31 July 2000 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 113(2):431-442. 2000. Cambarus (Cambarus) davidi, a new species of crayfish (Decapoda: Cambaridae) from North Carolina John E. Cooper North Carolina State Museum of Natural Sciences, Research Lab, 4301 Reedy Creek Road, Raleigh, North Carolina 27607, U.S.A. Abstract.—Cambarus (Cambarus) davidi is a new species of crayfish from the eastern Piedmont Plateau of North Carolina, where it is restricted to inter- mittent streams, seepage areas, springs, and burrows. Although the ranges of the two species appear to be broadly disjunct, C. (C.) davidi is most closely related to the ecologically more tolerant Cambarus (C.) bartonii (s.l.), and especially resembles some members of the controversial subspecies C. (C.) b. cavatus. Cambarus (C.) davidi is distinguished by a suite of characters that includes a vaulted carapace; a deeply excavate, ladlelike rostrum; a very nar- row, sparsely punctate areola; and an obtuse to nearly obsolete suborbital angle that almost always bears a small tubercle.

On 18 August 1993, David G. Cooper highway; NCSM, North Carolina State Mu- collected several specimens of a Cambarus seum of Natural Sciences, Raleigh; PCL, from under large rocks in a shallow, inter- postorbital carapace length; R, river; SR, mittent tributary of the Neuse River, Wake state secondary (county) road; TCL, total County, North Carolina. In the field they carapace length; US, United States high- appeared to be aberrant individuals of the way; USGS, United States Geological Sur- burrowing species, Cambarus (Depressi- vey; and UTM, Universal Transverse Mer- cambarus) reduncus Hobbs, 1956, which is cator coordinates. not uncommon in such habitats in the upper Neuse River watershed. In the laboratory, however, I was surprised to discover that Cambarus (Cambarus) davidi, the specimens belonged to the subgenus new species Cambarus, and to some species previously Fig. 1, Table 1 unknown from the Neuse River basin, Diagnosis.—Body and eyes pigmented, whose crayfish fauna is well documented eye small (X adult diam 1.7 mm, n = 30). (Cooper & Ashton 1985, Cooper & Bra- Carapace vaulted, thoracic section averag- swell 1995, Cooper & Cooper 1995). Since ing 1.3 times wider than deep (n = 52). that time, many additional specimens from Rostrum acarinate; margins elevated, sub- the Neuse and Cape Fear river basins have parallel, caudally thickened, strongly to either been collected or have been recog- moderately constricted at base of acumen, nized in prior collections. They belong to lacking marginal spines or tubercles; floor an undescribed species of Cambarus that (dorsal surface) of rostrum deeply concave, seems to have its closest affinities with con- ladlelike; acumen 24.5 to 49.1% (X = geners that occur in the Tennessee and Ohio 33.6%, n = 52) length of rostrum, latter river drainages. 13.0 to 19.3% (X = 16.2%, n = 52) of Abbreviations used in the text are as fol- TCL. Areola 5.2 to 14.5 (X = 8.1, n = 76) lows: j, juvenile; NC, North Carolina State times longer than wide, constituting 35.4 to 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 41.8% (X = 37.6%, n = 76) of TCL and like proximomesial apophyses; proximola- 42.2 to 47.3% (X = 44.1%, n = 45) of teral portion of gonopod set off from rest PCL; areola sparsely punctate, with 2 {n — of shaft by weak groove; in lateral aspect 22) to 3 (n = 64) punctations across nar- (Fig. IB), central projection curved over rowest part. Cervical spines reduced to 90° to plane of shaft, untapered, with prox- multiple tubercles. Branchiostegal spine re- imally directed subapical notch; mesial pro- duced to small tubercle; hepatic and sur- cess inflated, symmetrically tapered, slight- rounding regions of carapace crowded with ly constricted at base of caudal third, with tubercles. Suborbital angle obtuse to nearly subacute, caudally directed apex extending obsolete, almost always bearing small tu- slightly farther caudally than apex of cen- bercle; postorbital ridge short, cephalic tral projection; caudal process reduced to margin rounded and usually devoid of tu- swelling at caudodistal margin of shaft; in bercle. Antennal scale 2.0 to 3.6 (X = 2.5, mesial aspect (Fig. 1C), distal portion of n = 50) times as long as broad, widest just gonopod with essentially flat surface, few distal to midlength, lateral margin thick- setae at midlength. ened and with long distal spine. Annulus ventralis of allotypic female Palm of chela of cheliped 1.5 to 1.8 (X (Fig. 1H) 1.3 times broader than long, gen- = 1.6, n = 51) times wider than deep, width erally symmetrical and subrhomboidal; ce- 1.3 to 1.7 (X = 1.5, n = 51) times length phalic margin convex and fused to sternum, of mesial margin; dorsolateral margin cos- caudal margin subangular, free and capable tate distally, without impression; mesial of slight movement; cephalic half of annu- margin of palm with 2, rarely 3, rows of lus depressed, sloping, with narrow median tubercles: mesial row of 6 to 8 (usually 7) trough, flanked each side by low, poorly de- large, generally adpressed tubercles, sub- fined ridge; ridges diverging caudally, si- tended dorsally by row of 1 to 5 (usually 4 nistral ridge continuing obliquely and ter- or 5) smaller tubercles. Fixed finger of che- minating before joining caudosinistral wall, la costate laterally, with well defined lon- dextral ridge curving caudodextrally to gitudinal ridges dorsally and ventrally; op- merge with upper arm of heavy, C-shaped posable surface of finger with row of 4 to caudodextral wall; transverse tongue prom- 11 (usually 5 or 6) tubercles in addition to inent, originating from sinistral wall near subconical tubercle; dactyl 1.7 to 2.6 (X — caudal midline, continuing cephalically and 2.0, n = 51) times as long as mesial margin slightly obliquely before turning dextrally of palm, with strong longitudinal ridge dor- and plunging into fossa beneath dextral sally, weaker ridge ventrally; mesial margin wall; sinus dissecting caudal wall near mid- with prominent tubercles; opposable sur- line. face with row of 6 to 14 (usually 7 to 9) Measurements of type specimens provid- tubercles. Carpus of cheliped generally ed in Table 1. lacking dorsomesial tubercles; merus with Description of holotypic male, form I.— prominent multiple dorsodistal tubercles Body and eyes pigmented, eye 1.8 mm and often row of small squamous tubercles diam. Cephalothorax (Fig. 1A, D) subcy- along dorsal ridge. lindrical, thoracic section 1.2 times wider Hook on ischium of third pereiopod of than deep. Areola 9.2 times longer than males, that of form I male (Fig. IK) uni- wide, constituting 39.3% of TCL (45.1% of ramous, overreaching basioischial articula- PCL), sparsely punctate, with 3 punctations tion and opposed by tubercle on basis; coxa across narrowest part. Rostrum acarinate, of fourth pereiopod of males with vertically with slightly thickened caudal margins disposed, caudomesial boss. moderately converging to base of acumen, In situ gonopods (Fig. 1G) symmetrical, where moderately constricted; margins of with abutted or slightly separated, tubercle- acumen slightly concave and converging to VOLUME 1 13, NUMBER 2 433

Fig. 1. Cambarus (Cambarus) davidi, new species (all from holotypic male, form I, except E, F, from morphotypic male, form II, and H, from allotypic female): A, lateral aspect of carapace; B, E, lateral aspect of gonopod (first pleopod); C, F, mesial aspect of gonopod; D, dorsal aspect of carapace; G, caudal aspect of in situ gonopods; H, annulus ventralis and postannular sclerite; I, dorsal aspect of distal podomeres of right cheliped; J, antennal scale; K, hook on ischium of third pereiopod; L, epistome. dorsally directed apical tubercle, which floor of rostrum excavate, ladlelike, mod- reaching just beyond midlength of penulti- erately punctate, and ascending caudally mate podomere of antennular peduncle; into broad dorsomedian depression of car- acumen comprising 30.2% of rostrum apace; subrostral ridge strong, visible to length, latter constituting 14.2% of TCL; base of acumen in dorsal aspect. 434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements (mm) of types of Cam- with prominent caudomedian spine on me- barus (Cambarus) davidi, new species. sial lobe; mesial ramus of uropod with me- Holotyp- Morpho- dian keel bearing strong caudal spine, tip of ic Allotypic typic male female male which reaching caudal margin of ramus, caudolateral margin with small spine; lat- Carapace eral ramus with submedian keel bearing ter- Total length 30.3 36.5 30.7 Postorbital length 26.4 31.6 26.6 minal spine at transverse flexure, latter with Length cephalic section 18.4 22.4 19.0 total of 17 fixed spines along margin and 1 Width 15.7 18.4 15.3 long, movable sublateral spine. Telson with Depth 12.6 15.1 12.2 1 long stationary and 1 long articulated Length rostrum 4.3 5.6 4.9 spine in each caudolateral corner of ce- Length acumen 1.3 2.2 1.2 Length areola 11.9 14.1 11.7 phalic section; caudal margin domelike. Width areola 1.3 2.1 1.9 Epistome (Fig. 1L) with symmetrical, Antennal scale subtriangular cephalic lobe bearing short Length 4.5 5.4 5.2 cephalomedian projection; margins of lobe Width 1.8 2.1 1.8 uninterrupted, moderately thickened, lateral Abdomen apices thicker than rest; floor (ventral sur- Length 29.6 37.7 30.6 face) of lobe punctate and slightly convex, Width 13.6 16.4 12.2 concave along lateral margins; transverse Cheliped basal sulcus complete; central depression of Length lateral margin chela 24.4 26.7 23.8 body broad, moderately deep, with cephal- Length mesial margin palm 7.3 8.8 7.5 omedian fovea; lamellae punctate, lateral Width palm 11.5 13.4 1 1.1 margins subtruncate, with 2 large subacute Depth palm 6.8 7.8 6.7 tubercles and 1 small rounded tubercle in Length dactyl 15.5 16.3 15.2 right caudolateral corner (1 large subacute Length carpus 10.0 11.1 9.5 Width carpus 7.2 8.0 6.7 and 1 small rounded tubercle on left); zy- Length dorsal margin merus 12.0 13.8 12.2 goma moderately arched, pits elongate. An- Depth merus 7.1 8.5 6.9 tennal peduncle with small cephalolateral tubercle on basis, small ventral tubercle on Gondopod length 8.4 N/A 8.0 ischium; antennular peduncle with small, laterally displaced subdistal spine on ven- Postorbital ridge strong, groove essen- tral surface of basal podomere. Antennal tially lateral, cephalic margin with vestigial scale (Fig. 1J) 2.5 times longer than wide, tubercle. Branchiostegal spine reduced to broadest just distal to midlength; lateral tubercle; suborbital angle obtuse, without margin thickened, terminating in long distal tubercle. Thoracic section of carapace dor- spine, tip of which reaching distal margin sally and dorsolaterally punctate, laterally of penultimate podomere of antennular pe- with large, scattered granules; cephalic sec- duncle; lamella ca. 1.3 times as wide as tion 1.5 times longer than areola and con- thickened lateral margin; distal margin of stituting 60.7% of TCL, laterally crowded lamella subtransverse for most of width, with large tubercles and with row of small then sloping to widest point; mesial margin tubercles along ventral margin of cephalic subparallel to lateral margin for most of section of cervical groove; gastric region length. mostly glabrous. Cervical spine region on Third maxilliped with tip of endopodite right with 4 large and 3 smaller tubercles reaching about midlength of penultimate (2 large and 2 smaller on left). Abdomen podomere of antennal peduncle; exopodite slightly narrower and shorter than cepha- hirsute, tip reaching base of distal two- lothorax. Proximal podomere of uropod thirds of merus of endopodite; cephalola- without spine or tubercle on lateral lobe, teral corner of ischium slightly produced, VOLUME 1 13, NUMBER 2 435 not spinelike; ventrolateral ridge flanked from base largest); denticles in 2 or 3 rows mesially by row of punctations bearing from near tip of finger to sixth tubercle moderately long setae; lateral half of ischi- from base, single row from there. Fixed fin- um with punctations bearing short setae, ger with very strong dorsomedian ridge punctations most abundant on proximal flanked mesially by deep punctate groove third; mesial half with long stiff setae large- and second narrower ridge, laterally by ly obscuring mesial margin, latter with 27 deep punctate groove; lateral margin with denticles on right. Right mandible with in- irregular row of large punctations; ventral cisor ridge bearing 7 denticles (6 on left). surface with very strong longitudinal ridge, Right chela (Fig. II) 2.1 times longer flanked both sides by row of large puncta- than wide; palm 1.7 times broader than tions; opposable margin with subconical tu- deep, width 1.6 times length of mesial mar- bercle ventral to denticles at base of distal gin; latter 29.9% of chela length, 47.1% of third of finger, and 5 tubercles proximal to dactyl length. Dorsal surface of palm punc- subconical one, third from base more prom- tate; distolateral margin of palm and lateral inent than others (4 tubercles on left finger, margin of fixed finger costate, area at junc- third from base very large); denticles in 2 ture of palm and finger with aggregation of or 3 rows from tip of finger to subconical large punctations creating slight impres- tubercle, single row from there. sion; lateral margin of palm rounded and Carpus (Fig. II) 1.4 times as long as with row of large punctations. Ventral sur- wide, 1.4 times as long as mesial margin of face of palm less punctate than dorsal, dis- palm; carpus dorsally with long, deep, tolateral area with moderate depression and slightly oblique sulcus, lateral and mesial to aggregation of large punctations; lateral which surface punctate; mesial margin with eminence of articular ridge with distal tu- large distal spine and prominent proximal bercle, none proximal to ridge. Mesial mar- tubercle; ventral surface with stout, rounded gin of right palm with mesial row of 7 ad- tubercle at lateral articular condyle, similar pressed tubercles, proximal 3 of which with distomedian tubercle, and 1 small tubercle elevated distal margins (same on left), me- proximomesial to latter. Right merus 1.7 sial row subtended dorsally by row of 4 times longer than deep; dorsal surface with smaller tubercles (5 on left) and ventrally 3 prominent and 2 smaller subdistal tuber- by 1 small, squamous distal tubercle (3 on cles (2 prominent and 1 smaller on left), left). and small squamous tubercles along much Fingers gaping in proximal three-fourths of dorsomedian ridge; ventrolateral ridge of length, greatest width of gape about four- with 2 small acute tubercles and 1 minus- fifths width of base of fixed finger; oppos- cule tubercle near articular condyle (1 small able base of fixed finger with tuft of short acute tubercle and minuscule distal tubercle setae; both fingers slightly curved distov- on left); ventromesial ridge with 10 spini- entrally in lateral aspect, dactyl very slight- form tubercles, distalmost one somewhat ly bowed in dorsal aspect. Mesial margin of larger than others; ischium with 3 small right dactyl bearing row of 4 prominent and ventral tubercles (4 on left). 4 weaker tubercles on proximal half, distal Palm and fingers of chela of second pe- half of margin punctate; dorsal surface of reiopod hirsute. Ventral margins of pleura dactyl with low, rounded longitudinal ridge, subtruncate or slightly rounded, caudoven- flanked mesially by punctate groove, later- tral corners slightly angular, caudal margins ally by row of large, spaced punctations; rounded; terga very punctate, except artic- ventral surface with similar longitudinal ular cephalic portions glabrous. Sternites ridge; opposable surface with 8 tubercles, between third and fourth coxae with very fourth from base very large and slightly dis- dense, matted setae, covering distal ends of placed ventrally (7 tubercles on left, third in situ gonopods. 436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gonopods (Fig. IB, C, G) as described trum less constricted at base of acumen, in "Diagnosis." Length of gonopod 27.7% converging at ca. 45°, acumen comprising of TCL (31.8% of PCL). 24.5% of rostrum length, latter constituting Description of allotypic female.—Except 16.0% of TCL. Suborbital angle obtuse and for secondary sexual characters, differing with very small tubercle. Cephalic section from holotypic male in following respects: of carapace 1.6 times longer than areola and Areola 6.7 times wider than long, consti- constituting 61.9% of TCL. Cervical region tuting 38.6% of TCL (44.6% of PCL). Acu- on both sides of carapace with 3 prominent men comprising 39.3% of rostrum length, and 1 moderate tubercle, ventralmost of latter constituting 15.3% of TCL. Suborbit- which on right side subacute. Transverse al angle nearly obsolete, with small tuber- flexure of lateral ramus of right uropod with cle. Cervical spine region on both sides of 16 spines in addition to sublateral one. Lat- carapace with 3 prominent tubercles, largest eral corners of lamellae of epistome with 1 of those on right side subacute. Caudolater- moderate tubercle on right, 2 on left; zy- al corner of cephalic section of telson with goma strongly arched. Antennal scale 2.9 2 spines on right, 3 on left. Cephalic lobe times wider than long, lamella ca. 1.5 times of epistome subcordiform; lamellae with width of thickened lateral margin, distal single tubercle each caudolateral corner; zy- margin moderately declivous. Length of goma strongly arched. Antennal peduncle palm of cheliped 31.5% of chela length, lacking tubercle on basis; antennal scale 2.6 49.3% of dactyl length. Mesial margin of times longer than wide, lamella about 1.5 palm with mesial row of 6 adpressed tu- times as wide as thickened lateral margin. bercles, subtended dorsally on right palm Incisor ridge of right mandible bearing 8 by row of 3 moderate tubercles (4 on left) denticles. Right chela 2.0 times longer than and ventrally by 4 small squamous tuber- wide, palm length 33.0% of chela length, cles (2 on left). Opposable surface of dactyl 54.0% of dactyl length. Opposable surface with 9 tubercles on right, 8 on left, basal- of right dactyl with 10 tubercles (9 on left); most largest. Opposable surface of fixed opposable surface of both fixed fingers with finger with 6 tubercles on right in addition 6 tubercles in addition to subconical one, to subconical one, 7 on left, third from base fourth from base largest. Carpus of cheliped largest. Carpus 1.3 times as long as palm. I.3 times as long as mesial margin of palm; Merus 1.8 times longer than deep; dorsal merus 1.6 times longer than deep, dorsal surface of right merus with patch of 5 surface with 2 prominent and 2 smaller sub- prominent subdistal tubercles and 5 smaller distal tubercles (same on left); ventrolateral tubercles proximal to them (2 moderate and ridge with 5 small tubercles (same on left), 6 smaller on left); ventrolateral ridge on ventromesial ridge with 8 spiniform tuber- right with 2 subacute and 2 other tubercles cles (9 on left). (2 small, and 1 minuscule articular tubercle, Annulus ventralis (Fig. 1H) as described on left); ventromesial ridge on right with 10 in "Diagnosis." In addition, first pleopods small tubercles (9 on left). short, reaching caudal margin of annulus Hook on ischium of third pereiopod when abdomen flexed; annulus about 3 weak, not overreaching basioischial articu- times wider than postannular sclerite, which lation, opposed by tubercle on basis; boss elongate, ventrally domed, punctate. on coxa of fourth pereiopod moderately de- Description of morphotypic male, form veloped. II.—Differing from holotypic male in fol- Gonopod (Fig. IE, F) length 26.1% of lowing respects: Thoracic section of cara- TCL. In situ gonopods with weak, separat- pace 1.3 times wider than deep. Areola 6.2 ed proximomesial apophyses; mesial pro- times as long as wide, constituting 38.1% cess noncorneous, bulbous, distal surface of TCL (44.0% of PCL). Margins of ros- creased near extruded tip; in lateral aspect, VOLUME 1 13, NUMBER 2 437 gonopod with weak juvenile suture; central these bars producing zigzag lateral stripe projection noncorneous, curved 90° to each side of abdomen. Juveniles generally shaft, tapered to subacute tip; mesial pro- paler than adults, color patterns in most re- cess tapered, triangular in outline, tip di- spects similar but more vivid. rected caudally and inclined slightly disto- Type locality.—North Carolina, Wake laterally. Setae on sternites between third County, small intermittent stream entering and fourth coxae not dense. cove along western shore of Falls Lake (im- Color notes.—Adult ground color usu- poundment of Neuse River), ca. 1.4 air km ally dark olivaceous, sometimes light NW of western end of NC 98 bridge & ca. brown with orange or tan overtones. Ce- 2.6 air km W of Stony Hill (Bayleaf 7.5' phalic section of carapace often lighter in USGS quadrangle, UTM zone 17, 3984850/ color than thoracic section. Hepatic region 712190). with orangish midlateral streak just cephalic The shallow stream, which lies at the to cervical groove. Margins of rostrum out- bottom of a steep ravine in a hardwood for- lined in tan; antennal scale pale orangish- est, is seasonally intermittent and has a tan with dark lateral margin, antennal fla- maximum width of about 1.2 m. All spec- gellae green. Most tubercles, spines, and imens from this locality were found in the granules of carapace and chelipeds tan to mud of shallow residual pools under large orangish. Dorsal surface of cheliped green- rocks when water levels were very low. ish or olivaceous; articular ridges of chela Disposition of types.—The holotypic pinkish or orangish, ventral surface of chela male, allotypic female, and morphotypic pale grayish-tan. Lateral surface of entire male are in the crustacean collection of the propodus (palm and fixed finger) strikingly NCSM (catalogue numbers NCSM C-4413, colored, varying from pinkish-tan to C-4414, and C-2656, respectively), as are creamy orange or yellow. Tips of fingers of paratypes consisting of 2 S I, 15 6 II, 6 j chelipeds pale orange or orangish-tan, color 6, 18 ?, and 5 j ?. not subtended by black band. Proximal po- Range and specimens examined.—Ap- domeres of other pereiopods pale tan to parently limited to the upper Neuse and light brown with darker mottling, distal po- Cape Fear river basins in the eastern Pied- domeres greenish or bluish, fingers of che- mont Plateau of North Carolina. Voucher lae of second and third pereiopods pale specimens (n — 107), all in the crustacean blue. Cephaloventral structures bluish-gray, collection at NCSM (catalogue numbers in except epistomal zygoma almost white. An- parentheses), have been collected at the fol- nulus ventralis usually pale, mottled with lowing localities. orange in one female. Neuse River Basin: Durham Co.; upper Cephalicmost tergite of abdomen with trib Little R near Durham, 1 6 II (C-4742), transverse dark brown or black rectangular 13 Feb 1995, coll. T. Cuffney. Orange Co.; band. Lighter colored adults and all juve- small intermittent stream in headwaters niles with dark, diagonal blotch each side West Fork Eno R at SR 1358, 2.4 air km E of caudal margin of thoracic section of car- of Carr, 3 j? (C-3425), 25 Jul 1995, coll. apace, blotches extending onto dorsolateral M.A. Hartman, M.E. Savacool. Wake Co.; surfaces of two adjacent tergites as short, seep entering small trib New Light Crk curved markings. Series of short, dark bars along SR 1918, 0.3 km SW of jet SR 1909, producing interrupted dorsolateral stripe ca. 7.4 air km NNE of Bayleaf, 2 janimals. Also in these one in the other. The lateral lobe of the smaller individuals, the setae on the oppos- proximal podomere of the uropod normally able surface of the fixed finger of the che- lacks a spine or tubercle, but 12 specimens liped are far longer and more dense than have a very small spine or acute tubercle they are in adults, often filling the space on this lobe. The spine on the mesial lobe between the fingers and obscuring the tu- of this podomere varies in size from very bercles on both. small to moderate, and is absent in one an- Size.—The largest specimens collected imal. are two females measuring 50.7 and 50.0 The chela of form I males is longer than mm TCL (44.4 and 44.5 mm PCL), both that of mature form II males and females, from the Haw River subdrainage of the averaging 81% of TCL. In form II males Cape Fear River basin. The next largest the average is 71.6% and in females it is specimen is a form I male, which measures 70.8%. The largest tubercle on the oppos- 42.5 mm TCL (37.3 mm PCL). The largest able surface of the fixed finger varies from form II male measures 33.4 mm TCL (28.7 the third to the fifth from the base, but in mm PCL). most specimens the fourth tubercle is much Life history notes.—A form I male was larger than the others. The largest tubercle collected at the type locality on 14 June on the comparable surface of the dactyl 1997, one was taken in Chatham County on varies from the first to the fifth, but in most 13 February 1998, and one was found in it is the fourth, and the largest tubercle is amplexus with a female in Alamance Coun- almost always offset toward the ventral sur- ty on 2 March 1976. No females with at- face. The dorsal surface of the merus bears tached ova or young have yet been seen, from one to eight prominent subdistal tu- but one measuring 32.5 mm TCL, taken on bercles, with the usual number being three 4 April 1997, had all cement glands highly or four. Most specimens have from two to developed. four weaker tubercles just proximal to the Crayfish associates.—Seldom have other more prominent ones, and many have squa- crayfishes been found in the same habitats mous to subsquamous tubercles along at with C. (C.) davidi. At a few localities, least part of the dorsal ridge. The number however, a number of specimens of Cam- of tubercles on the ventrolateral ridge of the barus (Depressicambarus) latimanus (Le- merus ranges from two to five (usually Conte, 1856), and of at least one of the spe- three), and the distalmost is very small or cies in the complex subsumed under Cam- vestigial. The number of tubercles on the barus (Puncticambarus) acuminatus Faxon, ventromesial ridge ranges from six to elev- 1884, have been found. At two sites, juve- en (usually nine or ten), and the distalmost nile C. (D.) reduncus were collected. While is seldom much larger than the largest of juvenile Cambarus (Lacunicambarus) di- the others. ogenes Girard, 1852, have been taken from In 17 adult females the width of the an- under cover near the mouth of the stream nulus ranges from 1.2 to 1.8 (X — 1.5) at the type locality, and chimneyed burrows times its length, and the cephalomedian of this species may be seasonally abundant trough varies from moderately wide to along the nearby shoreline of Falls Lake, nearly obliterated. The thick, C-shaped wall this burrowing species has not been found of the annulus, beneath which lies the deep- with C. (C.) davidi. est part of the fossa, is dextral in 22 fe- Relationships.—It appears to me that C. males, sinistral in eight others. (C.) davidi has its strongest affinities with The floor (dorsal surface) of the rostrum Cambarus (Cambarus) bartonii (Fabricius, of juveniles and some subadults, while no- 1798). Students of American crayfishes, tably excavate, is often less ladlelike than however, have long been cognizant of the 440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON taxonomic perplexities presented by the signed to C. (C.) b. bartonii (s.l.). Camba- broadly distributed and highly variable pop- rus (C.) davidi differs from C. (C.) b. ca- ulations currently assigned to this species. vatus and most populations of C. (C.) b. Two subspecies have been described, Cam- bartonii (s.l.) in its obtuse to nearly obso- barus (Cambarus) bartonii cavatus Hay, lete suborbital angle, the shape of its anten- 1902, and Cambarus (Cambarus) bartonii nal scale, and the presence of multiple tu- carinirostris Hay, 1914, but opinions anent bercles on the dorsal surface of the merus their validity have for years fluctuated (see of the cheliped. In some montane popula- Faxon 1914, Ortmann 1931, Hobbs 1972, tions of C. (C.) b. bartonii (s.l.) the cara- 1989, Bouchard 1976, Thoma & Jezerinac pace is relatively vaulted and the rostrum is 1982, Jezerinac 1985, Fitzpatrick 1983, Jez- deeply excavate and somewhat ladlelike. erinac & Thoma 1984, Jezerinac et al. These populations, however, characteristi- 1995, Cooper 2000). The status of C. (C.) cally have areolae that are broader and have b. cavatus remains controversial, but C. more punctations across the narrowest part (C.) b. carinirostris has been elevated to than does the areola of C. (C.) davidi. Also, species status (Thoma & Jezerinac 1999). in nearly all C. (C.) bartonii (s.l.) the la- Although it is still difficult at this time to mella of the antennal scale is much narrow- establish precise diagnostic parameters for er than it is in C. (C.) davidi, and its distal these taxa throughout their ranges (what- margin is usually quite declivous from the ever those ranges might be), current diag- base of the lateral spine to the mesial mar- noses must be used in assessing the rela- gin. In addition, C. (C.) bartonii (s.l.) sel- tionships of C. (C.) davidi. dom displays prominent multiple tubercles In his brief description of C. b. cavatus, on the dorsal surface of the merus. whose type locality is the Powell River Roger F. Thoma, whose knowledge of (Tennessee-Ohio river drainage) near Ta- subgenus Cambarus is extensive, has sug- zewell, Claiborne County, Tennessee, Hay gested that a comparison of C. (C.) davidi (1902:435) emphasized its "deeply exca- with the burrower, Cambarus (Cambarus) vated rostrum," an areola that is "narrower ortmanni Williamson, 1907, would be use- and more thickly punctate than in C. bar- ful. That species differs from C. (C.) davidi toni bartoni," and a carapace that is "more in many ways, including the following: nearly cylindrical." Except for the "more Areola of C. (C.) ortmanni generally oblit- thickly punctate" areola, a number of the erated or nearly so, constituting 41.0 to characters displayed by C. (C.) davidi in- 44.0% of TCL; suborbital angle obsolete dicate a possible relationship with the pro- and lacking tubercle; cephalothorax mark- genitors of "topotypic" C. (C.) b. cavatus, edly longer than abdomen; subdistal spine although their similarities could just as well on mesial margin of carpus of cheliped be a result of convergence. In Ohio and thick and knoblike rather than long and West Virginia, this putative subspecies ap- acute; cervical region of carapace lacking parently lacks the deeply excavate rostrum, strong, multiple tubercles; ventrolateral and displays other characters that differ ridge of merus of cheliped usually lacking from those of the "typical" form (Jezerinac tubercles; and annulus ventralis and form I 1985, Jezerinac et al. 1995). male gonopod quite different in configura- The combination of a narrow (but not tion. obliterated), sparsely punctate areola, a la- Remarks.—Current evidence indicates dlelike rostrum, and a vaulted, unflattened that the range of C. (C.) davidi, limited to carapace will serve to separate C. (C.) dav- parts of the eastern Piedmont Plateau in two idi from all other members of the subgenus endemic North Carolina river basins, is dis- except some C. (C.) b. cavatus, and some junct from that of other members of the upland southern populations currently as- subgenus. The nearest known North Caro- VOLUME 1 13, NUMBER 2 441 lina populations of C. (C.) bartonii (s.l.) are berland Mountains, Kentucky, Virginia, Tennes- see, Georgia, and Alabama, Part II: The genera in the mountains and eastern foothills. Fallicambarus and Cambarus. Pp. 585—605 in Whether or not C. (C.) davidi is indeed al- J. W. Avault, Jr., ed., Freshwater crayfish. Lou- lopatric, however, will only be revealed by isiana State University Division of Continuing more extensive field work throughout the Education, Baton Rouge, Louisiana, 676 pp. Piedmont Plateau. Specimens of subgenus Cooper, J. E. 2000. A new species of crayfish of the Cambarus from seeps and burrows in the genus Cambarus, subgenus Cambarus (Deca- poda: Cambaridae), from the Broad River basin Dan River subdrainage of the Roanoke Riv- of North Carolina.—Journal of the Elisha er basin resemble C. (C.) davidi in many Mitchell Scientific Society 116(1): 1-12. respects, differ in others. The sample size , & R. E. Ashton, Jr. 1985. The Necturus lewisi of available adult specimens is far too small study: Introduction, selected literature review, for accurate assignment of the Roanoke ma- and comments on the hydrologic units and their terial at this time. faunas.—Brimleyana 10:1 — 12. , & A. L. Braswell. 1995. Observations on Cambarus (C.) davidi has yet to be found North Carolina crayfishes (Decapoda: Cambar- in the Tar-Pamlico River basin, whose cray- idae).—Brimleyana 22:87-132. fish fauna is as well known as that of the , & M. R. Cooper. 1995. A new species of cray- Neuse and is nearly identical (Cooper & fish of the genus Orconectes, subgenus Procer- Braswell 1995). Unfortunately, almost no icatnbarus (Decapoda: Cambaridae), endemic sampling has been done in appropriate hab- to the Neuse and Tar-Pamlico river basins, North Carolina.—Brimleyana 23:65-87. itats within the Tar-Pamlico basin. Fabricius, J. C. 1798. Supplementum entomologiae Etymology.—I take great pleasure in systematicae. Hafniae: Proft et Storch, 572 pp. naming this species for my son, David Faxon, W. 1884. Descriptions of new species of Cam- George Cooper, an avid naturalist who barus, to which is added a synonymical list of brought the species to my attention and who the known species of Cambarus and Astacus.— spent many hours tromping in mud and Proceedings of the American Academy of Arts and Sciences 20:107-158. muck to collect quite a few of the existing . 1914. Notes on the crayfishes in the United specimens. States National Museum and the Museum of Suggested vernacular name: Carolina la- Comparative Zoology with descriptions of new dle crayfish. species and subspecies.—Memoirs of the Mu- seum of Comparative Zoology at Harvard Col- Acknowledgments lege 40(8):351-427. Fitzpatrick, J. F., Jr. 1983. How to know the freshwater I am indebted to David G. Cooper for his crustaceans. William C. Brown Company Pub- enthusiastic and persistent field work, as lishers, Dubuque, Iowa, 227 pp. well as to those other biologists who col- Girard, C. 1852. A revision of the North American lected specimens (their names are provided Astaci, with observations on their habits and in the section on "Range and specimens ex- geographical distribution.—Proceedings of the amined"). I also express my sincerest grat- Academy of Natural Sciences of Philadelphia 6: 87-91. itude to Alvin L. Braswell, John E. Cooper, Hay, W. P. 1902. Observations on the crustacean fauna Jr., Martha Riser Cooper, and Don Howard, of Nickajack Cave, Tennessee, and vicinity.— for their unstinting assistance. The manu- Proceedings of the United States National Mu- script was greatly improved by the reviews seum 25(1292):417-439. of Roger F. Thoma, Rafael Lemaitre, and . 1914. Cambarus bartonii carinirostris Hay. an anonymous referee. Nancy Childs, Pp. 384-385 in W. Faxon, Notes on the cray- NCSM, provided technical assistance in the fishes in the United States National Museum and the Museum of Comparative Zoology with final preparation of Figure 1. descriptions of new species and subspecies.— Memoirs of the Museum of Comparative Zo- Literature Cited ology at Harvard College 40(8):351-427. Bouchard, R. W. 1976. Geography and ecology of Hobbs, H. H., Jr. 1956. A new crayfish of the genus crayfishes of the Cumberland Plateau and Cum- Cambarus from North Carolina and South Car- 442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON olina (Decapoda, Astacidae).—Journal of the ia.—Bulletin of the Ohio Biological Survey Elisha Mitchell Scientific Society 72(1 ):61-67. New Series 10:1-193. . 1972. Crayfishes (Astacidae) of North and LeConte, J. 1856. Descriptions of new species of As- Middle America. Biota of freshwater ecosys- tacus from Georgia.—Proceedings of the Acad- tems identification manual No. 9. U.S. Environ- emy of Natural Sciences of Philadelphia 7:400- mental Protection Agency, Washington, D.C., 402. 173 pp. Ortmann, A. E. 1931. Crawfishes of the southern Ap- . 1989. An illustrated checklist of the American palachians and the Cumberland Plateau.—An- crayfishes (Decapoda: Astacidae, Cambaridae, nals of the Carnegie Museum 20(2):61-160. and Parastacidae).—Smithsonian Contributions Thoma, R. F., & R. F. Jezerinac. 1982. New distribu- tional records of crayfish (Cambarus and Fal- to Zoology 480:1-236. licambarus) from Ohio, including a new sub- Jezerinac, R. F. 1985. Morphological variations of species record.—Ohio Journal of Science 82(3): Cambarus (Cambarus) bartonii cavatus (Deca- 136-138. poda: Cambaridae) from Ohio, with a diagnosis , & . 1999. The taxonomic status and of the Ohio form.—Ohio Journal of Science zoogeography of Cambarus bartonii cariniros- 85(3): 131-134. tris Hay, 1914 (Crustacea: Decapoda: Cambar- , & R. F. Thoma. 1984. An illustrated key to idae).—Proceedings of the Biological Society the Ohio Cambarus and Fallicambarus (Deca- of Washington 112:97-105. poda: Cambaridae) with comments and a new Williamson, E. B. 1907. Notes on the crayfish of Wells subspecies record.—Ohio Journal of Science County, Indiana, with a description of a new 84(3): 120-125. species. Pp. 749-763 in 31st Annual Report of , G. W. Stacker, & D. C. Tarter. 1995. The cray- the Department of Geology and Natural Re- fishes (Decapoda: Cambaridae) of West Virgin- sources of Indiana, 1906.