Cambarus latimanus (Le Conte 1856)

Cambarus (Depressicambarus) latimanus (LeConte)

Astacus latimanus LeConte, 1856:402*.—Hagen, 1870:9*, 10*, 79*.

Cambarus latimanus.—Hagen, 1870:78, 80, 82–84*, 98*, 100*, 105–107*, pl. I: figs. 43–46; pl. III: fig. 162.—Cope and Packard, 1881:881*, 882.—Faxon, 1884:144*; 1885a:(63, 69, 159, 167, 173),* pl. II: fig. 3; 1914:395*, 425*.—Underwood, 1886:370*.—Packard, 1888:40*, 41*.—Harris, 1903a:(59, 106, 138, 143, 152)*.—Goodnight, 1941: 72*.—Hobbs, 1942b:14, 20, 21, 158–161*; 1952b:172*; 1958a:74; 1959:897*; 1968a:272*; 1968b:K–16*, fig. 32j; 1969b:343.—Hall, 1957:(3–34, 51–53)*, pls. 1, 2, 5, map 1.—Hobbs and Hart, 1959:148*, 151, 159–161, 164, 172*, 185, 186*, 187.—Anonymous, 1967e, tab. 3*; 1969a:C–31*; 1969b:30–33*, 35–38*; 1969c:(61, 65, 66, 72, 74, 77, 82, 85)*; 1970b:(168, 172, 191, 211, 226)*; 1971:(159, 170, 179, 197)*; 1972b:10*; 1972d:81*, 85*, 99*; 1972f:169*, 178*, 179*.—Holt, 1968a:(302, 305, 310, 312)*; 1968b: 26*, 32*.—Boyce, 1969:(1, 6, 7, 73, 74, 76, 83, 85, 88, 89, 90, 92–101, figs. 31–32)*.—Sullivan and Heard, 1969: 307*.—Hobbs and Hall, 1969:286*; 1974:(199, 202, 205, 206)*.—Hart and Hart, 1971:107*, 108*; 1974:(21, 31, 32, 61, 73, 79, 88, 90, 131, 134, 136)*.—Holsinger and Peck, 1971:30*.—Bouchard, 1978:27–29*, 39, 43–44, 46, 47.—Wharton, 1978:(37, 46, 220)*.

Cambarus obesus var. latimanus.—Packard, 1880:222.

Cambarus Jordani Faxon, 1884:119–120*, 145* [“Holotype”, MCZ 3561 (II). Type-locality, Etowah River, Rome, Floyd Co., Georgia.]; 1885a:59, (83, 84, 160, 167, 173, 178)*, pl. 3: fig. 3.—Underwood, 1886:370*.

Cambarus jordani.—Hay, 1899b:959*, 963; 1902a:436–437.—Ortmann, 1902:277; 1931:97*−99, 103*, 104.—Steele, 1902:7.—Harris, 1903a:(59, 106, 144, 152)*.—Faxon, 1914:423*.—Creaser, 1931a:6.—Hobbs, 1956c:115, 120*; 1969a:104*.—Hall, 1959:221.—Anonymous, 1967a, tab. 6*; 1967i, tab. 3*.—Bouchard, 1978:27, 29*.

Cambarus latimus.—Steele, 1902:7 [erroneous spelling].

Cambarus (Bartonius) jordani.—Ortmann, 1905a:118 [by implication], 120, 121*, 130.

Cambarus (Bartonius) latimanus.—Ortmann, 1905a:120, 122*.

Cambarus (Cambarus) latimanus.—Fowler, 1912:341 [by implication].—Ortmann, 1931:124*, 125*.

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

Cambarus (Cambarus) extraneus.—Ortmann, 1931:97*, 98*, 104.

Cambarus sp.—Hobbs and Walton, 1960a:18*.—Holsinger and Peck, 1971:30*.—Hart and Hart, 1974:88*, 134*.

Cambarus (Depressicambarus) latimanus.—Hobbs, 1969a:104, 138*, figs. 1f, 8*, 13g, 14g, 18f; 1972b:114*, 146*, figs. 8b, 89o, 90e, 92b, 100b; 1974b:13*, fig. 33.—Hobbs and Hall, 1972:159*.—Holt, 1973a:246, 248*.—Bouchard, 1978:30, 34–37*, fig. 1 1.

Cambarus species F.—Hobbs, 1969a:(104, 136–138, fig. 8).*.

Cambarus (Depressicambarus) jordani.—Hobbs, 1969a:104* [by implication]; 1972b:114*, 146*, figs. 97d, 99d; 1974b:13*, fig. 32.—Hobbs and Hall, 1972:160*, 161.

Cambarus halli.—Hart and Hart, 1971:107*.

Cambarus (Depressicambarus) sp.—Hart and Hart, 1974:(21, 61, 73, 79, 88, 134)*.

Cambarus (Depressicambarus) sp. nov.—Anonymous, 1975a: 142*, 143*, 147.*

The synonomy presented here is believed to be complete only for Georgia.

SUMMARY OF LITERATURE PERTAINING TO GEORGIA.—The earliest record of the occurrence of this crayfish in Georgia was that of LeConte (1856:402), who described it from “Georgia superiore.” In his monograph of the Astacidae, Hagen (1870:84) cited two specific localities: Athens (Clarke County) and Milledgeville (Baldwin County), and a third locality, Roswell (Fulton County), was added by Faxon (1885a:69). The description of Cambarus jordani Faxon (1884:119) was based on a juvenile male, and, as pointed out by Bouchard (1978:29) and below, this crayfish differs in no important respect from C. (D.) latimanus. Although the latter was reported to occur in several localities in neighboring states, no additional records or information on it in Georgia were reported for more than 60 years. Between 1950 and 1960, Hobbs (1952b:172) added one locality, Wesleyan College at Rivoli, Bibb County, and he and Hart (1959:186) cited this crayfish from 0.75 mile east of Fort Gaines in Clay County. They indicated that in the lower Flint-Chattahoochee-Apalachicola system, C. (D.) latimanus

…appears to be confined to the small sand-bottomed tributaries…flowing in deep shaded ravines. Here they are found in the debris littering the stream beds and in burrows excavated in the banks of the stream. Many of the complex, highly branching burrows have openings above the water level in addition to those in the stream.

They also noted that first form males had been collected in April. Until recently the most outstanding contribution to our knowledge of this species is that of Hall (1957), who summarized previous published data, adding new information on its range and habits. Except for differences in interpretations of the limits of variation (involving the recognition of subspecies), his account of the species in Georgia is still apropos.

During the next decade several additional localities were added: (Anonymous, 1969a, b, c), Hobbs (1968a), Holt (1968a, b), and Sullivan and Heard (1969). Boyce (1969) also conducted a comparative ecological study of C. (D.) latimanus and P. (Pe.) spiculifer in the Yellow River with emphasis on respiration and tolerance to low oxygen concentrations. He found no significant difference in the respiration rate in the mature males of the two (p. 59), and both occur throughout the basin upstream from the Annistown Bridge.

Until recently I was convinced that the recognition by Hall (1957) of two subspecies of C. (D.) latimanus was tenable (Hobbs, 1969a) and referred to that segment of the species occupying the Savannah River Basin as “Cambarus species F.” Larger series from additional localities have shown that the populations in the basin are highly variable in most respects and should not be recognized as distinct.

Within the current decade, an additional 38 localities were cited for this crayfish: Anonymous (1971, 1972b, d, f), Hobbs and Hall (1972), and Hart and Hart (1974). On the basis of my erroneous identifications of a female and juvenile male, Holsinger and Peck (1971:30) reported this species from Byers and Hurricane caves in Dade County. Both specimens should be referred to C. striatus.

Despite the comparatively large list of references cited in the above synonomy, our understanding of this crayfish is still limited. The contributions of Holt and of Hart and Hart are primarily concerned with the commensal branchiobdellids and entocytherids, respectively, providing no data other than localities where this crayfish serves as host to one or more of these animals. As indicated above, among the most valuable contributions was that of Hall (1957). The anonymous references consist of reports of water quality surveys, and in them, not only are there a number of new localities cited, but also the physical and chemical data provided for the stations investigated broaden our appreciation of the habitats exploited by Cambarus (D.) latimanus. In his review of the subgenus Depressicambarus, Bouchard (1978) presented a detailed description and illustrations of topotypes; a discussion of its range and variations precede a brief statement of size and notes on its life history. The remaining references—the early ones largely repetitious—are concerned with relationships or the distribution of the species.

DIAGNOSIS.—Eyes of moderate size. Rostrum with or without marginal spines and almost always lacking median carina. Carapace usually with cervical spine or tubercle. Areola 3.4 to 10.0 (average 6.3) times as long as broad, constituting 30.1 to 37.1 (average 34.3) percent of entire length of carapace, and 38.2 to 44.9 (average 41.4) percent of postorbital carapace length, with 2 to 6 punctations across narrowest part. Suborbital angle obtuse to obsolete (rarely subacute). Postorbital ridge terminating cephalically in spine, more frequently in tubercle, or merging almost imperceptibly with cephalic area of carapace. Antennal scale approximately 2.5 times as long as broad, broadest distal to midlength. Palm of chela with 5 to 9 (usually 7) tubercles in mesialmost row, squamous tubercles scattered over at least mesial half of dorsal surface of palm, frequently over entire surface. First pleopod of first form male with moderately long to short terminal elements; central projection very variable, strongly arched or bent at angle only slightly greater than 90 degrees, usually tapering and with or without subapical notch; mesial process likewise markedly variable, usually inflated, truncate or tapering distally, occasionally overreaching distal extremity of central projection; caudal knob rarely present even as vestige. Color olive gray to orange tan; abdominal terga frequently with longitudinal stripes. Female with first pleopod present.

COLOR NOTES (Figure 37c,d).—(Two phases: 1, dark olive and greenish gray; 2, brownish olive and orange tan; latter described here). Cephalic region of carapace olive with lighter olive tan dorsomedian area sharply delimited laterally by darker olive dorsolateral area; latter continuing caudally on branchiostegite to caudal margin of carapace; margins of rostrum and postorbital ridges pale tan; hepatic region with reticulate pattern of dark brown on olive; mottled pattern of similar colors overlying mandibular adductor region and continuing mesially in band along cephalic side of cervical groove, flanking caudal margin of submedian pale shield in caudomedian gastric region; thoracic region pale tan dorsally with irregular dark brown splotches on dorsolateral part of branchiostegites, latter somewhat concentrated in paired longitudinal bands; lateral surface of carapace fading to pale tan ventrally. Ground color of abdomen dark olive; terga with paired dorsolateral very dark brown spots, latter conspicuous on first three terga, becoming progressively smaller and less obvious on caudal ones; pleura with similarly colored ventrally convex markings at bases and with ventral portions pale tan to cream; caudolateral margin of sixth abdominal pleuron dark brown. Telson and uropods somewhat mottled, former frequently with lateral margins of cephalic section dark brown. Cheliped distal to midlength of merus with dorsal surface very dark olive and bearing tan to cream spines, tubercles, and knobs; lateral costa of propodus also light in color. Remaining pereiopods with dark olive mottlings on cream background proximally and on olive tan distal to midlength of merus.

Occasional individuals occur from almost all parts of the range in Georgia in which the dark spots of the carapace and abdomen are sufficiently large to fuse, forming broad longitudinal stripes extending from the cervical groove caudally almost to the base of the telson (Figure 37d). This striped pattern has been observed more frequently in the Savannah Basin than elsewhere. Other specimens have been collected that are virtually concolorous; usually, these are individuals that were found on a light sandy substrate and adapted to a light background.

Boyce (1969:83) also noted two color patterns indicating that both occur in males and females.

TYPES.—Syntypes, MCZ 3378 (1 dry), ANSP 329 ().

TYPE-LOCALITY.—Athens, Clarke County, Georgia (by subsequent restriction, see below). LeConte (1856:402) listed a single source, “Georgia superiore,” for the specimens that were available to him. A no more precise locality was suggested until 1914 when Faxon (p. 395), in mentioning a collection of 19 specimens from Athens, stated: “These are essentially paratypes, and are of interest as fixing the type locality, Athens, Ga. which was not specified in LeConte's original description of the species nor on the labels accompanying the type specimens in Cambridge and Philadelphia.” In the same publication (p. 425) he cited the type-locality as “Athens, Clarke Co., Georgia.”

RANGE.—Piedmont and coastal plain from the Tar and Cape Fear basins in North Carolina southward to the Altamaha and Apalachicola basins in Georgia and Florida, westward to the Coosa Basin in Alabama.

In Georgia (Figure 48), it occurs in most of the counties upstream from and along the southern flank of the fall line. Only in Bulloch, Effingham, and Jenkins counties (along the Ogeechee and Savannah rivers), Pulaski County (along the Ocmulgee River), and Clay, Quitman, Randolph, and Stewart counties (along the Chattahoochee River) is it known to invade far into the coastal plain. In the Chattahoochee drainage system, its range, although apparently interrupted in southwestern Georgia, extends southward into the Florida panhandle (Hobbs, 1942b:159). Considerable field work in northwestern Georgia failed to reveal its presence in the Tennessee Basin except in tributaries of the Hiwassee River. Most surprising was my failure to find it in so much of the Conasauga drainage system, particularly in view of the occurrence of an apparently large population in Minnewauga Creek, a tributary stream in Polk County, Tennessee, that was sampled by R. W. Bouchard.

GEORGIA SPECIMENS EXAMINED.—I have examined 2424 specimens from approximately 400 localities (Figure 48).

VARIATIONS.—Cambarus (D.) latimanus is one of the most variable crayfishes in the state. Almost every characteristic that has been evaluated has a broader range of variability than has been noted in other species; furthermore, few of the variations can be correlated with a restricted part of the range within the state. The unusual lack of stability is clearly evidenced in the illustrations of the first pleopods of the first form male (Figures 49, 50). Most noticeable perhaps are the differences existing in the central projection. A long, strongly recurved element lacking a subapical notch seems to be typical of those populations occurring in the Hiwassee drainage system and in the Coosawattee segment of the Coosa Basin, but, elsewhere in the latter, the projection is less strongly recurved and not infrequently possesses a subterminal notch that is usually shallow. Similar to the appendage characteristic of populations in the Hiwassee and Coosawattee basins are those of members of the species occurring in the Flint, Chattahoochee, Ocmulgee, and Oconee drainage systems and of some individuals in the Tallapoosa. Generally, in rather marked contrast, the corresponding pleopods of individuals in the Ogeechee and Savannah basins possess a shorter central projection that is not so strongly recurved (the tip rarely reaching proximally beyond the level of the distal margin of the base of the mesial process) and almost invariably provided with a distinct subapical notch. The extreme variation of this type of central projection occurs in the Broad River in Banks and Madison counties, but downstream, in particular, and elsewhere in tributaries of the Savannah and Ogeechee, the subapical notch is less distinct and the element is longer. Throughout most of the range of the species within the state, the caudal knob is not recognizable; however, it is moderately prominent in four individuals from one locality each in Bartow (Figure 49h), Pickens (Figure 49i), Polk (Figure 49j) and Douglas (Figure 49s) counties (Coosa and Chattahoochee basins). The most heavily ornamented mesial process occurs in individuals in the Chattahoochee Basin.

Among the most conspicuous variations noted are those of the rostrum. In most populations, the smaller juvenile members possess marginal spines that, as the individual increases in size, seem progressively, with each molt, to become more atrophied until in the adult there is little or no trace of their earlier presence. In some isolated individuals this juvenile trait is retained in the form of marginal tubercles that may or may not be acute. In some populations, virtually all of the members have such tubercles or at least distinct angles at the base of the acumen (Figures 47a,j, 51a,c,e); this is the variant that was described by Faxon (1884:119) as Cambarus Jordani. In a few localities in Georgia, for example, 5.5 miles northeast of Molena on U.S. Highway 18, Pike County, and 6.5 miles south of the Chattahoochee River on U.S. Highway 219, and in several localities in the Oconee Basin, no marginal spines or tubercles occur on the rostra of even the smallest individuals (see Hall, 1957:18 and Hobbs, 1958a:74). Occasionally there is a weak median carina on the rostrum, but typically the upper surface is shallowly to moderately deeply concave.


Cervical spines are frequently present, particularly well developed in small individuals, and often accompanying them are spines on the postorbital ridges. Such spines are conspicuous in many adult specimens from the Broad River and adjacent smaller tributaries to the Savannah. Elsewhere they are less well developed in adults, sometimes reduced to tubercles, and in occasional individuals, there is not a trace of either cervical or postorbital spines. The suborbital angle is usually broadly obtuse to obsolete, but infrequently it is subacute.

Some populations in the streams just mentioned possess an areola that is decidedly broad, the width constituting as much as one-third the total length, and the narrowest observed in crayfishes frequenting these tributaries of the Savannah is 8.8 times as broad as long; the average there is 4.7 as compared with 7.2 elsewhere in Georgia, where the ratio of length to width of the areola ranges from 4.1 to 10.0 although seldom greater than 9.4.

While the spines and tubercles on the chelipeds do not exhibit marked variation, there are differences in the actual numbers and in their size. As indicated in the “Diagnosis,” for example, the number of tubercles in the mesialmost row on the palm varies from five to nine; similarly the adjacent row consists of four to eight.

Finally, variations in color patterns are noted in “Color Notes.”

SIZE.—The largest specimen available from Georgia is a female from the Chattahoochee River drainage in Fulton County, which has a carapace length of 52.9 (postorbital carapace length 44.7) mm. The smallest and largest first form males have corresponding lengths of 29.5 (24.6) and 40.9 (35.0) mm, respectively.

Cambarus (Depressicambarus) latimanus (LeConte)

Astacus latimanus LeConte, 1856:402.

Cambarus latimanus.—Hagen, 1870:83, pl. 1: figs. 43–46; pl. 3: fig. 162.—Faxon, 1885a:69, pl. 2: 014;Hobbs, 1942b: 158, figs. 186–190.

Cambarus obesus var. latimanus.—Packard, 1880:222.

Cambarus (Bartonius) latimanus.—Ortmann, 1905c: 120.

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

Cambarus (Depressicambarus) latimanus.—Hobbs, 1969b: 104, figs. 1f, 8, 13g, 14g, 18f.

TYPES.—Syntypes, MCZ 3378 (I dry), MCZ, ANSP 329 ().

TYPE-LOCALITY.—Athens, Clarke County, Georgia.

RANGE.—North Carolina to Alabama, chiefly in the piedmont but extending into western Florida along the Apalachicola River.

HABITAT.—Small streams and burrows (secondary burrower).

Cambarus (Depressicambarus) latimanus (Le Conte)

Astacus latimanus LeConte, 1856:402.

Cambarus latimanus.—Hagen, 1870:83, pl. 1: figs. 43–46; pl. 3: fig. 162.—Faxon, 1885a:69, pl. 2: fig. 3.—Hobbs, 1942b:158, figs. 186–190.

Cambarus obesus var. latimanus.—Packard, 1880:222.

Cambarus Jordani Faxon, 1884:119; 1885a:83, pl. 3: fig. 3. [Type: MCZ 3561 (male II). Type locality: Etowah River, near Rome, Floyd County, Georgia]

Cambarus jordani.—Hay, 1899b:959.

Cambarus latimus.—Steele, 1902:7 [erroneous spelling].

Cambarus (Bartonius) latimanus.—Ortmann, 1905c: 120.

Cambarus (Bartonius)jordani.—Ortmann, 1905c: 120 [by implication].

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

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

Cambarus (Depressicambarus) latimanus.—Hobbs, 1969b:104, figs. If, 8, 13g, 14g, 18f; 1974b:13, fig. 33; 1981:109, figs. 23f, 37c, d, 39e-g, 46–51, 202.—Bouchard 1978a:34, figs. 11, 2a, b, 5–8.

Cambarus (Depressicambarus) jordani.—Hobbs, 1969b:104 [by implication]

TYPES.—Syntypes, MCZ 3378 (male I, dry), MCZ 236 (listed by Faxon, 1914:425, as paratypes), ANSP 329 (female).

TYPE LOCALITY.—Athens, Clarke County, Georgia (by subsequent designation, Faxon 1914:395).

RANGE.—Piedmont and coastal plain from the Tar and Cape Fear basins in North Carolina southward to the Altamaha and Apalachicola basins in Georgia and Florida, westward to the Coosa Basin in Alabama.

HABITAT.—Small to moderately large streams and burrows (secondary burrower).

Cambarus (D.) latimanus

B Sathodrilus megadenus Holt (1968a:302): Haralson (Holt, 1968a:302)