Cambarus (Puncticambarus) hazardi, Loughman, Zachary J., Henkanaththegedara, Sujan M., Fetzner Jr, James W. & Thoma, Roger F., 2017

Loughman, Zachary J., Henkanaththegedara, Sujan M., Fetzner Jr, James W. & Thoma, Roger F., 2017, A case of Appalachian endemism: Revision of the Cambarus robustus complex (Decapoda: Cambaridae) in the Kentucky and Licking River basins of Kentucky, USA, with the description of three new species, Zootaxa 4269 (4), pp. 460-494 : 477-482

Cambarus (Puncticambarus) hazardi , new species

Figures 6–8 , Table 5

Cambarus robustus Girard, 1852:90 [in part];— Taylor and Schuster, 2004:103, Figs. 74 A, B, 75. Cambarus (Puncticambarus) robustus .—Hobbs, 1969:101, Figs. 1 c, 13a, 17o [in part]; 1974:21, Fig. 87 [in part]; 1989:27, Fig. 104 [in part].

Diagnosis. Body and eyes pigmented. Rostrum broad, moderately to deeply excavated, and deflected anteriorly, margins not thickened, subparallel to slightly converging at acumen. Acumen distinctly triangular with prominent dorsally directed spiniform tubercle at terminus. Areola 3.4–7.3 (x ‾ = 5.1, n = 27, SE = 1.06) times as long as wide with 5–9 (usually 6) punctations across narrowest point. Lacking cervical spines; occasionally 3–4 (x ‾ = 2.1, SE = 0.3) tubercles present (27% of individuals). Mandibular, branchiostegal, and orbital regions of carapace with welldeveloped tubercles. Postorbital ridges short; truncated, dorsally directed tubercle present in juveniles and subadults; adult postorbital ridge terminating in rounded tubercle. Suborbital angle acute. TCL 1.8–2.3 (x ‾ = 1.9, n = 27, SE = 0.10) 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 to three rows of tubercles; mesial most row with 6–11 (x ‾ = 9.1, n = 27, SE = 2.7) tubercles, second dorsal row with 8–11 (x ‾ = 7.2, n = 26, SE = 1.9) tubercles. Tubercles extend onto upper body of palm forming 1–2 additional disorganized rows of tubercles. Ventral surface of chelae with 2–6 (x ‾ = 3.1, n = 26, SE = 1.6) subpalmar tubercles scattered across chelae ventral surface. Dorsal longitudinal ridge of dactyl consisting of several well developed, highly pronounced scattered tubercles. Dorsomedian ridge of fixed finger of propodus pronounced. Well defined lateral impression at the junction of the fixed finger with the propodus. Dactyl and fixed finger with sharp corneous terminal tip. Form I male palm length 61.8–76.6% (x ‾ = 67.4%, n = 4, SE = 6.6%) of palm width, Form I male palm length 28.3–30.9% (x ‾ = 29.4%, n = 4, SE = 1.1%) of total propodus length; female dactyl length 57.8–85.9% (x ‾ = 63.8%, n = 16, SE = 6.5%) of total propodus length. First pleopod of Form I male with short terminal elements. Central projection not tapering distally; recurved>90° to main shaft of gonopod, with weak 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. ( Fig. 6 A–D, H, G, J, K; Table 5).––Body compressed dorsoventrally ( Fig. 6 A); carapace posterior to cervical groove wider than abdomen. Carapace depth 75.3% carapace width at caudodorsal margin of cervical groove. TCL 44.0 mm; PCL 38.0 mm. Areola 5.9 times longer than wide, with 7 punctations across narrowest part ( Fig. 6 H); length of areola 37.7% of TCL (43.7% of PCL). Rostrum excavated along entire length; margins not thickened, subparallel and continuous to base of acumen; floor of rostrum with numerous punctations. Rostrum 1.6 times longer than wide. Acumen broad and triangular, ending in dorsally directed corneous tip ( Fig. 6 H). Postorbital ridges well developed, terminating in rounded tubercles. Suborbital angle acute, with tubercle ( Fig. 6 A). Cervical spine absent. Mandibular, branchiostegal, and orbital regions of carapace ornamented with well-developed tubercles; greatest tubercle density in hepatic region. Lateral margin of terga angulate; lateral margin of second pleuron 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 11 immovable, small spines and 1 lateral, large, movable spine. Cephalomedian lobe of epistome subtriangular, zygoma moderately arched ( Fig. 6 G); cephalolateral margins thickened, forming sharp angle at junction with endostyle ( Fig. 6 G). Body of epistome not possessing prominent cephalomedian fovea. Antennal scale broadest anteriorly; lateral margin thickened, terminating in large corneous spine; setiferous. Right antennal scale 4.43 times as long as wide (6.2 mm × 1.4 mm) ( Fig. 6 D). Tip of right antenna reaching middle of telson when adpressed. Mesial surface of right chela with 2 well-formed rows of tubercles and additional tubercles scattered across dorsal palm; mesial most row with 7 tubercles, second dorsal row with 6 tubercles ( Fig. 6 K). Palm length 60.7% of palm width; depth 92.5% width. Ventral surface of palm with 5 subpalmar tubercles scattered across ventral surface. Dorsal longitudinal ridge of dactyl well developed and possessing large tubercles ( Fig. 6 K); dactyl terminating in large corneous spine. Dorsomedian ridge of fixed finger of propodus well pronounced. Well defined lateral impression at the junction of fixed finger with the propodus. Dactyl and fixed finger of propodus with sharp, corneous tip. All measurements and counts from right chela. Carpus with prominent dorsal furrow ( Fig. 6 K) and 5 weak dorsomesial tubercles; mesial margin with large, procurved spine at about mid-length, and reduced proximal spine. Distodorsal surface of merus with 8 spiniform tubercles; ventrolateral ridge with 3 small spines and large, corneous distal spine; ventromesial ridge with 3 well-developed spines. Carapace depth less than width. Hook on ischium of third pereopods and Form I gonopod as described in diagnosis ( Fig. 6 B–D); tip reaching anterior margin of fourth caudomesial boss.

Description of Allotypic Female. ( Fig. 6 I, Table 5).––Differing from holotype in following respects: carapace height (22.0 mm) 82.1% of carapace width (26.8 mm); TCL 49.2 mm, PCL 41.1 mm. Areola length 37.4% of TCL (44.7% of PCL), 4.7 times as long as wide. Posterior portion of rostrum more excavated than anterior portion; rostrum 1.4 times longer than wide. Abdomen length 47.8 mm. Mesial surface of chelae mesial most row of tubercles 8, second dorsal row 6. Palm length 64.5% of palm width; depth 58.6% of width. Right antennal scale 2.21 times as long as wide (6.4 mm × 2.9 mm). Annulus ventralis as described in diagnosis ( Fig. 6 I); width of postannular sclerite half total width; first pleopods uniramous, reaching central region of annulus ventralis when abdomen flexed.

Description of Morphotypic Male, Form II. ( Fig. 6 E–F, Table 5).––Differing from holotype in the following respects: carapace height (17.4 mm) 72.2% width (24.1 mm); areola length 36.5% of TCL (43.7% of PCL), 4.4 times longer than wide. Rostrum 1.8 times as long as wide. Mesial most row of tubercles on palm of chela 7; second dorsal row 7. Palm length (10.7 mm) 60.8% of palm width (17.6 mm). Antennal scale length (7.5 mm) 2.1 times width (3.6 mm). Central projection rounded ( Fig. 6 E–F), mesial process tapered, bulbous, directed caudolaterally. Hook on ischium of third pereopod small, not reaching basioischial joint.

Size. Form I male (n = 4) mean TCL 36.5 mm, ranging from 26.8–45.0 mm (PCL 22.4–37.6 mm). Form II male (n = 6) mean TCL 42.6 mm, ranging from 40.2–49.2 mm (PCL 32.9–40.3 mm). Female (n = 17) TCL mean 43.9 mm, ranging from 32.5–51.9 mm (PCL 26.3–42.9 mm). The largest specimen examined was a Form I male with TCL of 51.9 mm (PCL 42.9 mm).

Color in life. Cambarus hazardi ( Fig. 7 ) coloration is similar to C. guenteri but differs in the following respects. Rostrum margins and acumen cream to red-brown or orange-brown, rarely red. Lateral surface of cephalothorax heavily reticulated with olivaceous-brown to dark brown mottling. Dorsal surface of chelae olivaceous-brown, green-brown, brown to tan with yellow, red-brown highlights; mesial, second dorsal row, and dorsal surface of dactyl tubercles yellow, tan or red-brown. Dorsal surface of abdomen heavily reticulated with dark brown mottling, and two pronounced chestnut or buff-brown dorsal abdominal stripes.

Type locality. Red River at KY 77 crossing in Red River Gorge National Geologic Area at the Powell / Menifee county line, KY, 37.83376/-83.65990 ( Fig. 8 , star). At this site, the Red River is 20–25 m wide, and consists of a series of riffles and runs coursing over a sand and gravel substrate. Boulders are present at the heads of both riffles and runs. Water depth ranged from 0.5–1.5 m deep. Here the type series was collected along with two additional specimens (Form I males) on 26 October 2014 by ZJL and C. G. Vopal. Orconectes cristavarius was also collected at this location; neither species was common and concerted effort was needed to collect the type series.

Disposition of types. The holotype, allotype, and morphotype are deposited in the North Carolina Museum of Science ( NCSM), Raleigh, N.C. (catalogue numbers NCSM 27212, 27213 and 27214, respectively) . Paratypes are deposited in the USNM, Washington D.C. ( USNM 1422180 ) .

Range and specimens examined. Of the three putative taxa described herein, C. hazardi appears to have the broadest distribution. Cambarus hazardi occurs throughout the North Fork of the Kentucky River basin in Breathitt, Knott, Letcher, Perry and Wolfe counties, in the Red River watershed in Estill, Powell, Menifee and Wolfe counties, and in the mid-upper portions of the Licking River basin in Bath, Fleming, Magoffin, Montgomery, and Morgan counties. Cambarus hazardi appears to occur within the Cumberland Plateau ( Fig. 8 ).

All of the following collections, with the exception of the previously discussed type series, are either housed in the Branley A. Branson Museum at Eastern Kentucky University (denoted with EKU), the United States National Museum (denoted with USNM), or the West Liberty University Astacology Collection (denoted with the prefix WLU). The following abbreviations occur in the text: Cr. = Creek; R. = River; Frk. = Fork ; UNT = Un-named tributary; ZJL14—Collectors included ZJL 2014 field crew which consisted of S. S. Bell, Z. W. Dillard, N. M. Sadecky, L. K. Sadecky, E. Tidmore and E. M. Tennant.

KENTUCKY: Breathitt Co. (1.) WLU 2083 , Wolf Cr., 37.3935/-83.3967 , 10 June 2014, 1 IIM, 1 JV, ZJL14. (2.) WLU 2086, Frozen Cr., 37.6417/-83.3199, 11 June 2014, 1 IIM, 1 JV, ZJL14. (3.) WLU 2157, S. Frk. Quicksand Cr., 37.5148/-83.2116, 11 June 2014, 2 F, 3 IIM, ZJL14. (4) EKU 1050, Buckhorn Cr., 37.4 544/- 83.1767, 4 April 1969, 3 F, D. L. Batch. (5.) USNM 147040, Buckhorn Cr., 37.5363/-83.2522, 20 October 1974, R. Bouchard. (6.) USNM 260427, Roadside Ditch, 37.4495/-83.18297, 20 October 1974, R. Bouchard. Knott Co. (7.) WLU 2118 , Troublesome Cr., 37.3349/-83.1016 , 10 June 2014, 1 Jv, ZJL14. (8.) WLU 2122, Carr Frk., 37.2764/- 82.8545, 9 June 2014, 1 IIM, 1 Jv, ZJL14. (9.) WLU 2156, Laurel Frk. Quicksand Cr., 37.4609/-83.0 434, 2 F, 1 IIM, 5 Jv, ZJL14. Letcher Co. (10.) WLU 2085 , N. Frk. Kentucky R., 37.1679/-82.7521 , 9 June 2014, 1 F, 1 IIM, ZJL14. (11.) WLU 2119, Rockhouse Cr, 37.1731/-82.9188, 9 June 2014, 1 F, 3 IIM, ZJL14. (12.) WLU 2120, Millstone Cr., 37.1872/-82.7444, 9 June 2014, 1 F, 5 IIM, 8 Jv, ZJL14. (13) WLU 2 123, Sandlick Cr., 37.1310/- 82.8305, 9 June 2014, 3 IIM, 7 Jv, ZJL14. Menifee Co. (14.) USNM 145711 , Red R., 37.8227/-83.6284 , 20 October 1974, 1 F, R. Bouchard. Morgan Co. (15.) WLU 2222 , Straight Cr., 37.9710/-83.1740 , 28 April 2012, 1 F, 1 IM, D. Foltz, C. Z. & Z. J. Loughman, K. T. Skalican. 16.) WLU 2295, Cow Cr., 37.4342/-83.5639, 25 October 2014, 10 F, 10 IIM, C. G. Vopal and Z. J. Loughman. (17.) WLU 2299, Halsey Branch, 37.8534/-83.3418, 27 April 2012, 1 F, D. Foltz, C. Z. & Z. J. Loughman, K. T. Skalican. Perry Co. (18.) WLU 2084 , Cambell Cr., 37.3047/- 83.3357 , 10 June 2014, 2 IIM, ZJL14. (19.) WLU 2087, Carr Frk., 37.2088/-83.1109, 9 June 2014, 1 Jv, ZJL14. (20.) WLU 2089, Lotts Cr., 37.2860/-83.1484, 10 June 2014, 1 F, 2 JV, ZJL14. (21.) WLU 2117, Big Cr., 37.2352/ -83.2592, 9 June 2014, 1 IIM, 1 Jv, ZJL14. (22.) WLU 2125, Bull Cr., 37.1379/-83.0705, 9 June 2014, 2 F, 1 Jv, ZJL14. (23.) EKU 435, Leatherwood Frk., 37.8896/-83.6804, 19 June 2000, 3 IM, B. Dickey. (24.) USNM 147037, Big Cr, 37.3693/-83.0689, 20 October 1974, 1 F, R. Bouchard. (25.) USNM 147038, L. Frk. Maces Cr., 37.1810/- 83.1495, 20 October 1974, 3 IM, R, Bouchard. (26.) USNM 310634, Troublesome Cr., 37.3373/-83.1278, 20 October 1974, 1 IIM, R. Bouchard. Powell Co. (27.) NCSM 27212, 27213 , 27214 ; USNM 1422180, Red R. TYPE LOCALITY, 37.8337/-83.65990, 26 October 2016, 2 IM, 1 IIM, 2 F, C. G. Vopal and Z. J. Loughman. (28.) USNM 177297, Red R., 37.8244/-83.6624, 20 October 1974, 1 F, R. Bouchard. Roane Co. (29.) EKU 664 , Flooded Field, 38.1213/-83.5454 , 21 May 1988, 2 F, 1 IM, D. R. Peak. (30.) USNM 1078924, E. Frk. Triple Cr., 38.3173/- 83.4429, 22 April 1961, 3 F.

Habitat and life history notes. Cambarus hazardi is a slab-rock, large boulder specialist as an adult, with all recently collected specimens taken from under boulders without exception (ZJL, personal observation). Juveniles and young of the year have been collected from leaf packs, coarse woody debris snags and smaller substrate items. Slow moving riffles and moderate to low velocity runs appear to be C. hazardi preferred stream velocities. Considerable effort was undertaken to sample all available habitats in 2014 by ZJL; C. hazardi was only collected from the aforementioned habitats. Like other members of the C. robustus complex in Kentucky, C. hazardi does occur in headwater situations in addition to large tributaries and mainstems. The presence of sand and silt in streams in the lower reaches of the Red River, Upper Kentucky River, and Licking River basins appears to limit the downstream distribution of C. hazardi .

Cambarus hazardi annual life history is currently unknown. Form I males have been collected sparingly in May and June, and dominated collections in October and November. Form II males were frequently encountered in June and July, and rarely collected in October. Females exhibited glair in June. Ovigerous females have never been collected. Young-of-the-year were encountered in leaf packs in pools in October. Animals collected in June exhibited exoskeletons that were vibrant and lacked excessive accumulation biofilms, indicating the possibility of a population wide spring molt.

Conservation status. The entire range of Cambarus hazardi falls within the anthracite coal fields of eastern Kentucky. Coal mining activities, especially within the Licking and North Fork Kentucky River basins are extensive, and have caused basin wide stream degradation via sedimentation and chemical inputs associated with mining practices. Efforts in the fall of 2016 by ZJL in the Licking River basin to obtain additional specimens for this description proved fruitless. All streams sampled were heavily impacted by sedimentation. Dedicated surveys are needed to determine this species’ status within Kentucky.

Cambarus hazardi should be listed as vulnerable (V) using the American Fisheries Society criteria ( Taylor et al. 2007), and assigned a G3 ranking using the Master (1991) global conservation criteria for conservation listing due to its limited range and possible range wide decline. Cambarus hazardi should be listed as vulnerable (VU) using the International Union for the Conservation of Nature (IUCN 2001) criteria due to its narrow distribution and a reduction in range caused by anthropogenic stressors, specifically coal mining activities (ZJL, personal observation).

Crayfish associates. Cambarus hazardi has been collected with Cambarus (Jugicambarus) aff. dubius and Orconectes (Procericambarus) cristavarius .

Variation. Ontogenic variation exist in this species. Juveniles and sub adults lack the heavily sculptured chelae exhibited by adults. Specifically, juveniles lack a deep lateral impression and usually have more organized palmer tubercles compared to adults. Juveniles also tend to be olivaceous brown to green in overall coloration, transitioning to coloration dominated by browns as adults. Overall coloration also differs among populations. Animals from the headwaters of the North Fork, the Licking, and the Red River normally maintain pronounced dorsal stripes on the abdomen. Cambarus hazardi from the North Fork of the Kentucky River exhibit weakly pronounced abdominal stripes, and are heavily mottled in various shades of brown and green.

Relationships and comparisons. Cambarus hazardi is placed in the subgenus Puncticambarus based on its elongate chelae, its broad, densely punctate areola that is 2.1 to 6.2 times as long as broad, and the presence of a subapical notch on the Form I male gonopod (Hobbs 1969; Cooper 2001). Among described members of the subgenus, C. hazardi is most similar to C. guenteri and C. taylori .

Differences between C. taylori and C. guenteri are discussed in the previous relationships and comparison section for C. guenteri . Morphologically C. hazardi can be differentiated from C. taylori by its broad rostrum and by the absence of cervical spines; C. taylori has a lanceolate weakly excavated rostrum and has cervical spines. Cambarus hazardi chelae are subrectangular and possess up to three mesial margin tubercle rows with disorganized tubercles scattered on the upper palmar surface; C. taylori maintains two well organized mesial margin tubercle rows. Finally, C. hazardi has striping on the dorsal surface of the abdomen which C. taylori lacks.

Cambarus hazardi is easily differentiated from C. buntingi , C. callainus and C. cumberlandensis . Cambarus buntingi has a narrow rostrum with rostral tubercles, which C. hazardi lacks. Cambarus callainus coloration is strikingly different than C. hazardi , has a single row of mesial margin tubercles of which C. hazardi has two to three rows, and C. callainus rostrum is lanceolate; C. hazardi has a broad, excavated rostrum. Cambarus cumberlandensis , like C. buntingi , has rostral tubercles which C. hazardi lacks, and a markedly different color pattern than C. hazardi .

Cambarus hazardi resembles central Appalachian members of the subgenus Depressicambarus, namely Cambarus striatus Hay, 1902 . Both species have strongly marked abdominal stripes and mottled brown color patterns. Additionally, C. hazardi chelae are more robust and tuberculate than other members of the C. robustus complex in Kentucky, a trait that also makes C. hazardi similar to C. striatus , in some parts of its range. Cambarus hazardi can be differentiated from C. striatus by its wide, open areola compared to C. striatus narrow areola. Form I gonopods in C. hazardi possess a subapical notch which is normally absent in C. striatus Form I gonopods.

Etymology. Cambarus hazardi is named after Commodore Oliver Hazard Perry, noted American Naval Commander, and the name sake of Perry County Kentucky and the County Seat, Hazard Kentucky. The common name Brawny Crayfish is in reference to the powerfully built stature of C. hazardi .

Common name. Brawny Crayfish

Figures

FIGURE 1. Non-metric Multi-Dimensional Scaling (NMDS) ordination plot showing morphometric variation among crayfish specimens (stress = 0.081; Non-metric fit R 2 = 0.993; linear fit R 2 = 0.974). Each number represents an individual specimen and the relative distance between two points represents the similarity (or dissimilarity).

FIGURE 6. Cambarus hazardi, new species; all from holotype male, Form I (NCSM 27212), except E and F from morphotype male, Form II (NCSM 27214), and I from allotype female (NCSM 27213): A. lateral aspect of carapace; B – D. mesial and lateral aspect of Form I male gonopod respectively; E – F. mesial and lateral aspect of Form II gonopod respectively; G. ventral aspect of epistome; H. dorsal aspect of dorsal carapace; I. ventral aspect of the annulus ventralis; J. dorsal aspect of the antennal scale; K. dorsal aspect of the distal podomere of the left cheliped. Plate by Guenter Schuster.

FIGURE 7. Cambarus hazardi, new species. Form I male. Photograph by Guenter Schuster.

FIGURE 8. Map showing the distribution of Cambarus hazardi, new species, in the Kentucky and Licking river basins. Green circles indicate sampling localities where the species was detected. Green circle with star indicates the type locality. Gray shaded polygons indicate the different river sub-basins discussed in the text. Colored lines indicate stream order (Strahler 1957). Map inset abbreviations: LKy = Lower Kentucky River, UKy = Upper Kentucky River, SF = South Fork Kentucky River, MF = Middle Fork Kentucky River, NF = North Fork Kentucky River, SL = South Fork Licking River, LR = Licking River. Map generated using Google MyMaps.