Cambarus longirostris Faxon 1885

Cambarus (Hiaticambarus) longirostris Faxon

Cambarus Bartonii, var. longirostris Faxon, 1885a:64.

Cambarus bartonii longirostris.—Faxon, 1890:623.

Cambarus bartonii spinirostris.—Faxon, 1890:623 [lapsus for C. b. longirostris].

Cambarus longulus longirostris.—Hay, 1899b:959.—James. 1966:3. 12 [not references to Georgia], 13, 17, 21, 22 [in part], figs. 2a, 2b, pl. lc,f,g,n,o,r,s.—Holt, 1968b:32*.—Bouchard. 1976a:574.

Cambarus (Bartonius) bartoni longirostris.—Ortmann, 1905a:135 [by implication].

Cambarus (Cambarus) bartonii longirostris.—Fowler, 1912:341 [by implication].

Cambarus (Cambarus) longulus longirostris.—Ortmann, 1931:121, ? 123*.—Bouchard, 1976a:572 [in part].

Cambarus longerosilis.—Brimley, 1938:503 [erroneous spelling].

Cambarus baritonii var. longirostris.—James, 1966:2.

Cambarus longirostris.—James, 1966:2.—Hobbs, 1968b:K-17*[in part].—Anonymous, 1970c:35*; 1973b:66*, 67*.—Bouchard. 1976a:574 [in part]; 1976b:588, 592 [in part].—Wharton, 1978:220*.

Cambarus (Hiaticambarus) longirostris.—Hobbs, 1969a:141*, 142* [in part], figs. 10*. 18m; 1972b: 120*, 146*, 154* [in part], figs. 104b, 105a; 1974b:16* [in part], fig. 48.—Bouchard, 1976a:574, 575 [in part].—Bouchard and Hobbs, 1976:13.

Cambarus girardianus.—Bouchard, 1976a:572–575 [in part].

Cambarus (Hiaticambarus) girardianus.—Bouchard, 1976a:572 [in part]; 1976b:588, 592* [in part].

The above synonomy is believed to include all of the synonyms and misidentifications of specimens from Georgia but does not include all records or erroneous determinations from elsewhere in the range of the species. Those citations marked by an asterisk include references to Georgia or to specimens from the state.

SUMMARY OF LITERATURE PERTAINING TO GEORGIA.—The earliest reference to the occurrence of this crayfish in the state was that of Ortmann (1931:123). His specimens, however, are believed to have been members of C. (H.) girardianus, for subsequent collections of representatives of the subgenus Hiaticambarus in South Chickamauga Creek in the vicinity of Ringgold contain only members of that species. James' (1966:12) record is based on a misidentification of C. (H.) manningi. The first notice of the presence of C. (H.) longrirostris in Georgia that I have been able to confirm is that of Holt (1968b), who cited it as a host of an unidentified branchiobdellid worm belonging to the genus Pterodrilus collected in Union County. Two additional localities were included in faunistic surveys (Anonymous, 1970c and 1973b), one each in Dade and Union counties. Specimens on which these three localities are based are included among the “Georgia Specimens Examined,” all of which are deposited in the National Museum of Natural History, Smithsonian Institution. The reference to the association of this crayfish with C. (J.) nodosus by Bouchard and Hobbs (1976) is based on specimens obtained at the second locality listed under “Georgia Specimens Examined.”

The most comprehensive work cited is that of James (1966). All other references include synonyms, misidentifications, or statements concerning the range of the species. (See “Remarks” under subgenus Hiaticambarus.)

DIAGNOSIS.—Rostrum strongly tapering and lacking marginal spines or tubercles. Postorbital ridge terminating cephalically with or without small tubercle. Cervical spine absent, usually not even represented by low tubercle. Areola 3.1 to 4.7 times as long as broad and constituting 34.6 to 38.2 percent of entire length of carapace (41.5 to 44.7 percent of postorbital carapace length). Chela with row of tubercles on mesial margin of palm so strongly adpressed (or lacking) that not evident in silhouette, never with part of second row dorsolaterally, and almost always with conspicuous setal tuft along opposable base of fixed finger; lateral margin of propodus never distinctly costate. Hook on ischium of third pereiopod sometimes opposed by tubercle on basis. Pleura of third through fifth abdominal segments more rounded than angular. First pleopods of first form male contiguous basally; terminal elements comparatively short; central projection bearing distinct subapical notch, not tapering, and rather strongly recurved with notch directed almost proximally; mesial process inflated, with acute to subacute tip disposed at about right angle to shaft of appendage. Terga of abdomen with narrow, dark marginal band posteriorly, first segment often with broad similarly colored band in anterior section; longitudinal dark stripe lacking, and pleura without scalloped stripe at base. Female with first pleopod present.

MALE, FORM I (from Lookout Creek, Dade County, Georgia).—Body subovate, depressed (Figure 64a,j). Abdomen narrower than cephalothorax (10.2 and 12.0 mm); maximum width of carapace greater than depth at caudodorsal margin of cervical groove (12.0 and 8.7 mm). Areola 3.1 times as long as broad, densely punctate with 7 punctations across narrowest part, length 36.5 percent of total length of carapace (43.8 percent of postorbital carapace length). Rostrum with strongly convergent thickened margins ending rather suddenly at base of acumen; latter distinctly upturned, reaching base of ultimate podomere of antennular peduncle; dorsal surface of rostrum concave with prominent punctations; subrostral ridges well developed and, although in dorsal aspect disappearing beneath rostral ridges just anterior to orbit, reappearing posterior to base of acumen and continuing to tip. Postorbital ridge short, deeply grooved dorsolaterally, terminating cephalically in small tubercle. Suborbital angle subacute, branchiostegal spine represented by angulation ventral to cephalic margin of cervical groove. Cervical spine or tubercle absent. Carapace densely punctate dorsally except for median gastric region and wedge-shaped area in anterior half of areola where punctations sparse, lateral part of cephalic section also mostly punctate, majority of punctations conspicuously large; posterior orbital area swollen and with few tubercles; lateral part of branchiostegal region granulate to weakly tuberculate. Abdomen and carapace subequal in length (23.0 and 22.5 mm); pleura rather long and rounded posteroventrally and with gently sloping cephaloventral margins. Cephalic section of telson with 2 spines in each caudolateral corner, usual lateral incision deep and transverse suture distinct. Uropod with proximal podomere bearing acute mesial lobe, lateral lobe rounded; mesial ramus with median ridge obsolete but with premarginai distomedian spine and small distolateral spine.

Cephalomedian lobe of epistome (Figure 64g) subtriangular with slightly elevated (ventrally) margins, surface convex posteriorly; main body with prominent fovea and arched epistomal zygoma, usual elongate pits flanking anterolateral side of zygoma reduced to rather shallow grooves. Ventral surface of proximal podomere of antennule with small spine at base of distal fourth or fifth. Antennal peduncle without spines or prominent tubercles; flagellum reaching third abdominal tergum; antennal scale (Figure 64i) about 2.4 times as long as wide, sides subparallel but broadest distal to midlength, distal spine strong and reaching almost to midlength of ultimate podomere of antennular peduncle. Ventral surface of ischium of third maxilliped with broad longitudinal band of long stiff setae, submarginal lateral row of smaller plumose setae, and additional short plumose setae dispersed between; distolateral angle subacute but not produced.

Right chela (Figure 64l) 2.2 times as long as broad, mesial margin of palm occupying about one-third of its length. Mesial surface of palm with 3 moderately well defined tubercles followed distally by row of 5 punctations; dorsal, ventral, and lateral surfaces punctate, latter surface with slight suggestion of costa. Fingers widely gaping and proximal half of opposable surface of fixed finger with prominent tufts of long plumose setae; neither finger with clearly defined longitudinal ridges dorsally or ventrally, both conspicuously punctate; opposable margin of fixed finger with row of 9 low, rounded corneous tubercles, another on lower level opposite seventh tubercle from base, and band of minute denticles situated between sixth tubercle and corneous tip of finger; opposable margin of dactyl with irregular row of 12 (left with 11) tubercles similar to those on fixed finger, third and fifth from base larger than others, and band of minute denticles present between ninth tubercle and corneous tip of finger.

Carpus of cheliped with distinct oblique furrow dorsally, single spikelike tubercle mesially, and 2 tubercles on ventrodistal margin (one submedian and other forming articular knob), podomere otherwise punctate. Merus of right cheliped with 2 (left with 1) premarginai tubercles dorsodistally; ventrolateral row of tubercles represented by 1 (2 on left) and ventromesial consisting of 6 (7 on left), only distal members of latter row spikelike. Mesial margin of ischium without tubercles.

Ischium of third pereiopod with simple hook (Figure 64h) overreaching basioischial articulation and opposed by small tubercle on basis. Coxa of fourth pereiopod with caudomesial boss bearing transverse ridge caudoventrally, that of fifth lacking boss but bearing scattered setae on ventral membrane.

First pleopods (Figure 64b,d), reaching coxae of third pereiopods, symmetrical, and with contiguous bases (see “Diagnosis” for description).

FEMALE (from Lookout Creek, Dade County, Georgia).—Excluding secondary sexual characters, differing from male, form I, in following respects: tubercle lacking from cephalic end of postorbital ridges; suborbital angle obtuse; several tubercles in area occupied by cervical spine in other crayfishes slightly larger than others nearby on branchiostegite; median ridge on mesial ramus of uropod evident but weak; cephalomedian lobe of epistome irregularly but broadly rounded cephalically; antennal scale broadest at about midlength; mesial surface of palm of chela with 4 tubercles followed distally by 3 punctations; fixed finger with weak median longitudinal ridge; opposable margin of fixed finger of right chela with row of only 8 tubercles; mesial surface of carpus of cheliped with additional small tubercle proximally; merus of cheliped with 1 premarginai tubercle dorsodistally, only 1 tubercle representing ventrolateral row.

Annulus ventralis (Figure 64k about 1.8 times as broad as long, almost quadrangular, and rather shallowly embedded in sternum; cephalic section, more weakly sclerotized than caudal, with median longitudinal furrow flanked by paired ridges; caudal region broadly excavate anteriorly, inflated, forming convex caudal wall dextrally and angular, somewhat concave one, sinistrally; sinus originating at caudodextral end of median furrow, and following tilted S-shaped course, ending on edge of protruding midcaudal wall. Postannular sclerite about 2.8 times as broad as long, little more than half as wide and almost one-third as long as annulus, bearing punctate oval elevation (ventrally). First pleopods reaching slightly anterior to midlength of annulus when abdomen flexed.

MALE, FORM II (from West Fork of Wolf Creek, Union County, Georgia).—Differing from first form male in following respects: areola with 5 punctations across narrowest part; rostral margins not ending abruptly at base of acumen and merging imperceptibly with subrostral ridges, latter evident in dorsal aspect to base of acumen where joining rostral ridges, postorbital ridges with well-developed corneous tubercles cephalically; suborbital angle obtuse; posterior part of orbital area less swollen and tubercles on it much reduced; cephalic section of telson with only 1 spine in caudodextral corner; cephalomedian lobe of epistome with broadly arched cephalic margin as in female; antennal peduncle with acute spine on lateral surface of basis; antennal scale broadest at about midlength, distolateral spine reaching end of antennular peduncle; mesial surface of palm of chela with row of 6 very low tubercles becoming progressively more squamous and inconspicuous distally; fixed finger with poorly developed median longitudinal ridge dorsally, opposable margin ofthat finger on right chela with row of 10 tubercles; corresponding margin of dactyl also with 10; merus of both chelipeds with 2 premarginai tubercles dorsodistally, ventrolateral row represented by 2 tubercles and ventromesial by 7 on right and 8 on left cheliped; ventromesial margin of ischium with 2 very small tubercles; hook on ischium of third pereiopod not reaching basioischial articulation and not opposed by tubercle on basis; boss on coxa of fourth pereiopod with much suppressed caudal ridge.

First pleopods (Figure 64c,e) reaching coxae of third pereiopods, symmetrical, and separated basally by very narrow gap. Noncorneous terminal elements contiguous at base; mesial process directed at about right angle to shaft of appendage and tapering to subacute tip; and central projection deflected caudoproximally at about 110 degrees, rounded apically but with suggestion of subapical notch. Juvenile oblique suture present on shaft.

COLOR NOTES (Figure 58d).—Dorsum of carapace olive to orange tan or brown; mandibular adductor regions with dark brown reticulations, these flanked caudally and joined dorsally across posterior gastric region by very narrow dark band along cervical groove; rostral ridges pale olive to brownish cream; broad, marginal pinkish cream band extending from antennal region caudoventrally across mandibular and anteroventral branchiostegal regions; hepatic area with oval dark patch continuing over orbital region but pale more caudally and ventrally. Thoracic region olive to orange tan, fading ventrally almost to cream and with pale orange or tan semiovate spot abutting dorsomedian caudal ridge; latter, including flange, dark greenish gray or dark burgundy. Tergum of first abdominal segment sometimes with dark brown to burgundy rectangular band, its cephalic section, median transverse ridges, and caudal margin red to burgundy; pleura pale. Remaining terga pale olive or orange tan with narrow red to burgundy caudal band extending onto posterior margin of pleura, latter pale cephaloventrally. Telson olive to orange tan bordered in brown to red. Uropods similarly colored but with lateral parts of both rami darker and distal parts suffused with reddish orange to brown. Chelipeds with basal podomeres cream, dorsodistal part of merus dark olive to brown; dorsal part of carpus pale olive or tan basally and darker distally; dorsal part of palm pale proximolaterally and with pale knob opposite mesial base of dactyl, olive tan over most of surface, and mesial margin dark olive to brown; both fingers dark olive with pale tips, and setal tuft at base of fixed finger gray. Remaining pereiopods with cream or pinkish cream basal podomeres and olive to tan distal to ischium; distal part of merus and carpus darker than propodus and dactyl.

TYPE.—Holotype, MCZ 3629 (II).

TYPE-LOCALITY.—Eastern Tennessee and West Virginia (Faxon, 1885a). Restricted to Doe River, Elizabethton, Carter County, Tennessee, by Ortmann (1931:121).

RANGE.—The range of this crayfish has not been accurately defined, and in view of the difference of opinion concerning its relationship to C. (H.) girardianus (see “Remarks” under “Subgenus Hiaticambarus”), a detailed study of the two, particularly in Tennessee, needs to be made. It is a common species in the Tennessee Basin, especially in the Holston and Watauga watersheds. From Knoxville southward what seems to me to be the typical form is less abundant. In Georgia, it has been found only in the headwaters of the Nottely River (Hiwassee Basin) in Union County, in Lookout Creek in Dade County, and in Cane Creek (introduced?), a tributary of the Chattooga River in the southern part of Walker County, thus being confined to the Appalachian Plateau, Ridge and Valley, and Blue Ridge provinces of the state.

Measurements

(mm; based on specimens described herein)

GEORGIA SPECIMENS EXAMINED.—I have examined 119 specimens from the following localities. Dade County: (1) Lookout Creek at St Rte 189, E of Rising Fawn (Anonymous, 1970c:35), 1I, 23 Apr 1968, E. T. Hall, Jr., HHH, collectors; 3, 1j, 3j, 29 Sep 1969, M. W. Walker, ETH; 2I, 9, 1j, 21 Oct 1976, T. A. English, Jr., HHH. Union County: (2) East Fork of Wolf Creek 0.5 mi NE of Vogel State Park on US Hwy 19 (Holt, 1968b:32; Bouchard and Hobbs, 1976:13), 3I, 2, 5 May 1958, K. W. Simonds; (3) East Fork of Wolf Creek at confluence with West Fork, near Self Mountain (Anonymous, 1973b:67), 5II, 1, 9 Aug 1972, ETH, W. D. Kennedy; 2I, 7, 1j, 24 Oct 1976, TAE, HHH; (4) West Fork of Wolf Creek at confluence with East Fork near Self Mountain (Anonymous, 1973b:66), 2II, 3, 9 Aug 1972, ETH, WDK; (5) Nottely River 1.0 mi N of jet of St Rte 180E on US Hwy 19, 5I, 1II, 1, 9j, 8j, 3 ovig , 27 Apr 1967, T. Unestam, HHH. Walker County: (6) Cane Creek at St Rte 151, 3II, 6j, Jun 1977, R. W. Bouchard, J. R. Weaver; 1, 5j, 6j, 8 Oct 1977, WDK, HHH; 2I, 7, 12j, 10j, 9 Oct 1977, WDK, HHH.

VARIATIONS.—The most conspicuous difference noted between the Georgia specimens assigned to this species and topotypes of C. (H.) longirostris from the Doe River in Carter County, Tennessee, is in the position of the first pleopods of the male: in the Georgia specimens the bases of these appendages are contiguous, whereas in the topotypes there is a comparatively broad gap between them. The anticipated importance of such a difference, however, is negated by the occurrence of contiguous pleopods in specimens from another tributary of the Watauga River in Carter County. Other differences noted in the pleopods from the two areas are almost certainly insignificant. The rostra of the Georgia specimens from the Hiwassee Basin are markedly similar to those of topotypes, but those in populations frequenting Lookout Creek and the Chattooga basins are distinctly different (see below). The suborbital angle in none of the Georgia representatives is so prominent as in those from the Doe River. The chelae of specimens from Lookout Creek, and especially those of individuals from the Chattooga River basin, are distinctly more robust than those of the topotypes. Only two tubercles are present on the ventral surface of the carpus of the cheliped in the Georgia specimens, but a third one situated proximomesial to the distomedian tubercle is present, although sometimes very poorly developed, in the topotypes; even the distomedian tubercle in some of the latter is almost obsolete. Most of the topotypes possess only one well-developed tubercle representing the ventrolateral row on the merus of the cheliped, in this respect resembling most members of the Lookout Creek population and differing from most specimens from the Hiwassee Basin that possess two. Among available material from the Chattooga watershed neither one nor two tubercles seem to predominate.

Many of the variations noted among the Georgia specimens seem to be associated with the age or stage in the molt cycle of the specimens. For example, the young usually have well-developed postorbital spines, whereas in the larger, presumably older, individuals they are frequently reduced to rounded tubercles that are sometimes hardly evident in specimens in the late intermolt stage.

Characteristics exist among the Georgia specimens (Figure 66) that in most instances seem to distinguish those representatives occurring in the three watersheds: in the Hiwassee Basin, the rostral margins are evenly tapered to the tip of the acumen; there are two spines or tubercles representing the ventrolateral row on the merus of the cheliped; and the caudal wall of the annulus ventralis is almost symmetrical. In specimens from Lookout Creek, the rostral margins become suddenly thinner, actually ending, at the base of the acumen; almost always there is only one tubercle (sometimes acute) representing the ventrolateral row on the merus of the cheliped; and the caudal wall of the annulus is distinctly asymmetrical, with the side from which the tongue arises somewhat concave instead of being evenly rounded. In specimens from the Chattooga Basin locality, the rostrum is not strongly tapered and the rostral margins end abruptly at the base of the acumen, the latter feature resembling that of the population in Lookout Creek; not only do the adults attain a greater size but the chelae appear to be more robust than those of topotypes and specimens from the other two basins in Georgia. The annulus ventralis is nearly symmetrical as it is in the populations occurring in the Hiwassee Basin.

The ranges and averages (in parentheses) of three ratios obtained from specimens from the type-locality and the three drainage basins in Georgia are presented in the accompanying tabulation. Otherwise, no consistent differences in individuals from the three basins have been observed. The ratios of areola length (AL) to carapace length (CL) and to postorbital carapace length (POCL) are expressed in percentages. The relationship of areola length to areola width (AW) is expressed directly as a ratio.

SIZE.—The largest specimen available is a second form male having a carapace length of 33.1 (postorbital carapace length 28.2) mm. The smallest and largest first form males have corresponding lengths of 21.1 (17.1) and 30.5 (26.0) mm, respectively, and those of the smallest ovigerous female are 20.0 (16.4) mm.

Cambarus (Hiaticambarus) longirostris Faxon

Cambarus Bartonii var. longirostris Faxon, 1885a:64.

Cambarus bartonii longirostris.—Faxon, 1890:623.

Cambarus bartonii spinirostris.—Faxon, 1890:623 [lapsus for C. b. longirostris].

Cambarus longulus longirostris.—Hay, 1899b: 959.—James, 1966:9, fig. 2a,b, pl. 1c,f,g,n,o,r,s.

Cambarus (Bartonius) bartoni longirostris.—Ortmann, 1905c: 135 [by implication].

Cambarus (Cambarus) bartonii longirostris.—Fowler, 1912: 341 [by implication].

Cambarus (Cambarus) longulus longirostris.—Ortmann, 1931: 121.

Cambarus longerosilis.—Brimley, 1938:503 [erroneous spelling].

Cambarus (Hiaticambarus) longirostris.—Hobbs, 1969b: 106, figs. 10, 18m.

TYPE.—Holotype, MCZ 3629 ( II).

TYPE-LOCALITY.—“Eastern Tennessee and West Virginia” (Faxon, 1885a). Doe River, Elizabethton, Carter County, Tennessee, designated by Ortmann, 1931.

RANGE.—Tennessee River drainage in eastern Tennessee, southwestern Virginia, and western North Carolina, in the Coosa drainage in northeastern Alabama and Georgia.

HABITAT.—Rocky streams.

Cambarus (Hiaticambarus) longirostris Faxon

Cambarus Bartonii, var. longirostris Faxon, 1885a:64.

Cambarus bartonii longirostris.—Faxon, 1890:623.

Cambarus bartonii spinirostris.—Faxon 1890:623 [lapsus for C. b. longirostris].

Cambarus longulus longirostris.—Hay, 1899b:959.—James, 1966:9, fig. 2a, b, pl. 1c, f, g, n, o, r, s.

Cambarus (Bartonius) bartoni longirostris.—Ortmann, 1905c:135 [by implication].

Cambarus (Cambarus) bartonii longirostris.—Fowler, 1912:341 [by implication].

Cambarus (Cambarus) longulus longirostris.—Ortmann, 1931:121.

Cambarus longerosilis.—Brimley, 1938:503 [erroneous spelling].

Cambarus longulus longerostris.—Schurr and Stamper, 1962:474 [erroneous spelling].

Cambarus bartonii var. longirostris.—James, 1966:2.

Cambarus spinirostris.—James, 1966:2.

Cambarus longirostris.—James, 1966:2.

Cambarus longulus longirosilis.—James, 1966:8 [erroneous spelling].

Cambarus (Hiaticambarus) longirostris.—Hobbs, 1969b:106, figs. 10, 18m; 1974b:16, fig. 48.

Cambarus (Hiaticambarus) girardianus.—Bouchard, 1976a:572 [in part]; 1976b:588 [in part].

TYPE.—Holotype, MCZ 3629 (male II).

TYPE LOCALITY.—“Eastern Tennessee and West Virginia” (Faxon, 1885a). Restricted to Doe River, Elizabethton, Carter County, Tennessee, designated by Ortmann, 1931:121.

RANGE.—Tennessee River Basin upstream from below the mouth of the Sequatchie River: in Georgia, Tennessee, North Carolina, and Virginia. Introduced into headwaters of the Savannah River in South Carolina.

HABITAT.—Rocky segments of streams.