT. s. asperis partial to lotic waters (Crenshaw and Hopkins,op. cit.:16). There is no evidence of intergradation or hybridization.Many characters ofTrionyx feroxthat 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,feroxis more closely allied to Old World forms of the genus than tomuticusorspinifer.Carr (1940:107) recordedferoxfrom 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.[486]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).MonroeandOkaloosa(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).

Size large; head wide; carapace relatively long and narrow; snout short; greatest width of skull at level of quadratojugal; often no suture between hypoplastra and hyoplastra; callosities on epiplastron and preplastra usually lacking.

Description.—Plastral length of smallest hatchling, 2.9 centimeters (UMMZ 95613), of largest male, 26.0 centimeters (AMNH 63642), of largest female, 34.0 centimeters (UMMZ 38123).

Septal ridges present; over-all coloration of carapace and plastron, and soft parts of body of hatchlings slate or blackish; carapace having blackish, circular blotches, usually fused at margin, often with pale centers on buff background forming coarse reticulum; pale, narrow rim of carapace not separated from ground color by dark marginal line; pale rim, coincident with marginal ridge, absent from anteriormost nuchal region; longitudinal rows of tubercles on carapace resembling ridges; undersurface blackish, usually having posterior part of carapace pale with irregular blackish marks; blackish soft parts of body dorsally having large, pale markings, most consistent of which are postocular mark that may contact orbit, postlabial mark that curves around angle of jaws, inverted Y on top of snout, and one or two streaks on side of neck.

Over-all coloration of adults grayish, paler than in hatchlings; carapace gray sometimes having slightly darker, large, irregular markings; mottled and blotched pattern on females not contrasting; sex of many large individuals not distinguishable on basis of pattern on carapace; pale rim of carapace obscure or absent; soft parts of body dorsally gray or brownish on large adults of both sexes, sometimes having slightly paler, large markings; small adult males usually having contrasting pattern on head; surface of carapace smooth (not "sandpaper") on adult males; undersurface whitish, throat often grayish; well-defined marginal ridge; anterior edge of carapace laterally to region of insertion of forelimbs studded with low, flattened tubercles resembling hemispheres, never conical; carapace usually having blunted tubercles, best developed anteriorly and posteriorly on midline, but sometimes linearly arranged, resembling ridges, especially at margins; anterolateral parts of plastron often extending farther forward than corresponding parts of carapace.

Range in length (in cm.) of plastron of ten largest specimens of each sex (mean follows extremes), males, 17.0-26.0, 20.0; females 23.3-34.0, 27.9; ontogenetic variation in PL/HW, mean PL/HW of specimens having plastral lengths 6.5 centimeters or less, 3.52, and exceeding 6.5 centimeters, 4.87; ontogenetic variation in CL/CW, mean CL/CW of specimens having plastral lengths 8.0 centimeters or less, 1.18, and exceeding 8.0 centimeters, 1.30; mean CL/PCW, 2.01; mean HW/SL, 1.44; mean CL/PL, 1.26.

Jaws of some skulls that exceed 75 millimeters in basicranial length having expanded alveolar surfaces; greatest width of skull usually at level of quadratojugal (72%); ventral surface of supraoccipital spine narrow proximally, usually having medial ridge; foramen magnum rhomboidal; opisthotic-exoccipital spur absent (82%), sometimes indicated by ridge (16%); distal part of opisthotic wing[481]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; usually a combination of seven neurals, seven pairs of pleurals, and contact of seventh pair of pleurals (56%), often eight pairs of pleurals (31%); angle of epiplastron forming approximate right angle; often no suture between hypoplastra and hyoplastra; callosities on preplastra and epiplastron usually lacking.

Variation.—Crenshaw and Hopkins (1955:19) stated that in specimens from Lake Okeechobee and southward the carapace is wider relative to the width of the head, and Neill (1951:19) quoted Allen's observations thatferoxfrom southern Florida "average larger and darker than those collected farther north."

Carr (1952:417) reported that the pale reticulum on the carapace is yellowish olive, the markings on head are yellow on an olive ground color, some markings more orange, and the plastron slate gray. Duellman and Schwartz (1958:271) mentioned that the carapace of hatchlings is edged in orange grading to yellow posteriorly and has a pattern of bluish-black blotches on a dull brown background, whereas the carapace is dull brown or blackish on adults. Neill (op. cit.:18) wrote "that the head stripes and the marginal ring of the 'carapace' are orange rather than yellow (yellow at the time of hatching, however)."

The transition from the dark coloration of hatchlings to the paler coloration of adults is gradual and subject to individual variation. The loss of dark color ventrally occurs first on the plastral area, then the hind limbs, forelimbs, posterior part of carapace and last on the neck and throat. The soft parts of the body dorsally are gray or dark gray, and do not become so pale as the ventral surface. The smallest specimen that I have seen displaying the dark features of the hatchlings is a male, 7.7 centimeters (UMMZ 100673); a female, 9.5 centimeters (UMMZ 110987), is the smallest specimen having a whitish plastral area. The change from dark to pale coloration on the ventral surface occurs at a size of 8.0 to 9.0 centimeters. The largest specimens I have seen having indistinct, dusky blotches of the underside of the carapace are a female, 11.3 centimeters (UMMZ 100836), and a male, 16.0 centimeters (UMMZ 106322). A contrasting pattern on head and limbs, and a dark throat are still evident in a female 19.2 centimeters (UMMZ 106302).

Comparisons.—Trionyx feroxcan be distinguished from all other species of the genus in North America by the presence of a marginal ridge, longitudinal ridges of tubercles on the carapace of juveniles (less evident in adults), and the unique juvenal pattern and coloration. The lack of a juvenal pattern and a smooth surface on the carapace (not gritty like sandpaper) distinguish adult males from those ofT. spinifer. Most adults of both sexes can be distinguished fromspiniferandmuticusby the extension of the plastron farther forward than the carapace (developed to a slight degree in some specimens ofT. s. emoryi). Both sexes of all ages can be distinguished frommuticusby the presence of knoblike tubercles on the anterior edge of the carapace, and septal ridges.

T. feroxis the largest species in North America; the maximum size of the plastron in adult males is approximately 26.0 centimeters (16.0 inspinifer) and of adult females, 34.0 centimeters (31.0 inspinifer). The head is wider inferoxthan inmuticusand most subspecies ofspinifer(closely approached[482]byasper,guadalupensis,emoryiandT. ater). The carapace is narrower inferoxthan inmuticusand most subspecies ofspinifer(closely approached byemoryiandT. ater). The snout is shortest inferox, but almost as short inT. s. emoryiandT. ater.T. feroxhas proportionately the longest plastron in relation to length of carapace.

Most skulls offeroxdiffer from those ofmuticusandspiniferin having the greatest width at the level of the quadratojugal (as do someT. s. asper; see account of that subspecies). In the skull,feroxresemblesspiniferbut differs frommuticusin 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. feroxresemblesmuticusbut differs from most individuals ofspiniferin lacking a well-developed opisthotic-exoccipital spur.T. feroxresemblesspiniferbut differs frommuticusin having the epiplastron bent at approximately a right angle;feroxdiffers from bothmuticusandspiniferin lacking a callosity on the epiplastron and probably in the more frequent fusion of the hyoplastra and hypoplastra.

Remarks.—The early taxonomic history ofTrionyx feroxhas been discussed in detail by Stejneger (1944:27-32), who explained that Dr. Alexander Garden of Charleston, South Carolina, sent a description and specimen ofT. feroxto Thomas Pennant, and at the same time sent another specimen with drawings to a friend, John Ellis, in London. Pennant presented one of the specimens and drawings and the description to the Royal Society of London in 1771; the description was published in 1772 and included Garden's drawings. Because two specimens were involved the possibility exists that the description (text, drawings and type specimen) is a composite based on two specimens.

I have not seen the type. Garden's original description (inPennant, 1772:268-271) leaves little doubt that the text subject is a large adult female offerox(see especially the statements, "fore part, [of carapace] just where it covers the head and neck, is studded full of large knobs, [and] The under, or belly plate, … is … extended forward two or three inches more than the back plate, …"). I am indebted to Mr. J. C. Battersby, British Museum (Natural History), Department of Zoology (Reptiles), for information concerning the type and for comparing it with the text description and three figures published by Pennant. The carapace of the type is approximately 16 inches long, 131/2inches wide, and has low, flattened, knoblike tubercles along the anterior edge. Some inaccuracies on the part of the artist (such as five claws on both feet on the right side ofFig. 3, and four claws on the left front foot ofFig. 2are evident), and slight changes in the proportions of the type would have occurred after death and preservation. It is the opinion of Mr. Battersby that the type, text description and three figures represent one specimen. Figures 1 and 2, dorsal and ventral views respectively, probably represent the same specimen from life; the neck is withdrawn and the tail tip is visible in dorsal view, but concealed beneath the posterior edge of the carapace in ventral view. Presumably the same specimen (probably drawn from dried and stuffed animal) is depicted inFigure 3(dorsal view); the neck is fully extended and a large part of the thick, pyramidal tail is visible in dorsal view. British Museum (Natural History) 1947.3.6.17 is considered a holotype. The three figures published[483]by Pennant have been duplicated by Schoepff (1795:Pl. 19) and Duméril and Bibron (1835:482). To my knowledge, the holotype was first specifically designated as the "(Type.)" ofT. feroxby Boulenger (1889:259). The skull of the holotype is figured by Stejneger (1944:Pl. 5).

Garden did not list a specific locality for the two specimens that he sent to London, but did mention that the turtle was common in the Savannah and Altamaha rivers (of Georgia), and rivers in east Florida. Boulenger (loc. cit.) stated that the locality of the holotype was "Georgia." Baur (1893:220) restricted the type locality to the "Savannah river, Ga." Neill (1951:17), who believedT. feroxto be absent from the Savannah River, changed the type locality offeroxto east Florida. Schwartz (1956:8) reappraised the status of softshells in Georgia and Florida and reëstablished the Savannah River (at Savannah), Georgia, as the type locality ofT. ferox.

Pennant failed to use binomial nomenclature when he published the type description of Garden. The first name-combination (Testudo ferox) was proposed by Schneider (1783:220).

Lacépède (1788:137, Pl. 7) referred to Garden's description in Pennant only as "The Molle" but on a folded paper chart entitled "Table Méthodique des Quadrupèdes ovipares," which is inserted after an introduction of 17 pages, listedT. mollis; this name is again listed on another folded chart, entitled "Synopsis methodica Quadrupedum oviparorum," which is inserted between pages 618 and 619 under the genusTestudo. The illustration (Pl. 7) was taken from Pennant (Duméril and Bibron,loc. cit.). The type locality has been designated "(following Stejneger, 1944) as eastern Florida" by Schmidt (1953:108).

Bartram failed to use a binomial name with his description of "the great soft shelled tortoise," which appeared in hisTravels(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'sTravelshas 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 wasTestudo(ferox?)verrucosaproposed in 1795 by Schoepff; it appeared simultaneously inThe Historia Testudinumand 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 hisVoyage(French translation), proposed the nameTestudo bartrami; Harper (op. cit.:717) restricted the type locality ofT. bartramifrom "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'sTravelsthat 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 ofChelys(binomial nomenclature not employed), and Duméril and Bibron (loc. cit.) suggested that the description was based partly on a "Chelyde Matamata."[484]The descriptive comments of Bartram are not clearly applicable toTestudo feroxSchneider;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?)verrucosaSchoepff,Testudo bartramiDaudin, andMesodeca bartramiRafinesque are junior synonyms ofTestudo feroxSchneider.

Schweigger (1812:285) referredferoxto the genusTrionyxfollowing the description of that genus by Geoffroy in 1809.Testudo feroxwas listed as a synonym by Geoffroy in the description ofTrionyx georgicus(1809a:17); Duméril and Bibron (1835:432) mentioned that the specific characters ofgeorgicuswere taken from Pennant. The nameTrionyx georgianuspresumably appears for this taxon in Geoffroy's earlier-published synopsis (1809:367).T. georgicuswas 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 namesgeorgicusandgeorgianusare regarded as substitute names and junior synonyms ofT. ferox.

Geoffroy (1809a:14-15) also describedTrionyx carinatus, a name-combination that hitherto has been considered a synonym ofTrionyx ferox. There is no indication from the description thatcarinatusis applicable toferox. 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 listedTestudo membranaceaandTestudo rostrataas synonyms ofcarinatus. Fitzinger (1835:127) listedT. membranacea,T. rostrataandT. carinatusas synonyms ofTrionyx javanicus(=T. cartilagineus), which was also described by Geoffroy (op. cit.:15). Duméril and Bibron (op. cit.:478, 482) consideredcarinatusto be the young ofspinifer(feroxas synonym). Gray (1844:48), however, referredT. membranaceaandT. rostratato the synonymy ofT. javanicus, but consideredT. carinatusto be a synonym ofT. ferox(op. cit.:50), an interpretation followed by all subsequent authors.Trionyx carinatusis questionably listed as a synonym offeroxby Stejneger (1944:27). Duméril and Bibron (op. cit.:482) wrote that the young type ofcarinatusis in the museum at Paris. Dr. Jean Guibé informs me in letter of September 24, 1959, that the type of Geoffroy'sT. carinatuscannot be found in the Natural History Museum at Paris. For the present,T. carinatusis considered anomen dubium. According to Stejneger (1944:27),Trionyx brongniartiSchweigger is a substitute name forT. carinatus.

I am unable to add anything to Stejneger's (op. cit.:32) account ofTrionyx harlani; the mention of its occurrence in east Florida indicates that it is indistinguishable fromTestudo feroxSchneider.

T. feroxwas considered to be indistinguishable from Lesueur'sTrionyx 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 ofT. spinifer asperas the young offerox. Consequently, the geographic range offerox, as envisioned by Agassiz, extended from Georgia and Florida west to Louisiana. Neill (1951:15) considered all[485]American forms conspecific. Crenshaw and Hopkins (1955) and Schwartz (1956) demonstrated thatferoxis a distinct species.

Fitzinger (1843:30) designated the speciesferoxas the type species of his genusPlatypeltisas follows: "Platypeltis. Fitz. Am[erica].Platypelt. ferox. Fitz. Typus." If populations of soft-shelled turtles that are referable toTestudo feroxSchneider are considered to comprise a distinct genus by future workers,PlatypeltisFitzinger, 1835, is available as a generic name withTestudo feroxSchneider, 1783, as the type species (by subsequent designation).

Trionyx feroxin the northern part of its range is sympatric withT. spinifer asper. In the region of overlap, the two species are nearly always ecologically isolated;feroxinhabits lentic waters, whereasT. s. asperis partial to lotic waters (Crenshaw and Hopkins,op. cit.:16). There is no evidence of intergradation or hybridization.

Many characters ofTrionyx feroxthat 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,feroxis more closely allied to Old World forms of the genus than tomuticusorspinifer.

Carr (1940:107) recordedferoxfrom 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.[486]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).MonroeandOkaloosa(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 spiniferLesueurSpiny 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[487]largest specimens of each sex (mean follows extremes), males, 13.8-16.0, 14.4; females, 26.0-31.0, 28.0.

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[487]largest specimens of each sex (mean follows extremes), males, 13.8-16.0, 14.4; females, 26.0-31.0, 28.0.

Fig. 19.Geographic distribution ofTrionyx spinifer.Guide to subspecies:T. s. spiniferT. s. hartwegiT. s. asperT. s. pallidusT. s. guadalupensisT. s. emoryi

Fig. 19.Geographic distribution ofTrionyx spinifer.

Guide to subspecies:

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 spinifercan be distinguished fromT. feroxandT. muticusby the presence of any one of the characters mentioned in the "Diagnosis." Both sexes and all sizes ofT. spiniferresembleferoxbut differ frommuticusin having septal ridges. Most individuals ofT. spinifer(except some large females) resemblemuticusbut differ fromferoxand large females ofaterin 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 subspeciesspinifer,hartwegi,asperandpallidusmay 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[488]ferox(flattened, knoblike prominences), andmuticusandater(smooth surface, no prominences). Large females of the subspeciesguadalupensisandemoryiresemblemuticusandater, and to some extentferox, in having low, scarcely elevated prominences along the anterior edge of the carapace. Some females ofemoryiresembleferoxin that the plastron extends farther forward than the carapace.T. spiniferis intermediate in size betweenferox(larger) andmuticus(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 ofspiniferis proportionately narrower than inferoxbut wider than inmuticus.In the skull,spinifermore closely resemblesferoxthanmuticus, but differs from bothferoxandmuticusin usually having a well-developed opisthotic-exoccipital spur. Skulls ofspiniferresemble those ofmuticusbut differ from those offeroxin being widest at the level of the squamosal. Skulls ofspiniferresemble those offeroxbut differ from those ofmuticusin 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. spiniferresemblesferoxbut differs frommuticusin having the epiplastron bent at an approximate right angle.T. spiniferdiffers fromferoxin having an epiplastral callosity, and frommuticusin that the callosity does not cover the entire surface of the epiplastron. The hyo-hypoplastral suture is present more often inspiniferandmuticusthan inferox.Remarks.—Gray (1869:221) proposed the generic nameCalliniaas a new name forAspidonectesas understood by Agassiz (1857:403). Gray referredTrionyx spiciferus(=spiniferus) Lesueur toCallinia. Stejneger (1907:514) designatedTrionyx spiniferusLesueur as the type species ofCallinia. IfTrionyx spiniferusLesueur is considered to be generically distinct from other soft-shelled turtles,CalliniaGray, 1869, is available as a generic name withTrionyx spiniferusLesueur, 1827, as the type species by subsequent designation.

Comparisons.—Trionyx spinifercan be distinguished fromT. feroxandT. muticusby the presence of any one of the characters mentioned in the "Diagnosis." Both sexes and all sizes ofT. spiniferresembleferoxbut differ frommuticusin having septal ridges. Most individuals ofT. spinifer(except some large females) resemblemuticusbut differ fromferoxand large females ofaterin 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 subspeciesspinifer,hartwegi,asperandpallidusmay 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[488]ferox(flattened, knoblike prominences), andmuticusandater(smooth surface, no prominences). Large females of the subspeciesguadalupensisandemoryiresemblemuticusandater, and to some extentferox, in having low, scarcely elevated prominences along the anterior edge of the carapace. Some females ofemoryiresembleferoxin that the plastron extends farther forward than the carapace.

T. spiniferis intermediate in size betweenferox(larger) andmuticus(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 ofspiniferis proportionately narrower than inferoxbut wider than inmuticus.

In the skull,spinifermore closely resemblesferoxthanmuticus, but differs from bothferoxandmuticusin usually having a well-developed opisthotic-exoccipital spur. Skulls ofspiniferresemble those ofmuticusbut differ from those offeroxin being widest at the level of the squamosal. Skulls ofspiniferresemble those offeroxbut differ from those ofmuticusin 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. spiniferresemblesferoxbut differs frommuticusin having the epiplastron bent at an approximate right angle.T. spiniferdiffers fromferoxin having an epiplastral callosity, and frommuticusin that the callosity does not cover the entire surface of the epiplastron. The hyo-hypoplastral suture is present more often inspiniferandmuticusthan inferox.

Remarks.—Gray (1869:221) proposed the generic nameCalliniaas a new name forAspidonectesas understood by Agassiz (1857:403). Gray referredTrionyx spiciferus(=spiniferus) Lesueur toCallinia. Stejneger (1907:514) designatedTrionyx spiniferusLesueur as the type species ofCallinia. IfTrionyx spiniferusLesueur is considered to be generically distinct from other soft-shelled turtles,CalliniaGray, 1869, is available as a generic name withTrionyx spiniferusLesueur, 1827, as the type species by subsequent designation.

Geographic variation.—T. spiniferis 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 inT. s. emoryi, which has small white spots on the carapace. Some of the characters at the western terminus of this geographical gradient are shared withT. aterandmuticus. Those subspecies comprising theemoryigroup also show proportional characters that correspond closely with those ofT. ferox.

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

Trionyx spinifer spiniferLesueurEastern Spiny Softshell

Plates33,34, and52

Trionyx spiniferusLesueur, Mém. Mus. Hist. Nat. Paris, 15:258, pl. 6, December, 1827.

T[rionyx] s[pinifer] spiniferSchwartz, Charleston Mus. Leaflet, No. 26:11, May, 1956.

Trionyx ocellatusLesueur, Mém. Mus. Hist. Nat. Paris, 15:261, December, 1827.

Apalone hudsonicaRafinesque, Atlan. Jour., Friend Knowledge, Philadelphia, 1(No. 2, Art. 12):64, Summer, 1832.

Trionyx annuliferWied-Neuwied, Riese Nord-Amerika, 1(pt. 3):140, 1838.

Tyrse argusGray, Cat. Tort. Croc. Amphis. Brit. Mus., p. 48, 1844.

Aspidonectes nuchalisAgassiz, Contr. Nat. Hist. United States, 1(pt. 2):406, 1857.

?G[ymnopus] olivaceusWied-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[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 subspecieshartwegi); 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. spinifercan be distinguished from all other subspecies ofT. spiniferby 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. spiniferresemblesasperin having ocelli or dots on the carapace but differs fromasperin having only one dark marginal line and larger ocelli.T. s. spiniferdiffers fromhartwegionly in the large size of the ocelli.T. s. spiniferresembleshartwegiandasperbut differs frompallidus,guadalupensisandemoryiin having blackish spots and ocelli on the carapace and lacking whitish dots.T. s. spiniferresembleshartwegi,asper, andpallidusand differs fromguadalupensisandemoryiin having conical or knoblike tubercles on the anterior edge of the carapace on large females.T. s. spiniferdiffers from the subspeciesasper,guadalupensisandemoryiin having a relatively narrower head, and fromemoryiin having a relatively wider carapace.T. s. spiniferresembleshartwegiandasperbut differs from the other subspecies in having the carapace widest at a plane approximately one-half way back on the carapace. The subspeciesspiniferandhartwegihave longer snouts thanpallidus,guadalupensis, andemoryi.T. s. spiniferdiffers fromasperbut resembles all the other subspecies in having a relatively longer plastron.Remarks.—Lesueur's description ofTrionyx 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 describedTrionyx ocellatus(op. cit.:261-263) as a variety ofT. spiniferushaving 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 ofocellatusas 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 ofT. ocellatusLesueur, 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" asApalone 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'sTravelsthat showed each limb with five, clawed digits. Perhaps this was the basis for Boulenger (1889:245, footnote) regardingApaloneas "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 toT. 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 (inBishop, 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 (inBishop,loc. cit.), and symbolized as an isolated locality by Conant (1958:318, map 35). The type locality ofApalone hudsonicais herein restricted to the Hudson River, near Baker's Falls, Saratoga County, New York.Gray (1844:48) proposed the nameTyrse argusfor a specimen reported to have come from Sierra Leone, West Africa; later (1855:68), he referred the species to the genusTrionyx. After comparison with a specimen ofT. spiniferusLesueur, 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 ofTyrse argus], and whether it is not more probably a North American species." The same author (1869:222; 1870:109) listedTyrse argusas a synonym ofCallinia spinifera(=Trionyx spiniferusLesueur).Agassiz (op. cit.:406-07) describedAspidonectes nuchalison 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 ofA. nuchalisrequired 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 thatnuchaliswas not distinguishable fromT. 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 thatnuchalis"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 ofAspidonectes nuchalisAgassiz. MCZ 1908 is one of the young syntypes mentioned by Agassiz, and is referable tospinifer. 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 asTrionyx ocellatus, and agreed with Lesueur that those turtles having occulated spots on the carapace were distinguishable fromT. spiniferusandT. muticus. But because Duméril and Bibron in theirErpétologie Généralfailed to recognizeT. 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 ofT. annulifer(1838:140); seemingly Wied-Neuwied consideredT. annuliferandG. olivaceaas conspecific, although there is no mention ofannuliferin the text proper. Stejneger (op. cit.:49) designated the type locality ofT. annuliferas the Ohio River at Pittsburgh, Pennsylvania, and ofGymnopus olivaceaas New Harmony, Wabash River, Illinois (lapsusfor Indiana).Trionyx spiniferuswas questionably considered distinct fromT. feroxby[493]Lesueur who listed "Testudo ferox Gm. Tortue de Pennant?" and "Trionyx georgicus Geoffr.?" as synonyms. Subsequently, most authors consideredT. spiniferussynonymous withT. feroxuntil Agassiz (1857) pointed out differences between the two species.The average size of the ocelli on the carapace of the subspeciesspiniferdecreases 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 tospinifer, recognizing that intergradation occurs withhartwegiover a broad area paralleling the Mississippi River. The type locality ofspiniferis 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"spiniferto the east. Intergradation withasperpossibly occurs in that part of the Tennessee River in eastern Tennessee as exemplified by UMMZ 59198.Published reports indicate thatT. s. spiniferis 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 theTrionyxpopulation 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 thatspinifer"is not common in Lake Champlain."T. s. spiniferprobably extended its geographic range into the Hudson River drainage of New Yorkviathe 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 ofspiniferto the Hudson River drainage, Lake Ontario and intervening waterways in New York (Mertens, 1928:199). Netting (1944:86-87), however, suggested thatspiniferoccupied 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 thatTrionyxoccurs in Maryland.The unsuccessful introduction ofT. s. spiniferin 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[494]introduced into the eastern part of Pennsylvania through the canal from the Western and Central part of New York," and Roddy (inNeill, 1951:21) suggested that the species may be found in the Susquehanna River. Babcock (1919:420) mentioned a young specimen ofspiniferin 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 ofT. s. spiniferdo not occur in rivers of the Atlantic Coast drainage, except probably the Hudson-Mohawk drainage.Stockwell (1878:401) wrote thatspiniferwas 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 7294,1/2mi. 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[495]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,1/2mi. 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 (KennicottinStejneger, 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: (HahninStejneger, 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 (Garmanin[496]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 (GarmaninCahn,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 (KirschinStejneger, 1944:45).Franklin: (HughesinStejneger,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 (McLaininStejneger,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 (ClarkinRuthven, Thompson and Thompson, 1912:133).Genesee: (MilesinRuthven, Thompson and Thompson,loc. cit.).Iosco: (Lagler, 1943:283, symbol on map).Kent: (Lagler,loc. cit.).Montcalm: (ClarkinRuthven, 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 (EightsinBishop, 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,1/2mi. offshore from mouth of Crane Creek; Maumee River at Maumee; Swan Creek, W of Toledo; "near" Waterville; Swan Creek "near" Whitehorse.Madison: London.Medina:[497]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-NeuwiedinStejneger, 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 (NettinginStejneger, 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.).

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[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 subspecieshartwegi); 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. spinifercan be distinguished from all other subspecies ofT. spiniferby 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. spiniferresemblesasperin having ocelli or dots on the carapace but differs fromasperin having only one dark marginal line and larger ocelli.T. s. spiniferdiffers fromhartwegionly in the large size of the ocelli.T. s. spiniferresembleshartwegiandasperbut differs frompallidus,guadalupensisandemoryiin having blackish spots and ocelli on the carapace and lacking whitish dots.T. s. spiniferresembleshartwegi,asper, andpallidusand differs fromguadalupensisandemoryiin having conical or knoblike tubercles on the anterior edge of the carapace on large females.

T. s. spiniferdiffers from the subspeciesasper,guadalupensisandemoryiin having a relatively narrower head, and fromemoryiin having a relatively wider carapace.T. s. spiniferresembleshartwegiandasperbut differs from the other subspecies in having the carapace widest at a plane approximately one-half way back on the carapace. The subspeciesspiniferandhartwegihave longer snouts thanpallidus,guadalupensis, andemoryi.T. s. spiniferdiffers fromasperbut resembles all the other subspecies in having a relatively longer plastron.

Remarks.—Lesueur's description ofTrionyx 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 describedTrionyx ocellatus(op. cit.:261-263) as a variety ofT. spiniferushaving 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 ofocellatusas 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 ofT. ocellatusLesueur, 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" asApalone 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'sTravelsthat showed each limb with five, clawed digits. Perhaps this was the basis for Boulenger (1889:245, footnote) regardingApaloneas "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 toT. 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 (inBishop, 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 (inBishop,loc. cit.), and symbolized as an isolated locality by Conant (1958:318, map 35). The type locality ofApalone hudsonicais herein restricted to the Hudson River, near Baker's Falls, Saratoga County, New York.

Gray (1844:48) proposed the nameTyrse argusfor a specimen reported to have come from Sierra Leone, West Africa; later (1855:68), he referred the species to the genusTrionyx. After comparison with a specimen ofT. spiniferusLesueur, 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 ofTyrse argus], and whether it is not more probably a North American species." The same author (1869:222; 1870:109) listedTyrse argusas a synonym ofCallinia spinifera(=Trionyx spiniferusLesueur).

Agassiz (op. cit.:406-07) describedAspidonectes nuchalison 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 ofA. nuchalisrequired 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 thatnuchaliswas not distinguishable fromT. 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 thatnuchalis"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 ofAspidonectes nuchalisAgassiz. MCZ 1908 is one of the young syntypes mentioned by Agassiz, and is referable tospinifer. 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 asTrionyx ocellatus, and agreed with Lesueur that those turtles having occulated spots on the carapace were distinguishable fromT. spiniferusandT. muticus. But because Duméril and Bibron in theirErpétologie Généralfailed to recognizeT. 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 ofT. annulifer(1838:140); seemingly Wied-Neuwied consideredT. annuliferandG. olivaceaas conspecific, although there is no mention ofannuliferin the text proper. Stejneger (op. cit.:49) designated the type locality ofT. annuliferas the Ohio River at Pittsburgh, Pennsylvania, and ofGymnopus olivaceaas New Harmony, Wabash River, Illinois (lapsusfor Indiana).

Trionyx spiniferuswas questionably considered distinct fromT. feroxby[493]Lesueur who listed "Testudo ferox Gm. Tortue de Pennant?" and "Trionyx georgicus Geoffr.?" as synonyms. Subsequently, most authors consideredT. spiniferussynonymous withT. feroxuntil Agassiz (1857) pointed out differences between the two species.

The average size of the ocelli on the carapace of the subspeciesspiniferdecreases 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 tospinifer, recognizing that intergradation occurs withhartwegiover a broad area paralleling the Mississippi River. The type locality ofspiniferis 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"spiniferto the east. Intergradation withasperpossibly occurs in that part of the Tennessee River in eastern Tennessee as exemplified by UMMZ 59198.

Published reports indicate thatT. s. spiniferis 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 theTrionyxpopulation 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 thatspinifer"is not common in Lake Champlain."

T. s. spiniferprobably extended its geographic range into the Hudson River drainage of New Yorkviathe 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 ofspiniferto the Hudson River drainage, Lake Ontario and intervening waterways in New York (Mertens, 1928:199). Netting (1944:86-87), however, suggested thatspiniferoccupied 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 thatTrionyxoccurs in Maryland.

The unsuccessful introduction ofT. s. spiniferin 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[494]introduced into the eastern part of Pennsylvania through the canal from the Western and Central part of New York," and Roddy (inNeill, 1951:21) suggested that the species may be found in the Susquehanna River. Babcock (1919:420) mentioned a young specimen ofspiniferin 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 ofT. s. spiniferdo not occur in rivers of the Atlantic Coast drainage, except probably the Hudson-Mohawk drainage.

Stockwell (1878:401) wrote thatspiniferwas 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 7294,1/2mi. 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[495]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,1/2mi. 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 (KennicottinStejneger, 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: (HahninStejneger, 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 (Garmanin[496]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 (GarmaninCahn,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 (KirschinStejneger, 1944:45).Franklin: (HughesinStejneger,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 (McLaininStejneger,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 (ClarkinRuthven, Thompson and Thompson, 1912:133).Genesee: (MilesinRuthven, Thompson and Thompson,loc. cit.).Iosco: (Lagler, 1943:283, symbol on map).Kent: (Lagler,loc. cit.).Montcalm: (ClarkinRuthven, 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 (EightsinBishop, 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,1/2mi. offshore from mouth of Crane Creek; Maumee River at Maumee; Swan Creek, W of Toledo; "near" Waterville; Swan Creek "near" Whitehorse.Madison: London.Medina:[497]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-NeuwiedinStejneger, 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 (NettinginStejneger, 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

Plates35and36

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

T[rionyx] s[pinifer] hartwegiSchwartz, Charleston Mus. Leaflet, No. 26:11, May, 1956.

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