Procambarus versutus (Hagen 1870)

Procambarus (Pe.) versutus: 96, 106

66. Procambarus (S.) howellae: 9, 11, 12, 45, 46, 53, 76, 80, 83, 87, 91, 94, 96, 98, 103, 116, 123, 124, 129, 134, 138, 143, 150, 153, 156, 158

67. Procambarus (S.) paeninsulanus: 4, 10, 13, 14, 19, 20, 24, 32, 35, 37, 43, 47, 49, 50, 65, 86, 88, 92, 100, 101, 125, 129, 135–137, 148, 159

68. Procambarus (S.) troglodytes: 15–17, 21, 25, 51, 62, 81, 82, 89, 91, 98, 121, 124, 132, 150

List of Counties

MAP

1. Appling: 42, 47, 52, 60, 64

2. Atkinson: 43, 44, 48, 54, 60, 64

3. Bacon: 42, 44, 48, 60, 64

4. Baker: 36, 61, 64, 67

5. Baldwin: 8, 59, 64

6. Banks: 2, 8, 63

7. Barrow: 8, 64

8. Bartow: 8, 10, 14, 26, 29, 37, 39, 64

9. Ben Hill: 44, 60, 66

10. Berrien: 43, 44, 48, 60, 64, 67

11. Bibb: 8, 64, 66

12. Bleckley: 27, 36, 64, 66

13. Brantley: 44, 54, 60, 67

14. Brooks: 44, 60, 64, 67

15. Bryan: 41, 43, 46, 52, 53, 56, 58–60, 62, 68

16. Bulloch: 8, 36, 45, 46, 52, 53, 56, 62, 68

17. Burke: 8, 9, 36, 50, 56, 59, 62, 63, 68

18. Butts: 8, 64

19. Calhoun: 27, 36, 67

20. Camden: 44, 47, 48, 60, 67

21. Candler: 43, 45, 46, 52, 56, 68

22. Carroll: 6, 8, 26, 64

23. Catoosa: 10, 15, 26, 30, 37, 38, 57

24. Charlton: 43, 44, 48, 50, 54, 60, 64, 67

25. Chatham: 41, 43, 46, 52, 53, 58, 68

26. Chattahoochee: 8, 64

27. Chattooga: 8, 10, 17, 25, 26, 29, 34, 37, 57

28. Cherokee: 8, 10, 14, 64

29. Clarke: 2, 8, 64

30. Clay: 8, 27, 64

31. Clayton: 8, 10, 64

32. Clinch: 43, 44, 48, 54, 60, 64, 67

33. Cobb: 3, 8, 64

34. Coffee: 42–44, 47, 48, 54, 60, 64

35. Colquitt: 44, 48, 60, 64, 67

36. Columbia: 8

37. Cook: 44, 48, 60, 64, 67

38. Coweta: 8, 10, 64

39. Crawford: 8, 61, 64

40. Crisp: 36, 44, 60, 64

41. Dade: 2, 8, 10, 15, 16, 22, 24, 25, 37

42. Dawson: 2, 8, 10, 14, 20, 64

43. Decatur: 21, 27, 64, 67

44. De Kalb: 3, 8, 26, 63, 64

45. Dodge: 36, 42, 50, 52, 66

46. Dooly: 10, 36, 44, 60, 64, 66

47. Dougherty: 36, 64, 67

48. Douglas: 3, 8, 64

49. Early: 10, 27, 35, 36, 64, 67

50. Echols: 44, 48, 54, 60, 64, 67

51. Effingham: 8, 9, 27, 36, 41, 43, 46, 53, 58, 68

52. Elbert: 8, 11, 50, 63

53. Emanuel: 36, 45, 46, 50, 52, 56, 59, 66

54. Evans: 41, 46, 52, 56

55. Fannin: 2, 8, 20, 26

56. Fayette: 8, 64

57. Floyd: 8, 10, 17, 26, 29, 37, 39, 64

58. Forsyth: 3, 8, 64

59. Franklin: 2, 8, 63

60. Fulton: 3, 8, 64

61. Gilmer: 2, 8, 13, 18, 20, 26, 64

62. Glascock: 8, 9, 50, 59, 62, 68

63. Glynn: 44, 47, 58, 60

64. Gordon: 8, 10, 26, 29, 39, 57, 64

65. Grady: 64, 67

66. Greene: 8, 64

67. Gwinnett: 8, 10, (59?), 63, 64

68. Habersham: 2, 8, 64

69. Hall: 2, 3, 8, 64

70. Hancock: 8, 59, 62, 64

71. Haralson: 5, 6, 8, 10, 64

72. Harris: 8, 10, 36, 64

73. Hart: 2, 8, 63

74. Heard: 8, 10, 64

75. Henry: 8, 64

76. Houston: 64, 66

77. Irwin: 44, 48, 60, 64

78. Jackson: 8, 64

79. Jasper: 8, 64

80. Jeff Davis: 36, 42, 43, 47, 60, 66

81. Jefferson: 8, 50, 59, 62, 68

82. Jenkins: 8, 36, 45, 46, 50, 53, 56, 59, 68

83. Johnson: 36, 50, 52, 64, 66

84. Jones: 8, 10, 27, 64

85. Lamar: 8, 64

86. Lanier: 44, 48, 54, 60, 64, 67

87. Laurens: 10, 12, 36, 42, (45?), 50, 52, 59, 64, 66

88. Lee: 8, 27, 36, 61, 64, 67

89. Liberty: 41, 43, 46, 56, 58, 68

90. Lincoln: 8

91. Long: 41, 43, 46, 52, 58, 66, 68

92. Lowndes: 44, 48, 54, 60, 64, 67

93. Lumpkin: 2, 3, 8, 14, 20, 23, 64

94. Macon: 10, 36, 64, 66

95. Madison: 2, 8, 63

96. Marion: 61, 64, 65, 66

97. McDuffie: 8, 59

98. McIntosh: 41, 43, 46, 52, 58, 66, 68

99. Meriwether: 7, 8, 50, 64

100. Miller: 10, 27, 36, 64, 67

101. Mitchell: 64, 67

102. Monroe: 8, 10, 64

103. Montgomery: 36, 41, 42, 47, 50, 52, 60, 66

104. Morgan: 8, 64

105. Murray: 2, 4, 8, 10, 17, 18, 20, 26, 29, 39, 64

106. Muscogee: 8, 64, 65

107. Newton: 8, 10, 63, 64

108. Oconee: 8, 64

109. Oglethorpe: 8, 11, 40, 50, 59, 63

110. Paulding: 6, 8, 29, 64

111. Peach: 64

112. Pickens: 8, 10, 14, 18, 20, 29, 64

113. Pierce: 44, 48, 60, 64

114. Pike: 8, 10, 64

115. Polk: 8, 10, 14, 26, 29, 39

116. Pulaski: 8, 27, 36, 64, 66

117. Putnam: 8, 64

118. Quitman: 8, 10, 27, 49, 64

119. Rabun: 2, 19, 23, 28, 31

120. Randolph: 8, 27, 64

121. Richmond: 8, 27, 50, 52, 59, 63, 68

122. Rockdale: 64

123. Schley: 27, 61, 64, 66

124. Screven: 8, 36, 41, 46, 50, 52, 53, 56, 59, 62, 63, 66, 68

125. Seminole: 35, 36, 64, 67

126. Spalding: 8, 64

127. Stephens: 2, 8, 19, 63, 64

128. Stewart: 8, 10, 27, 64

129. Sumter: 27, 36, 61, 64, 66, 67

130. Talbot: 8, 27, 64

131. Taliaferro: 8, 10, 59, 62

132. Tattnall: 41, 46, 52, 56, 58, 64, 68

133. Taylor: 64

134. Telfair: 36, 42, 43, 52, 64, 66

135. Terrell: 8, 27, 36, 64, 67

136. Thomas: 44, 64, 67

137. Tift: 36, 43, 44, 48, 60, 64, 67

138. Toombs: 41, 46, 52, 60, 64, 66

139. Towns: 2, 23, 32, 33

140. Treutlen: 45, 50, 52

141. Troup: 8, 27, 64

142. Turner: 36, 44, 60

143. Twiggs: 10, 36, 50, 64, 66

144. Union: 2, 16, 23, 32, 39, 64

145. Upson: 8, 64

146. Walker: 2, 8, 10, 15, 16, 25, 26, 29, 30, 34, 37, 57

147. Walton: 8, 26, 63, 64

148. Ware: 43, 44, 48, 60, 64, 67

149. Warren: 8, 9, 59, 62

150. Washington: 8, 10, 36, 45, 50, 52, 59, 62, 64, 66, 68

151. Wayne: 42, 43, 47, 60, 64

152. Webster: 64

153. Wheeler: 36, 42, 52, 66

154. White: 2, 8, 23, 64

155. Whitfield: 8, 10, 15, 17, 26, 29, 30, 37, 39, 57, 64

156. Wilcox: 44, 60, 64, 66

157. Wilkes: 2, 8, 11, 40, 50, 59, 63

158. Wilkinson: 8, 10, 12, 36, 50, 52, 59, 64, 66

159. Worth: 36, 43, 44, 60, 64, 67

Map of Georgia, showing counties.

Appendix 2

Symbionts of Georgia Crayfishes

Only nine references have been found that treat the symbionts of the crayfishes occurring in Georgia, and extracted from them is the following list, consisting of one microsporidian, one trematode, three branchiobdellid worms, and 25 entocytherid ostracods.

Hoff (1944) described four new entocytherids from the southern part of the state, and, 15 years later, Hart (1959), treating the same group in the lower Chattahoochee-Flint watershed, reported the occurrence of a fifth species and added new locality records for two of those described by Hoff. Hoffman (1963) described a new branchiobdellid worm from the southwestern part of Georgia, and Holt (1968a,b) added two additional ones from the Tallapoosa and Hiwassee watersheds. Hobbs and Walton (1968) reported the occurrence of a new entocytherid that infests the troglobitic crayfish in Climax Cave in Decatur County. The only parasites, of which I am aware, that have been reported to occur in Georgia crayfishes are those described by Sprague (1950) and Sullivan and Heard (1969). (The latter author kindly advised me of the change in the generic assignment of this fluke from Macroderoides to Allogossidium.) Hobbs III (1970) added a new entocytherid record for the Chattahoochee Basin. The remaining 18 ostracods known to occur in Georgia were reported in the studies of Hart and Han (1971, 1974).

In the following compilation, the crayfishes are listed alphabetically, and their symbionts (currently employed combinations are used throughout) are alphabetized in each of the four groups (B = branchiobdellid worms; E = entocytherid ostracods; S = sporozoans; T = trematode worms). The author, publication date, and page reference for each symbiont follow the name the first time it appears in the list. In addition, county records are included, and the bibliographic citation to the first reported locality in the county (or counties) is given in parentheses.

Carnbarus (C.) bartonii

S Thelohania cambari Sprague (1950:46): Towns (Sprague, 1950:46)

B Pterodrilus simondsi Holt (I968b:23): Fannin (Holt, 1968b: 23)

Procambarus (Pennides) versutus (Hagen)

Cambarus versutus Hagen, 1870:28, 31, 34, 51–52, 97, 101, 106, 107, pl. I: figs. 55–58, pl. III: fig. 150.—Faxon, 1884:138; 1885a:8, 17–19, 31, 33–34, 158, 167, 168, 173; 1885b:358; 1890:619; 1898:646; 1914:367, 412.—Underwood, 1886: 373.—Lönnberg, 1895:4.—Hay, 1899b:959, 962—Ortmann, 1902:277.—Harris, 1903a:58, 131, 138, 144, 151, 152.—Creaser, 1936:125.—Goodnight, 1941:72, 73—(?)Penn, 1941:8—Hobbs, 1942b:8, 9; 1972a:2.—Hoff, 1944:349.

Cambarus (Cambarus) versutus.—Ortmann, 1905a:101, 128.

Cambarus (Ortmannicus) versutus.—Fowler, 1912:341 [by implication].—Creaser, 1934:4 [by implication).—(?)Lyle, 1937:31, 37, 61–63; 1938:76.

Procambarus versutus.—Hobbs, 1942b: 13–15, 20, 21, 32, 92, 95, 103, 109, 118–121, 123, 126–129, 166, figs. 136–140; 1951: 272, 275; 1952a:218; 1953a: 173, 178; 1953b:412, fig. 13; 1956a: 117; 1959:884*; 1962:273, 281, 283, 286, fig. 18; 1968b:K-l1*, fig. 21g; 1969b:344, fig. 34; 1976, fig. 1c,j,k—Penn, 1946:29; 1953:1.—Villalobos, 1959:312.—Hobbs and Hart, 1959:148, 151, 159, 160, 165, 167, 169, figs. 9, 22—Black, 1968:5.—Hobbs III, 1969:22, 41, 42, 55, tab. 3—Fitzpatrick and Hobbs, 1971:95.—Holt, 1973b:99—Hart and Hart, 1974:21, 30, 61, 87, 131, 134—Bouchard, 1976a:577.

Procambarus (Pennides) versutus.—Hobbs, 1972a: 10; 1972b:67, 152*, 154*, fig. 54a; 1974b:64, fig. 267—Fitzpatrick, 1976:57,—Bouchard, 1976c: 14.

The above is believed to be a complete bibliography for the species. References to Georgia are indicated by asterisks.

SUMMARY OF LITERATURE.—Hagen (1870) described this crayfish on the basis of two lots of specimens, one from Spring Hill, Mobile County, Alabama, and the other from 10 miles east of Mobile, presumably in Baldwin County. Faxon (1884) reported its occurrence in the neighborhood of Mobile and at Cape Barrancas, Florida. In his revision of the Astacidae, Faxon (1885a) assigned it to his Group I, and, in 1890, 1898, and 1914, cited five new localities in Alabama: Escambia River at Flomaton, at Pollard (both in Escambia County), Greenville (Butler County), Calera (Shelby County), and Auburn (Lee County). In 1914, he cited the type-locality as Spring Hill, the first locality listed by Hagen. Although Ortmann correctly recognized the relationships of the crayfishes in assigning it to his subgenus Cambarus, he overlooked the fact that the type-species of the genus Cambarus (Astacus Bartonii Fabricius) had been selected by Faxon (1898:644). As pointed out by Fowler (1912), the name Cambarus had to replace Ortmann's Bartonius, thus leaving the group that Ortmann had assigned to his subgenus Cambarus without a name, and Fowler proposed Ortmannicus as a substitute.

No new data were offered subsequently until Lyle (1937, 1938) and Penn (1941) reported the occurrence of this species in Mississippi and Louisiana, respectively. There is every reason to believe that both records were based on erroneous identifications. Hobbs (1942a), in elevating Ortmann's subgenera to generic rank, transferred this crayfish to the genus Procambarus and, in his study of the crayfishes of Florida (1942b), he presented an account of its presence in Florida and cited two new county records (Conecuh and Elmore) in Alabama. In 1953 (b), he presented a spot map including all of the localities known for the species at the time. The first record of its occurrence in Georgia was noted by Hobbs (1959), and, although no locality was cited, the record was based on the only three specimens known from the State at that time. In their summary of the crayfishes of the lower Apalachicola River basin, Hobbs and Hart (1959) utilized the same data presented by Hobbs (1942b). Other references cited contain a few additional locality records, lists of crayfish associates, and statements concerning its affinities with other species.

DIAGNOSIS.—Rostrum with marginal spines and median carina. Carapace with 2 pairs of cervical spines. Areola 2.1 to 2.7 times as long as broad and constituting 23.8 to 27.7 percent of entire length of carapace (35.0 to 39.3 percent of postorbital carapace length). Antennal peduncle with spine on ischium. Ventral surface of basis and ischium of third maxilliped not densely bearded. Basis of cheliped with ventromesial spine. Mesial surface of palm of chela bearing crowded, irregularly arranged or sublinear rows of 9 or 10 tubercles. Male with hooks on ischia of third and fourth pereiopods. First pleopods asymmetrical, suddenly contracted distally, and reaching coxae of third pereiopods; cephalic surface with moderate shoulder; subapical setae abundant, many arising from distolateral surface of appendage; mesial process slender, extending caudodistally at angle of approximately 30 degrees to shaft of appendage; cephalic process also slender, similarly disposed, and situated on lateral side of central projection; central projection somewhat beaklike, corneous, exceeded distally only by cephalic process, its cephalic base situated distinctly distal to base of caudal process; caudal element consisting of (1) short, corneous, subacute caudal process, (2) prominent caudal knob, noncorneous lobe lateral to base of caudal process, and (3) adventitious process in form of caudomesial ridge somewhat rudimentary. Female with sternum cephalic to annulus ventralis bearing paired tuberculate projections extending far over ventral surface of cephalic part of annulus; first pleopod present.

COLOR NOTES (Figure 173e; based on population of Conecuh County, Alabama, Escambia River drainage).—Basic color of carapace tan to brown, with black and cream markings. Rostrum with black margins, and black median line extending from carina caudally, dividing at base of rostrum, with branches diverging over cephalic part of gastric area; orbital, antennal, and much of hepatic area black and continuing in narrow band, flanking cervical groove in posterior gastric region; cream-colored stripe extending from posterior parts of antennal and mandibular areas caudally just below cervical spines and continuing over branchiostegite almost to caudal flange; stripe flanked dorsally on branchiostegite by large right-triangular black mark, apex of triangle continuous with black marking in hepatic region and broad base lying on caudal margin of carapace, posterior acute apices of triangle almost reaching median line of carapace. (Black triangle representing horn and part of bar of saddle in postulated primitive color pattern (see Hobbs, 1958a: 74).) Tergum of first abdominal segment cream posteriorly, with reduced pleura almost white; succeeding segments tan to brown dorsally, with similar black markings arranged serially on ventrolateral part of each tergum and expanding onto cephalic part of pleuron, latter with broad cream tan band along oblique caudal margin continuous with that along posterior margin of tergum. Telson and uropods with prominent dark, often black splotches, those at base and on posterior part of telson and on uropods most conspicuous. Antennule and antenna dark brown to almost black, and antennal scale with black line on lateral margin. Cheliped cream from base through half of merus, latter with dark patch dorsally just distal to midlength and olive to tan with black markings distally; carpus and chela tan with black tubercles dorsally, fingers tipped with scarlet. Dorsal surface of remaining pereiopods mostly olive tan from merus distally, fading to cream ventrally and proximally; sternal areas cream to pinkish cream.

TYPES.—Syntypes, MCZ 190 (3II, ), USNM 4963 (I), MHNP (2 specimens), AMS (lost.)

TYPE-LOCALITY.—Spring Hill, Mobile County, Alabama (see Faxon, 1914:412).

RANGE.—From the Mobile River drainage (as far north as Choctaw, Tuscaloosa, Blount, and Tallapoosa counties) in Alabama eastward to the Chattahoochee-Apalachicola drainage in Alabama, Florida, and Georgia.

GEORGIA SPECIMENS EXAMINED.—Only five specimens from two localities are available from Georgia. Marion County: (1) Pine Knot Creek 11.4 airmi NNW of Buena Vista on St Rte 355, 1I, 1, 29 Apr 1978, Keith Floyd and Win Seyle, collectors. Muscogee County: (2) Fort Benning, 2, 1j, Jan 1951, Joseph Pollock.

VARIATIONS—Although this crayfish demonstrates considerable variation in certain features throughout its range, the Georgia specimens differ little from the syntypic first form male in the Smithsonian except as follows: a median carina is present on the rostrum; there is a lateral shoulder-like prominence at the base of the more spiniform caudal process of the pleopod of the first form male; the cephalic process extends slightly distal to the central projection; the ratio of the areola to the length of the carapace is perhaps slightly greater, but there are too few specimens available to be certain as to whether or not this is a consistent feature of the populations in the Georgia localities. The specimens from Marion County did not reach me until after this study had gone to press.

SIZE.—The largest specimen of the species available is a first form male, with a carapace length of 39.2 (postorbital carapace length, 27.4) mm. Corresponding lengths of the smallest first form male are 16.3 (11.5) mm, and those of the smallest female carrying eggs or young are 17.1 (12.3) mm.

Procambarus (Pennides) versutus (Hagen)

Cambarus versutus Hagen, 1870:51, figs. 55–58, 150.

Cambarus (Cambarus) versutus.—Ortmann, 1905c:101.

Cambarus (Ortmannicus) versutus.—Fowler, 1912:341 [by implication].—Creaser, 1934b:4 [by implication].

Procambarus versutus—Hobbs, 1942a:342 [by implication]: 1942b:126, figs. 136–140.

Procambarus (Pennides) versutus.—Hobbs, 1972a: 10.

TYPES—Syntypes, MCZ 190 (3, II, ), USNM 4963 ( I), MHNP (2 specimens), AMS (lost).

TYPES-LOCALITY.—Spring Hill, Mobile County, Alabama.

RANGE.—From western Alabama to the Apalachicola River in Florida.

HABITAT.—Streams.

Procambarus (Pennides) versutus (Hagen)

Cambarus versutus Hagen, 1870:51, figs. 55–58, 150.

Cambarus (Cambarus) versutus.—Ortmann, 1905c:101.

Cambarus (Ortmannicus) versutus.—Fowler, 1912:341 [by implication].—Creaser, 1934b:4 [by implication].

Procambarus versutus.—Hobbs, 1942a:342 [by implication]; 1942b:126, figs. 136–140; 1981:463, figs. 12f, 173e, 175, 181, 183e, 259.

Procambarus (Pennides) versutus.—Hobbs, 1972a:10; 1974b:64, fig. 267.

TYPES.—Syntypes, MCZ 190 (3 male II, female), USNM 4963 (male I), MHNP (2 specimens), AMS (no longer there).

TYPE LOCALITY.—Spring Hill, Mobile County, Alabama (see Faxon, 1914:412).

RANGE.—“From the Mobile River drainage (as far north as Choctaw, Tuscaloosa, Blount, and Tallapoosa counties) in Alabama eastward to the Chattahoochee-Apalachicola drainage in Alabama, Florida, and Georgia” (Hobbs, 1981:465).

HABITAT.—Streams.