A New Crayfish of the Genus Cambarus Erichson, 1846 (Decapoda: Cambaridae) from the Cape Fear River Basin in the Sandhills of North Carolina John E

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116(4):920-932. 2003. A new crayfish of the genus Cambarus Erichson, 1846 (Decapoda: Cambaridae) from the Cape Fear River basin in the Sandhills of North Carolina John E. Cooper and David G. Cooper (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]; (DGC) North Carolina State Museum of Natural Sciences, 11 West Jones Street, Raleigh, North Carolina 27601 U.S.A., e-mail: [email protected] Abstract.—Cambarus (Puncticambarus) hystricosus is a new species of crayfish that appears to be endemic to certain streams of the Cape Fear River basin that originate in or flow through the xeric Sandhills district of the Atlantic Coastal Plain of North Carolina. It shares a number of unusual characters with C. (P.) spicatus Hobbs of the Broad River basin, and is in many ways similar to an undescribed Cambarus of the same subgenus with which it is syntopic. It is readily distinguished from these and other members of the subgenus by a combination of multiple cervical spines, hepatic spines, an obtuse to subacute suborbital angle that is almost always devoid of a tubercle or spine, and the absence of marginal spines or tubercles on the rostrum.

The crayfish whose description follows is Plain, the district is characterized by dis- another of the undescribed species in the sected topography, consisting of broad, dis- complex previously subsumed under the continuous ridges and adjacent slopes. In name Cambarus (.Puncticambarus) acumi- North Carolina it is traversed by streams in natus Faxon, 1884 (Cooper 2001). The parts of the Cape Fear, Lumber, and Yad- North Carolina populations of this complex kin-Pee Dee river basins. The distinguish- have also been referred to as Cambarus ing terrestrial floral community of the Sand- (.Puncticambarus) sp. C (Hobbs and Peters hills is the Pinus palustris/Quercus cates- 1977, Cooper and Braswell 1995), since byi/Aristida stricta association. Aquatic an- probably none are legitimately referable to imals known to be endemic to the district, C. (P.) acuminatus. This new crayfish is in addition to the crayfish described herein, currently known only from certain streams within or near that part of the Cape Fear are the fishes Etheostoma mariae (Fowler), River basin that lies in the xeric Sandhills which appears to occur only in the Lumber district of the Inner Atlantic Coastal Plain. basin, and Semotilus lumbee Snelson and This is a physiographically distinctive re- Suttkus, which occurs in all three river ba- gion of Eocene sediments (Pinehurst For- sins. An undescribed salamander of the ge- mation) that sit atop Cretaceous strata (Mid- nus Eurycea is also endemic to the Sand- dendorf Formation) at the eastern margin of hills, where it is known from in and near the Fall Zone of the Piedmont Plateau, bor- streams in both the Cape Fear and Lumber dered on the east by the Orangeburg Scarp, river basins. a marine escarpment that "is generally con- Since at least one other member of the sidered to be an early Pliocene wave-cut C. (P.) species C complex, ostensibly an- feature." (Cabe et al. 1992:177). In contrast other undescribed species, occupies some to the flatness of the rest of the Coastal of the same streams as the new species, no VOLUME 116, NUMBER 4 921 reliable synonymy can at this time be pre- Cervical spines strong, 2-4 each side of sented. carapace, plus one to several tubercles. Measurements of crayfish structures were Postorbital ridge strong, cephalic margin ta- made to the nearest 0.1 mm with a Fowler pered and with strong spine or acute tuber- precision dial caliper, following the meth- cle. Suborbital angle obtuse to subacute, al- ods of Hobbs (1981:9-10) unless otherwise most always without tubercle or spine; noted. Abbreviations used in the text are as branchiostegal spine strong. Antennal pe- follows: j, juvenile; NC, North Carolina duncle with 2 strong spines; antennal scale State highway; NCSM, North Carolina 2.4-2.9 (X = 2.6, n — 24) times as long as State Museum of Natural Sciences, Ra- wide, greatest width at midlength; lateral leigh; PCL, postorbital carapace length; R, margin thickened and terminating distally river; SR, state secondary (county) road; in long spine. TCL, total carapace length; US, United Palm of chela 1.5-1.8 (X = 1.7, n = 22) States highway; USGS, United States Geo- times wider than deep; width of palm 1.2— logical Survey; USNM, National Museum 1.7 (X = 1.4, n— 19) times length of mesial of Natural History, Smithsonian Institution, margin, latter constituting 29.0-32.2% (X Washington, D.C.; UTM, Universal Trans- = 30.4%, n = 19) of total chela length; verse Mercator coordinates; and WRC, mesial margin with subserrate mesial row North Carolina Wildlife Resources Com- of 7-9 (usually 8) large, semierect tuber- mission. cles, subtended dorsally by row of 6-8 (usually 7) tubercles of same size or slightly Cambarus (Puncticambarus) hystricosus, smaller, and other smaller but produced tu- new species bercles immediately dorsal to this row; 1— Fig. 1, Table 1 3 small tubercles ventral to mesial row. Fixed finger of chela costate laterally, with Diagnosis.—Body and eyes pigmented, moderately developed dorsal and ventral eye large (X adult diam 2.3 mm, n = 21). ridges; opposable surface of finger with row Rostrum with narrow, moderately elevated of 6-9 (usually 7 or 8) tubercles, in addition margins, slightly or strongly converging to to subconical tubercle ventral to denticles; base of acumen, which is not delimited by dactyl length 1.8-2.3 (X = 2.0, n = 20) tubercles or spines; margins not constricted times length of mesial margin of palm, with at base of acumen, converging and concave moderately to poorly developed ridges dor- from there to acute or subacute, cephalo- sally and ventrally; mesial surface of dactyl dorsally directed apical tubercle or spine; with strong tubercles on proximal half or acumen comprising 36.3 to 47.3% (X = more; opposable surface with row of 9-12 43.2%, n — 26) of rostrum length, latter (usually 10) tubercles. Carpus with strong constituting 21.8 to 26.9% (X = 22.7%, n dorsomesial tubercles, mesial surface with = 26) of TCL; floor (dorsal surface) of ros- massive subdistal spine, ventral surface trum broadly concave. Areola 3.3 to 4.9 (X usually with 2 strong distal spines; merus — 4.0, n = 26) times as long as broad with pair of prominent subdistal dorsal (broader in juveniles), constituting 30.3 to spines. 34.0% (X = 32.7%, n = 26) of TCL and Hook on ischium of third pereiopod of 38.8 to 43.3% (X = 41.8%, n = 26) of PCL, males, that of form I male uniramous, mas- with 5 or 6 punctations across narrowest sive, tapered, overreaching basioischial ar- part. Thoracic section of carapace dorsally ticulation by most of length, opposed by punctate, dorsolaterally granulate, laterally large tubercle on basis; coxa of fourth pe- cobbled with tubercles; cephalic section lat- reiopod of males with vertically disposed, erally with many tubercles, hepatic region caudomesial boss. with strong spines or spiniform tubercles. In situ gonopods (first pleopods) of form 922 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cambarus (Puncticambarus) hystricosus, new species. All from holotypic male, form I (NCSM 5115), except E, F, from morphotypic male, form II (NCSM 5120), and J, from allotypic female (NCSM 5004): A, lateral aspect of carapace; B, E, mesial aspect of left gonopod (first pleopod); C, F, lateral aspect of left gonopod; D, dorsal aspect of carapace; G, caudal aspect of in situ gonopods; H, right antennal scale; I, epistome; J, annulus ventralis and postannular sclerite; K, dorsal aspect of distal podomeres of left cheliped. Setae not illustrated. Scale line = 2 mm.

I male (based on holotypic male; Fig. 1G) apical notch, tip not extending as far cau- symmetrical; proximomesial apophyses dally as tip of mesial process; latter inflated, moderately developed, subglobose, slightly tapered, tip acute; distal margin convex separated; central projection directed cau- near shaft and inclined caudoproximally, dally; mesial process directed caudally and proximal margin inclined slightly caudo- slightly laterally, with acute tip recurved distally; in mesial aspect (Fig. IB), distal caudally; in lateral aspect (Fig. 1C), central portion of gonopod relatively plane, with projection curved over 90° to plane of shaft, sparse setae at midlength. Total length of untapered, bearing proximally directed sub- gonopod 22.6% of TCL (28.3% of PCL). VOLUME 116, NUMBER 4 923 Table 1.—Measurements (mm) of types of Cambarus (Puncticambarus) hystricosus, new species.

Holotypic male, Allotypic Morphotypic form I female male, form II Carapace Total length 46.8 40.7 39.7 Postorbital length 37.4 32.3 31.7 Width 24.7 20.8 20.3 Depth 14.2 13.8 14.0 Length rostrum 10.2 9.1 8.7 Length acumen 3.7 4.1 3.7 Length areola 15.9 13.2 13.1 Width areola 3.7 2.7 3.5 Antennal scale Length 8.9 8.6 7.9 Width 3.4 3.1 3.0 Abdomen Length 44.6 43.8 41.5 Width 22.0 20.2 18.6 Cheliped Length lateral margin chela 42.4* 28.5 31.6 Length mesial margin palm 12.3 8.9 9.5 Width palm 20.5 12.4 14.3 Depth palm 11.5 7.7 8.5 Length dactyl 27.8 17.5 19.1 Gonopod length 10.6 N/A 8.9 Regenerated.

Annulus ventralis (based on allotypic fe- Mirror image of this configuration present male; Fig. 1J) about 1.6 times wider than in 29 of 52 females. long, subsymmetrical, subrhomboidal in Measurements of type specimens provid- shape; cephalic half depressed, sloping, ed in Table 1. with narrow median trough not dissecting Description of holotypic male, form I.— cephalic margin, trough flanked each side Body and eyes pigmented, eye 2.6 mm di- by low ridge; dextral ridge straight, pro- ameter. Cephalothorax (Fig. 1A, D) subcy- ceeding caudally to meet cephalic margin lindrical, thoracic section wider than deep. of caudodextral wall; sinistral ridge less Areola 4.3 times longer than wide, consti- well defined, straight, proceeding caudally tuting 34.0% of TCL (42.5% of PCL), quite to about midlength of annulus before mak- punctate, 6 punctations across narrowest ing abrupt turn to join upper arm of reverse part. Rostrum with narrow margins slightly C-shaped caudosinistral wall; caudodextral converging nearly to base of acumen, where wall fairly thick, straight, slanted about BO- not constricted, but more convergent and SS0 from horizontal plane; caudosinistral concave to apical tubercle, which reaching wall rounded, margin directed about 55° base of distal two-thirds of ultimate podo- from horizontal; transverse tongue originat- mere of antennular peduncle; acumen com- ing from caudodextral wall, proceeding sin- prising 36.3% of rostrum length, latter con- istrally, curving slightly before plunging stituting 21.8% of TCL; floor of rostrum into deep fossa beneath sinistral wall; sinus broadly concave, proximal three-fourths of nearly dissecting caudal margin in midline, surface punctate; usual row of deep punc- caudal wall with weak median labiellum. tations along inner surface of dorsal ridge; 924 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON subrostral ridge weak, narrowly visible to central depression of body broad, deep, base of acumen in dorsal aspect; ventral with cephalomedian fovea; lamellae punc- keel of rostrum devoid of tubercles or tate, lateral corners subtruncate, without tu- spines. bercles; zygoma moderately arched. Anten- Postorbital ridge strong, straight, laterally nal peduncle with strong cephalolateral grooved, cephalic margin with acute tuber- spine on basis, similar spine on ventral sur- cle, caudal terminus somewhat inflated. face of ischium; antennular peduncle with Suborbital angle rounded, without spine or small subdistal median spine on ventral sur- tubercle; branchiostegal spine strong. Tho- face of basal podomere. Antennal scale racic section of carapace dorsally punctate, (Fig. 1H) 2.6 times longer than wide, dorsolaterally granulate, laterally cobbled broadest near midlength; lateral margin with tubercles; row of small tubercles along thickened, straight, terminating in long dis- ventral margin of anterior part of cervical tal spine, tip of which reaching midlength groove; cephalic section of carapace 1.9 of ultimate podomere of antennular pedun- times longer than areola, constituting 66.0% cle; lamella about 1.5 times as wide as of TCL, laterally covered with small to thickened lateral margin, with declivous large tubercles, most acute or subacute; gas- distal margin and broadly curved mesial tric region punctate. Cervical spine region margin. on each side of carapace with 2 strong Third maxilliped with tip reaching slight- spines, 1 smaller spine dorsal to them, and ly beyond midlength of penultimate po- some tubercles. domere of antennal peduncle; exopodite Abdomen slightly narrower and shorter hirsute, tip reaching nearly to midlength of than cephalothorax; abdominal pleura trun- merus of endopodite; distolateral corner of cate ventrally, subangular caudoventrally, ischium produced as acute tubercle; lateral slightly rounded caudally; terga very punc- half of ischium with some punctations, tate, except articular surfaces glabrous. short setae present along inner margin of Proximal podomere of uropod with spine lateral ridge; mesial half of ischium with on lateral lobe, larger spine on mesial lobe; eroded bristles, mesial margin on right with mesial ramus of uropod with weak median 21 denticles, 22 on left. Incisor ridge of ridge bearing submarginal caudomedian right mandible with 6 denticles, 5 on left. spine; caudolateral spine of ramus moder- Both chelipeds regenerated (left more ate; cephalic section of lateral ramus of uro- nearly normal; deviations noted in follow- pod with broad, curved median ridge bearing). Left chela (Fig. IK) 2.1 times longer ing terminal spine at transverse flexure, lat- than wide; palm 1.8 times broader than ter on right with total of 14 fixed spines and deep, width 1.7 times length of mesial mar- 1 long, articulated sublateral spine along gin (normal range 1.2-1.7, X = 1.4); latter caudal margin. Telson with 2 strong spines 29.0% of total chela length, 44.2% of dactyl in each caudolateral corner of cephalic sec- length. Dorsal surface of palm punctate, tion, mesialmost articulated (tip of fixed those punctations on mesial third with small spine on left side bifurcate); caudal margin tubercles at proximal bases; longitudinal domelike. dorsomesial sulcus shallow, with row of Epistome (Fig. II) with cephalic lobe ba- moderate, produced tubercles; distolateral sically subcordiform in ventral outline, but margin of palm costate for about a third of cephalic margin slightly curved, not peak- length (continuous onto fixed finger), with ed, and lateral margin on each side with moderate impression and aggregation of weak concavity; most of margin crenulate, large punctations with tubercles at proximal with tuberclelike protuberances; lateral cor- bases; lateral eminence of articular ridge ners inflated; floor (ventral surface) of lobe with several small distal protuberances; lat- moderately convex, punctate, with setae; eral margin of palm rounded, with large, VOLUME 116, NUMBER 4 925 widely spaced punctations. Ventral surface stout, curved distal spine and minuscule of palm less punctate than dorsal, some proximal tubercle; ventral surface with 2 punctations on mesial half with minuscule strong distal spines (tip of distolateral spine basal tubercles; distolaterally with broad broken) and 1 proximomesial tubercle. Left depression (continuous onto fixed finger) merus 1.7 times longer than deep, dorsal and scattered punctations without tubercles; surface with 2 strong, suberect subdistal lateral eminence of articular ridge with 1 spines and 1 minuscule tubercle; ventrolat- subdistal tubercle and 1 distal protuberance; eral ridge with 1 small tubercle, 2 stout 2 tubercles proximal to ridge. Mesial mar- spines, and moderate distal spine; ventro- gin of left palm with subserrate mesial row mesial ridge with 4 subacute tubercles, 4 of 8 tubercles, row subtended dorsolaterally spines, and broken distal spine; ischium by row of 6 tubercles, and others dorsal to with 5 small ventral tubercles. this row; 2 tubercles ventral to mesial row. Gonopods as described in "Diagnosis." Fingers gaping most of length, greatest Description of allotypic female.—Except width of gape about equal to width of base for secondary sexual characters, differing of dactyl. Left fixed finger costate laterally; from holotypic male in following respects: dorsal surface with moderately strong lon- Areola 4.9 times wider than long, consti- gitudinal median ridge, flanked each side by tuting 32.4% of TCL (40.9% of PCL), with row of punctations; proximolateral area 4 or 5 large punctations across narrowest with moderate impression and large punc- part. Acumen long, spiniform, comprising tations, some with basal tubercles; lateral 45.1% of rostrum length, tip reaching mid- margin with row of large punctations; ven- length of ultimate podomere of antennular tral surface with weak, rounded longitudi- peduncle; rostrum comprising 22.4% of nal ridge, flanked each side by row of punc- TCL; subrostral ridge narrowly visible only tations; opposable margin with usual sub- along proximal half of rostrum. Cephalic conical tubercle (worn) ventral to denticles, margin of postorbital ridge with strong located at base of distal fourth of finger, and spine. Cephalic section of carapace 2.1 another smaller ventral tubercle distal to times longer than areola, constituting 67.6% subconical one (abnormal); row of 11 tu- of TCL; hepatic region with spiniform tubercles dorsal to denticles (normal range 6— bercles. Cervical spine region on left side 9, usually 7 or 8), first of these very small, of carapace with 1 long spine, 2 smaller fifth from base largest; denticles eroded, in spines dorsal to it, and several tubercles; 2 rows distally. Dorsal surface of dactyl right side with 2 long and 1 small spine. with low ridge, flanked each side by row of Telson with 1 fixed and 1 articulated punctations; mesial surface with strong tu- spine in caudolateral corner of cephalic sec- bercles on proximal half, basal tubercles in tion on right, 1 fixed and 2 articulated 2—3 rows, encroaching dorsally; ventral sur- spines on left, all spines strong. face with obsolete longitudinal ridge, de- Floor of cephalic lobe of epistome with fined by 2 rows of punctations; opposable long setae; lamellae with single caudal tu- surface with 13 tubercles (normal range 9— bercle each corner. Antennal scale 2.8 times 12, usually 10), first and fourth from base longer than wide; lamella about 1.2—1.3 largest; denticles eroded, in 2 rows on distal times as wide as thickened lateral margin. third of finger. Third maxilliped with cephalolateral corner Left carpus (Fig. IK) 1.4 times as long of ischium produced as spine. as wide, 1.3 times as long as mesial margin Both chelipeds appear normal. Right che- of palm; dorsal surface of carpus with shal- la 2.3 times longer than wide; palm 1.6 low, oblique sulcus, lateral to which surface times broader than deep, width 1.4 times punctate, mesial to which surface with 8 or length of mesial margin, which is 31.2% of 9 dorsomesial tubercles; mesial margin with total chela length and 50.9% of dactyl 926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON length. Lateral eminence of ventral articular duncle; lamella about 1.6 times width of ridge with subacute distal tubercle; single thickened lateral margin. Incisor ridge of tubercle proximal to ridge. Mesial margin right mandible with 7 denticles. of palm with staggered mesial row of 7 tu- Right cheliped replaced by regenerating bercles, row subtended dorsolaterally by bud. Left chela 2.2 times longer than wide; staggered row of 8 tubercles (6 on left), and palm 1.7 times broader than deep, width 1.5 several small tubercles dorsal to this row. times length of mesial margin; latter 30.1% Fingers only slightly gaping in proximal of total chela length, 49.7% of dactyl half, opposable base of fixed finger with length. Lateral eminence of ventral articular sparse long setae. Opposable surface of ridge with 2 distal tubercles; 2 tubercles right fixed finger with small subconical tu- proximal to ridge. Mesial margin of palm bercle ventral to denticles at base of distal with subserrate mesial row of 7 tubercles, fourth of finger, and row of 8 tubercles dor- row subtended dorsolaterally by staggered sal to denticles, fourth tubercle from base row of 7 tubercles, and others dorsal to this largest (on left finger, third and fourth from row; 2 produced tubercles ventral to mesial base largest and same size). Mesial surface row. Fingers gaping in proximal three- of right dactyl with strong tubercles on fourths of length, greatest width of gape proximal two-thirds of finger; opposable about four-fifths width of base of dactyl. surface with 11 tubercles, basalmost one Dorsal surface of fixed finger with strong largest, fourth from base offset ventrally median ridge, flanked mesially by punctate (left dactyl with 12 tubercles). groove; opposable margin with strong sub- Dorsal surface of carpus with row of 5 conical tubercle ventral to denticles, and dorsomesial tubercles; mesial surface with row of 8 tubercles dorsal to denticles, long, curved spine at about midlength, no fourth tubercle from base largest. Oppos- proximal tubercle. Right merus 1.9 times able surface of dactyl with 12 tubercles, longer than deep; ventrolateral ridge with 3 basalmost largest. Mesial surface of carpus strong spines; ventromesial ridge with 8 with long, curved subdistal spine and small spines or spiniform tubercles (7 on left mer- proximal tubercle; ventral surface with 2 us) and strong distal spine. strong distal spines and large proximome- Annulus as described in "Diagnosis." In sial tubercle. Merus 1.8 times longer than addition, first pleopods present, fairly long; deep; ventrolateral ridge with 3 strong postannular sclerite about one-third width spines and moderate distal spine; ventro- of annulus, ridgelike, ventral surface pitted. mesial ridge with 4 small tubercles and 6 Description of morphotypic male, form spines. II.—Differing from holotypic male in fol- Hook on ischium of third pereiopod lowing respects: Areola 3.7 times longer small, tuberclelike, not overreaching bas- than wide, constituting 33.0% of TCL ioischial articulation, not opposed by tuber- (41.3% of PCL). Acumen comprising cle on basis. Gonopod (Fig. IE, F) length 42.5% of rostrum length. Cephalic margin 22.4% of TCL (28.1% of PCL). In situ gon- of postorbital ridge with strong spine. Ce- opods with small, separated proximomesial phalic section of carapace with several apophyses; central projection directed cau- strong hepatic spines on each side of body. dally; mesial process inflated, directed cau- Cervical spine area with 3 spines and a tu- dolaterally and inclined slightly proximally; bercle. in lateral aspect, central projection curved Cephalic lobe of epistome subcordiform 90° to plane of shaft, broad, tapered; mesial in outline, margins crenulate, with some process tapered, tip slightly extruded. short setae; floor plane, with setae. Tip of Color notes.—Ground color of carapace spine of antennal scale reaching distal mar- and abdomen (except tailfan) gray-green to gin of ultimate podomere of antennular pe- olive. Thoracic section of carapace with VOLUME 116, NUMBER 4 927 large, pale greenish blotches, one such tones, each with large, diagonal yellowish blotch in median third of carapace at caudal or tannish blotch near ventral border and end of areola; laterally darker, with small, dark oblique dorsolateral mark. Telson and light blotches and white granules; cervical uropods aquamarine dorsally and ventrally, spines ivory or roseate. Cephalic section of dorsal surface of telson with tan overtones; carapace nearly uniform olivaceous dorsal- both rami of uropods with narrow trans- ly, darker laterally, with large tan or whitish verse subcaudal band of orange-tan, sub- blotch along cephalolateral margin between tended caudally by narrow blue band; trans- suborbital area and white branchiostegal verse flexure of lateral ramus of uropod also spine; hepatic spines and tubercles on lat- with narrow orange-tan band. eral surface white. Postorbital ridge pale Ventrally, all cephalic structures of body grayish-green, cephalic spine tan to yellow- blue-green or bluish, except zygoma white, ish. Antennal scale bluish, but mesial mar- and mandible and parts of third maxilliped gin of thickened lateral portion tan; anten- lighter in color, sometimes pale yellowish; nal and antennular flagellae variable in col- other ventral surfaces of body white or or, with dark brown, reddish tan, or tan ar- translucent blue, ventral abdomen usually ticles, and brown or pale green interstices. with blue markings. Dorsal surfaces of palm and fingers of Juveniles with same general pattern, but cheliped blue-green; lateral margin of entire paler. Ventral margins of carapace often dis- propodus creamy orange, or yellowish with tinctly blue; dorsal abdomen pale greenish, blue-gray areas; ventral surfaces of palm terga with very prominent dark banding. and fingers pale grayish-green or blue- Type locality.—North Carolina, Hoke green, but mesial third of palm and most of County, Puppy Creek at SR 1409, ca. 2.4 dactyl usually darker; tips of fingers orange air km E of Wayside (Nicholson Creek 7.5' or tannish-orange, color not subtended by USGS quadrangle, UTM Zone 17, coordi- black band; tubercles on mesial margin of nates 670662E/3876690N). palm and dactyl pale green, tubercles on Disposition of types.—The holotypic opposable surfaces of both fingers ivory, male, form I, allotypic female, and morpho- pale green, or blue-gray. Dorsal and mesial typic male, form II, are in the NCSM crus- surfaces of carpus gray-green, dorsal tuber- tacean collection (catalogue numbers 5115, cles same color but paler; proximal and dis- 5004, and 5120, respectively), as are para- tal margins of carpus with narrow russet topotypes consisting of 3j 8, 3 j? (5055); band. Dorsal surface of merus gray-green 2 2 (5060); 1 $ (5117); 1 6 II (5119); 4 on distal half, fading to white on proximal jc?,2j$ (5118); 1 $ (5254); 1 jc3 (5288); half, distal membrane blue, spines ivory; 5 janimal (TCL 43.7 vier), Notropis cummingsae Myer, and No- mm) from Stewarts Creek is gynandromor- turus insignis (Richardson); the neotenic phic. It has a normal annulus ventralis, fe- salamander Necturus punctatus (Gibbes); male first and second pleopods, and postan- and the watersnake Nerodia taxispilota nular sclerite, but lacks oviduct apertures in (Holbrook). the coxae of the third pereiopods. It has Crayfish associates.—The only other male sperm duct openings in the coxae of crayfishes yet found at sites occupied by C. the fifth pereiopods. (P.) hystricosus are an undescribed member Size.—The largest specimen is a female of the subgenus, and Procambarus (Ort- measuring 52.6 mm TCL (43.0 mm PCL). mannicus) acutus (Girard, 1852). Camba- The largest form I male measures 46.8 mm rus (Depressicambarus) latimanus (Le- TCL (37.4 mm PCL); the only other form Conte, 1856) occurs in some other streams I males have TCLs of 30.8, 35.0, and 38.0 of the area, but has not yet been found in mm (25.4, 27.7, and 30.2 mm PCL, respec- those occupied by C. (P.) hystricosus. tively). The largest form II male measures Affinities.—On the basis of its extensive 39.7 mm TCL (31.7 mm PCL). spinal armature, which we consider a ple- Life history and ecological notes.—Form siomorphic character state (and see Hobbs I males were found in early June and July, 1962:276, 1969:116-118), C. (P.) hystri- and in October and November. No oviger- cosus appears to have strong affinities with ous females or those with attached young Cambarus (Puncticambarus) spicatus have yet been seen. Hobbs, 1956. Both species characteristical- On 21 May 1998, Puppy Creek was a ly have hepatic spines, or strong spiniform "clear blackwater stream with a white sand or very acute tubercles, a rare feature in bottom" (B. Tracy, pers. comm.). Specific epigean Cambarus; strong cervical spines conductance on that date was 18 |xmhos/ that are multiple in C. (P.) hystricosus and cm, pH 4.9, water temperature 22°C, and sometimes so in C. (P.) spicatus; long bran- dissolved oxygen content 6.9 ppm. In Feb- chiostegal spines; and strong, multiple ruary and March 1999, the creek at the type spines on the antennal peduncle, ventral locality was about 3 m wide, with fairly carpus, and dorsal and ventral merus. Major shallow water and a few deeper sections. differences between the two species are the The bottom was sandy, with some sticks presence of marginal spines or strong tu- and detritus, several large submerged logs, bercles on the rostrum of C. (P.) spicatus, and dense growths of long, slender sub- which are never normally present in C. (P.) merged vegetation in which the crayfish hystricosus; a spine or tubercle on the sub- were found. At some points the banks were orbital angle of C. (P.) spicatus, which is undercut and covered with overhanging rarely seen in C. (P.) hystricosus; the struc- roots, but passing a dipnet through these ture of the central projection of the form I produced no crayfish. In June of the same male gonopod, which in C. (P.) spicatus is year, water levels had dropped significantly, scarcely curved beyond 90° to the shaft and the vegetation was not completely sub- has a very weak subapical notch; and the merged, and the few crayfish collected were more symmetrical annulus of that species. found only among sticks and detritus in ar- The other member of the subgenus that eas with the greatest current. occurs in some streams with C. (P.) hystri- VOLUME 116, NUMBER 4 931 cosus lacks hepatic spines and multiple cer- ical to exploitation by more cold- and lotic- vical spines, although it may display acute adapted cambaroid stocks. The later radia- or subacute cervical tubercles along with a tion into the Piedmont of more highly com- single strong spine. The suborbital angle in petitive and closely related forms could this species is almost always acute or sub- have fragmented the ranges of the earlier acute and bears a tubercle or spine. Only inhabitants into a number of allopatric pop- two of forty-five specimens examined had ulations that evolved into distinctive relicts. an obtuse suborbital angle, and in only one Other such populations may await discov- of them did the angle lack a tubercle. The ery. postorbital ridge of adults seldom bears The "residual" genetic influences of a more than a small cephalic tubercle, al- once widespread, presumably plesiomorph- though some smaller specimens have a ce- ic Puncticambarus could help explain the phalic spine. This species also has a shorter occasional appearance of hepatic spines, acumen and a broader areola than C. (P.) marginal spines, and even instances of mul- hystricosus, and the base of its acumen is tiple cervical spines, observed in some in- more angular. dividuals from a number of populations of Remarks.—Both C. (P.) hystricosus and more derived forms in the subgenus (JEC, C. (P.) spicatus occupy limited, circum- pers. obs.). scribed ranges, the former in streams of the Etymology.—Species hystricosus, mean- Cape Fear River basin in the Sandhills of ing prickly, from the Latin hystrix, porcu- North Carolina, the latter in streams of the pine, in reference to its extremely spinose Broad River basin in the Piedmont Plateau body parts. of North and South Carolina. The ranges of Suggested vernacular name: Sandhills both are surrounded by the ranges of other, spiny crayfish. presumably more derived, members of Puncticambarus. In the Sandhills, this is Acknowledgments comparable to the distributions of two other endemic animals. Populations of Semotilus We are very grateful to the biologists lumbee there are surrounded by its more who collected specimens that contributed to broadly distributed relative, Semotilus atro- our study of this species; their names are maculatus Mitchill, and populations of the provided in the section on specimens ex- undescribed Eurycea are surrounded by its amined. Helpful comments on the general close relative, Eurycea cirrigera (Green). biology and geology of the Sandhills were It seems reasonable that the forerunner of provided by J. C. Beane and C. Tacker, re- C. (P.) spicatus and C. (P.) hystricosus was spectively. JEC expresses his gratitude to one of the few members of the genus Cam- A. L. Braswell and D. Howard for their barus to expand its range eastward from the generosity. Very helpful reviews of the Blue Ridge escarpment—which then lay manuscript were provided by Horton H. farther east than it does today—or from far- Hobbs, III, and Rudolph Prins. ther west. Colonization of the Piedmont Plateau and the eastern extremity of the In- Literature Cited ner Coastal Plain would have met with little Cabe, S., J. G. Nickerson, C. W. Hoffman, & K. M. opposition, but continued eastward expan- Farrell. 1992. Cretaceous and Tertiary stratig- sion would have brought this putative pro- raphy of the northern Sandhills area, North Car- genitor into conflict with procambaroid olina. Pp. 177-188 in J. M. Dennison and K. G. Stewart, eds., Geologic field guides to North stocks. The latter crayfishes were adapted Carolina and vicinity. Geologic Guidebook No. to the warm, lentic or low-gradient habitats 1, University of North Carolina at Chapel Hill. of the broad, base-leveled Coastal Plain, Cooper, J. E. 2001. Cambarus (Puncticambarus) hobb- habitats that likely would have been inim- sorum, a new crayfish (Decapoda: Cambaridae) 932 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from North Carolina.—Proceedings of the Bi- cidae).—Proceedings of the Biological Society ological Society of Washington 114(1): 152— of Washington 69:115-122. 161. . 1962. Notes on the affinities of the members , & A. L. Braswell. 1995. Observations on of the Blandingii Section of the crayfish genus North Carolina crayfishes (Decapoda: Cambar- Procambarus (Decapoda, Astacidae).—Tulane idae).—Brimleyana 22:87-132. Studies in Zoology 9(5):273-293. Erichson, W. F. 1846. Uebersicht der Arten der Gattung . 1969. On the distribution and phylogeny of Astacus.—Archiv fur Naturgeschichte 12(1): the crayfish genus Cambarus. Pp. 93-178 in P. 86-103. C. Holt, R. L. Hoffman, and C. W. Hart, Jr., Faxon, W. 1884. Descriptions of new species of Cam- eds., The distributional history of the biota of barus, to which is added a synonymical list of the southern Appalachians, Part I: invertebrates. the known species of Cambarus and Astacus.— Research Division Monograph 1, Virginia Poly- Proceedings of the American Academy of Arts technic Institute, Blacksburg, 295 pp. and Sciences 20:107-158. . 1981. The crayfishes of Georgia.—Smithson- Girard, C. 1852. A revision of the North American ian Contributions to Zoology 480:1-236. Astaci, with observations on their habits and , & D. J. Peters. 1977. The entocytherid ostra- geographical distribution.—Proceedings of the cods of North Carolina.—Smithsonian Contri- Academy of Natural Sciences of Philadelphia 6: butions to Zoology 247:1-73. 87-91. LeConte, J. 1856. Descriptions of new species of As- Hobbs, H. H., Jr. 1956. A new crayfish of the Extra- tacus from Georgia.—Proceedings of the Acad- neus Section of the genus Cambarus with a key emy of Natural Sciences of Philadelphia 7:400- to the species of the Section (Decapoda, Asta- 402.