North American Recent Soft-shelled Turtles (Family Trionychidae) Part 11

_Rapides_: TU 14040, Red River at Rapides. _Richland_: OU 25082. _St.

Bernard_: TU 16170, Delacroix Island. _St. Charles_: TU 4539, 4579, 5224, 5990, 11928 (12), 13698 (16), Bayou Gauche between Paradis and Des Allemands; TU 5863, 11580, Bonnet Carre Spillway at Norco.

_Tensas_: TU 5762, Lake St. Joseph near Newellton. _Union_: USNM 138946, Meridian Creek, 1 mi. E Conway; USNM 138947, Ouachita River, Alabama Landing. _Parish unknown_: MCZ 1622, Lake St. John (Concordia or Tensas Parish); USNM 029266, Louisiana?

MINNESOTA: _Hennepin_: AMNH 4759-60, Fort Snelling. _Lesueur_: KU 46742-43, Waterville, Lake Tetonka. _Winona_: USNM 59263-66, Homer.

MISSOURI: _Carter_: UMMZ 70737, "near" Van Buren. _Chariton_: UI 17509, Triplett. _Franklin_: USNM 55689. _Gasconade_: UMMZ 95900, Bourbeuse Creek, 8 mi. S Owensville. _Jefferson_: USNM 95405, Glaize Creek. _Lewis_: USNM 59279-80, Canton. _Miller_: UMMZ 91929, Barren Fork Tavern Creek, 5 mi. NW Iowna. _Newton_: UMMZ 82822, Shoal Creek, 12 mi. W Momit. _Phelps_: UMMZ 91930, Bourbeuse River, 10 mi. N St.

James. _Reynolds_: CNHM 35392, Black River at Warner Bay Spring; USNM 55688. _Ripley_: UMMZ 90435. _Shannon_: INHS 6223, Alley Spring State Park. _St. Charles_: USNM 93089-94, Dardenne Creek, St. Peters. _St.

Louis_: USNM 55685-87, Mississippi River at St. Louis. _Stone_: USNM 55684. _Washington_: USNM 55690. _Wayne_: UI 16554, Sam A. Baker State Park; UMMZ 95879, St. Francis River at Lodi. _County unknown_ (Wayne or Butler): UMMZ 83264, Clark National Forest, St. Francis River.

MONTANA: _Big Horn_: USNM 54421, Crow Agency. _Roosevelt_: USNM 58, Fort Union (locality reads "Yellowstone, Fort Union"; probably the Yellowstone River near Fort Union). _Wheatland_: UMMZ 92005, Musselsh.e.l.l River near Shawmut. _Yellowstone_: USNM 14535, Custer.

OKLAHOMA: _Alfalfa_: OU 9316, 2 mi. S Cherokee. _Cleveland_: OU 22973, Norman. _Delaware_: UMMZ 81476, Spavinaw. _LeFlore_: OU 16802, 1.5 mi.

E Zoe. _Osage_: UMMZ 89628, Big Hominy Creek. _Pottawatomie_: OU 25175, 5 mi. SW Shawnee. _Rogers_: OU 7317, Verdigris River, 5 mi. W Claremore; UMMZ 81473-74, near Garnett, Tulsa County; UMMZ 81475, 4 mi. NE Inola. _Sequoyah_: OU 9008, 2 mi. NE Gore; TU 13885, Little Vian Creek, 1 mi. E Vian. _Texas_: OU 5005, 5 mi. SE Guymon. _Tulsa_: TU 17061, Bird Creek "near" Skiatook, Osage County. _Woods_: CHNM 11809, Waynoka; OU 9432, 2.5 mi. W Waynoka; OU 9579, 9581-82, 1 mi. S Waynoka.

TEXAS: _Hansford_: TTC 719, 10 mi. S, 2 mi. W Gruver. _Hutchinson_: TTC 1090, Carson Creek, Turkey Track Ranch.

WYOMING: _Goshen_: USNM 7648, Fort Laramie. _Weston_: UMMZ 78080, Beaver Creek.

NO DATA: CNHM 21687-88, 22925. SM 142 (locality of Waco, McLennan County, Texas, believed in error). USNM 7649, 11625, 19622-23, 36412 (Illinois River).

_Records in the literature._--ARKANSAS: _Benton_: (Dowling, 1957:37).

_Chicot_: Lake Chicot. _Clark_: Terre Noir Creek, 13 mi. W Arkadelphia. _Garland_: Ouachita River, Mountain Pine (Conant and Goin, 1948:7). _Hempstead_: _Jefferson_: (Dowling, _loc. cit._).

_Lawrence_: Black Rock (Dellinger and Black, 1938:46). _Madison_: _Scott_: _St. Francis_: (Dowling, _loc. cit._). _Washington_: near Greenland (Dellinger and Black, _loc. cit._).

COLORADO: _Boulder_: Boulder Creek, E Boulder; Boulder Creek, 6 mi. S and 1 mi. E Longmont. _Larimer_: Cache la Poudre River. _Logan_: 8 mi.

NE Sterling. _Morgan_: Platte River "near" Fort Morgan. _Otero_: Purgatoire River at Higbee. _Prowers_: Arkansas River at Lamar.

_Weld_: Poudre River "near" Greeley; Evans. _Yuma_: Bonny Dam, Republican River (Maslin, 1959:24-25).

IOWA: _d.i.c.kinson_: Little Sioux River, Okoboji Twp. (Blanchard, 1923:24). _Story_: Skunk River, 5 mi. NNE Ames (Conant and Goin, 1948:9).

KANSAS: _Allen_: Petrolia (KKA). _Barber_: 7 mi. S Sun City. _Butler_: 3 mi. SE Augusta (Burt and Hoyle, 1934:198). _Chase_: 10 mi. SW Olpe; 7 mi. SW Saffordville (Breukelman and Smith, 1946:112). _Cherokee_: tributary of Spring River, 1 mi. N Riverton (Hall and Smith, 1947:451). _Coffey_: (Smith, 1956:160, symbol on map). _Cowley_: 11 mi. SE Winfield (Stejneger, 1944:55). _Crawford_: Pittsburg (Hall and Smith, _loc. cit._). _Doniphan_: "near" Geary (Linsdale, 1927:81).

_Elk_: (Smith, _loc. cit._). _Ellis_: Big Creek (Brennan, 1934:190); Ellis (Conant and Goin, 1948:2). _Franklin_: Middle Creek, SE part of county (Gloyd, 1928:135). _Greenwood_: (Stejneger, _op. cit._:54).

_Leavenworth_: Missouri River "near" Fort Leavenworth (Brumwell, 1951:208). _Lyon_: 5 mi. E Emporia (Breukelman and Smith, _loc.

cit._). _Marion_: (Smith, _loc. cit._). _Meade_: Meade County State Park, _ca._ 13 mi. SW Meade (Tihen and Sprague, 1939:505). _Ness_: 5.5 mi. NW Ness (Breukelman and Smith, _loc. cit._). _Osage_: Marais des Cygnes River; Long and Jordan Creeks (Clarke, 1958:21). _Reno_: 6 mi. E Turon. _Sedgwick_: 2 mi. NE Cheney (Burt, 1935:321). _Sheridan_: State Lake 7 mi. NE Quinter, Gove County (Breukelman and Smith, _loc.

cit._). _Wabaunsee_: Dragoon Creek at Harveyville (Clarke, 1956:215).

_Wallace_: (Burt, 1933:208). _Wilson_: Fall River, 1/2 mi. S Neodesha (Clarke, _loc. cit._).

MINNESOTA: _Anoka_: _Benton_: _Chisago_: (Breckenridge, 1944:184, symbols on map). _Crow Wing_: (Breckenridge, _op. cit._:185).

_Dakota_: (Hedrick and Holmes, 1956:126). _Goodhue_: (Breckenridge, _op. cit._:184, symbol on map). _Hennepin_: Minneapolis; Lake Minnetonka (Breckenridge, _op. cit._:187); 5 mi. N. Minneapolis (Breckenridge, 1955:5). _Houston_: Root River near Hokah. _Lesueur_: Lake Washington (Hedrick and Holmes, _loc. cit._). _Meeker_: Swan Lake (Breckenridge, 1957:232). _Pine_: (Breckenridge, 1944:185). _Ramsey_: _Rice_: _Sherburne_: _Stearns_: (Breckenridge, _op. cit._:184, symbols on map). _Washington_: just north of Stillwater (Hedrick and Holmes, _loc. cit._). _Winona_: Winona (Breckenridge, _op. cit._:187). _Yellow Medicine_: (Breckenridge, _op. cit._:185). _County unknown_ (Goodhue or Wabasha): Lake Pepin (Breckenridge, _op. cit._:184).

MISSOURI: _Boone_: east of Ashland (Henning, 1938:92). _Jackson_: Missouri River "near" Atherton (Anderson, 1942:219). _Jefferson_: Mississippi River "near" mouth Glaize Creek at Sulphur Springs; Glaize Creek at Barnhart (Boyer and Heinze, 1934:199). _St. Clair_: Osage River "near" Osceola. _Vernon_: Marmaton River, 7 mi. N Moundville (Conant and Goin, 1948:9).

MONTANA: Yellowstone River (Conant and Goin, 1948:9).

NEBRASKA: _Adams_: 1 mi. N Ayr (Hudson, 1942:101). _Dawson_: 2 mi. SE Gothenburg (Gehlbach and Collette, 1959:142). _Franklin_: 2 mi. SW Naponee. _Gage_: 1 mi. W Barnston. _Hitchc.o.c.k_: 3 mi. E Stratton.

_Holt_: Elkhorn River "near" Atkinson. _Lancaster_: Lincoln (Hudson, _loc. cit._). _Lincoln_: 1 mi. S Sutherland (Gehlbach and Collette, _loc. cit._). _Red Willow_: 14 mi. NW McCook. _Richardson_: 2 mi. S Rulo. _Wheeler_: 2 mi. W Ericson (Hudson, _loc. cit._).

OKLAHOMA: _LeFlore_: Wister (Conant and Goin, 1948:9); Shady Pointe (KKA); Poteau River, 6.5 mi. W Heavener (Trowbridge, 1937:301).

_Tulsa_: Arkansas River "near" Tulsa (Force, 1930:38).

WYOMING: _Goshen_: Platte River (Conant and Goin, 1948:10).

=Trionyx spinifer asper= (Aga.s.siz)

Gulf Coast Spiny Softsh.e.l.l

Plates 37 and 38

_Aspidonectes asper_ Aga.s.siz, Contr. Nat. Hist. United States, 1(Pt. 2):405; 2(Pt. 3):pl. 6, fig. 3, 1857.

_Trionyx spinifer asper_ Schwartz, Charleston Mus. Leaflet, No. 26:17, pls. 1-3, map 2, May, 1956.

_Platypeltis aga.s.sizii_ Baur, Amer. Nat., 22:1121, 1888.

_Type._--Lectotype, MCZ 1597; alcoholic female; locality designated as Pearl River, Columbus, Marion County, Mississippi; received from Mr.

Winthrop Sargent of Natchez, Mississippi.

_Range._--Southeastern United States except peninsular Florida from the Florida Parishes of Louisiana east to southern North Carolina; Gulf Coast drainage including that of Lake Pontchartrain, Louisiana, eastward to the Apalachicola River system, and Atlantic Coast drainage including that of the Altamaha River in Georgia northward to the Pee Dee River drainage in South Carolina (see map, Fig 19).

_Diagnosis._--Juvenal pattern of black ocelli and spots, and two or more black, interrupted, lines paralleling rear margin of carapace; pale postocular and postl.a.b.i.al stripes often united on side of head; length of plastron short.

_Description._--Plastral length of smallest hatchling, 2.9 centimeters (USNM 134244); of largest male, 13.2 centimeters (TU 17117); of largest female, 27.0 centimeters (TU 13474).

Blackish marginal rings on carapace number two, three or four posteriorly, but decrease in number anteriorly; segments of marginal rings may extend to nuchal region; marginal rings increasingly interrupted inwardly; pattern of hatchlings having well-defined marginal rings that are not extensively interrupted (often males), or having marginal rings broken into small segments or series of dots, and pale outer margin of carapace marked by ill-defined, hazy, inner border (often females); conspicuous marginal rings often lacking on hatchling females; pale rim of carapace not four or five times wider posteriorly than laterally; carapace having blackish dots, spots, small ocelli or a combination thereof; marks on carapace of slightly varying sizes, some occasionally barlike (usually males); some hatchling females showing pale, irregular blotching on carapace, often characterized by small lichenlike figures superimposed on blackish dots.

Striping on snout variable; pale, dark-bordered stripes usually unite in front of eyes and form right or acute angle; medial dark borders of pale stripes on snout not joined anteriorly, broken into segments or dots, reduced to single median line, united to form straight line connecting anterior margins of orbits (usually with slight medial indentation), or absent; pale postocular and postl.a.b.i.al stripes often joined, relationship variable and on either side of head; side of head with or without dark markings, sometimes a pale subocular blotch bordered below by a dark line; pattern on dorsal portions of soft parts of body contrasting, less so on limbs of hatchlings; pattern of irregular dark marks, dark streaks usually coincident with digits; longitudinal streaks often occur on neck; elongate tail of adult males usually having well-defined, dorsolateral, pale bands with dark lower border more diffuse than upper border.

Underparts whitish often with dusky markings on rear of carapace or in region of bridge; blackish marks often on webbing and portions of soles and palms, and chin and throat.

Small conical tubercles along anterior edge of carapace on adult males; remnants of juvenal pattern usually present on carapace of large females; conical or k.n.o.blike tubercles on anterior edge of carapace of large females; accessory k.n.o.blike tubercles in nuchal region (a paravertebral pair usually most prominent), and posteriorly in middle of carapace on large females.

Ontogenetic variation in PL/HW, mean PL/HW of specimens having plastral lengths 7.0 centimeters or less, 3.87, and exceeding 7.0 centimeters, 4.94; ontogenetic variation in CL/CW, mean CL/CW of specimens having plastral lengths 8.5 centimeters or less, 1.11, and exceeding 8.5 centimeters, 1.16; mean CL/PCW, 1.71; mean CL/PL, 1.45.

_Variation._--The s.e.x of some hatchlings can be distinguished by the pattern on the carapace (see Plate 37 for different patterns), but the s.e.x of many hatchlings cannot be distinguished on the basis of pattern.

In the early stages of this study, I thought that the pattern on the carapace differed in eastern and western populations, and that the zone of intergradation was in Alabama. Adult males from the Tombigbee-Alabama river drainage and westward were noted to have blackish spots (some slightly ocellate) intermixed with few, if any, smaller blackish dots, whereas the adult males from east of the Tombigbee-Alabama river drainage had many small, black dots intermixed with slightly larger, mostly ocellate marks (see Plate 38, left, top and bottom, for contrast); also, hatchlings from western populations were never observed to have four marginal rings. On the basis of pattern, I would have thought that the individual having many ocelli, that lacks correct locality data and that is photographed by Stejneger (1944:Pl. 26), came from Georgia or South Carolina; but, the pattern (_op. cit._:Pl. 27) of a specimen, probably an adult male, from South Carolina, resembles the pattern on adult males from Louisiana. The differences noted above are probably due to individual variation rather than geographic variation.

Color notes taken from life of a freshly-killed adult male (TU 16071, Louisiana) are: carapace olive, spots blackish, outer rim buff; top of head olive, postocular and postl.a.b.i.al stripes yellow with blackish borders, stripes on snout buff with blackish borders; dorsal ground color of soft parts of body pale olive-green, larger marks blackish, ground color laterally toward juncture of pattern and immaculate undersurface, and toward insertions of neck and limbs becoming yellowish; webbing on hind limbs having reddish tinge; dorsolateral bands on tail yellow with blackish borders; undersurface whitish; chin and throat olive-green with blackish marks; becoming buff then whitish posteriorly.

[Ill.u.s.tration: FIG. 20. Basicranial length and ratio of greatest diameter of internal choanae to least width of maxillary bridge (IC/MB) on 30 skulls of _T. ferox_ (open circles), 26 of _T. spinifer_ (crosses), and 12 of the _aga.s.sizi_-form (solid circles; half shaded circle represents holotype of _aga.s.sizi_). Skulls of the _aga.s.sizi_-form tend to have slightly smaller internal choanae than those of _spinifer_ or _ferox_.]

Occasional specimens have only one definite dark line paralleling the rear margin of the carapace. Schwartz (1956:16) reported that Charleston Museum No. 55.159.26 has only one solid line at the margin of the carapace, and I received an adult male (KU 47120) reported to have come from the Pearl River that is aberrant in not having more than one dark marginal line. USNM 95191, a large stuffed female from the Pearl River is mentioned by Stejneger (1944:59, Pl. 17) as having marks that "a.s.sume the form of short lines parallel with the submarginal ring"; I examined this specimen and noted that it had only one dark marginal line. Stejneger (_op. cit._:64) mentioned another from the Pearl River drainage, and Crenshaw and Hopkins (1955:20) wrote that some individuals from Georgia have only one dark marginal line. Presumably MCZ 1606 (now in the Albany Museum) recorded by Stejneger (_op. cit._:52) as _Amyda s. spinifer_ from Columbus, Georgia, is another specimen.

[Ill.u.s.tration: FIG. 21. Basicranial length and greatest width of alveolar surface of maxilla on 52 skulls of _T. spinifer_ (open circles) and 11 of the _aga.s.sizi_-form (solid circles; half shaded circle represents holotype of _aga.s.sizi_). Most skulls of the _aga.s.sizi_-form that exceed 43 mm. in basicranial length have a more expanded, alveolar surface of the maxilla than skulls of _spinifer_ of approximately the same size. All skulls exceeding 50 mm. are those of females.]

Some skulls of soft-sh.e.l.led turtles from streams of the Atlantic Coast drainage, including the skull of the holotype of _Platypeltis_ (= _Trionyx_) _aga.s.sizi_ Baur (MCZ 37172, Pl. 54), show at least two differences from other skulls of _asper_ and from those of other subspecies of _T. spinifer_. Figure 20 shows that skulls of _aga.s.sizi_ tend to have slightly smaller internal choanae (ratio IC/MB) than those of _T. spinifer_ and _T. ferox_; there is seemingly little difference between skulls of _ferox_ and _spinifer_, and little, if any, ontogenetic variation. Figure 21 shows that most skulls of the _aga.s.sizi_-form that exceed 43.0 millimeters have a more expanded, alveolar surface of the maxilla than skulls of _spinifer_ of approximately the same size; most skulls exceeding a basicranial length of 43.0 millimeters, and certainly all skulls exceeding 50.0 millimeters are those of females. Stejneger (1944:Pl. 30) also has provided photographs of a skull of the _aga.s.sizi_-form. It is of interest that of the 12 _aga.s.sizi_-form skulls (MCZ 37172; USNM 8708, 029034, 51981, 66859, 71681, 91282, 91310-11, 92521, 92583-84) that I examined some resemble _ferox_ (Neill, 1951:9) in having the alveolar surfaces of the jaws broadened, and the greatest width at the level of the quadratojugal (Table 3, Plate 54); also, the localities of all 12 skulls are within the geographic range of _ferox_. Skulls of _ferox_, however, have conspicuously broadened alveolar surfaces of the jaws only when they exceed in length the largest skulls of _aga.s.sizi_. The differences of skulls of the _aga.s.sizi_-form possibly reflect isolation in the Atlantic Coast drainage, and an adaptation in feeding habits. So far as I can ascertain, individuals occurring in rivers of the Atlantic Coast drainage in Georgia and South Carolina (referable to _aga.s.sizi_) do not differ consistently in external characters from individuals of _T. s. asper_ that occur westward in the Apalachicola drainage.

_Comparisons._--_Trionyx s. asper_ can be distinguished from all other subspecies of _T. spinifer_ by usually having more than one black line paralleling the rear margin of the carapace. This character and the frequent fusion of the postl.a.b.i.al and postocular stripes on the side of the head distinguish _asper_ from _spinifer_ and _hartwegi_. _T. s.