PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116(3):827-838. 2003. A new burrowing of the Cambarus Erichson, 1846 (: ) from the lower Flint River basin in the Dougherty Plain of Georgia, with notes on C. (D.) harti Hobbs, 1981

John E. Cooper and Christopher E. Skelton

(JEC) North Carolina State Museum of Natural Sciences, Research Lab, 4301 Reedy Creek Road, Raleigh, North Carolina 27607, U.S.A., e-mail: [email protected]; (CES) Georgia Department of Natural Resources, Wildlife Resources Division, Nongame Wildlife & Natural Heritage Section, Georgia Natural Heritage Program, 2117 U.S. Highway 278 S.E., Social Circle, Georgia 30025-4714, U.S.A.; (Present address for CES) Department of Biological and Environmental Sciences, Georgia College & State University, CBX 081, Milledgeville, Georgia, 31061, U.S.A., e-mail: [email protected]

Abstract.—Cambarus (Depressicambarus) doughertyensis is a new species of obligate burrowing crayfish known from a single locality in the lower Flint River basin in the Dougherty Plain of the East Gulf Coastal Plain, Dougherty County, Georgia. It appears to be most closely related to Cambarus (D.) harti Hobbs, which is known from two localities in the western Piedmont Plateau, Meriwether County. Although the two species are morphologically similar in many respects, C. (D.) doughertyensis differs from C. (D.) harti in having a longer areola; a plethora of tubercles on the carpus and ventral surface of the palm; more tubercles on the opposable surfaces of both fingers of the chela, and differences in the morphology of those surfaces; a lack of spines or tu- bercles on the proximal podomere of the uropod; a radically different color pattern; and in a number of other characters. Spines on the ventral keel of the rostrum of other than certain Mexican crayfishes of the genus Pro- cambarus Ortmann are reported for the first time. Inadvertent errors in the description of C. (D.) harti are corrected.

Five similar species of highly special- Hills District; Cambarus (D.) strigosus ized, obligate burrowing crayfishes of the Hobbs, 1981, is limited to the Savannah genus Cambarus Erichson, 1846, subgenus River basin in the eastern Piedmont Pla- Depressicambarus Hobbs, 1981, latimanus teau; and Cambarus (D.) truncatus Hobbs, Group Bouchard 1978, are known to oc- 1981, is known only from the Oconee River cupy limited, widely separated ranges in basin in the Fall Line Hills District. Of Georgia (Hobbs 1981). Cambarus (D.) cy- these five crayfishes, C. (D.) harti, which is matilis Hobbs, 1970, occurs in the Cona- cobalt blue, occupies the southwesternmost sauga-Coosa River basin in the Ridge and range, and is known only from wetlands at Valley physiographic province; Cambarus two localities in Meriwether County. Its (D.) harti Hobbs, 1981, is known from the type locality is within the Cold Spring Flint-Chattahoochee River basin in the Creek subdrainage of the upper Flint River, western Piedmont Plateau; Cambarus (D.) the other is within the Flat Shoal Creek sub- reflexus Hobbs, 1981, occupies parts of the drainage of the Chattahoochee River, about Savannah and Ogeechee River basins in the 18.4 air km northwest of the type locality. Atlantic Coastal Plain and eastern Fall Line On 20 May 1999, CES excavated a num- 828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ber of chimneyed burrows in a seasonally verging to base of short acumen, which not flooded swamp forest adjacent to the flood- delimited by tubercles or spines; margins plain of Kiokee Creek, a tributary of the constricted at base of acumen, strongly con- lower Flint River in the Dougherty Plain of verging and concave from there to small, the East Gulf Coastal Plain in Dougherty dorsally directed apical tubercle; acumen County. This locality is about 147 air km comprising 28.6-42.9% (jc = 34.6%) of south-southeast of the type locality of C. rostrum length, latter constituting 11.7— (D.) harti. The initial digging efforts pro- 13.3% (jc = 12.5%) of TCL; floor (dorsal duced four specimens of a relatively drab, surface) of rostrum broadly concave, ce- brownish crayfish that superficially resem- phalic half cuplike; ventral keel of rostrum bled C. (D.) harti except in color pattern. bearing 1-4 corneous spines. Areola oblit- Additional specimens have since been col- erated or nearly so, length constituting lected, and it is apparent that this is 40.9-45.6% (jc = 43.7%) of TCL and 46.2- an undescribed species related to C. (D.) 51.2% (jc = 48.9%) of PCL. Thoracic sec- harti and the four other previously men- tion of carapace dorsally crowded with tioned species. Examination of these spec- punctations, dorsolaterally and laterally imens also revealed the presence of spines granulate or with small tubercles; cephalic on the ventral keel of the rostrum, a char- section laterally with many, usually small, acter that has been reported only in some tubercles. Cervical spines reduced to tuber- Mexican crayfishes of the genus Procam- cles, one to several each side of carapace. barus, subgenera Paracambarus Ortmann, Postorbital ridge moderate, cephalic margin 1906, and Villalobosus Hobbs, 1972 (Vil- rounded and without spine or tubercle. Sub- lalobos 1955, 1983). Confusing statements orbital angle obsolete to broadly obtuse, in the original description of C. (D.) harti without tubercle or spine; branchiostegal are clarified, based on a recent examination spine vestigial or a small tubercle. Antennal of the primary types. scale 2.5-3.1 (jc = 2.8) times as long as Measurements of crayfish structures were wide, greatest width at midlength; lateral made to the nearest 0.1 mm with a Fowler margin thickened and terminating distally precision dial caliper, following the meth- in long spine, lamella narrow, cephalic mar- ods of Hobbs (1981:9—10) except where gin moderately or strongly declivous, me- noted. Abbreviations used in the paper are: sial margin subparallel to lateral margin; GMNH, University of Georgia Museum of antennal peduncle without tubercles or Natural History, Athens; j, juvenile; spines. NCSM, North Carolina State Museum of Palm of chela inflated, 1.7-1.9 (jc = 1.8) Natural Sciences, Raleigh; PCL, postorbital times wider than deep, depth 89.5-98.6% carapace length; TCL, total carapace length; (jc = 95.2%) of length of mesial margin, USGS, United States Geological Survey; width 1.6—1.9 (jc = 1.7) times length of me- USNM, National Museum of Natural His- sial margin; latter constituting 26.3-31.3% tory, Smithsonian Institution, Washington, (jc = 28.9%) of total chela (propodus) D.C.; UTM, Universal Transverse Mercator. length, bearing subserrate mesial row of 6 (rarely 5 or 7) large, semierect tubercles, Cambarus (Depressicambarus) row subtended dorsally by second row of 5 doughertyensis, new species or 6 (rarely 3 or 4) smaller ones, and other Fig. 1, Table 1 smaller but produced tubercles dorsolateral to this row; 1 or 2 small, produced tubercles Diagnosis.—Body and eyes pigmented, ventral to mesial row. Fixed finger of chela eye small (x adult diam. 1.2 mm, n = 15). costate laterally, with strong median ridge Rostrum with slightly thickened, elevated dorsally and weak submedian ridge ven- margins, subparallel or moderately con- trally; opposable surface of finger with row VOLUME 116, NUMBER 3 829

Fig. 1. Cambarus (Depressicambarus) doughertyensis, new species; all from holotypic male, form I (NCSM 7997), except E, F, from morphotypic male, form II (NCSM 7999), and I from allotypic female (NCSM 7998): A, lateral aspect of carapace; B, E, mesial aspect of gonopod (first pleopod); C, F, lateral aspect of gonopod; D, dorsal aspect of carapace; G, caudal aspect of in situ gonopods; H, epistome; I, annulus ventralis and postannular sclerite; J, antennal scale; K, ventral aspect of left palm; L, dorsal aspect of distal podomeres of left cheliped; M, basis and ischium of third pereiopod. Scale line = 2 mm. 830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1 .—Measurements (mm) of types of Cambarus (Depressicambarus) doughertyensis, new species.

Holotypic male, Morphotypic male, form I Allotypic female form II

Carapace Total length 28.5 29.0 29.4 Postorbital length 25.7 25.8 26.2 Width 14.5 14.5 14.7 Depth 11.0 11.4 11.2 Length rostrum 3.4 3.7 3.8 Length acumen 1.2 1.4 1.3 Length areola 12.7 12.7 13.0 Width areola 0.2 oblit oblit Antennal scale Length 3.2 3.2 3.4 Width 1.1 1.3 1.1 Abdomen Length 25.5 26.4 26.0 Width 10.1 10.7 10.3 Cheliped Length propodus 22.1 21.5 24.4 Length mesial margin palm 6.5 6.4 7.0 Width palm 11.5 11.1 12.6 Depth palm 6.0 6.3 6.9 Length dactyl 14.6 14.0 15.8 Gonopod length 6.8 N/A 7.1

of 5-7 (rarely 8) tubercles, in addition to ramus with small caudolateral and caudo- subconical tubercle ventral to denticles, median spines, latter submarginal. third tubercle from base usually very large; Hook on ischium of third pereiopod of dactyl length 2.2-2.6 (x = 2.3) times length males, that of holotypic male, form I (Fig. of mesial margin of palm, with very strong 1M) uniramous, tapered, oblique, over- median dorsal ridge and weak submedian reaching basioischial articulation by most ventral ridge; mesial surface of dactyl with of length, opposed by very weak tubercle strong, broad tubercles on proximal two- on basis; coxa of fourth pereiopod of males thirds or more; opposable surface with row with vertically disposed, caudomesial boss. of 9-12 (usually 9 or 10) tubercles, fourth In situ gonopods (first pleopods) of form from base usually very large. Carpus with I male (based on holotypic male; Fig. 1G) weak dorsomesial tubercles; large, stout, symmetrical; proximomesial apophyses conical spine at distal margin of mesial sur- moderate, partly rounded; pair of bulbosi- face; 2 low, subconical distal mounds at ties on base below apophyses; central pro- distal margin of ventral surface; and mul- jection directed caudally; mesial process in- tiple strong tubercles on mesial and ventral flated at base, tip extruded, directed cau- surfaces; merus with 2-4 small subdistal dally and inclined slightly proximally; in dorsal tubercles, 4-6 (usually 5) tubercles lateral aspect (Fig. 1C), central projection on ventrolateral ridge, and 7—11 (usually 9 curved over 90° to plane of shaft, slightly or 10) tubercles on ventromesial ridge. tapered, not recurved, with moderate, prox- Abdomen shorter and considerably nar- imally directed subapical notch; tip extend- rower than carapace; proximal podomere of ing proximally nearly to distal margin of uropod without tubercles or spines; mesial mesial process but not as far caudally as tip VOLUME 116, NUMBER 3 831 of latter; mesial process subglobose, distal dorsal aspect; ventral keel of rostrum with margin arched; tip of process extruded, sub- 2 corneous spines. acute, caudally directed; caudal process rep- Postorbital ridge fairly strong, poorly de- resented by small, rounded protuberance; in fined ventrally, inflated caudally; groove mesial aspect (Fig. IB), setae at midlength shallow, lateral; cephalic margin rounded. of shaft and some near proximocaudal mar- Suborbital angle obsolete, margin without gin of caudal curvature. tubercle or spine; branchiostegal spine re- Annulus ventralis (based on allotypic fe- duced to small tubercle. Thoracic section of male; Fig. II) about 1.5 times wider than carapace dorsally crowded with deep punc- long, subovate in ventral outline; cephalic tations, dorsolaterally and laterally granu- half moderately depressed, with broad, un- late; row of small tubercles along ventral even median trough flanked each side by margin of anterior section of cervical strong ridge; cephalomedian margin domed, groove; cephalic section of carapace 1.2 with median concavity; sinistral ridge of ce- times longer than areola, constituting 55.5% phalic half descending obliquely to join up- of TCL; laterally with many scattered, per arm of reverse C-shaped caudosinistral small to moderate, tubercles, gastric region wall, dextral ridge slightly curved, descend- with scattered punctations. Cervical spine ing to cephalic margin of caudodextral region with 1 tubercle and several granules. wall; caudosinistral wall inflated, rounded Antennal peduncle devoid of spines or caudally and laterally, caudodextral wall tubercles; tip of adpressed antennal flagel- long, nearly horizontal, less inflated but lum reaching caudal margin of third tergite; broad; transverse tongue originating at antennular peduncle with very small, sub- broad median end of caudodextral wall, distal median spine on ventral surface of proceeding horizontally to plunge into deep basal podomere. Antennal scale (Fig. 1J) fossa beneath sinistral wall; sinus not dis- 2.9 times as long as wide, broadest near secting caudal margin. Mirror image of this midlength; lateral margin slightly convex, configuration observed in 7 of 10 females. thickened, terminating in long spine, tip of Measurements of type specimens provid- which reaching distal margin of penultimate ed in Table 1. podomere of antennular peduncle; lamella Description of holotypic male, form I.— 1.2 times width of thickened lateral portion, Body and eyes pigmented, eye 1.4 mm di- distal margin strongly declivous from base ameter. Cephalothorax (Fig. 1A, D) subcy- of spine, mesial margin subparallel to lat- lindrical, thoracic section 1.3 times wider eral. than deep. Areola nearly obliterated, 1 Abdomen narrow, greatest width 69.7% punctation in narrowest part, length consti- of greatest carapace width; length 89.5% of tuting 44.6% of TCL (49.4% of PCL). Ros- TCL; abdominal pleura fairly short, round- trum with relatively thick, strongly elevated ed ventrally and caudally; terga with many margins slightly converging to base of short large punctations, except articular surfaces acumen, where sharply constricted, more glabrous. Proximal podomere of uropod convergent and concave to small, dorsally without spines or tubercles; mesial ramus of directed apical tubercle; latter reaching uropod with weak median ridge bearing midlength of penultimate podomere of an- very small, submarginal caudomedian tennular peduncle; acumen comprising spine; caudolateral spine of ramus very 35.3% of rostrum length, latter constituting small; cephalic section of right lateral ra- 11.9% of TCL; floor of rostrum excavate, mus of uropod with weak ridge, caudal cephalically cuplike, sparsely punctate, margin of transverse flexure bearing total of with usual row of deep punctations along 19 fixed spines and 1 long, articulated sub- inner surface of dorsal ridge; subrostral lateral spine (15 fixed and 1 sublateral on ridge strong, visible to base of acumen in left). Telson with 2 spines in right caudo- 832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lateral corner of cephalic section, innermost with weak depression (continuous onto articulated, left corner with single fixed and fixed finger); lateral eminence of strong ar- 2 articulated spines; transverse flexure ticular ridge with large, depressed, subcon- strong, caudal margin broadly rounded. ical subdistal tubercle; 3 strong and 3 weak- Epistome (Fig. 1H) with subcordiform to er tubercles proximal to ridge (total of 13 subtriangular cephalic lobe bearing short obvious, produced tubercles on ventral sur- cephalomedian projection; margins of lobe face). Mesial margin of palm with subser- relatively narrow, moderately elevated, lat- rate mesial row of 6 strong tubercles, row eral corners produced as tuberclelike pro- subtended dorsolaterally by row of 6 gen- tuberances; floor (ventral surface) broadly erally smaller tubercles, and 4 others dorsal convex, very punctate; central depression to this row; 1 strong distal tubercle ventral shallow, with deep median fovea; lamellae to mesial row. very broad, moderately punctate, laterally Fingers gaping except at tips, greatest truncate, with 1 small tubercle and 1 strong width of gape slightly more than width of caudal tubercle each side; zygoma well dactyl base, largest tubercles on opposable arched, cephalolateral margins flanked by surfaces of fingers separated when fingers deep pit. closed. Fixed finger costate laterally, inner Third maxilliped with tip reaching mid- margin of proximal fourth scalloped; dorsal length of penultimate podomere of antennal surface with very strong, glabrous median peduncle; exopodite very hirsute, tip reach- ridge, flanked each side by deep punctate ing nearly to midlength of merus of endop- groove; lateral surface with row of large odite; distolateral corner of ischium sub- punctations; ventral surface with fairly acute, lateral half apunctate, glabrous; me- strong ridge, closer to opposable margin sial section very broad, with clumps of long than median in position, flanked mesially bristles, mesial margin of right ischium by row of coalescing punctations, laterally with 21 denticles, 20 on left; incisor ridge by slanted surface crowded with large of right mandible with 8 denticles, 7 on left. punctations; opposable surface with rela- Right cheliped regenerated; left chela tively small subconical tubercle ventral to (Fig. 1L) 1.9 times longer than wide, palm denticles at base of distal third or so of fin- 1.9 times as wide as deep, width 1.8 times ger, and row of 5 other tubercles dorsal to length of mesial margin; latter 29.4% of to- or interrupting denticles, second from base tal chela (propodus) length, 44.5% of dactyl largest; denticles in 1 or 2 rows. Dorsal sur- length. Dorsal surface of palm covered with face of dactyl with very strong, glabrous mostly deep punctations, those on mesial median ridge, flanked each side by deep, half and some on lateral half with small punctate groove; mesial surface with very basal tubercles; longitudinal dorsomesial strong, broad tubercles on proximal three- sulcus weak, with row of small, produced fourths, basal tubercles in 3 rows, strongly tubercles; dorsolateral margin of palm cos- encroaching dorsally, weakly encroaching tate for much of length (continuous onto ventrally; ventral surface with moderate, fixed finger), with slight impression and glabrous ridge, flanked each side by row of large punctations, some with tubercles at punctations; opposable surface with 9 tu- proximal bases; articular ridge strong, lat- bercles, third from base largest, and exci- eral eminence with dense, short setae on sion in surface proximal to this tubercle; distolateral margin, continuing onto base of denticles in 1 row. opposable surface of fixed finger; lateral Carpus (Fig. 1L) 1.4 times as long as margin of palm with row of large puncta- wide, 1.4 times as long as mesial margin of tions. Ventral surface of palm (Fig. IK) palm; dorsal surface of carpus with narrow, very punctate, mesial portion sharply set off deep sulcus, lateral to which surface punc- from greatly inflated portion; distolaterally tate, mesial to which surface with 3 dor- VOLUME 116, NUMBER 3 833 somesial tubercles; mesial margin with short, stout distal spine. Right merus with short, stout distal spine, 1 moderate tubercle 1 moderate and 4 weak, rounded distodorsal close to its proximal margin, and 9 other tubercles (3 moderate tubercles on left); strong, often subacute, tubercles; ventral ventromesial ridge with 10 tubercles (9 on surface with 2 subconical distal mounds, left), in addition to distal spine; ischium and 1 strong and 1 smaller proximomesial with 2 small ventral tubercles each limb. tubercle. Merus 1.7 times longer than deep, First pleopods strong, distally hirsute; distodorsal surface with 1 large, broad ad- preannular sternite relatively narrow, walls pressed tubercle and 1 smaller tubercle; steep (see "Affinities"); ventral surface of ventrolateral ridge with 6 small, rounded tu- postannular sclerite (Fig. II) with large, bercles, none on articular condyl, which subconical papilla. rounded, glabrous; ventromesial ridge with See "Diagnosis" for description of an- 8 subacute tubercles and short, stout distal nulus ventralis. spine; ischium with 4 small, subacute tu- Description of morphotypic male, form bercles on ventral ridge. II.—Differing from holotype in following See "Diagnosis" for description of gon- respects: Areola obliterated, constituting opods. 44.2% of TCL (49.6% of PCL). Ventral Description of allotypic female.—Except keel of rostrum with single spine near base. for secondary sexual characters, differing Branchiostegal spine obsolete. Cephalic from holotypic male in following respects: section of carapace 1.3 times longer than Areola obliterated, constituting 43.8% of areola, constituting 55.8% of TCL. Greatest TCL (49.2% of PCL). Acumen comprising width of abdomen 69.7% of greatest width 37.8% of rostrum length, latter constituting of carapace, abdomen length 88.4% of 12.8% of TCL. Postorbital ridge fairly TCL. Telson with 1 very small fixed spine weak, nowhere sharply defined. Cephalic and 1 minuscule articulated spine in right section of carapace 1.3 times longer than caudolateral corner of cephalic section, all areola, constituting 56.2% of TCL. Cervical spines congenitally absent from left. Lateral spine region with 5 small tubercles and sev- corner of lamella of epistome devoid of tu- eral granules. Greatest width of abdomen bercles; zygoma moderately arched. 73.8% of greatest carapace width, length Right cheliped regenerated; palm of left 91.0% of TCL. Telson with single fixed chela 1.8 times broader than deep; length of spine in each caudolateral corner of ce- mesial margin 28.7% of total chela length, phalic section, transverse flexure weak, cau- 44.3% of dactyl length; ventral surface of dal margin domed. palm covered with tubercles of various siz- Palm of chela 1.8 times broader than es; subconical tubercle on lateral eminence deep, width 45.7% of length of mesial mar- of articular ridge originating at proximal gin, latter 29.8% of total chela length, margin of ridge, not adpressed; 3 large tu- 45.7% of dactyl length; dorsomesial sulcus bercles proximal to ridge. Largest tubercle of palm obliterated; ventral surface with to- on opposable surface of fixed finger second tal of 10 obvious, produced tubercles. Op- from base, largest on opposable surface of posable surface of right fixed finger with dactyl fourth from base. Carpus length 1.5 broken subconical tubercle ventral to den- times length of mesial margin of palm; dor- ticles at base of distal fourth of finger, row sal sulcus short, flanked mesially by row of of 8 other tubercles (6 on left finger), third 3 tubercles; mesial surface with total of 11 from base massive. Opposable surface of tubercles in addition to usual stout subdistal right dactyl with 10 tubercles, fourth from spine; ventral surface with single proxi- base massive. Carpus 1.3 times as long as momesial tubercle. Distodorsal surface of wide, mesial surface with 10 tubercles of left merus with 1 moderate, subconical and varying sizes and shapes in addition to 2 smaller, rounded tubercles; ventrolateral 834 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ridge with 5 very small tubercles, none on access road near western boundary Albany distal condyl; ventromesial ridge with 11 Nursery Wildlife Management Area, ca. small, conical tubercles. 12.8 km W of Albany (Pretoria 7.5" USGS Hook on ischium of third pereiopod quadrangle, UTM Zone 16, coord. moderate, not overreaching articulation, op- 751779E, 3496706N). Locality within sub- posed by extremely weak tubercle on ven- drainage of Kiokee Creek, Flint River ba- tral surface of basis. In situ gonopods with sin, Apalachicola River drainage. moderate, nearly abutted proximomesial Disposition of types.—The holotypic apophyses; in lateral aspect (Fig. IF), cen- male, form I, allotypic female, and morpho- tral projection broad, slightly tapered, not typic male, form II, are in the NCSM crus- reaching as far caudally as tip of mesial tacean collection (7997, 7998, and 7999, re- process; latter inflated nearly entire length, spectively). The following are designated directed caudally, tip subtruncate and with paratopotypes: 1 jS, 3 9 (5253), 22 Jul small, spiniform protuberance directed cau- 1999, coll. CES, S. E. Cammack, E. E. Van domesially; juvenile suture absent; in me- De Genachte; 4 2 j$ (7987), 25 Feb sial aspect (Fig. IE), short setae at mid- 2001, coll. CES; 1 6 I (7988), 13 Aug 1999 length of shaft. (was 6 II when collected), coll. CES, S. K. Color notes.—Most specimens discol- Berckman; 1 6 I, 1 $ (USNM 1004610), ored by ferrous deposits; the following 2 Jun 1999, coll. CES, M. C. Freeman based on a live adult female. (MCF); 1 $ (GMNH 6239), 2 Jun 1999, Ground color of carapace brown, fading coll. CES, MCF; 1 $ (23692); 14 May ventrally to pale greenish-brown, ventral 2002, coll. CES. margin orangish; cephaloventral area of lat- Variations.—Some variations other than eral surface of cephalic section of carapace those previously addressed have been ob- creamy orange. Coxa, basis, and ischium of served. In dorsal aspect, the margins of the cheliped orangish; merus brownish, fading rostrum are usually rather straight, and sub- to orangish ventrally and proximally; car- parallel or only moderately convergent to pus brown dorsally, orangish ventrally, with the base of the acumen. In several speci- orangish dorsomedian furrow. Large distal mens, though, the margins are slightly con- tubercle on mesial surface of carpus orange, cave between the orbits, and in two very all others orange-cream. Dorsal surface of small females the rostrum is triangular. chela greenish-brown, articular ridge or- These juveniles also have marginal spines ange; tips of both fingers orange, not sub- on the rostrum. In all specimens, the mesial tended by black band; tubercles on mesial (lamellar) portion of the antennal scale is margin of palm and dactyl orangish; lateral only slightly wider than the thickened lat- surface of propodus orangish. Largest tu- eral portion, and the distomesial margin is bercle on opposable surface of dactyl and moderately or strongly declivous from the fixed finger cream; other tubercles on both base of the spine to the widest point. In fingers orange-cream. Proximodorsal part some specimens the distomesial margin is and merus of second through fifth pereio- deeply incised, bearing decidedly spinelike pods orangish, fading to brownish on basis protuberances. In ventral outline, the ce- and ischium; ventral surfaces of these pe- phalic lobe of the epistome varies in shape reiopods orangish. Dorsal surface of abdo- from subcordiform or subtriangular to sub- men brown; cephalic part of all pleura pink- pentagonal, and in some specimens the ish-orange, color most obvious on cephalic margins are erose, with small angular pro- two-thirds of second abdominal segment; trusions. The central depression of the epi- uropods and telson light brown. stome is obsolete in 10 specimens, very Type locality.—Georgia, Dougherty shallow in the others, and always displays County, burrows in wetland just south of a deep fovea. VOLUME 116, NUMBER 3 835

The greatest width of the abdomen varies mens yet collected are two females with from 69.1-74.8% (x = 72.2%) of carapace TCLs of 7.0 and 7.7 mm. width, and is only slightly wider in females Life history notes.—Form I males were than in males. The length of the abdomen collected in May and July. A form II male, varies from 86.5-95.2% (pc = 90.3%) of collected in August 1999, underwent two TCL. The number of spines in the caudo- molts in the laboratory, became form I in lateral corner of the cephalic section of the January 2000. No ovigerous females or telson is highly variable. Four specimens those with attached young have yet been have a single fixed lateral spine and a single found. The two very small females, possi- articulated inner spine in each corner. Four bly representative of recruitment size, were have two spines in one corner, and either a dug from the same burrow on 25 February single fixed or articulated spine in the other. 2001. One specimen has no spines in one corner, Ecological notes.—The type locality of two in the other; another has two spines in C. (D.) doughertyensis is in a swamp forest one corner, three in the other; and two spec- dominated by an overstory of Quercus spp., imens have a single fixed spine in each cor- Nissa biflora, and Acer rubrum, while the ner. In most specimens the articulated understory has abundant Crataegus aesti- spines are very small. valis, Sebastiana fruticosa, Rhus radicans, and a variety of graminoids. The area is In most specimens the punctations on at seasonally flooded, and in very wet periods least the mesial half or third of the ventral produces a shallow outflow to Kiokee surface of the palm have small basal tuber- Creek. Burrows are typically found just cles, but the surface also bears from 6-13 above the standing water mark among roots prominent, produced tubercles, several of of small trees and Serenoa repens. The soils which may be very large. The largest tu- in the area are Grady clay loams and pro- bercle on the opposable surface of the fixed vide easy digging for the first 25 cm. Below finger is almost always the third (rarely the that, the soil becomes much more dense and second or fourth) from the base; the largest is more difficult to excavate. When the wa- tubercle on the corresponding surface of the ter table is high, burrows are anastomosed dactyl is usually the fourth from the base, and usually have four or five openings, of- but often is the third, and occasionally the ten marked by well developed chimneys ap- first and fourth tubercles are equally large. proximately 10 cm high and 15 cm across. The total number of tubercles or spines on In this situation, the are most often the carpus, excluding the stout distal spine found in one of the horizontal passages of the mesial surface and the two stout dis- about 30 cm below the surface. As the wa- tal tubercles or mounds of the ventral sur- ter table drops, burrows are marked by only face, ranges from 7-13 (usually 10-13). two or three openings, with chimneys, that The ventral surface of the ischium of the angle down to a single subvertical passage cheliped bears from 2-4 (usually 3 or 4) that penetrates the water table. The animals small, subacute tubercles. The total length are then found in the vertical passage, 5 to of the chela of adult males averages about 10 cm below the water level. When a bur- 85% of TCL (94% of PCL), while the av- row is excavated, the crayfish can rarely be erage for adult females is about 75% of induced to come to the air-water interface, TCL (84% of PCL). and when encountered they are unmoving Size.—The largest specimen is a female and rarely try to escape. measuring 34.5 mm TCL (31.0 mm PCL). Crayfish associates.—Faxonella clypea- The largest form I male measures 31.0 mm ta (Hay, 1899) was collected in open water TCL (27.5 mm PCL), the smallest 28.5 mm in the swamp forest when flooded, and was TCL (25.7 mm PCL). The smallest speci- dug from simple burrows when the water 836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON table dropped. Two female Cambarus (De- IK), proximal to the articular ridge, bears pressicambarus) cf. C. (D.) striatus Hay, many prominent, produced tubercles, 1902 (5157, 23691) and a female Camba- whereas only a few such tubercles are pre- rus (Lacunicambarus) diogenes Girard, sent in C. (D.) harti. The opposable surface 1852 (23690) were dug from separate bur- of the fixed finger of C. (D.) dougherty ensis rows in the floodplain of Little Kiokee bears a row of from 5-8 tubercles in addi- Creek, not far from the C. (D.) dougher- tion to the usual subconical tubercle ventral tyensis site. to the denticles, and there is a wide space Affinities.—Cambarus (D.) doughertyen- between the base of the finger and the prox- sis shares so many similarities with C. (D.) imalmost tubercle. In C. (D.) harti, the harti and several of the other small Georgia number of tubercles that lie dorsal to the burrowers that, although some of them can denticles is 4 or 5 (6 in one specimen). The undoubtedly be attributed to convergence opposable surface of the dactyl of C. (D.) and the channeling effect of adaptation to doughertyensis bears 9—12 (usually 9 or 10) an obligate burrowing existence, it seems tubercles, whereas that of C. (D.) harti reasonable these species have descended bears 5—7 tubercles. When the fingers of the from a common proximate ancestor. The chela are closed in C. (D.) doughertyensis, annuli ventrales of some of the species are the largest tubercles on the opposable sur- very similar, as are their narrow abdomens, faces are separated from each other, but in many aspects of their chelipeds, rostra, an- C. (D.) harti the largest tubercles either tennal scales, suborbital angles, and bran- abut or overlap. The carpus of C. (D.) chiostegal spines, and other features. Some, doughertyensis usually bears from 7—13 including C. (D.) harti, also have spines on spines or tubercles in addition to the large, the ventral keel of the rostrum. They differ stout distomesial spine and the two broad significantly, however, in many ways. distoventral structures, whereas that of C. In C. (D.) doughertyensis the areola con- (D.) harti has 3 or 4 tubercles in addition stitutes 40.9-45.6% (jc = 43.7%) of TCL to the usual. The former species has a short- and 46.2-51.2% (jc = 48.9%) of PCL, while er, deeper merus, in which the length av- in C. (D.) harti it constitutes 38.2-40.3% erages 1.7 times the greatest depth, whereas (jc = 39.4%) of TCL and 44.2-45.8% (jc = in C. (D.) harti the merus averages twice as 44.8%) of PCL. Correspondingly, the ce- long as deep. phalic section of the carapace in the former The following species differ from C. (D.) species is 1.2-1.4 times as long as the are- doughertyensis as indicated: C. (D.) cyma- ola, while in the latter it is 1.5—1.6 times as tilis has a strong, usually subacute subor- long. The proximal podomere of the uropod bital angle, the caudomedian spine on the in C. (£>.) harti usually bears a small spine mesial ramus of the uropod extends well on both the lateral and mesial lobes, but beyond the caudal margin, and the color is such spines are lacking in C. (£>.) dough- blue; C. (D.) reflexus has a long, strongly erty ensis. In C. (D.) harti, the preannular recurved central projection and a well-de- sternite of the female is broader and flatter fined caudal knob, both on the form I male than that of C. (£).) dougherty ens is, which gonopod, and the color is reddish or blue; is narrow and steep. The width of this ster- C. (D.) strigosus has a row of plumose se- nite, i.e., the distance between the ventral tae on the caudal border of the mesial pro- angles of the walls at their articulations cess of the form I male gonopod, and a sin- with the mesial surfaces of the fourth cox- gle spine in each cephalolateral corner of ae, is about 38% of carapace width in the the telson; and C. (D.) truncatus usually former species, about 31% in the latter. lacks both caudomedian and caudolateral The ventral surface of the palm of the spines on the mesial ramus of the uropod cheliped of C. (D.) dougherty ensis (Fig. and a transverse flexure in the telson, and VOLUME 116, NUMBER 3 837

the form I male gonopod has a very short vea. . . . On page 107, column 1, paragraph central projection. 1, line 10 says "ventrolateral row of setae The members of some Georgia popula- on carpus reduced to 2. . . ." This should tions of C. (D.) striatus and Cambarus (De- read: ventrolateral row of tubercles on mer- pressicambarus) latimanus (LeConte, us reduced to 2. The next line says "mesial 1856) display multiple strong tubercles on surface of ischium of cheliped devoid of tu- the carpus and the ventral surface of the bercles," which should read: mesial and chela. In these species, however, there is no ventral surface of ischium. . . . These latter excision in the opposable surface of the statements (p. 107) apply to the "morpho- dactyl; there is a tubercle or spine on the typic male, form II," but the specimen ac- ischium of the antennal peduncle; both tually appears to be a juvenile rather than lobes of the proximal podomere of the uro- an adult male of the second form. pod bear a spine; the rostrum is longer, and the abdomen is longer and much broader; Acknowledgments the antennal scale is subquadrate, with a Our sincerest appreciation is expressed to broader lamella; and the annulus is sub- S. K. Berckman, S. E. Cammack, E. E. Van rhomboidal rather than subovate in outline. De Genachte, and M. C. Freeman for as- Remarks—As far as we can determine, sisting CES in the field, to T. S. Patrick for the presence of spines or acute tubercles on characterizing the vegetation at the type lo- the ventral keel of the rostrum, while a not cality, and to G. Henry for providing access uncommon feature in some , to the type locality. The USNM types of C. has never been reported in freshwater cray- (D.) harti were examined through the cour- fishes except, as mentioned, some Mexican tesy of K. J. Reed and R. Lemaitre. The of the subgenera Paracam- manuscript was improved by the comments barus and Villalobosus. An investigation of of R. Franz, G. Schuster, C. A. Taylor and the presence or absence of this character in R. Lemaitre, and publication was facilitated other U.S. species is just getting underway, with funds provided by the Georgia De- but it has to date been found in several oth- partment of Natural Resources, Wildlife er members of subgenus Depressicamba- Resources Division. rus, including C. (D.) harti, and in one spe- cies of subgenus Erebicambarus. Literature Cited Etymology.—This species is named for both the physiographic province and the Bouchard, R. W. 1978. , ecology, and phy- county in which it appears to be endemic. logeny of the subgenus Depressicambarus, with Suggested vernacular name: Dougherty the description of a new species from Florida burrowing crayfish. and redescriptions of Cambarus graysoni, Cam- barus latimanus, and Cambarus striatus (De- capoda: Cambaridae).—Alabama Museum of Cambarus (Depressicambarus) harti Natural History Bulletin 3:26-60. Hobbs, 1981 Erichson, W. F. 1846. Uebersicht der Arten der Gattung Astacus.—Archiv fur Naturgeschichte 12(1): Some inadvertent errors appeared in the 86-103. Girard, C. 1852. A revision of the North American description of this species (Hobbs 1981: Astaci, with observations on their habits and 104-109). On page 104, column 2, para- geographical distribution.—Proceedings of the graph 2, part of the first sentence has been Academy of Natural Sciences of Philadelphia 6: omitted. It should read: Cephalic lobe of 87-91. epistome (Figure 45g) subrhomboidal with Hay, W. P. 1899. Description of two new species of crayfish.—Proceedings of the United States Na- short cephalomedian projection and scal- tional Museum 22(1187): 121-123. loped margins; main body with broad, shal- . 1902. Observations on the fauna low central depression lacking distinct fo- of Nickajack Cave, Tennessee, and vicinity.— 838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Proceedings of the United States National Mu- LeConte, J. 1856. Descriptions of new species of As- seum 25( 1291 ):417-439. tacus from Georgia.—Proceedings of the Acad- Hobbs, H. H., Jr. 1969. On the distribution and phy- emy of Natural Sciences of Philadelphia 7:400- logeny of the crayfish genus Cambarus. Pp. 93- 402. 178 in P. C. Holt, R. L. Hoffman, and C. W. Ortmann, A. E. 1905. Procambarus, a new subgenus Hart, Jr., eds., The distributional history of the of the genus Cambarus.—Annals of the Car- biota of the southern Appalachians, Part I: In- negie Museum 3(3):435-442. vertebrates. Research Division Monograph 1, . 1906. Mexican, Central American and Cuban Virginia Polytechnic Institute, Blacksburg, 295 Cambari.—Proceedings of the Washington pages. Academy of Sciences 8:1-24. . 1970. New crayfishes of the genus Cambarus Villalobos, A. 1955. Cambarinos de la Fauna Mexi- from Tennessee and Georgia (Decapoda, Asta- cana: Crustacea Decapoda. Tesis, Facultad de cidae).—Proceedings of the Biological Society Ciencias, Universidad Nacional Autonoma de of Washington 83(23):241-259. Mexico, 290 pp. . 1972. The subgenera of the crayfish genus . 1983. Crayfishes of Mexico (Crustacea: De- Procambarus (Decapoda: Astacidae).—Smith- capoda). Translation of Villalobos 1955 by H. sonian Contributions to Zoology 117:1-22. H. Hobbs, Jr. Smithsonian Institution Libraries . 1981. The crayfishes of Georgia.—Smithson- and National Science Foundation, Washington, ian Contributions to Zoology 318:1-549. D.C., 276 pp.