Procambarus barbatus (Faxon 1890)

Procambarus (L.) barbatus

E Ankylocythere ancyla: Liberty (Hart and Hart, 1974:21)

Ankylocythere hobbsi: McIntosh (Hart and Hart, 1974:28)

Entocythere elliptica: Liberty, McIntosh (Hart and Hart, 1974:88)

Procambarus (Leconticambarus) barbatus (Faxon)

Astacus penicillatus LeConte, 1856:401 [not Astacus penicillatus Olivier, 1791].

Cambarus penicillatus.—Hagen, 1870:16, 53, figs. 93, 94, 149 [not 95 and 96, which are P. (O.) ancylus].

Cambarus barbatus Faxon, 1890:621 [substitute name; in part].

Cambarus (Cambarus) barbatus.—Ortmann, 1905c:102.

Cambarus (Ortmannicus) barbatus.—Fowler, 1912:341 [by implication].

Procambarus barbatus.—Hobbs, 1942a:340; 1942b:39, figs. 1–5.

Procambarus (Leconticambarus) barbatus.—Hobbs, 1972a:8, figs. 1m, 2f, 9a–g; 1974b:50, fig. 204; 1981:342, figs. 18b, 130a, b, 131, 132, 240.

TYPE.—MCZ 279 (male I), designated neotype of A. penicillatus LeConte by Hobbs, 1974b:50. Specimens from Flomaton, Alabama, cited by Faxon (1890) and listed as paratypes by him (1914:414) are paratypes of Procambarus (L.) escambiensis Hobbs.

TYPE LOCALITY.—“Habitat in fossis Georgiae inferioris” (LeConte, 1856:401). Restricted by Hobbs (1974b:50) to 2.5 miles (4 km) west of Riceboro, Liberty County, Georgia, in pine flatwoods.

RANGE.—In the coastal plain from the Altamaha River in Georgia to the Edisto River in South Carolina.

HABITAT.—Lentic situations and burrows (secondary burrower).

Procambarus (Leconticambarus) barbatus (Faxon)

Astacus penicillatus LeConte, 1856:401*.—Hagen, 1870:16.—Faxon, 1890:621; 1914:414*.—Hobbs, 1972a:2*; 1974b:50 [not Astacus penicillatus, Olivier, 1791].

Cambarus penicillatus.—Hagen, 1870:31 (?), 53, 54* [in part], 55 (?), 97* [in part], 107* [in part], pl I: figs. 93, 94 [not figs. 95, 96 = P. (O.) ancylus Hobbs, 1958b:164].—Faxon, 1884:138*[in part: not Mississippi and South Carolina]; 1885a:36–38* [in part], 158*, 167* [in part], 173* [in part], [not 168].—Underwood, 1886:371* [in part].—Ortmann, 1902:277.—Harris, 1903a:72, 118.—Hobbs, 1942b:39, 40*; 1972a:2.

Cambarus barbatus Faxon, 1890:621 [in part: not specimens from Escambia River (= P. (L.) escambiensis Hobbs, 1942b:46)]; 1914:367, 370, 414*.—Hay, 1899b:959*, 964.—Harris, 1903a:58* [in part], 72* [in part], 118, 137 [in part], 138, 150 [in part], 152*, 154 (?).—Wolf, 1934:104 [name only; probably error for Cambarus (Erebicambarus) laevis Faxon (1914:391) in Indiana and for C. (E.) tenebrosus Hay (1902b:232) in Kentucky].—Hobbs, 1940a:389, 410, 414, 418; 1940b:3; 1942c: pl. II: figs. 2, 3; 1962:273; 1972a:2; 1974b:50.

Cambarus (Cambarus) barbatus.—Ortmann, 1905a:102, 105*; 1906a:18.

Cambarus (Ortmannicus) barbatus.—Fowler, 1912:341 [by implication].

(?) Astacus barbatus.—Wolf, 1934:104 [name only].

Procambarus barbatus.—Hobbs, 1942a:340, figs. 6, 9; 1942b:34*, 35–42*, 44*, 48, 54, 70, 108, 109, pl. II: figs. 1–5; 1943b:203; 1959:887*, fig. 31.28; 1962:273 [by implication]; 1966b:70; 1968b:K–8*, fig. 17a; 1974a:15—Hoff, 1944:349, 356*—Pennak, 1953, fig. 288c.—Hart, 1959:203.—Hart and Hart, 1974:21*, 28*, 63, 88*, 137.—Wharton, 1978:46*.

Procambarus (Leconticambarus) barbatus.—Hobbs, 1972a:8, figs. 1m, 2f, 9a–g; 1972b:53*, 150*, 154*, figs. 25b, 26f, 29c, 43h, 44f; 1974b:50,* fig. 204.—Hart and Hart, 1974:21*, 28*, 63, 88*, 137.—Hobbs III, Thorp, and Anderson, 1976:3, 5, 13, 32*–33, fig. 13.

The list of references cited above is believed to be a complete bibliography for the species.. Those citations noting the occurrence of this crayfish in Georgia are indicated by asterisks.

SUMMARY OF LITERATURE.—Perhaps the nomenclature of no other American crayfish has had as much confusion involved in its early history. Not realizing that the combination Astacus penicillatus had been used by Olivier in 1791 for a spiny lobster, LeConte (1856) applied it to a new crayfish from Georgia, the whereabouts of the types of which is unknown. In his monograph of the Astacidae, Hagen (1870:54), who did not have the opportunity to examine LeConte's types, applied the name Cambarus penicillatus to three lots of specimens that later were found to consist of representatives of three species: lot 250 from South Carolina contains members of Procambarus (Ortmannicus) ancylus; lot 254, representatives of P. (Scapulicambarus) troglodytes; and only lot 279 contains a specimen that is believed to be a member of LeConte's A. penicillatus. Faxon (1884:138) pointed out the error in Hagen's identification of the specimens from South Carolina and questioned his determination of lot number 250. Further, he tentatively identified a second form male from eastern Mississippi as belonging to this species (the specimen is apparently lost but probably was a member of Procambarus (O.) hybus Hobbs and Walton, 1957:39). Faxon's (1885a) references to the species consist of an expanded discussion of the materials mentioned by him in 1884, comparing the specimen from eastern Mississippi with those from Georgia and South Carolina. In 1890, Faxon (p. 621) pointed out that LeConte's Astacus penicillatus must be rejected because of the priority established by Olivier, and he proposed the substitute name Cambarus barbatus. Unfortunately, he assigned 11 specimens from the “Escambia River at Flomaton,” on the Alabama-Florida line, to the species, citing them in 1914 (p. 414) as paratypes of his C. barbatus. Some years later, Hobbs (1942b:48) referred Faxon's paratypes to a new species Procambarus escambiensis (Hobbs, 1942b:46). Thus the only extant specimen of the species was Faxon's “type” of C. barbatus, which was the “young” male mentioned by Hagen (MCZ 279). In order to establish the synonomy of Faxon's Cambarus barbatus with LeConte's Astacus penicillatus, Hobbs (1974b:50) designated this specimen as the neotype of LeConte's species. He (1974b:50) also restricted the rather vague type-locality implied by LeConte. The latter (1856:400) stated that of his new species “two of them [were] from the upper part of the State of Georgia, the rest from the low country.” Astacus spiculifer and A. latimanus are noted as occurring in “Georgia superiore,” thus the source of his specimens of the remaining species must be assumed to have been in “Georgia inferiore.”

In his study of the genus Cambarus, Ortmann (1905a, 1906a) proposed several subgenera, introducing the combination Cambarus (Cambarus) barbatus. His error in assigning this species to the nominate subgenus was pointed out by Fowler (1912:341), and, in correcting Ortmann's mistake, implied that the name should be Cambarus (Ortmannicus) barbatus. No subsequent authors have used either combination, perhaps because Faxon (1914), in his checklist of the crayfishes of the family Astacidae, did not recognize Ortmann's subgenera. In the generic revision proposed by Hobbs (1942a), the combination Procambarus barbatus was introduced (p. 340) and was later modified by him (1972a:8) when he assigned the species to his subgenus Leconticambarus.

Until 1940, no reliable specific locality had been cited for the species; all previous references pertain to the specimens just discussed. In describing Cambarus lunzi (= P. (O.) lunzi), Hobbs (1940b:3) stated that P. (L.) barbatus had been dug from burrows in a roadside ditch 1.4 miles southeast of Early Branch on State Highway 28, Hampton County, South Carolina, nearby those of C. lunzi. Although no specific localities were cited, Hobbs (1942b) reported its occurrence in Bulloch, Effingham, Jenkins, Liberty, Long, and Screven counties, Georgia, and referred to the record from South Carolina given by him in 1940. Only two additional specific localities have been recorded, these by Hart and Hart (1974): one each in Liberty (pp. 21, 88) and McIntosh (p. 28) counties, where it served as host to Ankylocythere ancyla and Entocythere dorsorotunda Hoff (1944:332), and to Ankylocythere hobbsi, respectively.

Almost all references subsequent to 1940 have to do with relationships of this to other crayfishes or to its range. No data concerning aspects of its biology were recorded until Hobbs III, Thorp, and Anderson (1976:32–33) presented data that were extracted from the manuscript of this report. No adequate description of the species has been presented. LeConte's account (in Latin) is so general that virtually all of it may be applied to any one of a number of species. Had he not stated that “mas latere interiore chelarum pilositatem densam habet spongiam referente,” it is doubtful that in the absence of the type it could be recognized. For this reason, a description of topotypes is offered here.

DIAGNOSIS.—Rostrum without marginal spines or tubercles. Carapace lacking cervical spine, sometimes with small tubercle. Areola 4.0 to 9.1 times as long as broad and constituting 31.9 to 35.7 percent of entire length of carapace (38.6 to 45.0 percent of postorbital carapace length). Ventral surface of basis and ischium of third maxilliped densely bearded. First form male with palm of chela almost always bearded; hooks on ischia of third and fourth pereiopods, those on third frequently much smaller than those on fourth; coxa of fourth pereiopod with prominent caudomesial boss; first pleopods slightly asymmetrical, subcontiguous, bearing proximomesial spur and prominent proximomedian lobe, reaching coxae of third pereiopods, subapical setae present on cephalically sloping cephalodistal region; terminal elements consisting of (1) subspiculiform mesial process curved cephalodistally and somewhat laterally and far overreaching other terminal elements, (2) short, distally directed cephalic process arising from mesial surface of appendage and extending distally only slightly beyond tip of central projection, (3) caudal element comprising broad, subtruncate caudal knob, appearing broadly rounded distally in caudal aspect, and (4) small subtriangular cephalodistally inclined corneous central projection situated immediately cephalic to caudal knob. Female with annulus ventralis bearing very prominent, subconical, cephalolaterally projecting prominences, resulting in greatest width of annulus occurring distinctly cephalic to midlength; postannular sclerite not markedly smaller than annulus ventralis; first pleopod present.

TOPOTYPIC MALE, FORM I.—Body (Figure 131a, i) subovate, compressed laterally. Abdomen narrower than thorax (11.1 and 11.6 mm). Width of carapace less than height (11.6 and 12.4 mm) at caudodorsal margin of cervical groove. Areola 6.5 times as long as wide, with 2 punctations across narrowest part. Cephalic section of carapace almost 1.9 times as long as areola, latter occupying 35.0 percent of entire length of carapace and 42.7 percent of postorbital carapace length. Rostrum excavate dorsally with weakly thickened convergent margins, lacking marginal spines or tubercles, with usual submarginal row of setiferous punctations, and with moderately deep punctations in caudal part; acumen not delimited basally from remainder of rostrum. Subrostral ridge moderately strong and evident in dorsal aspect along basal third of rostrum. Postorbital ridge well developed, grooved dorsolaterally, and lacking spine or tubercle. Suborbital angle distinct. Branchiostegal spine clearly defined. Carapace punctate dorsally and granulate laterally, granules in hepatic and anteroventral branchiostegal region tuberculiform, those in area usually occupied by cervical spine no larger than others nearby. Abdomen longer than carapace (25.6 and 24.3 mm); pleura truncate ventrally. Cephalic section of telson with 2 spines in each caudolateral corner. Uropod with both lobes of proximal podomere bearing spine; mesial ramus with distolateral and median preapical spines, latter far removed from distal margin; lateral ramus with row of short spines immediately proximal to transverse suture, that immediately adjacent to lateralmost member rather long. Cephalic lobe of epistome (Figure 131h) subovate and bearing small cephalomedian projection; posteromedian part of surface slightly elevated, margins not thickened, and fovea not clearly defined, area of usual occurrence shallowly excavate. Antennular peduncle with very small spine on midventral surface near midlength of basal podomere, mesial border of proximal podomere not strongly setose. Antennae extending caudally to third abdominal tergum, peduncle with small tubercle on basis and ischium. Antennal scale (Figure 131g) about twice as long as broad, greatest width distal to midlength, lamellar portion about 1.7 times as broad as thickened lateral portion, latter terminating in short spine. Third maxilliped with ventral surface of proximal podomeres through merus and peduncle of exopod conspicuously hirsute, bearing clusters of plumose setae.

Right chela (Figure 131l) moderately short, somewhat depressed, and subovate in cross section. Mesial surface of palm with prominent brushes of long plumose setae obscuring mesial rows of tubercles (mesialmost consisting of 8 or 9); lateral surface with row of squamous tubercles, and dorsal and ventral surfaces similarly tuberculate, tubercles extending onto basal parts of both fingers, ventral surface with prominent tubercle distolateral to articular condyle at base of dactyl. Fixed finger with rounded longitudinal ridge dorsally and ventrally, dorsal ridge flanked proximally by squamous tubercles replaced more distally by setiferous punctations, lateral surface weakly costate, and opposable margin with row of 6 tubercles along proximal three-fourths, second from base largest, and large tubercle projecting from lower level at base of distal fourth; clusters of minute denticles present along entire ventromesial margin except on corneous tip. Dactyl with dorsal and ventral surfaces similar to those of fixed finger; mesial surface bearing proximal subserrate row of 4 tubercles, decreasing in size distally, along proximal third of finger; opposable margin with row of 5 low, rounded tubercles along proximal third, and row of minute denticles between tubercles and continuing distally to corneous tip of finger.

Carpus of right cheliped longer than broad (7.0 and 5.1 mm), with mesial, dorsomesial, and ventromesial surfaces tuberculate, otherwise punctate; dorsal surface with shallow oblique depression; mesial surface with 1 tubercle larger than others; ventral surface with large tubercle at base of condyle on distolateral angle and row of 3 others mesially on distal margin.

Merus of right cheliped tuberculate dorsally and ventrally, those tubercles on dorsal surface increasing in size distally, with 2 preapical ones larger than others; mesial and lateral surfaces mostly punctate; ventral surface with mesial and lateral rows of 13 and 12 tubercles, respectively, distal members of lateral row forming arc bent distinctly mesially toward distalmost tubercle in mesial row. Ischium with 2 rows of tubercles, 6 ventromesially and 4 dorsolaterally.

Hook on ischium of third and fourth pereiopods (Figure 131k), that on third much smaller than that on fourth and not overreaching basioischial articulation, latter hook distinctly overreaching corresponding articulation. Coxa of fourth pereiopod with very prominent, often subacute, caudomesial boss; boss on fifth much less conspicuous but well developed, thin, and rounded distally.

Sternum between third, fourth, and fifth pereiopods deep and bearing prominent fringe of plumose setae on ventrolateral margins.

First pleopods (Figure 131 b,f,j) as described in “Diagnosis.”

TOPOTYPIC FEMALE.—Differing from first form male, other than in secondary sexual characters, as follows: areola with 3 punctations across narrowest part; merus of third maxilliped devoid of plumose setae and those on ischium not so prominent; mesial surface of palm of chela (Figure 131m) without conspicuous brush of plumose setae obscuring irregular rows of tubercles, mesialmost row consisting of 8 or 9; opposable margins of fixed finger and dactyl with rows of 8 and 11 tubercles, respectively, and minute denticles arranged in single row interrupted by tubercles. (See “Measurements.”)

Annulus ventralis (Figure 131d) moderately deeply embedded in sternum, comparatively small, its median length three-fifths maximum width, and, disregarding prominent cephalolateral projections, only slightly broader than postannular sclerite, taking projections into account about 1.5 times as broad; cephalic area studded with conical projections flanking broad, shallow median longitudinal trough; caudal part subplane and bearing tilted S-shaped sinus. Postannular sclerite, about two-thirds as wide as long, rather broadly rounded anteriorly, and with ventral surface bearing arc of small tubercles. First pleopod extending over posterior third of annulus when abdomen flexed.

MALE, FORM II (From McIntosh County, 4.9 miles SE of Eulonia).—Differing from first form male in following respects: rostrum slightly more acuminate; areola with 3 punctations across narrowest part; tubercle on ventral surface of ischium of antennular peduncle spiniform; opposable margins of fixed finger and dactyl of chela with rows of 8 and 10 tubercles, respectively; ventral surface of merus of cheliped with mesial and lateral rows consisting of 12 and 13, respectively; ischium of cheliped with 5 tubercles in both dorsolateral and ventromesial rows; hooks on ischia of third pereiopods reduced to small tubercles, neither overreaching basioischial articulation; boss on coxa of fourth pereiopod decidedly reduced, that on fifth not markedly so. (Also see “Measurements.”)

First pleopod (Figure 131c,e) with subapical setae reduced in length; mesial process more robust and shorter but distinctly overreaching other terminal elements; caudal process not markedly smaller, but cephalic process and central projection rather poorly defined and neither protruding as in first form male.

COLOR

Procambarus (Leconticambarus) barbatus (Faxon)

Astacus penicillatus LeConte (not Astacus penicillatus Olivier, 1791), 1856:401.

Cambarus penicillatus.—Hagen, 1870:16, 53, figs. 93, 94 (not 95 and 96, which are P. ancylus), 149.

Cambarus barbatus Faxon, 1890:621 [in part].

Cambarus (Cambarus) barbatus.—Ortmann, 1905c: 102.

Cambarus (Ortmannicus) barbatus.—Fowler, 1912:341 [by implication].

Procambarus barbatus.—Hobbs, 1942a:342 [by implication]; 1942b:39, figs. 1–5.

Procambarus (Leconticambarus) barbatus.—Hobbs, 1972a:8, figs. 1m, 2f, 9a–g.

TYPE.—Neotype of A. penicillatus LeConte, MCZ 279 ( I), here designated. Specimens from the Escambia River at Flomaton cited by Faxon (1890) and listed as paratypes by him (1914:414) are paratypes of Procambarus escambiensis Hobbs.

TYPE-LOCALITY.—“Habitat in fossis Georgiae inferioris” (LeConte, 1856:401). Here restricted to 2.5 miles west of Riceboro, Liberty County, Georgia, in pine flatwoods.

RANGE.—In the coastal plain, from north of the Altamaha River in Georgia to the Edisto River in South Carolina.

HABITAT.—Burrows (secondary burrower).