Procambarus advena (Le Conte 1856)

Procambarus (H.) advena

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

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

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

Procambarus (Hagenides) advena (LeConte)

Astacus advena LeConte, 1856:402.

Cambarus advena.—Hagen, 1870:87, pl. 3: fig. 164; pl. 7.—Faxon, 1884:140.

Cambarus Carolinus.—Hagen, 1870:87–88 [latter in part], pl. 1: figs. 51–54; pl. 3: fig. 165.

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

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

Procambarus advena.—Hobbs, 1942a:340.

Procambarus (Hagenides) advena.—Hobbs, 1972a:7.

TYPES.—Syntypes, MCZ 3379 (), ANSP 321 ().

TYPE-LOCALITY.—“Georgia inferiore.” Here restricted to 2.5 miles west of Riceboro, Liberty County, Georgia.

RANGE.—Lower coastal plain of Georgia, between Savannah and Altamaha rivers.

HABITAT.—Burrows (primary burrower).

Procambarus (Hagenides) advena (LeConte)

Astacus advena LeConte, 1856:402.

Cambarus Carolinus—Hagen, 1870:31, 87–88 [latter in part], pl. 1: figs. 51–54, pl. 3: fig. 165.

Cambarus advena.—Hagen, 1870:87, pl. 3: fig. 164, pl. 7.—Faxon, 1884:140.

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

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

Procambarus advena.—Hobbs, 1942a:342 [in part].

Procambarus (Hagenides) advena.—Hobbs, 1972a:7 [in part]; 1974b:48 [in part].fig. 194; 1981:311, figs. 17f, 118a, 119–121, 235.

TYPES.—Syntypes, MCZ 3379 (female), ANSP 321 (female).

TYPE LOCALITY.—“Georgia inferiore.” Restricted to 2.5 miles (4 km) west of Riceboro, Liberty County, Georgia, by Hobbs, 1974b:48.

RANGE.—Lower coastal plain of Georgia, between Savannah and Altamaha rivers.

HABITAT.—Burrows (primary burrower).

Procambarus (Hagenides) advena (LeConte)

Astacus advena LeConte, 1856:402.—Hagen, 1870:9, 10.—Faxon, 1885a:12, 54.—Hobbs, 1972a:2, 7; 1974a:15.

Cambarus advena.—Hagen, 1870:31, 53, 86 [name only], 87, 98 [in part, Georgia only], 100 [in part], 107, pl. VII [ instead of , form I]; not 86 [description = P. (G.) hagenianus], 101, 105, 106, pl. I: figs. 90–92 [figures, P. (G.) hagenianus].—Brocchi, 1875:27.—Faxon, 1884:113, 140; 1885a:8, 9, 17, 47–49, 54–56, 58, 158, 167, 173, 178; 1885b:358; 1914:412.—Underwood, 1886:366, 368.—Hay, 1899b:959, 961.—Ortmann, 1902:277, 279; 1905c: 403; 1905d:438.—Harris, 1903a:58, 68, 129, 150, 152.—Graeter, 1909:470.—Spandl, 1926:96.—Hobbs, 1938:65; 1940a:389, 393; 1942a:335; 1972a:2.—Hobbs and Villalobos, 1964:321.

Cambarus Carolinus.—Hagen, 1870:32, 74, 75, 87 [description only], 88 [in part, excluding specimens from Alabama and South Carolina], pl. III: fig. 165 (?).

Cambarus (Cambarus) advena.—Ortmann, 1905a:98, 100, 101, 104 [in part, by implication].

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

Procambarus advena.—Hobbs, 1942a:340; 1942b:73–76, 84, 86, 92 [all in part]; 1954:110, 116; 1959:887 [in part], fig. 31.26; 1968b:K-8 [in part]; 1969a:118.—Villalobos, 1955:38.—Hart and Hart, 1974:21 [in part], 28 [in part].

Procambarus (Hagenides) advena.—Hobbs, 1972a: 7, figs. 2e, 8a–g; 1972b:50, 150 [in part], 154 [in part], figs. 5g, 41d; 1974b:48, fig. 194.—Hobbs and Bouchard, 1973:52 [by implication], 63.

This synonomy is believed to represent a complete bibliography for the species, and inasmuch as this crayfish is endemic in Georgia all references concern the state.

SUMMARY OF LITERATURE.—LeConte's (1856) description of Astacus advena, together with Hagen's (1870) concept of the species, left no doubt in the mind of the latter as to its distinctive features, and, following Girard (1852) in assigning the American species to the genus Cambarus, he employed the combination Cambarus advena. As pointed out by Faxon (1884, 1885a), unfortunately a transposition of the descriptions and most of the illustrations of his Cambarus advena and C. Carolinus (Erichson, 1846:96) occurred. Furthermore, in describing the first form male of the former, he chose a specimen purportedly (although highly unlikely—see Fitzpatrick, 1978b: 59) from Charleston, South Carolina, which was later designated as the “type” of Cambarus hagenianus (Faxon, 1884:141; 1914:366). It seems clear that when Hagen prepared his account of C. advena and C. carolinus, he had obviously misidentified specimens of the former as C. carolinus, and representatives of the species later described as C. hagenianus as C. advena, inadvertently transposing the Latin descriptions and all of his illustrations except that on plate VII. The synonomy cited above reflects this interpretation.

In addition to the female type in the Philadelphia Academy of Natural Sciences (there is no indication that he saw the female syntype that is now in the Museum of Comparative Zoology), the specimens available to Hagen consisted of

[1] a first form male (M.C.Z. No. 232) from Charleston, South Carolina…[2] No. 3368, dry female from Georgia, L. Agassiz…[3] No. 3367…a young female, also from Georgia…[4] No. 230, young female specimens from Mobile, Ala…[5] No. 275, a very young male from the same locality…(Faxon, 1884:141), [6] No. 282, Georgia, Dr. Jones. Male. Fem. Spec. 6…[and 7] No. 1850, Georgia. Male. Spec. 1 (Hagen, 1870:87, 88).

Of these, the first is the type of Procambarus (Girardiella) hagenianus; the second and third are in all probability members of P. (H.) advena; the fourth and fifth are clearly members of the genus Cambarus; the sixth includes representatives of two species, P. (H.) talpoides and P. (H.) pygmaeus; and the seventh is apparently no longer extant. Thus, as indicated by Faxon (1885a), P. (H.) advena was known to occur only in Georgia. Uncertainty surrounds the source of the specimens from which Hagen's figures 51–54 were illustrated; however, I suspect that they were from the first form male of Procambarus (H.) pygmaeus included in the sixth lot just mentioned, “No. 282, Georgia….”

All of the references from 1885 until 1942 are based on Faxon's identifications of the material that was available to Hagen and add nothing except notes on the affinities of P. (H.) advena to other species. No new localities were cited for it until Hobbs (1942b:77) recorded its presence in Bryan County, Georgia (although the species occurs in the county, this report was based on the misidentification of specimens of P. (H.) pygmaeus; all other locality records cited by him for P. advena are referable to P. (H.) talpoides). Subsequent to that time, the only reliable records that have been reported are the following based on collections and identifications made by me: 7.6 miles southwest of Midway on State Route 38, Liberty County, Georgia (Hart and Hart, 1974:21), and 16.9 miles north of Darien on U.S. Highway 17, McIntosh County, Georgia (Hart and Hart, 1974:28), where it served as host to the entocytherid ostracods, Ankylocythere ancyla Crawford, A. hobbsi (Hoff, 1944), and Entocythere elliptica.

The several definitions of the range of the species offered by Hobbs (1942–1972) that included that part of Georgia south of the Altamaha River and northern Florida encompass that of P. (H.) talpoides, which is described herein. Not until 1974 did he note a range consistent with that recorded below.

The only record of the habitat of this species is that of LeConte (1856), who stated, “Hymeme vitam degit subterraneam. Aestate in fossis invenitur.” This statement led Graeter (1909) to conclude that this crayfish is an inhabitant of cave waters, an assumption that was negated by Spandl (1926).

DIAGNOSIS.—Rostrum without marginal spines or tubercles. Carapace lacking cervical spines. Areola obliterated to 19.8 times as long as broad and constituting 35.6 to 39.7 (average 38.0) percent of entire length of carapace (40.5 to 45.3, average 43.6, percent of postorbital carapace length). Ventral surface of basis and ischium of third maxilliped not densely bearded. Male with mesial surface of palm of chela lacking beard but bearing cristiform row of 7 to 9 tubercles; hooks on ischia of third pereiopods only; coxa of fourth pereiopod with caudomesial boss. First pleopods of first form male asymmetrical, subcontiguous, and reaching coxae of second pereiopods, cephalic surface without prominent shoulder; preapical setae limited to few on cephalodistal border; mesial process tapering to acute apex and directed caudodistally; cephalic process consisting of short, straight, rounded to subacute corneous blade at cephalic base of prominent, corneous, subtriangular central projection, latter arising from distal surface of appendage and directed caudodistally; caudal knob somewhat inflated but falling far short of central projection. Female with chela as in male; annulus ventralis subovate in outline, longer than broad, and with cephalolateral elevations frequently, if not usually, tuberculate; first pleopod present.

MALE, FORM I.—(From restricted type-locality, Liberty County, Georgia.) Body (Figure 119a,i) subovate, compressed laterally. Abdomen narrower than thorax (11.2 and 15.2 mm). Width of carapace greater than depth at caudodorsal margin of cervical groove (15.2 and 13.6 mm). Areola 29 times as long as broad, with 1 punctation in narrowest part. Cephalic section of carapace 1.6 times as long as areola, length of latter 38.6 percent of total length of carapace (44.1 percent of postorbital carapace length). Rostrum excavate dorsally, with rather slender convergent margins lacking spines or tubercles; upper surface with few punctations between submarginal rows; acumen not clearly defined basally and reaching anteriorly to base of ultimate podomere of antennular peduncle; subrostral ridges moderately well developed and evident in dorsal view to base of acumen. Postorbital ridges rather strong, grooved dorsolaterally, and cephalic extremity subangular but lacking spines or tubercles. Suborbital angle absent. Branchiostegal spine very small. Carapace punctate dorsally and weakly tuberculate laterally except in hepatic, mandibular, and anteroventral branchiostegal regions, in them tubercles more conspicuous; cervical spine absent. Abdomen shorter than carapace (28.3 and 30.3 mm); pleura broadly rounded and subtruncate ventrally. Cephalic section of telson with 2 spines in caudolateral corner; caudal section rounded and narrow. Uropod with short acute prominence on both lobes of proximal podomere; mesial ramus with conspicuous distolateral spine and less prominent premarginal tubercle on median ridge; lateral ramus with usual row of spines flanking proximal side of transverse suture, lateralmost spine no larger than others in row. Cephalic lobe of epistome (Figure 119h) subpentagonal, with small cephalomedian projection; ventral surface weakly convex and margins slightly thickened and elevated ventrally; fovea indistinct, replaced by shallow median depression, and epistomal zygoma strongly arched. Antennular peduncle with strong acute tubercle near midlength of ventral surface of basal podomere. Antenna almost reaching caudal margin of carapace, peduncle without spines; diameter of renal tubercle greater than half maximum width of coxal podomere bearing it. Antennal scale (Figure 119d) about 2.4 times as long as broad, greatest width distal to midlength, lamellar portion about 1.6 times as wide as thickened lateral part, latter terminating in strong spine. Third maxilliped with basis bearing conspicuous long, stiff, simple setae; ischium with distolateral tubercle, ventral surface with submarginal lateral row of short plumose setae, few clusters of similar setae in mostly naked proximolateral half, and 2 irregular rows of long stiff setae lateral to crista dentata.

Right chela (Figure 119k) subovate in cross section, strongly depressed. Mesial surface of palm with cristiform row of 9 (7 on left) tubercles subtended dorsolaterally by row of 5 squamous tubercles and similar ones scattered over dorsal surface and lateral margin of palm; ventral surface also with scattered squamous tubercles and punctations (latter replacing tubercles proximally and laterally), and strong tubercle opposite base of dactyl. Fixed finger with conspicuous median longitudinal ridge dorsally and ventrally, ventral one flanked by setiferous punctations, and dorsal ridge by squamous tubercles proximally and punctations along most of distal three-fourths; opposable margin of finger with row of 5 tubercles, third from base largest, along proximal two-thirds and tubercle present on lower level at base of distal fourth of finger; minute denticles, arranged in single row except proximal to second tubercle from base, extending from base of finger to corneous tip. Dactyl with dorsal and ventral surfaces similar to those of fixed finger although ridges less prominent; mesial surface with row of tubercles, decreasing in size distally, along proximal third of finger, giving way to punctations; opposable surface with row of 9 (left with 7, perhaps due to injury) tubercles, first and third from base larger than others, otherwise decreasing in size distally; single row of minute denticles extending between tubercles (beginning at third tubercle from base) and continuing to corneous tip of finger.

Carpus of right cheliped punctate dorsally, laterally, and ventrally, and tuberculate mesially; dorsal surface with shallow oblique furrow; dorsomesial margin with row of 7 tubercles; mesial surface with cluster of 7; and ventral surface with 2 on distal margin: 1 on ventrolateral articular knob and other on ventromesial angle.

Merus with dorsal surface bearing slightly irregular subserrate row of 15 (11 on left) tubercles, more distal member of which larger than more proximal ones; mesial and lateral surfaces sparsely punctate; and ventral surface with mesial row of 14 tubercles, and lateral one of 13 (left with 12 and 11, respectively), single tubercle between subdistal ends of rows. Ischium with row of 3 tubercles ventromesially.

Hook on ischium of third pereiopod (Figure 119j) only; hook inflated and extending proximally over distal part of corresponding basis. Coxa of fourth pereiopod with moderately strong boss caudomesially; that on fifth small and inconspicuous.

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

First pleopods (Figure 119b,g,l) as described in “Diagnosis.”

FEMALE.—(Liberty County, Georgia.) As in male, form I, except for secondary sexual characteristics and following: rostrum reaching midlength of ultimate podomere of antennular peduncle; ventrolateral part of branchiostegites more strongly tuberculate; lateral half of ventral surface of third maxilliped with short plumose setae more abundant; mesial margin of palm of chelae with row of 9 tubercles on right and 8 on left; opposable margin of dactyl of chela with row of 6 tubercles; carpus with cluster of 10 tubercles on mesial surface of right cheliped and 9 on left; merus with dorsal row of 12 and 15 tubercles, respectively, and ventrolateral row of 10; ischium with only 2 tubercles on ventromesial margin. (See “Measurements.”)

Annulus ventralis (Figure 119e) slightly longer than broad, subovate, with broad, deep median depression (occupying three-fourth of its length) flanked by high, caudally diverging, anterolateral ridges, left ridge weakly tuberculate; caudal part of annulus forming somewhat flattened shelf; sinus originating dextrally in caudal extremity of depression and, following S-shaped course over shelf, ending on median line anterior to caudal margin of annulus. Postannular sclerite unadorned, strongly arched anteriorly and truncate posteriorly, its width approximately two-thirds and its length little less than half that of annulus.

MALE, FORM II.—(Liberty County, Georgia.) Differing from first form male in only few minor respects: mesial margin of palm of chela with row of 8 tubercles; opposable margin of dactyl with 6; mesial surface of carpus with 10 tubercles in cluster; dorsal surface of merus with row of about 12, ventrolateral row of 7 on right and 6 on left, and ventromesial row of 10 and 13, respectively; ischium bearing row of 4 tubercles. Hook on ischium of third pereiopod greatly reduced, its free end not nearly reaching basioischial articulation, and bosses on coxae of both fourth and fifth pereiopods rudimentary. (See “Measurements.”) First pleopod (Figure 119c,f) with mesial process and central projection markedly similar in shape to those of first form male, but cephalic process not evident although cephalodistal part of appendage produced in broad prominent excrescence; juvenile suture on basal part of shaft clearly defined.

COLOR NOTES (Figure 118a).—Carapace almost uniformly mauve brown or olive brown, with small darker brown spots and flecks, fading ventrally to bluish gray, caudal flange dark blue. Abdomen mauve to brown with transverse darker brown bands (completely covering first tergum) on caudal fourth of second through fifth terga; pleura also brown without conspicuous markings although margins darker than central area of each. Sixth tergum with posteromedian dark spot. Telson with brown triangular area basally, flanked by paired, small, oblique cream spots abutting caudolateral margins of triangle; more caudal part becoming pale greenish blue matching uropods; ridges and articular areas of both telson and uropods dark blue. Antennular and antennal peduncles bluish gray, with darker lateral margins and bands at bases of articulations; flagella gray to pale tan. Chelipeds distal to ischium deep lavender to bluish gray dorsally, with most tubercles gray to black, fingers usually more brightly colored in shades of blue or lavender; major tubercles on carpus and propodus often pale lavender or blue and nearly always tipped with cream; ventral and ventrolateral surfaces of cheliped ranging from cream proximally to orange cream on fingers; distal extremities of fingers orange to yellowish tan. Remaining pereiopods with basal 2 podomeres cream, more distal ones bluish to lavender gray; color intensifying adjacent to articulations, and dactyls of fourth and fifth pereiopods distinctly blue or lavender. Ventral surface of body cream.

Measurements (mm)

Male, form I Female Male, form 11

Carapace

Height 13.6 14.0 10.5

Width 15.2 14.3 10.7

Entire length 30.3 29.7 23.0

Postorbital length 26.5 25.7 20.0