Cambarus (Puncticambarus) theepiensis, Loughman & Foltz & Garrison & Welsh, 2013

Loughman, Zachary J., Foltz, David A., Garrison, Nicole L. & Welsh, Stuart A., 2013, Cambarus (P.) theepiensis, a new species of crayfish (Decapoda: Cambaridae) from the coalfields region of Eastern Kentucky and Southwestern West Virginia, USA, Zootaxa 3641 (1), pp. 63-73 : 64-72

Cambarus (Puncticambarus) theepiensis new species

Figures 1–3 , Tables 1–2

Diagnosis. Body and eyes pigmented. Posterior region of rostrum concave and deflected anteriorly. Rostrum margins thickened, parallel to slightly converging at acumen. Acumen distinctly triangular with prominent dorsally deflected spiniform tubercle at terminus. Areola 1.8–6.7 (x = 4.3, n = 60, SD = 2.4) times as long as wide with 3–9 (usually 6) punctations across narrowest point. Cervical spines rarely present (3% of individuals; n = 60); usually (97% of individuals; n =60) 1–3 (x = 2, SD = 1.3) weakly developed tubercles present. Mandibular, branchiostegal, and orbital regions of carapace with well-developed tubercles. Postorbital ridges short; spiniform, laterally deflected tubercle present in juveniles and subadults; adult postorbital ridge terminating in truncated tubercle; rarely spiniform tubercles present in adults. Suborbital angle acute. Antennal scale widest in middle, 1.2–2.9 (x = 2.2, n = 60, SD = 0.03) times as long as wide. Total carapace length (TCL) 1.7–2.2 (x = 2.0, n = 60, SD = 0.30) times longer than width. Form I and II males possessing hook on ischium of third pereopods only; hook gently curved at apex, overarching basioischial joint in form I males, not reaching basioischial joint in form II males; hooks not opposed by tubercle on basis. Mesial surface of chelae with two rows of tubercles; mesialmost row with 6–10 (x = 9, n = 60, SD = 0.92) tubercles, second dorsal row with 4–8 (x = 6, n = 60, SD = 0.89) tubercles. Occasionally tubercles extend from mesial surface to body of palm. Dorsal longitudinal ridge of dactyl consisting of several well developed scattered tubercles. Dorsomedian ridge of fixed finger of propodus pronounced. Well defined lateral impression at the junction of the fixed figure with the propodus. Dactyl and fixed finger with sharp corneous subterminal tip. Form I male palm length 60.1–72.1% (x = 64.1.%, n = 20, SD = 4.0%) of palm width, form I male palm length 25.1–34.0% (x = 28.3%, n = 20, SD = 2.2%) of total propodus length; female dactyl length 55.7–70.0% (x = 64.1, n = 20, SD = 3.5%) of total propodus length. First pleopod of form I male with short terminal elements. Central projection not tapering distally; recurved> 90º to main shaft of gonopod, with distinct subapical notch. Mesial process directed 90° to shaft, bent cephalolaterally; inflated cephalically, tapering to distinct caudal point at or slightly beyond terminance of central projection. Annulus ventralis immovable; distinctly asymmetrical posteriorly; cephalic portion with median trough leading to strongly sculptured central fossa; exaggerated “S” bend in sinus terminating at caudal edge.

Description of holotypic male, form I. ( Figs. 1 A,B,D,E,F,G,I,J ).––Body somewhat compressed dorsoventrally ( Fig. 1A ); carapace posterior to cervical groove slightly wider than abdomen. Carapace depth less than carapace width at caudodorsal margin of cervical groove. Total carapace length 46.6 mm; post orbital carapace length 38.9 mm. Areola 5.5 times longer than wide, with 8 punctations across narrowest part ( Fig 1E ); length of areola 35.1 % of TCL (42.4 % of postorbital carapace length (PCL). Rostrum moderately excavated, moreso anteriorly than posteriorly; margins thickened, subparallel and continuous to base of acumen; floor of rostrum with numerous punctations. Rostrum 1.6 times longer than wide. Acumen distinctly triangular, ending in dorsally deflected corneous tip ( Fig. 1A ). Postorbital ridges well developed, short, terminating in weak cephalic tubercles. Suborbital angle acute, with tubercle ( Fig. 1A ). Cervical spine absent; single weakly developed tubercle present. Mandibular, branchiostegal, and orbital regions of carapace punctated with well-developed tubercles; greatest tubercle density in hepatic region.

Abdomen supraequal in length to carapace, pleura rounded cephaloventrally, angled distoventrally. Lateral margin of terga angulate; lateral margin of second pleluron deeply furrowed. Cephalic section of telson with 2 large spines in each caudolateral corner. Proximal podomere of uropod with distal spine on mesial lobe; mesial ramus of uropod with median ridge ending distally in distomedian spine not overreaching margin of ramus; laterodistal spine pronounced. Distal margin of proximal segment of lateral ramus of right uropod having 18 immovable, small spines and 1 lateral, large, movable spine. Cephalomedian lobe of epistome subtriangular, zygoma moderately arched ( Fig. 1J ); cephalolateral margins thickened, forming sharp angle at junction with endostyle ( Fig. 1J ). Body of epistome possessing prominent cephalomedian fovea. Antennal scale broadest in middle; lateral margin thickened, terminating in large corneous spine; setiferous. Right antennal scale 6.9 mm long, 3.1 mm wide ( Fig. 1D ). Tip of right antenna reaching middle of telson when adpressed.

Mesial surface of right chela with 2 rows of tubercles; mesialmost row with 9 tubercles, second dorsal row with 7 tubercles ( Fig. 1F ). Palm length 63.0 % of palm width; depth of palm 11.2 mm. Ventral surface of palm containing 3 subpalmar tubercles. Dorsal longitudinal ridge of dactyl developed and possessing moderate sized tubercles ( Fig. 1F ); dactyl terminating in large corneous spine. Dorsomedian ridge of fixed finger of propodus pronounced. Well defined lateral impression at the junction of fixed figure with the propodus. Fixed finger junction with defined lateral impression; numerous setiferous punctations present. Dactyl and fixed finger of propodus with sharp, corneous tip. All measurements and counts from right chela.

Carpus with prominent dorsal furrow ( Fig. 1F ) and 4 weak dorsomesial tubercles; rest of surface with some setiferous punctations; mesial margin with large, procurved spine at about midlength, and reduced proximal spine. Distodorsal surface of merus with 2 spiniform tubercles; ventrolateral ridge with 10 small spines and large, corneous distal spine; ventromesial ridge with 2 well-developed spines; ventrolateral margin of ischium with 3 small, spiniform tubercles. Carapace depth less than width. Hook on ischium of third pereopods only ( Fig. 1I .); hook gently curved at apex, overarching basioischial joint, not opposed by tubercle on basis. Form I gonopod as described in diagnosis ( Fig. 1B ); tip reaching anterior margin of fourth caudomesial boss ( Fig. 1G ).

Description of allotypic female. ( Fig. 1H , Table 1 ). Differing from holotype in following respects; carapace height less than carapace width (17.6 and 19.0 mm, respectively); TCL 42.6 mm, PCL 35.7 mm. Areola 36.2% of TCL (43.1% of PCL), 3.6 times as long as wide. Posterior portion of rostrum more excavated than anterior portion; rostrum 1.4 times longer than wide. Abdomen length 44.7 mm. Mesial surface of chelae with 2 rows of tubercles; mesialmost row with 7 tubercles, second dorsal row with 5. Palm length (9.4 mm) 61.4% of palm width (15.3 mm); depth of palm 9.1 mm. Antennal scale 7.0 mm long, 2.7 mm wide. All measurements and counts from right chela. Annulus ventralis as described in diagnosis ( Fig. 1H ); width of postannular sclerite half total width of annulus ventralis; first pleopods uniramous, reaching central region of annulus ventralis when abdomen flexed.

Description of morphotypic male, form II. ( Fig. 1C , Table 1 ). Differing from holotype in the following respects: Carapace wider than abdomen (25.6 and 21.5 mm, respectively); carapace height less than carapace width (19.9 and 25.6 mm respectively); TCL 47.3 mm and PCL 39.6 mm. Areola length 48.0% of TCL (57.3% of PCL), 5.6 times longer than wide. Rostrum margins subparallel to base of acumen; rostrum ventrally deflected; rostrum 1.3 times as long as wide. Abdomen 47.5 mm long. Mesial row of tubercles on palm of chela with 8 tubercles; second dorsal row with 6 tubercles. Palm length (12.5 mm) 64.1% of palm width (19.4 mm). Antennal scale 7.0 mm long, 3.2 mm wide. Gonopods reaching anterior margin of 4 th pereopod caudomesial boss; 21.0% of TCL length. Central projection curved 90° to shaft, with complete apex; rounded ( Fig. 1C ). Mesial process tapered, bulbous, directed caudolaterally. Hook on ischium of third pereopod small, not reaching basioischial joint. All measurements and counts from right chela.

Size. Form I male (n = 21) TCL ranges in size from 29.8–60.7 mm (PCL 24.9–50.1 mm) with a mean TCL of 41.8 mm. Form II male (n = 21) mean TCL is 40.2 mm and ranges in size from 33.5–50.0 mm (PCL 27.9–42.5 mm). Non-ovigerous female (n = 21) TCL mean is 40.7 mm and ranges from 32.6–51.0 mm (PCL 27.8–42.2 mm). Ovigerous female (n = 4) TCL ranges in size from 25.0– 36.1 mm (PCL 19.7–30.0 mm) with a mean of 31.7 mm. The largest specimen examined was a form I male with TCL of 60.7 mm (PCL 50.1 mm).

Color. Carapace ground color red-brown to pink-brown; posterior margin of carapace dark chestnut brown. Hepatic and antennal region of carapace punctuated with yellow, white, or tan tubercles. Postorbital ridge color same as carapace, to orange-brown. Rostrum margins and acumen distinctly orange to red-orange or red-brown, rarely red. Cephalic section of carapace immediately anterior to and including cervical groove black; mandibular abductor scars ranging from grey to brown. Lateral margin of antennal scale gray-pink to blackish red; body of antennal scale pink to red-brown. Antennal flagellum and antennules red-brown, with olivaceous hue; dorsal surface of lamellae green-brown to pink-brown; ventral surface tan to olivaceous. Dorsal surface of chelae olive green to brown-orange with light green highlights, olivaceous to brown mottling; mesial, second dorsal row, and dorsal surface of dactyl tubercles amber, orange or red-brown. Denticles on opposable surfaces of fingers yellow, white, or tan. Ventral surface of chelae pink-orange or pink-brown. Dorsal surface of carpus brown, olivaceous or green-brown; occasionally orange; region adjacent to and including furrow olivaceous orange to green; carpus spine cream. Merus orange-brown, cream, or olivaceous brown. Podomeres of pereopods light blue, blue-green, or green-brown; joints of pereopod podomeres pink. Dorsal and dorsalateral surface of abdomen same color as carapace; anterior region of abdomen with slight olivaceous tint; tergal margins brown to reddish brown. Uropods brown, red, red-brown to pink-brown, with olivaceous tint; margins gray to brown. Ventral surface of abdomen and carapace tan. Dorsal ridge of form I gonopod central projection amber; body of central projection, gonopod, and mesial process tan. Form II gonopod and all associated processes cream. Cephalic portion of annulus ventralis pink to pink-cream; ridge of fossa pink; caudal region of annulus ventralis ranges from pink to cream colored.

Type locality. Left Fork of Millers Branch, 2.2 miles (3.54 km) from CR 22 / CR 27 intersection on CR 22, 0.05 mi (0.08 km) north of Crockett , Wayne County, West Virginia (38.23766º N, - 82.37685ºW). The holotype was collected underneath a large slab boulder isolated on a bedrock glide. The allotype and morphotype were also collected under isolated slabs in a low velocity reach of stream with sandy substrate. Left Fork of Millers Branch between 3.0–8.0 m wide, and 0.2–1.2 m deep when the type series was collected. Stream substrate was composed primarily of sands and gravels with numerous slabs and boulders. Leaf packs in pools and coarse woody debris provided additional habitat. The holotype, morphotype, and allotype were collected on 1 December 2012 by DAF,. K. R. Loughman, C. Z. Loughman, and ZJL.

Disposition of types. The holotype, allotype, and morphotype are deposited in the National Museum of Natural History ( USNM), Smithsonian Institution, Washington, D. C. (catalogue numbers USNM 1197339, 1197340, 1197341, respectively). Paratypes are deposited in the following museums and collections: Carnegie Museum of Natural History, Pittsburgh, PA ( CMNH 38224), and West Liberty University Astacology Collection, West Liberty, WV ( WLU 1210) .

Range and specimens examined. The range of Cambarus theepiensis is limited to the junction of the Appalachian Plateau with the northern Cumberland Mountains in southwestern West Virginia and eastern Kentucky. In West Virginia it occurs in the Lower Ohio, Guyandotte, Twelve Pole, and Big Sandy basins. In Kentucky it is limited to the Levisa Fork, Little Sandy, and Big Sandy basins. Cambarus theepiensis is most frequently encountered in moderate to large (3–6 order) streams possessing runs with boulders or slabs. As stream gradient increases, C. theepiensis numbers decline (ZJL personal observation). In the Big and Little Sandy basins, where slabs and boulders are rare substrates, C. theepiensis burrows extensively in stream banks.

All West Virginia collections in 2009 and 2011 were made by ZJL, DAF, NLG and associated West Liberty University Astacology Laboratory summer field crews and are housed in the West Liberty University Astacology Collection, denoted with the prefix WLU. Field crew members for 2009 included Evan I. Hewitt, and Mathew I. M c Kinney. Summer 2011 field crew members were Nate D. Taylor and Kinsey T. Skalican. Kentucky specimens were either collected by ZJL or loaned from the Eastern Kentucky University Branley A. Branson Museum of Zoology (EKU) or were part of the United States National Museum (USNM) collection. The following abbreviations are defined as follows: appx = approximately; CR = county road; I = interstate; KY = Kentucky state highway; mi = miles; RD = road; US = U. S. route; WV = West Virginia state highway; Rd = road

A total of 485 specimens was examined from the following 85 localities. KENTUCKY: FLOYD CO: (1.) EKU 497, Right Fork of Beaver Creek near Eastern High School , 15 May 1978, 1 II ♂. H. D. Bryant et al. (2.) EKU 2652, Small drainage ditch 8.5 mi E of Salyersville beside Mountain Parkway, 2 Jan 1968, 1 I ♂, D. L. Batch. (3.) USNM 359217, Right Fork of Beaver Creek. on KY 80 at confluence with Wilson Creek , 2.1 mi WSW of Langley, 21 Jun 1978, 1 II ♂. S. Call. JOHNSON CO: (4.) USNM 204997, Paint Creek in Paintsville , 1 mi N of Carbon, 24 Jun 1933, 2 ♀. MARTIN COUNTY: (5.) EKU 565, Unnamed tributary due N of Warfield along KY 292 , 26 Aug 1981, I ♂. J. R. Macgregor et al. LAWRENCE COUNTY: (6.) WLU 1208, Hood Creek , at crossing of KY 201, 29 Apr 2012, 5 ♀, 2 I ♂, 7 II ♂ (7.) WLU 1209, Blaine Creek, falls in Fallsburg , 29 Apr 2012, 1 ♀, 2 Juv. (8.) WLU 1207, Caney Fork, adjacent to KY 1, 1.1 mi W of Webbville , 29 Apr 2012, 3 ♀, 2 II ♂. (9.) USNM 313737, Hood Creek at KY 32 In Blaine , 22 Oct 1974, 4 I ♂, 2 juv ♂, 4 ♀, R.W. & J. W. Bouchard (10.) USNM 359217, Rockastle Creek Rd, 0.16 Km W 0f KY 3, 3 Oct 1978, 1 II ♂, 1 ♀. S. Call. (11.) WLU 1207, Caney Fork , adjacent to KY 1, 1.1 mi W of Webbville, 29 Apr 2012, 3 ♀, 2 II ♂. (12.)WLU 1208, Hood Creek , at crossing of KY 201., 29 Apr 2012, 2 OF, 5 ♀, 2 I ♂, 7 II ♂ (13.)WLU 1209, Blaine Creek, falls in Fallsburg , 29 Apr 2012, 1 ♀, 2 Juv. WEST VIRGINIA: CABELL COUNTY (14.) WLU 1020, Sevenmile Creek, at intersection of CR 11 & Blue Sulphur Rd ., 11 Aug 2012, 3 ♀, 2 II ♂. (15.) WLU 1027, Heath Creek (Guy), crossing of SR 10, at intersection with CR 42, 12 Jul 2011, 3 ♀, 1 II ♂. (16.) WLU 1028, Merritt Creek , parallel to CR 68, 1.1 mi SW of SR 10 in Sarah, 12 Jul 2011, 4 ♀, 1 II ♂. (17.) WLU 1029, Ninemile Creek , crossing of CR 7, 1.0 mi N of SR 2 in Lesage, 12 Jul 2011, 1 ♀. (18.) WLU 1030, Sevenmile Creek , adjacent to CR 11, at intersection w/ CR 12 in Lesage, 12 Jul 2011, 2 ♀, 2 II ♂. (19.) WLU 1031, Big Cabell Creek , adjacent to CR 18, 0.8 mi E of intersection w/ CR 1, 12 Jul 2011, 3 ♀, 6 II ♂. (20.) WLU 1038, Madison Creek , crossing of CR 49, 2.0 mi S of SR 10 in Salt Rock, 13 Jul 2011, 1 II ♂. (21.) WLU 1047, Bowen Creek , parallel to CR 49, 1.8 mi SE of junction w/ CR 43, 15 Jul 2011, 1 Juv. (22.) WLU 1060, Tyler Creek , parallel to CR 31, 0.1 mi NE of junction w/ WV 10 in Salt Rock, 16 Jul 2011, 3 Juv. (23.) WLU 1061, Fudges Creek , crossing of CR 29/2 at intersection w/ CR 29, 16 Jul 2011, 1 II ♂ (24.) WLU 1062, Lee Creek , at intersection of CR 60/19 and I- 64 in Milton, 16 Jul 2011, 1 II ♂. (25.) WLU 1063, Balls Gap Branch, parallel to CR 27, 1.8 mi S of US 60 (Main St.) in Culloden, 16 Jul 2011, 3 ♀, 1 II ♂. (26.) WLU 1064, Coon Creek , parallel to CR 2, 3.2 mi N of WV 34 in Nye, 16 Jul 2011, 3 II ♂. LINCOLN COUNTY: (27.) WLU 1032, Mud River headwater, crossing of CR 7 in Myra, 13 Jul 2011, 5 ♀, 1 II ♂. (28.) WLU 1033, Big Creek , crossing of CR 7, 1.0 mi S of CR 45 & CR 7 in Sias, 13 Jul 2011, 2 ♀, 4 II ♂. (29.) WLU 1034, Parsoner Creek , adjacent to CR 44, 0.7 mi E of intersection w/ CR 46 in Palermo, 13 Jul 2011, 7 ♀, 1 I ♂, 6 II ♂. (30.) WLU 1035, Merritt Creek , adjacent to CR 3/ 7, 1.3 mi SSW of WV 3 in Griffithsville, 13 Jul 2011, 3 II ♂. (31.) WLU 1036, Trace Creek, adjacent to CR 47, 0.2 mi E of intersection w/ CR 29 ( Trace Creek Rd. ), 13 Jul 2011, 2 II ♂. (32.) WLU 1037, Buffalo Creek (Guy), parallel to CR 6, 1.0 mi SW of intersection w/ CR 1, 13 Jul 2011, 2 ♀. (33.) WLU 1039, Fourmile Creek , crossing of CR 11, 1.8 mi WSW of WV 10 in Branchland, 14 Jul 2011, 3 ♀, 2 II ♂. (34.) WLU 1040, Sixmile Creek headwater, crossing of CR 22, 0.9 mi W of intersection with CR 72, appx. 3.7 mi W of Sias, 14 Jul 2011, 3 ♀, 3 II ♂. (35.) WLU 1041, Ninemile Creek , at intersection of CR 48 & CR 54, 2.3 mi E of Midkiff, 14 Jul 2011, 9 ♀, 5 II ♂. (36.) WLU 1042, Furnett Creek , crossing of CR 10/10, 3.1 mi N of WV 10 in Ranger, 14 Jul 2011, 3 ♀, 2 II ♂, 31 Juv. (37.) WLU 1043, Lick Branch, crossing of CR 10/14, 0.4 mi SW of junction w/ SR 10 in Ranger , 14 Jul 2011, 1 II ♂. (38.) WLU 1044, Fourteenmile Creek , WV 37, 0.8 mi SW of junction w/ WV 10, 14 Jul 2011, 1 ♀, 5 II ♂. (39.) WLU 1045, East Fork Fourteenmile Creek , parallel to WV 10, 2.5 mi S of Ranger, 14 Jul 2011, 1 II ♂, 1 Juv. LOGAN COUNTY: (40.) WLU 720, Elklick Branch, parallel to unnamed road, 0.2 mi SE from CR 16, 1.9 mi drive NE of Lorado , 10 Jun 2009, 2 ♀, 5 II ♂. (41.) WLU 721, Rum Creek , crossing of CR 14, 0.5 mi W of Yolyn, 10 Jun 2009, 2 ♀, 3 II ♂. (42.) WLU 722, Rich Creek , parallel to CR 10/4, appx. 2.5 mi WNW of Man, 10 Jun 2009, 1 ♀, 1 II ♂. (43.) WLU 723, Green Branch, adjacent to CR 55, 0.2 mi S of WV 10, 1.9 mi W of Mallory , 10 Jun 2009, 4 ♀, 3 II ♂. MASON COUNTY: (44.) WLU 1065, Right Fork Eighteenmile Creek , crossing of CR 39/9 at intersection w/ CR 41 in Upland, 17 Jul 2011, 3 ♀, 3 II ♂. (45.) WLU 1066, Eighteenmile Creek , crossing of CR 80, at intersection w/ CR 41, 17 Jul 2011, 2 II ♂. (46.) WLU 1067, White Pine Creek, adjacent to CR 8, 1.5 mi SSW of CR 37 ( CR 9) in Pliny , 17 Jul 2011, 1 ♀. RALEIGH COUNTY: (47.) WLU 724, Mill Branch, adjacent to WV 16, 0.3 mi S of Ury , 12 Jun 2009, 1 ♀, 1 I ♂, 1 II ♂. (48.) WLU 725, Mill Branch , adjacent to WV 16, 0.8 mi S of Ury, 12 Jun 2009, 1 ♀, 7 II ♂. (49.) WLU 726, Winding Gulf , adjacent to CR 3/21, 0.6 mi NW of Winding Gulf, 12 Jun 2009, 11 ♀, 12 II ♂. WAYNE COUNTY: (50.) WLU 741, Mill Creek , crossing of CR 36/2, 0.9 mi N of Delta Road 22, appx. 2.5 mi SW of Radnor , 11 Jul 2009, 3 ♀, 1 II ♂. (51.) WLU 1021, Mill Creek , crossing of CR 36 in Eddie, 11 Jul 2011, 2 II ♂. (52.) WLU 1022, Upper Tabors Creek , crossing of CR 37/2, 0.6 mi N of intersection with SR 37, 11 Jul 2011, 6 Juv. (53.) WLU 1023, Gragston Creek , crossing of CR 18, 1.9 mi SE of US 52 intersection, 11 Jul 2011, 1 ♀, 1 II ♂. (54.) WLU 1024, Whites Creek , crossing of CR 20 at intersection with CR 19, 11 Jul 2011, 5 ♀. (55.) WLU 1025, Balangee Branch, adjacent to intersection of CR 7 & CR 7/35, 11 Jul 2011, 3 ♀, 3 II ♂. (56.) WLU 1026, Buffalo Creek , crossing of CR 14/2, 0.1 mi W of intersection with CR 7, 11 Jul 2011, 1 ♀. (57.) WLU 1046, Butler Adkins Branch, adjacent to Delta Road 9, 0.3 mi N of CR 17/8, appx. 13.7 mi E of Lavalette, 14 Jul 2011, 7 ♀, 6 II ♂. (58.) WLU 1048, Raccoon Creek , adjacent to CR 43, 0.6 mi S of junction with CR 49 & CR 17/ 8, 15 Jul 2011, 4 ♀, 6 II ♂ (59.) WLU 1049, Little Lynn Creek , at intersection of CR 22 & CR 26, 0.7 mi NE of WV 37 in East Lynn, 15 Jul 2011, 1 ♀. (60.) WLU 1050, Wilson Creek , at intersection of CR 7/4 & CR 17/ 5 in Wayne, 15 Jul 2011, 1 ♀, 1 II ♂. (61.) WLU 1051, East Fork Twelvepole Creek , adjacent to WV 37, 1.7 mi W of intersection w/ CR 24, 15 Jul 2011, 2 II ♂. (62.) WLU 1052, East Fork Twelvepole Creek , crossing of CR 37/ 13 in East Lynn, 15 Jul 2011, 1 II ♂. (63.) WLU 1053, Camp Creek , crossing of CR 25, 1.4 mi E of WV 37 in East Lynn, 15 Jul 2011, 1 ♀, 1 I ♂, 2 II ♂. (64.) WLU 1054, Cove Creek , parallel to WV 37, 1.2 mi W of junction w/ CR 37/ 20, 15 Jul 2011, 1 ♀, 3 II ♂. (65.) WLU 1055, Lynn Creek , at intersection of CR 30 & CR 30/1, 15 Jul 2011, 4 ♀, 1 I ♂, 3 II ♂, 2 Juv. (66.) WLU 1056, West Fork Twelvepole Creek , crossing of CR 34 at intersection with WV 152 in Genoa, 15 Jul 2011, 2 ♀, 2 II ♂. (67.) WLU 1057, Moses Fork, parallel to SR 152, 1.6 mi S of junction w/ CR 35, 15 Jul 2011, 1 ♀. (68.) WLU 1058, Patrick Creek , at intersection of CR 7/5 & CR 7, 1.0 mi W of WV 37 ( WV 152), 15 Jul 2011, 5 II ♂, 3 Juv. (69.) WLU 1059, Big Creek , parallel to CR 1/6, 0.2 mi W of CR 52/8, 0.3 mi W of WV 152 in Wayne, 15 Jul 2011, 6 ♀, 6 II ♂. (70.) WLU 1205, Twelvepole Creek at Dixon Dam , 09 Oct 2004, 2 I ♂ Z. J. Loughman and C. G. Vopal. (71.) WLU 1206, Butler Adkins Branch , 0.5 mi from intersection of CR 17, 24 Apr 2010, 3 ♀, 5 II ♂. (72.) TYPE SERIES ( USNM 1197339 holotype, USNM 1197340 allotype, USNM 1197341 morphotype; CMNH 38224 paratypes ,, WLU 1210 paratypes, Left Fork of Millers Branch , 2.2 miles from CR 22 & CR 27 intersection on CR 22, 0.05 mi N of Crockett, 01 Dec 2012, 1 ♀, 2 II ♂, 3 Juv. (73.) WLU 742, West Fork Twelvepole Creek, crossing of Turkey Creek Rd , at junction with CR 41 in Wilsondale, 11 Jul 2009, 3 ♀, 5 II ♂. WYOMING COUNTY: (74.) WLU 727, Backers Creek , parallel to SR 10, 1.2 mi S of Bud, 1.1 mi N of Montecarlo, 12 Jun 2009, 3 ♀, 3 II ♂. (75.) WLU 728, Pinnacle Creek crossing of CR 18, 1.6 mi SW of Herndon Heights, 12 Jun 2009, 4 ♀, 3 II ♂. (76.) WLU 729, Pinnacle Creek , crossing of CR 18, 1.6 mi SW of Herndon Heights, 12 Jun 2009, 6 ♀, 1 I ♂, 2 II ♂. (77.) WLU 730, Whiteoak Branch, crossing of CR 18, 1.9 mi SW of Herndon Heights , 12 Jun 2009, 5 ♀, 2 II ♂. (78.) WLU 731, Cabin Creek , adjacent to CR 10/13, 3.4 mi drive NE of New Richmond, 12 Jun 2009, 4 ♀, 2 II ♂. (79.) WLU 732, Little Indian Creek , parallel to SR 16, 0.8 mi drive S of CR 12/ 4 in Wolf Pen, 13 Jun 2009, 4 ♀, 3 II ♂. (80.) WLU 733, Little Indian Creek, parallel to CR 12/4, 2.2 mi NW of WV 16 in Wolf Pen , 1.4 mi SE of Shop Branch Rd. in Fanny, 13 Jun 2009, 6 ♀, 2 I ♂. (81.) WLU 734, Huff Creek , adjacent to WV 10, 4.0 mi drive W of Cyclone, 13 Jun 2009, 3 II ♂. (82.) WLU 735, Huff Creek, parallel to WV 10, 5.1 mi drive W of Cyclone , 1.3 mi E of Gillman Bottom, 13 Jun 2009, 1 ♀, 2 II ♂. (83.) WLU 736, Guyandotte River , adjacent to WV 97, 0.4 mi E of Pump Creek Rd. in Baileysville, 09 Jul 2009, 2 ♀, 2 II ♂. (84.) WLU 737, Pinnacle Creek , crossing of CR 18, 1.6 mi SW of Herndon Heights, 09 Jul 2009, 14 ♀, 5 I ♂, 13 II ♂. (85.) WLU 743, Clear Creek , parallel to CR 6, 0.8 mi W of Hanners Branch Rd. in Simon, 09 Jul 2009, 1 ♀, 1 II ♂.

Conservation status. Cambarus theepiensis should be listed as vulnerable (V) according to the American Fisheries Society criteria ( Taylor et al. 2007), and assigned a G3 ranking according to the global conservation criteria ( Masters 1991) for conservation listing due to its limited range. Cambarus theepiensis should be listed as near threatened (VU) using the International Union for the Conservation of Nature (IUCN 2001) criteria due to its narrow distribution and extreme land use practices occurring within the central portion of its range.

Habitat and life history notes. Cambarus theepiensis occurs among boulders and slabs during all months of the year, and utilizes depositional habitats when available. Several streams in the Big Sandy , Little Sandy , and Lower Guyandotte basins have limited amounts of slabs and boulders and are dominated by sand substrates. In these situations C. theepiensis utilize simple burrows in hardpan glides and banks as refuge. Cambarus theepiensis populations decrease as gradient increases from main stems to headwater streams. Based on surveys performed in 2009 and 2011, C. theepiensis populations are most robust in the Twelvepole , Lower Guyandotte , and Lower Ohio basins, and reduced in the Big Sandy and Little Sandy basins (ZJL, unpublished data). Male form lacks seasonality in this species, with form I and II males present throughout the year. Eleven ovigerous females have been collected; two from Hood Creek , Lawrence Co. Kentucky, and nine from the type locality, Left Fork of Millers Branch , Wayne Co. West Virginia ( Table 2 ). Based on available data, egg extrusion occurs from late April through June. First through third instars are present from June through October, and molt to fourth stage instars by September. Young of the year appear in streams en masse independent of females in September (DAF, personal observation). Cambarus theepiensis growth rates and maximum age are unknown.

Crayfish associates. Cambarus (P.) theepiensis has been collected with Cambarus (C.) bartonii cavatus Hay ; 1902, Cambarus (J.) distans Rhoades; 1944, Cambarus (J.) dubius Faxon ; 1884, Cambarus (P.) veteranus Faxon ; 1914, Cambarus (T.) thomai Jezerinac; 1993, Orconectes (Procericambarus) cristavarius Taylor ; 2000 and Orconectes (Procericambarus) rusticus (Girard; 1852).

Variation. Ontogenetic variation in rostrum morphology and tubercle shape exists in C. theepiensis . Adult rostral margins are subparallel and inflated, with the anterior terminus of the margin distinct from the origin of the acumen. Juvenile rostra are weakly convergent, slightly inflated, and grade directly into the acumen without a distinct demarcation. Postorbital ridges of juveniles frequently (68%; n = 44) terminate in small spinose tubercles; adult postorbital ridges terminate in ablated tubercles (95%; n = 63), rarely in spinose tubercles (5 %; n = 63).

Geographic variation exists in the shape of adult rostra and the number of palmar tubercles. In the eastern portion of its range (i.e., headwaters of the Guyandotte River), rostral margins of C. theepiensis are mostly subparallel and occasionally convergent. Convergent rostra were only observed in form II males and females; they were always subparallel in form I males. Across the central and western portion of the range of C. theepiensis adult rostra are consistently broad and subparallel. Western populations occasionally exhibit tubercles extending from the second mesial row onto the dorsal surface of the palm, a condition most frequently observed in form I males. Palmar tubercles rarely were observed in eastern and central populations.

Relationships and comparisons. Cambarus theepiensis is placed in the subgenus Puncticambarus based on its elongate chelae with two distinct rows of tubercles on the mesial margin of the palm; its broad, densely punctate areola that is 2.1 to 6.2 times as long as broad; the presence of a subapical notch on the form I male gonopod; and a distinct lateral impression at the base of the propodus ( Hobbs 1969; Cooper 2001). Among described members of the subgenus, C. theepiensis is most similar to C. robustus . It can be distinguished from the latter species by its subparallel, broad rostrum, compared to the convergent, narrow rostrum of C. robustus . In addition, in C. theepiensis the rostral margins are thickened from their base to the base of the acumen, which lacks thickened margins. The rostral margins of Cambarus robustus can be thickened compared to the acumen margins, but usually are the same width as the margins of the acumen, and are never as thick as observed in C. theepiensis . The latter also has a greater areola width/ length ratio (26 %; n = 60; SE ± 2.9%) than C. robustus (22 %; n = 30; SE ± 1.9 %). The color pattern of Cambarus theepiensis is a mottled reddish/chestnut brown ( Figure 2 ) along the lateral surfaces of the cephalothorax and abdomen. Cambarus robustus from across the northern portion of its range near the type locality, New England, and the Upper Ohio Valley lack this mottled pattern, and are a uniform tan to olive brown.

Cambarus buntingi Bouchard 1973 and Cambarus cumberlandensis Hobbs and Bouchard 1973 are puncticambarids that occur in Kentucky in eastern portions of the Cumberland Plateau in the Cumberland River system and are not syntopic with C. theepiensis . Cambarus theepiensis can be differentiated from C. buntingi and C. cumberlandensis easily through rostrum morphology. Cambarus buntingi has a lanceolate rostrum with small rostral tubercles at the acumen/rostrum junction. Cambarus cumberlandensis has pronounced tubercles and parallel rostral margins. Cambarus theepiensis differs from both taxa by having a broad, subparallel rostrum that lacks rostral tubercles.

The rostrum of Cambarus theepiensis is similar to that of C. sciotensis . The rostrum of the latter can be differentiated from that of C. theepiensis by the thickness and extent of the margins and the shape. The thickness of the rostral margin in Cambarus theepiensis is equal from its base to the base of the acumen. In Cambarus sciotensis , the rostrum lacks uniform margins, tapering and decreasing in thickness as it joins with the acumen. The margins of New River C. sciotensis rostra are parallel, while those of C. theepiensis are subparallel. There is considerable variation in the convergence of the rostrum in Scioto River C. sciotensis . In the majority of specimens rostral margins are parallel (ZJL personal observation). Rostrum width/ length ratios also differ between C. theepiensis (63.1 %; n = 60; SE ± 3.8 %) and C. sciotensis (57.2 %; n = 30; SE 4.1 %).

Differences between C. theepiensis and C. sciotensis also exist in the sculpturing of the chelae and the morphology of the gonopod. Cambarus theepiensis chelae have a pronounced lateral impression at the base of the propodus, well developed tubercles that extend along the dorsal surface of the dactyl, and two to four well developed subpalmar tubercles. Cambarus sciotensis from both the Scioto and New River basins lack a pronounced lateral impression at the base of the immovable finger and have moderate to reduced tubercles on the dorsal surface of the dactyl. Cambarus sciotensis from the Scioto River basin have one to two subpalmar tubercles, while animals from the New River system rarely possess subpalmar tubercles, and if present usually have a single tubercle. The central projection of C. theepiensis form I gonopod is equal to or slightly longer than the mesial process, while in C. sciotensis the tip of the mesial process extends past the tip of the central projection.

Cambarus theepiensis can be differentiated from sympatric C. b. cavatus by the shape of the rostrum and chelae morphology. Cambarus theepiensis has a long, subparallel rostrum compared to the short, converging rostrum of C. b. cavatus, and the rostral margins of Cambarus b. cavatus are not noticeably thickened like those of C. theepiensis . Chelae shape can also be used to differentiate between the two species, with C. b. cavatus having short, squamous chelae compared to the elongate chelae of C. theepiensis .

Etymology. Latinized form of the Shawnee word for river, theepi. The Shawnee were among the first settlers of the Big Sandy, Lower Ohio and Guyandotte watersheds.

Common name. Coalfields Crayfish.

Figures

FIGURE 1. Cambarus theepiensis n. sp.: A. lateral view of carapace; B. lateral (left) and mesial (right) view of left first gonopod of form I male; C. lateral (left) and mesial (right) view of form II male gonopod; D, antennal scale; E. dorsal view of carapace; F. dorsal view of distal podomere of right cheliped of form I male; G. caudal view of in situ form I male gonopods; H. annulus ventralis; I. ischial hook of form I male; J. epistome; A, B, D–G, J, I from holoype; H from allotype; C from morphotype.

FIGURE 2. Cambarus theepiensis n. sp., holotype (USNM 1197339) in life.

FIGURE 3. Known range of Cambarus theepiensis n. sp. in Kentucky and West Virginia. Virginia.