From the Waccamaw River Basin, North and South Carolina

Home , Procambarus

6 April 1998 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(1):81-91. 1998. A new species of crayfish of the genus Procambarus, subgenus Ortmannicus (Decapoda: Cambaridae), from the Waccamaw River basin, North and South Carolina

John E. Cooper

North Carolina State Museum of Natural Sciences, P. O. Box 29555, Raleigh, North Carolina 27626, U.S.A.

Abstract.—Procambarus (Ortmannicus) braswelli is a new species of crayfish from the Waccamaw River basin in North and South Carolina. A primitive member of the Pictus Group, P. braswelli has its closest affinities with P. (0.) chacei, P. (O.) enoplosternum, and P. (O.) pictus. The new species is less closely related to P. (O.) epicyrtus, and distantly related to its geographically nearest relative, P. (O.) lepidodactylus, with which it has been confounded. The occurrence of P. lepidodactylus in North Carolina is currently uncon- firmed. Procambarus braswelli may be distinguished from the other members of the Pictus Group by its combination on the form I male gonopod (first pleopod) of a prominent, truncated, distally directed caudal knob; a large, somewhat bulbous adventitious process; and a long, caudodistally directed mesial process; and by a long acumen.

The Waccamaw River basin of south- maw River, apparently 7.5 air mi. [12.0 air eastern North Carolina and northeastern km] south of Lake Waccamaw, Columbus South Carolina has long been known as County (1949)" (Cooper & Cooper 1977b: home to a number of animal species that 206). This record was based on a female in are either endemic or are shared with a sin- the collections of the National Museum of gle other river basin. The endemic fauna Natural History, Smithsonian Institution includes several fishes and mollusks. Infor- (USNM 129841), collected on 29 March mation on the fishes can be found in Hubbs 1949 by E. C. Raney. It is the only North & Raney (1946), Frey (1951), Shute et al. Carolina specimen identified as P. lepido- (1981), Menhinick (1991), and Rohde et al. dactylus in the catalogued collections of (1994). The mollusks are discussed in Ful- that institution, and the locality is obviously ler (1977), Johnson (1984), and Porter & the same one referred to as "Columbus Horn (1984). Teulings & Cooper (1977: County" by Hobbs (1968:K9, 1972:61, 414-415) provided a list of the endemic 1974:57, 1989:68). None of the North Car- species. olina specimens later assigned to P. lepi- Included in the Waccamaw invertebrate dodactylus had received critical examina- fauna is a crayfish previously assigned to tion. Prompted by a form I male collected Procambarus (Ortmannicus) lepidodactylus by David R. Lenat, North Carolina Division Hobbs, 1947a, a species considered the of Water Quality, I determined that these northernmost representative of the primitive specimens represent an undescribed species Pictus Group (Hobbs 1958a, 1962, 1968, of the Pictus Group that appears to be en- 1972, 1974, 1989; Cooper & Cooper 1977a, demic to the Waccamaw River basin and 1977b). Few specimens of this Waccamaw that is only distantly related to P. lepido- crayfish have been collected, and the only dactylus. This new species is currently published locality was "canal off Wacca- known only from Columbus County, North 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Carolina, and from a single locality in Hor- Hobbs, 1968.K9 (p.p.; Columbus Coun- ry County, South Carolina, but it may be ty, North Carolina).—Hobbs, 1972:151 found elsewhere within the Waccamaw ba- (p.p., North Carolina).—Franz & Lee, sin. 1982:61 (p.p.; North Carolina).—Le- Because P. lepidodactylus (s.s.) occurs in Grand & Hall, 1997:32 (p.p.; Waccamaw the Pee Dee River basin in South Carolina, drainage, North Carolina). Cooper & Cooper (1977b:207) speculated Procambarus (Ortmannicus) lepidodacty- that it "may yet be discovered in tributaries lus.—Cooper & Cooper, 1977a: 198, of the Lumber River of North Carolina . . 1977b:206, 207 (p.p.; North Carolina).— . . " A distribution map showing localities Cooper & Cooper, 1977a:200 (p.p.; by for members of the Pictus Group (Hobbs implication, North Carolina).—Hobbs, 1958a:72) contains a single North Carolina 1972:61, 1974:57, 1989:68 (p.p.; Colum- site for "P. lepidodactylus." The site indi- bus County, North Carolina).—Hobbs & cated by a dot in the map lies just west of Peters, 1977:8 (p.p.; North Carolina).— the 79th meridian, which would place the Hobbs et al„ 1977:19 (p.p.; North Caro- locality in the Lumber River basin. How- lina).—'Teulings & Cooper, 1977:415 ever, the dot undoubtedly was meant to rep- (p.p.; North Carolina).—Fitzpatrick, resent the Columbus County record in the 1983:214 (p.p.; Carolinas, viz. North Waccamaw basin (USNM 129841). I know Carolina).—Hobbs & Franz, 1986:516 of no specimens of Procambarus from (p.p.; North Carolina).—Hobbs, 1989:86 North Carolina that incontrovertibly can be (p.p.; North Carolina). assigned to P. lepidodactylus. Several spec- Procambarus leptodactylus.—Williams et imens that could belong to this species have al., 1989:26, 64 (p.p.; by implication, been collected in the Lumber basin (Cooper North Carolina; erroneous spelling).— & Braswell 1995:120), but form I males LeGrand, 1993:23, LeGrand & Hall, have yet to be seen. 1995:31 (p.p.; Waccamaw drainage, Abbreviations used in the text are: j = North Carolina; erroneous spelling). juvenile; NC = North Carolina state high- "Undescribed species."—Cooper & Bras- way; NCSM = North Carolina State Mu- well, 1995:120. seum of Natural Sciences, Raleigh; PCL = postorbital carapace length; R = river; SR Diagnosis.—Body and eyes pigmented, = state secondary road; TCL = total cara- eyes large (X adult diam 2.4 mm, n = 11). pace length; UNC = University of North Rostrum acarinate, margins narrow, parallel Carolina; USGS = United States Geologi- to subparallel near base, slightly convex be- cal Survey; US = United States highway; tween orbits, then recurving and tapering to USNM = United States National Museum base of long acumen, which delineated by of Natural History, Smithsonian Institution, strong marginal spines; acumen comprising Washington, D. C.; and UTM = Universal 41.9 to 49.1% (X = 44.8%, n = 12) of ros- Transverse Mercator coordinates. trum length, latter comprising 31.7 to 37.9% (X = 34.4%) of TCL. Areola 2.1 to 4.2 (X = 2.9, n = 13) times as long as broad Procambarus (Ortmannicus) braswelli, and constituting 26.3 to 28.2% (X = 26.9%, new species n = 12) of TCL and 39.4 to 42.8% (X = Fig. 1 41.3%, n = 13) of PCL, and with 6 to 9 Procambarus lepidodactylus.—Hobbs, (usually 7-8) punctations across narrowest 1958a:72 (p.p.; map locality for North part. Carapace densely granulate, cephalic Carolina), 75, 76 (p.p.; southeastern section constituting 71.8 to 74.8% (X = North Carolina).—Hobbs, 1962:284 73.1%, n = 12) of TCL. Cervical spines (p.p.; southeastern North Carolina).— strong, 1 each side; cervical groove inter- VOLUME 111, NUMBER 1 83

Fig. 1. Procambarus (Ortmannicus) braswelli, new species (all from holotypic male, form I, except C, E, from morphotypic male, form II, and G, from allotypic female; setae not illustrated): A, lateral aspect of cephalothorax; B, C, mesial aspect of left gonopod (first pleopod); D, dorsal aspect of cephalothorax; E, F, lateral aspect of left gonopod; G, annulus ventralis and associated structures; H, epistome; I, basal podomeres of third, fourth, and fifth pereiopods; J, antennal scale; K, caudal aspect of in situ gonopods; L, M, caudomesial and caudolateral aspects, respectively, of tip of left gonopod; N, dorsal aspect of distal podomeres of right cheliped. 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rupted. Branchiostegal spine strong; hepatic caudally, and with expanded base forming area with some weak tubercles. Suborbital cowl around cephalic base of central pro- angle obtuse to obsolete. Postorbital ridge jection; latter subtriangular, directed cau- moderately strong, with nearly obliterated dodistally at much greater angle than ce- groove and strong cephalic spine. Antennal phalic process; caudal element consisting scale 2.9 to 3.4 (X = 3.2, n = 13) times as of: prominent distolateral caudal knob, di- long as broad, widest proximal to mid- rected distally and delimited cephalically by length; lateral margin thickened and termi- groove; small, toothlike caudal process, nating in acute spine, mesial (lamellar) mar- originating on mesial surface of central pro- gin subangular. jection and directed caudodistally; and in- Palm of chela of cheliped 1.1 to 1.7 (X flated adventitious process, originating at = 1.4, n = 13) times wider than deep, 1.3 proximomesial base of cephalic process, ly- to 1.8 (X = 1.6, n = 13) times longer than ing wholly mesial or cephalomesial to cau- wide, and constituting 40.2 to 46.7% (X = dal process, and obscuring part of proxi- 42.5%, n = 12) of chela length; mesial mar- momesial bases of central projection and gin of palm with staggered row of 5 to 13 caudal process. small, subconical tubercles. Fingers without Annulus ventralis (Fig. 1G) symmetrical, gape, without dense setae at opposable bas- subovate, 2.4 times as wide as long, mov- es; dactyl 1.1 to 1.4 (X = 1.2, n = 13) times able; cephalic margin broadly arched, cau- length of mesial margin of palm. dal margin mildly convex and with weak Hooks on ischia of third and fourth per- caudomedian labiellum; median xh of an- eiopods of male; in form I male (Fig. II), nulus ventrally elevated, moundlike; dextral hook on third pereiopod oblique, subconi- half of mound hemitubular, C-shaped, sin- cal, overreaching basioischial articulation istral half slightly narrower, following con- by most of length, not opposed by tubercle tours of dextral half; both parts of central on basis; hook on fourth pereiopod smaller, mound descending cephalically as pair of vertically disposed, not reaching articula- short, narrow, curved ridges, which taper- tion, and opposed by prominent protuber- ing cephalically and terminating sinistral to ance on basis, center of protuberance with midline of cephalic margin of annulus; nar- curved, setiferous depression; coxa of row, somewhat C-shaped ridge dextrolater- fourth pereiopod with low, vertically dis- al to central mound at about midlength of posed caudomesial boss, that of fifth pe- annulus, and short horizontal ridge sinistro- reiopod with compressed ridgelike boss at lateral to mound at same level; deep sub- caudomesial angle (Fig. II). triangular depression cephalolateral to cen- Gonopods (first pleopods) of form I male tral mound on either side of curved ridges, (Fig. IB, F, K, L, M) asymmetrical, proxi- each depression with prominent protuber- momesial apophyses strong, tapering, gen- ance near cephalolateral margin. erally rounded but with subacute tip cau- Measurements of type specimens provid- dally, overlapping; total length of gonopod ed in Table 1. 22.1 to 25.6% (X = 23.8%, n = 4) of TCL; Description of holotypic male, form I.— distal XA of shaft weakly inclined caudo- Body and eyes pigmented, eye 2.9 mm distally, cephalic surface with narrow con- diam. Cephalothorax (Fig. 1A, D) subcy- vexity subjacent to base of cephalic pro- lindrical; maximum width of carapace 1.1 cess; mesial process long, slender, noncor- times depth, cephalic section 2.8 times neous, directed caudodistally and inclined length of areola and constituting 73.4% of laterally, tip acute to subtruncate; central TCL. Areola 3.3 times as long as wide, con- projection and cephalic process corneous, stituting 26.6% of TCL (39.5% of PCL), subequal in length; cephalic process with densely punctate, with 7 to 8 punctations acute apex, directed distally and slightly across narrowest part; branchiocardiac VOLUME 111, NUMBER 1 85

Table 1.—Measurements (mm) of types of Procam- ing, floor (dorsal surface) slightly concave, barus (Ortmannicus) braswelli, new species. punctations most abundant in cephalic half;

Morpho- acumen comprising 42.2% of rostrum Holotype Allotype type length, apex corneous, directed cephalically Carapace and extending to distal margin of second Total length 27.4 29.6* 20.2 article of antennular flagellum; ventral keel Postorbital length 18.5 19.4 13.8 of acumen bladelike, broadly subangular in Length cephalic section 20.1 21.3* 14.5 lateral aspect; subrostral ridge visible in Width 11.8 13.5 9.0 dorsal aspect only at base of rostrum. Post- Depth 10.4 12.4 9.1 orbital ridge moderately strong, with thin Rostrum dorsal crest and narrow, nearly obliterated Length 9.0 9.7* 6.4 lateral groove bearing small punctations; Width at base 4.0 4.6 3.4 Length acumen 3.8 off 2.7 cephalic margin with strong spine. Subor- Areola bital angle obsolete, orbital rim subrectili- Length 7.3 8.3 5.7 near and with concavity near base of anten- Width 2.2 2.0 2.1 nal peduncle. Cervical spines strong, 1 each Antennal scale side, and area also with dense granulations Length 9.1 9.8 7.4 dorsal to spine; cervical groove interrupted Width 2.7 3.4 2.3 just dorsal to spine, with short, broad sulcus Abdomen cephalic to groove; ventral margin of ce- Length 28.3 34.6 22.0 phalic portion of groove with row of small Width 10.5 12.9 7.3 tubercles. Carapace with thoracic section Cheliped densely granulate laterally and dorsally, Length lateral margin granules ascending to margins of branchio- chela 19.8 15.9 10.5 cardiac grooves; cephalic section of cara- Length mesial margin pace with scattered granules laterally; cau- palm 8.6 6.6 4.9 dal mandibular region broadly convex, de- Width palm 5.2 5.2 2.7 Depth palm 3.9 3.0 2.0 limited by moderate groove; gastric region Length dactyl 9.5 8.3 5.5 with crowded punctations, caudal margin of Length carpus 6.4 6.2 4.1 region in form of low, arcuate ridge. Width carpus 3.9 4.1 2.3 Branchiostegal spine long, slightly pro- Length dorsal margin merus 10.6 9.9 6.8 curved. Antennal peduncle with long dis- Depth merus 3.7 4.1 1.9 tolateral spine on basis and similar spine on

Gonopod length 6.2 NA 4.8 ventral surface of ischium; antennular peduncle with strong, semierect ventral spine * estimated; acumen damaged. situated near mesial margin at about midlength of basal podomere, which hirsute grooves strong, flaring caudolaterally from mesially and with sparse setae ventrally; tip about midlength. Rostrum with narrow, el- of antennal flagellum reaching cephalic evated margins, extending caudally nearly margin of telson when flagellum adpressed. to caudal margin of postorbital ridges; mar- Antennal scale (Fig. 1J) 3.4 times as long gins of rostrum parallel near base, slightly as wide, broadest proximal to midlength; convex between orbits, then recurving and lateral margin thickened and terminating in converging to base of long, spiculiform strong distal spine, tip reaching slightly be- acumen which delineated by strong margin- yond proximal margin of third article of an- al spines; rostrum deeply excavate, margins tennular flagellum; lamella approximately flanked medially by continuous row of se- twice as wide as thickened lateral portion, tiferous punctations; walls of rostrum slop- distal margin slightly sloping for short dis- 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

tance then strongly declivous to widest part; ite; basal podomere of exopodite not hir- mesial margin subangular. sute; ventrolateral margin of ischium with Abdomen slightly longer and narrower row of setiferous punctations at base of lon- than carapace; pleura of most abdominal gitudinal ridge, distolateral corner produced segments with rounded cephalolateral mar- as acute spine; ventrolateral half of ischium gins, and slightly rounded to subangular with scattered punctations bearing short se- caudoventral margins. Proximal podomere tae; ventromesial half with longitudinal of right uropod (left uropod regenerate, de- rows of long setae, moderately obscuring formed) with very strong caudolateral spine mesial margin; basis of ischium with on lateral lobe, and slightly weaker, some- clumps of long setae, forming brushes. what laterally situated spine on mesial lobe; Right mandible with incisor ridge bearing mesial ramus of uropod with long caudo- 8 denticles, third from distal end largest, lateral spine, and strong median ridge ter- penultimate one small, left incisor ridge minating in spine situated well cephalic to with 6 denticles, distalmost largest. caudal margin; lateral ramus with broad Palm of chela of cheliped (Fig. IN) sub- median ridge on cephalic section, and poor- ovate in cross section, 1.3 times wider than ly defined ridge lateral to it; transverse flex- deep, 1.7 times longer than wide; mesial ure of ramus with margin bearing row of margin of right palm with mesial row of 11 11 fixed spines, and large movable spine, subconical tubercles (9 on left) of varying with small spine at dorsal base, near lateral sizes, most with distal margin elevated; oth- margin of ramus. Telson with 3 spines in er obvious tubercles dorsal and ventral to each caudolateral corner of cephalic sec- mesial row; distal margin of mesial surface tion, middle one movable, mesial one on developed as 12th (10th on left) tubercle, left bifurcate; transverse suture strong; cau- with large spiniform tubercle and smaller dal margin of telson truncate and with slight rounded tubercle proximoventral to it; dor- median concavity. Uropods and telson with sal, mesial, and lateral surfaces of palm few setae dorsally. crowded with strong squamous tubercles of Epistome (Fig. 1H) with subtriangular varying sizes, and recumbent setae; ventral cephalic lobe bearing long, triangular ceph- surface of palm less densely tuberculate, alomedian projection; lobe strongly con- many tubercles subspiniform; articular stricted at base, transverse sulcus indistinct; ridge of palm, dorsally and ventrally, poorly margins of lobe slightly thickened, elevated defined, lateral eminence especially weak; (ventrally), somewhat undulant, incomplete lateral eminence ventrally with very strong at base of projection, and with long, dense subdistal spine. Fingers narrow, with op- setae; lateral apices thicker than rest of mar- posable surfaces contiguous and lacking gin, rounded, somewhat flared; floor (ven- prominent setae at bases; fingers slightly tral surface) of lobe slightly convex, very curved distoventrally in lateral aspect, dor- punctate, setiferous; body of epistome with sal surfaces studded with punctations and broad central depression bearing shallow tufts of stiff setae. Right dactyl 48.0% of cephalomedian fovea; lamellae punctate, ta- chela length, 1.1 times as long as palm; dor- pering laterally to subtruncate margin de- sal surface of dactyl with narrow, weak lon- void of tubercles; zygoma moderately gitudinal ridge, ventral surface rounded and arched, flanked cephalolaterally by usual without ridge; proximal Vz of ventral surface elongate pits. with 5 or 6 small tubercles; mesial margin of dactyl with 2 (4 on left) large, semierect Third maxilliped with tip of endopodite tubercles in row near base, and several -of ischium extending nearly to distal mar- smaller tubercles dorsal and ventral to gin of penultimate podomere of antennal them, rest of margin punctate; opposable peduncle, tip of exopodite extending to just margin with dense pad of denticles in 6 to beyond distal margin of merus of endopod- VOLUME 111, NUMBER 1 87

7 rows throughout length of dactyl, pad nar- between distal extremity of ridge and large rower near base; proximal half of finger distal spine; ventromesial ridge with large with 8 to 9 (5 on left) small tubercles dorsal distal spine, patch of 3 large spines just to denticles, basal 2 largest; ventral to den- proximal to distal spine, and 11 or 12 spi- ticles, surface with 1 small, subacute tuber- niform tubercles; other obvious tubercles on cle near base and 2 large contiguous tuber- dorsomesial surface, and patch of 5 to 6 cles situated at base of distal 34 of finger, tubercles between distal extremities of both interrupting denticles (on left, single large ridges; ventral surface of merus between tubercle at this site). Fixed finger with nar- ridges with some small tubercles and dense row longitudinal ridge dorsally, and mod- setae. Ischium with row of 4 (5 on left) sub- erate ridge ventrally; lateral surface of fin- spiniform tubercles on ventral ridge; suffla- ger rounded, with rows of deep punctations men obsolete on right, short on left. Merus and clumps of stiff setae; opposable margin of second through fourth pereiopods with with dense pad of denticles in 7 to 8 rows prominent distolateral spine. throughout length of finger, pad narrower Hooks on ischia of third and fourth per- near base; large subconical tubercle ventral eiopods simple (Fig. II), that on third long, to denticles just distal to midlength; proxi- subconical, slightly curved and overreach- mal V3 of finger with 5 (4 on left) small, ing basioischial articulation by most of rounded tubercles dorsal to denticles, third length, not opposed by tubercle on basis; from base largest. hook on fourth pereiopod short, vertically Carpus of cheliped (Fig. IN) 1.6 times as disposed, not reaching articulation, and as long as wide, length 74.4% of palm length; in "Diagnosis." Coxae of fourth and fifth dorsal surface with very shallow, slightly pereiopods also as in "Diagnosis." Ster- oblique sulcus, surface lateral to which nites between third and fourth pereiopods punctate, mesial to which with 2 to 3 rows with long setae. of subconical tubercles extending onto dor- For description of gonopod see "Diag- somesial surface; large spine mesial to dor- nosis." In addition, intact subapical setae sal articular eminence; ventral surface with (not illustrated) flanking mesial, cephalic, very strong distolateral spine, strong distal and lateral bases of cephalic process and spine mesial to distolateral one, smaller central projection, largely obscuring both spine just proximal to distomedian one, 2 elements. small acute tubercles proximal to both of Description of allotypic female.—Differ- the latter, and 2 weak spines and 4 tubercles ing from holotypic male, except in second- near mesial margin; latter with strong, ary sexual characters, as follows: Areola curved subdistal spine and 4 to 5 proximal 4.2 times as long as broad, constituting ap- tubercles. Merus of cheliped 2.9 times lon- proximately 28.0% of TCL (acumen dam- ger than deep, depth fairly uniform aged) and 42.8% of PCL, with 6 puncta- throughout length, latter 38.3% of TCL; tions across narrowest part. Cervical area dorsal surface with 2 strong, contiguous with 2 tubercles ventral to cervical spine. subdistal spines, and row of spiniform tu- Cephalolateral corners of cephalic section bercles along dorsomedian ridge; dorsal of telson with 4 spines on left, 3 on right. spines and tubercles flanked mesially and Antennal scale 2.9 times as long as broad. laterally by other subspiniform to squamous Chela of cheliped 1.7 times wider than tubercles; distal half of mesial surface tu- deep; mesial margin of palm with mesial berculate, lateral surface punctate and with row of 5 to 6 tubercles. Longitudinal ridges some minute, scattered tubercles; ventrolat- on fingers of cheliped well developed. Me- eral ridge with large distal spine near artic- sial surface of dactyl with single prominefit ular eminence, 3 other strong spines; 8 to 9 tubercle and other squamous to subsqua- small tubercles, and row of small tubercles mous tubercles near base; opposable sur- 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON face with 4 rounded tubercles on proximal tubercles. Hooks on ischia of third and V3 to V4, and several smaller tubercles dis- fourth pereiopods reduced; boss on coxa of tally; denticles in single row; opposable fourth pereiopod not pronounced, that on margin of fixed finger with row of 6 tuber- fifth pereiopod narrow. cles, denticles in single row. Carpus of che- Gonopods (Fig. 1C, E) with proximo- liped 1.5 times as long as wide, length mesial apophyses separated; mesial process 93.9% of length of mesial margin of palm; stout, tapering, tip directed caudally; gon- merus 2.4 times as long as deep, length opod in lateral aspect with "juvenile su- 33.4% of TCL; ventrolateral ridge with row ture"; cephalic convexity apparent; all ter- of 7 spines or spiniform tubercles in addi- minal elements blunter and thicker than in tion to distal spine, ventromesial ridge with form I male, not corneous, all except caudal 13 spines or tubercles and large distal spine. process identifiable and relationships clear- For description of annulus ventralis (Fig. ly visible; subapical setae sparse; gonopod 1G) see "Diagnosis." In addition, postan- in mesial aspect with poorly defined caudal nular sclerite nearly twice as wide as long, process, and adventitious process reduced about half as wide as annulus. Preannular to narrow ridge. sternite with broadly flared walls, deep me- Disposition of types.—The holotypic dian cleft in caudal half, and 5 to 6 protu- male, allotypic female, and morphotypic berant lobes on either side of cleft, caudal- male are in the crustacean collections of the most pair overhanging cephalic margin of NCSM (catalogue numbers NCSM C-2507, annulus. First pleopods uniramous, extend- C-2549, and C-2550, respectively), as are ing just beyond caudal margin of preannu- paratypes consisting of 3 8 I, 3 j 6,3 $, lar sternite when abdomen flexed. 3j 9. Description of morphotypic male, form Type locality.—North Carolina, Colum- II.—Differing from holotypic male in fol- bus County, Waccamaw River at NC 130 lowing respects: Areola 2.7 times as long near Brunswick County line, 8.0 air km as broad, constituting 28.2% of TCL (5.6 air mi) SSE of Old Dock (Freeland (41.3% of PCL), sparsely punctate; apex of USGS 7.5' quadrangle, UTM coordinates acumen reaching to proximal base of first 377521 ON/726190E). article of antennular flagellum. Antennal Range and specimens examined.— scale 3.2 times as long as wide. Mesial mar- Known only from the Waccamaw River ba- gin of palm of cheliped with mesial row of sin in North and South Carolina, where the 8 tubercles; lateral eminence of ventral ar- following collections have been made: ticular ridge with small tubercle. Mesial North Carolina. Columbus Co.—Wacca- margin of dactyl with 3 tubercles; opposa- maw R at NC 130 (type locality); 1 6 I ble margin with 3 minuscule tubercles ven- (NCSM C-2507), 17 Jun 1991, 1 $ (NCSM tral to denticles in proximal V4 of finger, and C-2234), ? Aug 1984, coll. D. R. Lenat; 2 small tubercles dorsal to denticles near Waccamaw R at Lake Waccamaw dam, S base of finger; denticles in single row. Op- end of SR 1967, ca. 7.4 air km S of town posable margin of fixed finger with 3 or 4 Lake Waccamaw; 1 6 I, 1 j 6, 1 j $ small tubercles, denticles in 2 to 3 rows. (NCSM C-316), 1 $ (NCSM C-2549), 22 Carpus of cheliped 1.8 times as long as Oct 1978, coll. A. L. Braswell, R. E. Ash- broad, length 83.7% of length of mesial ton, Jr., P. S. Ashton; Waccamaw R between margin of palm; merus 3.4 times as long as spillway & SR 1928; 1 6 I, 1 2 j 9 deep, length 33.7% of TCL; ventrolateral (NCSM C-2066), 29 Mar 1978, coll. W. S. ridge with 2 strong spines in addition to Birkhead; Waccamaw R below Bogue large distal spine, ventromesial ridge with Swamp, "near dam" at Lake Waccamaw; 1 11 small tubercles and large (broken) distal j

Waccamaw R, N of NC 130 (probably 2.8 of both fingers of the chela are densely river km N); 15 I (NCSM C-963), 1 6 II packed with denticles, arranged in five to (NCSM C-2550), 22 Apr 1979, coll. UNC- nine somewhat irregular rows. Females and Wilmington biologists; canal off Wacca- juvenile males have a single row of denti- maw R, 1.6 km N of river, apparently 12.0 cles on these surfaces, and the morphotypic air km S of Lake Waccamaw; 1 $ (USNM male has two to three rows on the fixed 129841), 29 Mar 1949, coll. E. C. Raney. finger and a single row on the dactyl. South Carolina. Horry Co.—Smith Lake at Size.—The largest specimen, a female end of Park Ave, northern Conway; 1 j 6 with a damaged acumen, has an estimated (NCSM C-3247), 10 May 1994, coll. R. G. TCL of 29.6 mm (PCL 19.4 mm). Four oth- Arndt & students. er females range from 18.5 to 25.5 mm Variations.—Significant variations are TCL (12.0-16.7 mm PCL). The four form addressed in the "Diagnosis," but others I males range from 19.5 to 27.4 mm TCL are also evident. The tubercle on the lateral (12.8-18.5 mm PCL). eminence of the ventral articular ridge of Life history notes.—Form I males have the palm of the chela varies from large and been found in March, April, June, and Oc- spiniform to small and subsquamous, and is tober. No ovigerous females or those with absent in one small female. The tubercles attached young have been collected. in the mesial row of the mesial margin of Crayfish associates.—Cooper & Bra- the palm range in number from five to thir- swell (1995:120-121) briefly discussed the teen, and in size from barely discernible to crayfishes of the Waccamaw River basin. large and obvious; in the four form I males The only species that have been taken with they number from ten to thirteen and are P. braswelli are Procambarus {Ortmanni- conspicuous, while in females they number cus) acutus (Girard, 1852), Procambarus from five to nine and generally are incon- (Ortmannicus) ancylus Hobbs, 1958b, and spicuous. The number of tubercles on the Procambarus (Ortmannicus) blandingii opposable margin of the fixed finger, exclu- (Harlan 1830). sive of the prominent subconical one, varies Relationships.—Based on the configura- from one to six, but the usual number is tion of the form I male gonopod, P. bra- two or three. The number of tubercles on swelli has its closest affinities with Procam- the opposable margin of the dactyl ranges barus (Ortmannicus) chacei Hobbs, 1958c, from two to thirteen, but usually is two to Procambarus (Ortmannicus) enoploster- five. In most specimens these tubercles are num Hobbs, 1947b, and Procambarus (Ort- very small, and in several they are scarcely mannicus) pictus (Hobbs 1940), is some- discernible. Some are dorsal to the denti- what more distantly related to Procambarus cles, but others are either ventral to them or (Ortmannicus) epicyrtus Hobbs, 1958c, and interrupt them ventrally. The tubercles is even more distantly related to its geo- along the dorsomedian ridge of the merus graphically nearest neighbor, P. lepidodac- vary in size and configuration, from barely tylus. visible, squamous tubercles to small spines. Procambarus braswelli may be distin- In all specimens there is a single cervical guished from all other members of the Pic- spine on each side of the carapace, but the tus Group by the combination of: a gono- allotype also has two tubercles ventral to pod whose distal one-fourth is only slightly the spine. There almost always are three caudally directed; a small but obvious ce- spines in each caudolateral corner of the cephalic convexity; an almost distally directed phalic section of the telson, but five speci- cephalic process whose caudal base is trans- mens have four spines in one corner, three versely expanded and forms a cowl or hood in the other. around the cephalic base of the slightly lon- In form I males the opposable surfaces ger, caudodistally directed central projec- 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tion; a long mesial process that is caudo- Literature Cited distally directed at about 45° to the shaft of the gonopod; a prominent, distally directed Cooper, J. E., & A. L. Braswell. 1995. Observations caudal knob that extends only slightly be- on North Carolina crayfishes (Decapoda: Cam- baridae).—Brimleyana 22:87-132. yond the proximocaudal bases of the ce- Cooper, M. R., & J. E. Cooper. 1977a. A comment phalic process and central projection; a on crayfishes. Pp. 198—199 in J. E. Cooper, S. large adventitious process that in mesial as- S. Robinson, & J. B. Funderburg, eds., Endan- pect obscures part of the proximomesial gered and threatened plants and animals of North Carolina. North Carolina State Museum bases of the central projection and caudal of Natural History, Raleigh, 444 pp. process; an acumen that on average com- , & . 1977b. Procambarus (Ortmanni- prises about 45% of the rostrum length; and cus) lepidodactylus Hobbs. Pp. 206-207 in J. E. a carapace that, caudal to the cervical Cooper, S. S. Robinson, & J. B. Funderburg, groove, is granulate both dorsally and lat- eds., Endangered and threatened plants and animals of North Carolina. North Carolina State erally. Museum of Natural History, Raleigh, 444 pp. Etymology.—Despite his being an unre- Fitzpatrick, J. F., Jr. 1983. How to know the fresh- pentant vertebrate zoologist who has always water crustacea. William C. Brown Company, "outcrayfished" me in the field, I take great Publishers, Dubuque, Iowa, 227 pp. Franz, R., & D. S. Lee. 1982. Distribution and evo- pleasure in naming this new species for Al- lution of Florida's troglobitic crayfishes.—Bul- vin L. Braswell, Curator of Lower Verte- letin of the Florida State Museum, Biological brates, NCSM, who has been a friend, col- Sciences 28:53-78. league, and congenial field companion for Frey, D. G. 1951. The fishes of North Carolina's bay many years. Suggested vernacular name: lakes and their intraspecific variations.—Journal of the Elisha Mitchell Scientific Society 67:1 — Waccamaw Crayfish. 44. Fuller, S. L. H. 1977. Freshwater and terrestrial mollusks. Pp. 143-194 in J. E. Cooper, S. S. Rob- Acknowledgments inson, & J. B. Funderburg, eds.. Endangered and threatened plants and animals of North Car- My thanks go to those collectors who olina. North Carolina State Museum of Natural History, Raleigh, 444 pp. provided the specimens of this new cray- Girard. C. 1852. A revision of the North American fish, and particularly to David R. Lenat. I Astaci with observations on their habits and am grateful, too, for the reviews of the geographical distribution.—Proceedings of the manuscript by Roger F. Thoma and Steve Academy of Natural Sciences of Philadelphia 6: Busack, and especially for the always astute 87-91. Harlan, R. 1830. Description of a new species of the attentions of Joseph F. Fitzpatrick, Jr. Nancy genus Astacus.—Transactions of the American Childs provided technical assistance in the Philosophical Society 3( 15):464-465. preparation of the figure. I also express my Hobbs, H. H., Jr. 1940. Seven new crayfishes of the sincerest gratitude to Alvin L. Braswell, genus Cambarus from Florida, with notes on other species.—Proceedings of the United John E. Cooper, Jr., Martha R. Cooper, Jes- States National Museum 89:387-423. se Perry, and particularly to Don Howard, . 1947a. A key to the crayfishes of the Pictus without whose unstinting assistance this pa- Subgroup of the genus Procambarus, with the per would never have been realized. My description of a new species from South Caro- greatest debt, which I cannot adequately ex- lina.—The Florida Entomologist 30(3):25—31. . 1947b. Two new crayfishes of the genus Pro- press, remains to the late Horton H. Hobbs, cambarus from Georgia, with notes on Procam- Jr., mentor and friend, whose outstanding barus pubescens (Faxon).—Quarterly Journal work on the members of the Pictus Group of the Florida Academy of Sciences 9:1-18. of Procambarus provided the framework . 1958a. The evolutionary history of the Pictus for understanding the relationships of P. Group of the crayfish genus Procambarus (De- capoda, Astacidae).—Quarterly Journal of the braswelli. Florida Academy of Sciences 21:71-91. VOLUME 111, NUMBER 1 91

. 1958b. Two new crayfishes of the genus Pro- silis) fullerkati (Bivalvia: Unionidae) from Lake cambarus from South Carolina.—Journal of the Waccamaw, Columbus County, North Carolina, Washington Academy of Sciences 48(5): 160- with a list of the other unionid species of the 168. Waccamaw River system.—Museum of Com- . 1958c. Two new crayfishes of the genus Pro- parative Zoology Occasional Papers on Mol- cambarus from South Carolina and Georgia.— lusks 4:289-298. Notulae Naturae 307:1-12. LeGrand, H. E., Jr. 1993. Natural Heritage Program . 1962. Notes on the affinities of the members list of the rare animal species of North Carolina. of the Blandingii Section of the crayfish genus North Carolina Natural Heritage Program, Ra- Procambarus (Decapoda, Astacidae).—Tulane leigh, 45 pp. Studies in Zoology 9(5):273-293. , & S. P. Hall. 1995. Natural Heritage Program . 1968. Crustacea: Malacostraca. Pp. K1-K36 list of the rare animal species of North Carolina. in F. K. Parrish, ed., Keys to water quality in- North Carolina Natural Heritage Program, Ra- dicative organisms (southeastern United States). leigh, 67 pp. Federal Water Pollution Control Administration, , & . 1997. Natural Heritage Program United States Department of the Interior, Wash- list of the rare animal species of North Carolina. ington, D. C., 36 pp. North Carolina Natural Heritage Program, Ra- . 1972. Crayfishes (Astacidae) of North and leigh, 82 pp. Middle America. Biota of freshwater ecosys- Menhinick, E. F 1991. The freshwater fishes of North tems identification manual no. 9. United States Carolina. North Carolina Wildlife Resources Environmental Protection Agency, Washington, Commission, Raleigh, 227 pp. D. C„ 173 pp. Porter, H. J., & K. J. Horn. 1984. Freshwater Mollusca . 1974. A checklist of the North and Middle of upper Waccamaw River, North and South American crayfishes (Decapoda: Astacidae and Carolina.—Journal of the Elisha Mitchell Sci- Cambaridae).—Smithsonian Contributions to entific Society 97:270. Zoology 166:1-161. Rohde, F. C„ R. G. Arndt, D. G. Lindquist, & J. F. . 1989. An illustrated checklist of the Ameri- Parnell. 1994. Freshwater fishes of the Caroli- can crayfishes (Decapoda: Astacidae, Cambari- nas, Virginia, Maryland, and Delaware. Univer- dae, and Parastacidae).—Smithsonian Contri- sity of North Carolina Press, Chapel Hill, 222 butions to Zoology 480:1-236. pp. , & R. Franz. 1986. New troglobitic crayfish Shute, J. R., P. W. Shute, & D. G. Lindquist. 1981. with comments on its relationship to epigean Fishes of the Waccamaw River drainage.— and other hypogean crayfishes of Florida.— Brimleyana 6:1-24. Journal of Crustacean Biology 6:509-519. Teulings, R. P., & J. E. Cooper. 1977. Cluster areas. , & D. J. Peters. 1977. The entocytherid ostra- Pp. 409-433 in J. E. Cooper, S. S. Robinson, cods of North Carolina.—Smithsonian Contri- & J. B. Funderburg, eds., Endangered and butions to Zoology 247:1-73. threatened plants and animals of North Caroli- , H. H. Hobbs, III, & M. A. Daniel. 1977. A na. North Carolina State Museum of Natural review of the troglobitic decapod crustaceans of History, Raleigh, 444 pp. the Americas.—Smithsonian Contributions to Williams, A. B„ L. G. Abele, D. L. Felder, H. H. Zoology 244:1-183. Hobbs, Jr., R. B. Manning, P. A. McLaughlin, Hubbs, C. L„ & E. C. Raney. 1946. Endemic fish & I. Perez Farfante. 1989. Common and sci- fauna of Lake Waccamaw, North Carolina.— entific names of aquatic invertebrates from the Miscellaneous Publications of the University of United States and Canada: Decapod crusta- Michigan Museum of Zoology 65:1-30. ceans. American Fisheries Society Special Pub- Johnson, R. I. 1984. A new mussel, Lampsilis (Uunp- lication 17, 77 pp.