Zootaxa 3750 (3): 223–236
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Copyright © 2013 Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3750.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:9F2C8AFC-A7C2-46A2-818F-2015ED70E5C8
Cambarus (C.) hatfieldi, a new species of crayfish (Decapoda:Cambaridae) from
the Tug Fork River Basin of Kentucky, Virginia and West Virginia, USA
ZACHARY J. LOUGHMAN1, RAQUEL A. FAGUNDO1, EVAN LAU2,
STUART A. WELSH1 & ROGER F. THOMA3
1
West Liberty University, Department of Natural Sciences and Mathematics, P.O. Box 295, West Liberty, West Virginia, 26074.
E-mail: zloughman@westliberty.edu; rfagundo@westliberty.edu; evan.lau@westliberty.edu
2
U.S. Geological Survey, West Virginia Cooperative Fish and Wildlife Research Unit, 322 Percival Hall, Morgantown, West Virginia,
26506. E-mail: swelsh@mail.wvu.edu
3
Midwest Biodiversity Institute, P.O. Box 21561, Columbus, Ohio, 43221-0561. E-mail: cambarus1@mac.com
Abstract
Cambarus (Cambarus) hatfieldi is a stream-dwelling crayfish that appears to be endemic to the Tug Fork River system of
West Virginia, Virginia, and Kentucky. Within this region, it is prevalent in all major tributaries in the basin as well as the
Tug Fork River’s mainstem. The new species is morphologically most similar to Cambarus sciotensis and Cambarus
angularis. It can be differentiated from C. sciotensis by its squamous, subtrinagular chelae compared to the elongate triangular chelae of C. sciotensis; its shorter palm length/palm depth ratio (1.9) compared to C. sciotensis (2.3); and a smaller
areola length/total carapace length ratio (30.4% vs.36.5% respectively). Cambarus hatfieldi can be differentiated from C.
angularis by its smaller areola length/total carapace length ratio (30.4% vs. 36.7% respectively); a smaller rostrum width/
rostral length ratio (59.4% vs. 67.2% respectively); its rounded abdominal pleura as compared to the subtruncated pleura
of C. angularis; the length of the central projection and mesial process of C. hatfieldi which both extend to the margin of
the gonopod shaft or slightly beyond the margin compared to the central projection of C. sciotensis and C. angularis where
both extend well beyond the margin of the gonopod shaft.
Key words: Crayfish, Cambarus, Kentucky, Virginia, West Virginia, Tug Fork River, Appalachian Mountains
Introduction
Cambarus sciotensis Rhoades, 1944, has one of the most disjunct ranges of any Cambarus species, with three
geographically isolated populations (Jezerinac et al. 1995; Taylor and Shuster 2004) (Jezerinac et al. 1995; Taylor
and Shuster 2004). The type population occurs in the Scioto River basin in Dublin Ohio, and shares morphological
characters with populations in central and southern Ohio. Cambarus sciotensis also occurs upstream of Kanawha
Falls in the New River system of West Virginia and Virginia; throughout the New River basin, C. sciotensis is the
dominant large Cambarus species. Cambarus sciotensis is replaced in the Ohio River mainstem between the Scioto
and eastern Kentucky populations and New River populations in the Big Sandy and Kanawha River system of West
Virginia by Cambarus robustus Girard, 1852. The third population occurs in the Tug Fork River system of
Kentucky, Virginia, and West Virginia.
The distribution of C. sciotensis in southwestern West Virginia and eastern Kentucky has long been
inadequately documented (Jezerianc et al. 1995; Z. J. Loughman personal obs.). Jezerinac et al. (1995)
documented C. sciotensis sporadically occurring throughout several watersheds in southwestern West Virginia
outside of the New River basin. Recently, ZJL and SAW initiated a statewide survey of crayfishes in West Virginia,
with special attention towards determining the ranges of C. robustus and C. sciotensis. Populations present outside
of the New River in West Virginia’s Guyandotte River basin of West Virginia, and the Big Sandy River basin of
West Virginia and Kentucky excluding the Tug Fork River system of West Virginia and Kentucky were found to be
an undescribed species since described as Cambarus theepiensis (Loughman et al. 2013).
Accepted by J. Goy: 21 Nov. 2013; published: 19 Dec. 2013
223
In 2009 independent crayfish surveys were performed in the Tug Fork River basin, a major tributary of the Big
Sandy River, in West Virginia by ZJL and SAW, and in Kentucky by RFT. Both efforts collected what initially was
considered C. sciotensis, and both noticed the distinct morphological differences from both Scioto and New
River C. sciotensis populations. What was referred to as C. sciotensis in the Tug Fork appeared to have closer
morphologic affinities with another species, Cambarus angularis Hobbs and Bouchard, 1994; which, prior to its
description, was included in the C. sciotensis complex (Hobbs and Bouchard 1994). In addition to differing from
both C. sciotensis and C. angularis, Tug Fork populations also were distinct from C. theepiensis, the Big Sandy
River system’s dominant tertiary burrowing Cambarus species. Subsequent meristic, morphologic and molecular
analyses supported the conclusion of an undescribed species of crayfish in the Tug Fork River system in West
Virginia, Virginia and Kentucky. This new species differs from both C. sciotensis and C. angularis, and is formally
described herein as Cambarus hatfieldi.
Material and methods
All measurements were taken with digital calipers to the nearest 0.1 mm; specimen morphometrics follow Cooper
2006. Total carapace length is abbreviated TCL; postorbital carapace length is abbreviated PCL. DNA was
extracted from crayfish pereiopods using DNeasy Blood & Tissue Kit (Qiagen, Inc.). PCR primer pairs for
amplifying the mitochondrial cytochrome oxidase I (COI) subunit I gene were previously published in Folmer et
al. (1994) modified with the addition of universal M13 forward and reverse primers on their 5’-ends: Forward
primer, GTAAAACGACGGCCAGGGTCAACAAATCATAAAGATATTG, Reverse primer, CAGGAAACAGCT
ATGACTAAACTTCAGGGTGACCAAAAAATCA (M13 primer regions underlined). PCR reaction mixtures
(from Fisher BioReagents exACTGene® PCR Kit, Fisher Scientific Inc.) contained 1X PCR buffer, 1.5 mM MgCl2,
250 µM final concentration of each dNTP, a 0.3 µM final concentration of each primer, 0.3 U of Taq polymerase,
and approximately 100 ng of template DNA in a final volume of 40 µl. PCR conditions were: 94°C for 3 min,
followed by 40 cycles for 50 secs, 45°C for 1 min 10 secs, and 72°C for 1 min, and a final 10 min extension at
72°C. PCR products were purified using QIAquick PCR Purification Kit (Qiagen Inc.), and sequenced through
SimpleSeqTM through Eurofins MWG Operon (http://www.operon.com).
The COI subunit I gene sequence of Cambarus (cf.) bartonii cavatus (GenBank Accession no. AY701190) was
used as an outgroup in all phylogenetic analyses. Partial COI gene sequences were aligned using ClustalW
(available on MEGA). Phylogenetic reconstruction was implemented using MEGA (Molecular Evolutionary
Genetics Analysis) version 5.1 (http://www.megasoftware.net). Statistical support for all trees was obtained from
1,000 bootstrap replicates (only bootstrap values >50% are reported). Pairwise base comparisons of partial COI
nucleotide sequences within and between taxa were determined using ClustalW2 (http://www.ebi.ac.uk/Tools/msa/
clustalw2/), with calculations for mean and standard error using Excel® ver. 12.3.2 (Microsoft® Corp.).
Phylogenetic analyses based on Maximum Likelihood (ML, using Tamura-Nei model, gamma distributed rates
among sites, nearest-neighbor-interchange for heuristic tree inference), Maximum Parsimony (MP, using subtreepruning-regrafting for tree inference) and Neighbor Joining (NJ, using transitions and transversions substitution
model based on the number of differences) were implemented on the alignment for partial COI nucleotide
sequences, which consisted of 599 nucleotides for 17 sequences.
Cambarus (Cambarus) hatfieldi n. sp.
Figures 1–5, Tables 1–4
Diagnosis. Body and eyes pigmented. Posterior dorsal region of rostrum concave and deflected anteriorly. Rostrum
margins thickened, parallel to base of acumen. Acumen distinctly triangular with prominent dorsally deflected
spiniform tubercle at terminus. Areola 2.7–4.5 ( x = 3.3, n = 52, SD = 2.5) times as long as wide with 7–9 (mode =
7) punctations across narrowest point. Cervical spines absent. Mandibular, branchiostegal, and orbital regions of
carapace with well-developed tubercles. Postorbital ridges short; spiniform, dorsally deflected tubercle present in
juveniles and subadults; adult postorbital ridges terminating in either spiniform or truncated tubercles. Suborbital
angle acutue. Antennal scale widest in middle, 0.9–2.6 ( x = 1.9, n = 52, SD = 0.8) times as long as wide. Total
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LOUGHMAN ET AL.
carapace length (TCL) 1.6–2.2 ( x = 1.9, n = 52, SD = 0.3) times longer than width. Form I and II males possessing
hooks 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
palm of chelae with single row of 5–9 ( x = 7.7, n = 52, SD = 0.8) weakly developed tubercles; deep punctations
found lateral to tubercles. Dorsal longitudinal ridge of dactyl consisting of several moderately-developed scattered
tubercles. Dorsomedian ridge of fixed finger of propodus pronounced. Poorly defined lateral impression at the
junction of the fixed finger portion of the propodus. Dactyl and fixed finger with sharp corneous tip. Form I male
palm length 63.4–70.8 % ( x = 67.5%, n = 12, SD = 2.3%) of palm width, form I male palm length 22.4–33.2% (
x = 29.8%, n = 12, SD = 2.8%) of total propodus length; female dactyl length 52.6–62.3% ( x = 58.4, n = 21, SD =
2.8%) of total propodus length.
First pleopod of form I male with short terminal elements. Central projection gently tapering distally; recurved >
90º to main shaft of gonopod, with distinct subapical notch. Mesial process directed 90° to shaft, bent
cephalolaterally, inflated at its base, tapering to distinct caudal point at or slightly beyond terminance of central
projection. Neither process of first pleopod projecting significantly beyond caudal margin of gonopod shaft.
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 formed by two
asymmetrical hardened ridges.
Description of holotypic male, form I. (Figs. 1 A–C, F–I, K–L, 2; Table 1). Body compressed dorsoventrally
(Fig. 1A); carapace posterior to cervical groove wider than abdomen. Total carapace length 38.9 mm; postorbital
carapace length 33.0 mm. Areola 3.8 times longer than wide, 7 punctations across narrowest part (Fig 1G); length
of areola 36.2 % of TCL (42.7 % of postorbital carapace length (PCL). Rostrum weekly excavated; margins
thickened, continuous to base of acumen, parallel at midpoint and flared at base; floor of rostrum with numerous
punctations. Rostrum 1.8 times longer than wide. Acumen distinctly triangular, ending in dorsally oriented
corneous tip (Fig. 1A). Postorbital ridges developed, short, terminating in weak dorsally oriented cephalic
tubercles. Suborbital angle acute, lacking tubercle (Fig. 1A). Cervical spine/tubercles absent. Mandibular,
branchiostegal, and orbital regions of carapace punctated with well-developed tubercles; highest 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 pleura 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 10 immovable, small
spines and 1 lateral, large, movable spine.
Cephalomedian lobe of epistome subtriangular, zygoma moderately arched (Fig. 1K); cephalolateral margins
thickened, forming sharp angle at junction with endostyle (Fig. 1K). Body of epistome possessing prominent
cephalomedian fovea. Antennal scale broadest in middle (Fig. 1F); lateral margin thickened, terminating in large
corneous spine; setiferous. Right antennal scale 5.5 mm long, 2.5 mm wide (Fig. 1F). Tip of right antenna reaching
middle of telson when adpressed.
Mesial surface of right chela palm with single row of 9 tubercles (Fig. 1H), length 69.5 % of width; depth 8.6
mm, ventral surface with 0 subpalmar tubercles. Dorsal longitudinal ridge of dactyl developed, mesial margin
possessing moderate equal-sized tubercles (Fig. 1F). Dorsomedian ridge of fixed finger of propodus pronounced.
Poorly defined lateral impression at junction of fixed finger with the propodus. Dactyl and fixed finger of propodus
with sharp, corneous tip.
Carpus with prominent dorsal furrow (Fig. 1H) and 6 weak dorsomesial tubercles; rest of surface with some
setiferous punctations; mesial margin with large, procurved spine at midlength, subtended by reduced proximal
spine. Distodorsal surface of merus with 8 spiniform tubercles; ventrolateral ridge with 3 small spines and large,
corneous distal spine; ventrolateral margin of ischium with 2 small, spiniform tubercles. Carapace depth less than
width. Hook on ischium of third pereopods only (Fig. 1L.); hook gently curved at apex, overarching basioischial
joint, not opposed by tubercle on basis. Form I gonopod as described in diagnosis (Fig. 1C–D); tip reaching
anterior margin of fourth caudomesial boss.
Description of allotypic female. (Fig. 1J, Table 1). Differing from holotype in following respects; carapace
depth less than carapace width (15.8 and 20.2 mm, respectively); TCL 37.7 mm, PCL 32.0 mm. Areola 37.1% of
CAMBARUS HATFIELDI DESCRIPTION
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TCL (43.8% of PCL), 3.7 times as long as wide. Rostrum 1.3 times longer than wide. Abdomen length 39.9 mm;
abdomen width 45% of abdomen length. Mesial surface of palm of chelae with single row of 8 weak tubercles.
Palm length (8.6 mm) 66.1% of palm width (13.0 mm); depth of palm 8.0 mm. Antennal scale 5.9 mm long, 2.7
mm wide. All measurements and counts from right chela. Annulus ventralis as described in diagnosis (Fig. 1J);
width of postannular sclerite 33.0% total width of annulus ventralis; first pleopods uniramous, reaching central
region of annulus ventralis when abdomen flexed.
TABLE 1. Measurements (mm) of Cambarus hatfieldi, new species.
Holotype
Allotype
Morphotype
Total carapace length
38.9
37.7
26.0
Postorbital length
33.0
32.0
21.8
Length cephalic section
24.7
18.0
16.8
Width
21.6
20.2
13.6
Carapace
Depth
14.9
15.8
10.6
Length rostrum
8.6
8.2
5.0
Length acumen
2.6
2.8
1.6
Length areola
14.2
14.0
9.2
Width areola
3.7
3.8
1.8
2.5
2.7
1.1
16.6
18.2
10.6
Length mesial margin palm
10.7
9.4
6.2
Width palm
15.5
9.3
8.9
Depth palm
8.6
8.0
5.4
Length dactyl
20.4
18.1
10.3
Length carpus
11.0
10.4
7.9
Width carpus
7.3
9.2
6.0
Length dorsal margin merus
13.0
11.3
10.8
Depth merus
7.7
7.4
5.9
Gonopod length
7.1
N.A.
5.1
Antennal scale
Width
Abdomen
Width
Cheliped (Right)
Description of morphotypic male, form II. (Fig. 1D–E, Table 1). Differing from holotype in the following
respects: TCL 26.0 mm and PCL 21.8 mm. Areola length 35.3% of TCL (42.2% of PCL), 5.1 times longer than
wide. Rostrum margins subparallel to base of acumen; rostrum 1.9 times as long as wide. Abdomen 14.0 mm long.
Mesial row of tubercles on palm of chela with 8 tubercles. Palm length (6.2 mm) 69.7% of palm width (8.9 mm).
All measurements and counts from right chela. Antennal scale 2.2 mm long, 0.9 mm wide. Gonopods 23.4% of
TCL length. Central projection with complete apex rounded (Fig. 1D–E). Mesial process bluntly tapered, bulbous
at base. Hook on ischium of third pereopod small, not reaching basioischial joint.
Size. Form I male (n = 12) TCL size range 19.0–40.1 mm (PCL 23.5–34.7 mm), mean TCL of 33.5 mm. Form
II male (n = 19) mean TCL 32.3 mm, size range 18.4–36.6 mm (PCL 18.5–31.7 mm). Non-ovigerous female (n =
21) mean TCL 34.3 mm, size range 26.6–42.8 mm (PCL 21.9–36.5 mm). Ovigerous female (n = 4) TCL size range
26.1–39.5 mm, mean TCL of 32.8 mm. The largest specimen examined was a form I male with a TCL of 40.9 mm
(PCL 34.7 mm).
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LOUGHMAN ET AL.
FIGURE 1. Cambarus hatfieldi n. sp.: A. lateral view of carapace; B. lateral and C. mesial view of first gonopod of form I
male; D. lateral and E. mesial view of form II male gonopod; F. antennal scale; G. dorsal view of carapace; H. dorsal view of
distal podomere of right cheliped of form I male; I. caudal view of in situ form I male gonopods; J. annulus ventralis; K.
epistome; L. ischial hook; A–C, F–I, K–L from holotype; J from allotype; D–E from morphotype.
CAMBARUS HATFIELDI DESCRIPTION
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FIGURE 2. Cambarus hatfieldi n. sp., holotype (USNM 1227861) in life.
Color. Carapace ground color orange-brown to pink-brown; posterior and anterior margins of carapace dark.
Hepatic and antennal region of carapace punctuated with yellow, white, or cream tubercles. Postorbital ridge
reddish to orange-brown. Rostrum margins and acumen distinctly orange to red-orange. Cephalic section of
carapace immediately anterior to and including cervical groove black; mandibular abductor scars ranging from
black to brown. Lateral margin of antennal scale gray-blue to gray; body of antennal scale gray to blue-gray.
Antennal flagellum and antennules blue-gray, with olivaceous hue; dorsal surface of lamellae green-brown to bluebrown; ventral surface tan to olivaceous. Dorsal surface of chelae green, olive green to brown-orange with green
highlights, olivaceous to orange mottling; mesial surface of dactyl tubercles amber, orange or red-brown. Denticles
on opposable surfaces of fingers yellow, white, or tan. Ventral surface of chelae orange or orange-brown. Dorsal
surface of carpus brown, olivaceous or green-brown; occasionally orange; region adjacent to and including furrow
olivaceous orange to green; carpus spine orange. Merus orange-brown, green-brown, or olivaceous brown.
Podomeres of pereopods light blue, blue-green, or blue-gray; joints of pereopod podomeres white. Dorsal and
dorsolateral surface of abdomen olivaceous, green-brown or brown; tergal margins brown, reddish brown or
crimson red. Uropods green-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. Mate Creek at CR 6 crossing in Red Jacket, Mingo County, West Virginia (37.64807º N, 82.13524º W). The holotype, allotype, and morphotype were all collected mid channel in a riffle underneath large
slab boulders in 0.75 m of water. Mate Creek ranged between 9.0–15.0 m wide, and 0.3–1.0 m deep when the typeseries was collected. Stream substrate was composed primarily of gravel, cobbles, boulders and slabs.
Anthropogenic impacts included siltation, channelization, and bank erosion. Coal fines were also prevalent in
situations with sands and gravels. Cambarus hatfieldi also was associated with coarse woody debris snags and leaf
packs. The holotype, morphotype, and allotype were collected on 20 Mar 2013 by K. R. Loughman, C. Z.
Loughman, and ZJL.
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LOUGHMAN ET AL.
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 122861 122862,
122863, respectively). Paratypes are deposited in the following museums and collections: Carnegie Museum of
Natural History, Pittsburgh, PA (CMNH 3829.1, 3829.2), and West Liberty University Astacology Collection, West
Liberty, WV (WLU 2000).
Range and specimens examined. Cambarus hatfieldi appears to be endemic to the Tug Fork River system
and its associated tributaries in Kentucky, Virginia and West Virginia. Kentucky’s population occurs in Pike and
Martin counties, with substantial populations occurring in Blackberry and Pond Creeks (Fig. 3). Virginia
populations are confined to the Dry Fork watershed in Tazewell Co. West Virginia’s populations are limited to
McDowell, Mingo and Wayne counties, with stable populations occurring in Dry Fork, Mate, Panther, and Horse
Creeks. All three states share the Tug Fork mainstem population. Cambarus hatfieldi appears to be replaced by
Cambarus theepiensis in lower reaches of the Tug Fork River where the river loses gradient and gains sandy
substrates.
FIGURE 3. Known range of Cambarus hatfieldi n. sp. in Kentucky, Virginia, and West Virginia. Tug Fork River basin is
highlighted in red; yellow circles represent C. hatfieldi collection locales.
All West Virginia collections are housed in the West Liberty University Astacology Collection, denoted with
the prefix WLU and were collected in the summer of 2009 unless otherwise noted. West Liberty University field
crew members for 2009 included ZJL, SAW, Nicole L. Garrison, David A. Foltz, Evan I. Hewitt, and Mathew I.
McKinney. All Kentucky and Virginia specimens examined were collected by RFT and are housed in the Ohio State
University Museum of Biological Diversity Crustacean Collection. Abbreviations are defined as follows: CR =
county road; I = interstate; KY = Kentucky state highway; mi = miles; RD = road; US = U. S. route; VA = Virginia
state highway; WV = West Virginia state highway; Rd = road; F = female; OF = ovigerous female; IM = Form I
male; IIM = form II male; JV = juvenile.
A total of 256 specimens were examined from the following 24 localities. KENTUCKY: Martin Co: (1.)
OSUM 7423, Knox Creek upstream of confluence with Tug Fork and intersection of Woodman Creek Road &
unnamed road, 1.2 mi. NW of Woodman, 18 Sep 2009, 1 F. Pike Co: (2.) OSUM 7423, Knox Creek upstream of
confluence with Tug Fork River and intersection of Woodman Creek Road & unnamed road 1.17 mi. NW of
Woodman, 17 Sep 2009, 1 F. (3.) OSUM 7428, Peter Creek at Freeburn adjacent to KY 194 1.2 mi. W of Vulcan,
CAMBARUS HATFIELDI DESCRIPTION
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17 Sep 2009, 1 JV. (4.) OSUM 7431, Blackberry Creek adjacent KY Rt. 1056 2.26 mi. S of Matewan, 17 Sep 2009,
1 F, 2 IIM. (5.) OSUM 7423, Pond Creek at Sherondale adjacent US. 119 3.9 mi N of Pinsonfork, 17 Sep 2009, 6 F,
2 IIM, 9 JV. (6.) OSUM 7441, Big Creek adjacent to KY 468 at confluence of Lick Branch 1.29 mi. SW of Nolan,
18 Sep 2009, 3 JV. VIRGINIA: Tazewll Co.: (7.) OSUM 8737, Dry Fork of Tug Fork at VA Rt. 636 bridge; 1.42
Km NNW of Rourkes Gap, 10.53 Km NW of Tazewell, 28 Jul 2011, 2 F, 3 IIM, 23 JV. WEST VIRGINIA:
McDowell Co: (8.) WLU 2, Crane Creek adjacent to CR 5/02, 2.0 mi SE of Avondale, 11 Jun 2009, 10 F, 5 IIM. (9.)
WLU 3, Elk Horn Creek 0.1mi NW of intersection of Coe Street and E Main Street in Kimball, 13 Jun 2009, 2 F, 2
II M. (10.) WLU 7, Horse Creek adjacent to CR 1–4, 0.5 mi from Johnny Cake Road 3.4 mi SE of Panther, 10 Jun
2009, 2 OF. WLU 26, 10 Jul 2009, 22 F, 24 IIM, 16 JV. (11.) WLU 9, Dry Fork adjacent to SR 80, 2.7 mi SE of
Iaeger, 1.0 mi SE of Apple Grove, 10 Jun 2009, 4 F, 2 IIM, 1 JV. (12.) WLU 10, Barrenshe Creek adjacent to CR
83–20/CR 83 intersection in Yukon, 10 Jun 2009, 1 OF, 2 F, 1 IM, 3 IIM, 2 JV. (13.) WLU 12, Tug Fork River
parallel to SR 103, 0.6 mi NW of Gary, 10 Jun 2009, 3 F, 1 IIM. (14.) WLU 13, Clear Fork parallel to CR 2, 1.1 mi
W of CR 2-1 in Coalwood, 10 Jun 2009, 2 F, 8 IIM, 15 JV. (15.) WLU 23, Tug Fork at Clear Fork confluence in
Roderfield, 13 Jun 2009, 9 F, 5 IIM. (16.) WLU 25, Spice Creek adjacent to US 52, 0.3 mi SE of Erin, 13 Jun 2009,
6 F, 10 IIM. (17.) Mingo Co: (17.) WLU 3, Left Fork of Gilbert Creek adjacent to CR 13 & CR 13-4 junction, 2.4
mi SE of Baisden, 11 Jun 2009, 1 F, 4 IIM. (18.) WLU 14, Pigeon Creek parallel to US 52 in Musick, 11 Jun 2009,
1 OF, 7 F, 4 IIM. (19.) WLU 16, Pigeon Creek at SR 65/ US 52 intersection 2.8 mi W of Varney, 11 Jun 2009, 1 F.
(20.) TYPE SERIES (USNM 12861 holotype, USNM 122862 allotype, USNM 12863 morphotype; CMNH
3829.1,3829.2 paratypes, WLU 2000 paratypes, Mate Creek at CR 6 crossing in Red Jacket, 20 Apr 2013, 12 F, 9
IM, 3 IIM, 11 JV. (21.) WLU 18, Mate Creek parallel to CR 6, 0.1 mi NW of Mark's Branch Rd in Newtown, 11
Jun 2009. 2 F. (22.) Sycamore Creek parallel to US 52, 0.2 mi N of SR 49 in Williamson, 11 Jun 2009, 4 F, 3 IIM.
(23.) WLU 20, Buffalo Creek adjacent to CR 14, at intersection with Mullberry Street in Chattaroy, 11 Jun 2009, 2
F, 1 IM, 5 IIM, 2 JV. Wayne Co: (24.) WLU 22, Mill Creek at intersection of CR 36-3/CR 34-3, 2.75 mi W of
Radnor, 11 Jun 2009, 2 F, 2 IIM.
Conservation status. It is recommended Cambarus hatfieldi 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 as a consequence of its limited range. Cambarus
hatfieldi should be listed as near threatened (NT) using the International Union for the Conservation of Nature
(IUCN 2001) criteria due to its narrow distribution. All of the range of C. hatfieldi occurs in portions of West
Virginia, Virginia, and Kentucky that currently undergo extensive amounts of surface mining. The majority of
streams harboring C. hatfieldi experience elevated siltation rates and high conductivity due to mine drainage issues
(Pond et al. 2008). At present, the impact of environmental change on C. hatfieldi populations remains unknown.
Habitat and life history notes. Cambarus hatfieldi occurs in small to large, moderate to high gradient streams
with substrates composed of cobbles, boulders and slabs. Slab boulders are the preferred refuge of C. hatfieldi,
though when absent course woody debris snags, leaf packs, and rootwads are readily used as shelter. As stream
gradient decreases and sandy substrates pervade, C. hatfieldi numbers decline (ZJL, personal observation). In
situations with these habitat parameters in lower reaches of the Tug Fork River, C. hatfieldi is replaced by C.
theepiensis.
Cambarus hatfieldi male reproductive form state is seasonal. Nine percent of males collected in June and July
were first form, compared to 89% collected in March and April. These results suggest the majority of males molt
from form II to form I in late summer/early fall, with an additional molting event occurring in May/Jun back to
form II. Females collected from Mate Creek, Mingo Co. West Virginia in March and April 2013 exhibited active
glair glands. Three ovigerous females were collected from Horse Creek, McDowell Co. West Virginia and a single
female with stage 1 instars and unhatched eggs was collected on 11 Jul 2009 from Barenske Creek, McDowell Co.
West Virginia (Table 4). Hatched eggs were still attached to the Barenske Creek females pleopods, indicating
hatching had occurred very close to the time of capture (ZJL, personal observation). All ovigerous females were
taken from embedded boulders in slack water environments with sandy substrates (ZJL, personal observation).
Young of the year were collected in September 2011 from the same environs ovigerous females were collected
from in Horse Creek during the preceding years.
Crayfish associates. Cambarus (Cambarus.) hatfieldi has been collected with Cambarus (C.) bartonii cavatus
Hay, 1902, Cambarus (Jugicambarus.) dubius Faxon, 1884, Cambarus (Puncticambarus.) veteranus Faxon, 1914,
and Orconectes (Procericambarus) cristavarius Taylor, 2000.
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LOUGHMAN ET AL.
Variation. Morphological variation was homeoplasic across Cambarus hatfieldi’s range, with little variation
observed between similar age cohorts occurring in different streams. Ontogenic morphologic variation does occur
in the following respects. Juvenile rostrums and chelae are noticeably elongate compared to adults. With age, both
structures become both broader and deeper in form. Mesial margin tubercles on chelae are reduced in juveniles and
become steadily more pronounced as adulthood is reached. Tubercles on the post-orbital ridge and acumen are
always spinose in juveniles, compared to adults where they can either be spinose or truncated. Coloration also
differs between juveniles and adults, with juvenile coloration dominated by various shades of brown. With
maturity, blues, greens, and grays begin to dominate the walking legs, cephalothorax, and abdomen as described in
the color section of the manuscript.
Relationships and comparisons. Cambarus hatfieldi is placed in the subgenus Cambarus based on the
presence of a subapical notch in the form I gonopod and the lack of a well-developed mesial second tubercle row
on the palm (Hobbs 1969). Among described members of the subgenus, C. hatfieldi is most similar to C. sciotensis
and C. angularis in overall body size and shape and thickening of the rostral margins.
Meristic percentages/ratios that distinguish C. hatfieldi from C. sciotensis include areola length/carapace
length, propodus length/areola length, two chelae ratios and size of the central projection and mesial process in
comparison to gonopod shaft in form I-males. Areola length on average represents 30.4 % (n = 52; SE ± 3.9%) of
the carapace length compared to 36.5% (n = 30; SE ± 0.1%) in C. sciotensis. Cambarus hatfieldi areola length is
2.3 (n = 52; SE ± 0.2) times the propodus length compared to 2.8 (n = 30; SE ± 0.2) times in C. sciotensis.
Cambarus hatfieldi’s palm length/dactyl length ratio is smaller ( x = 1.9; n = 52; SE ± 0.3) than C. sciotensis’s ( x =
2.3; n = 48; SE = 2.1–2.6); palm depth to palm length ratio is also smaller in C. hatfieldi ( x = 1.4; n = 52; SE ± 0.1)
compared to C. sciotensis ( x = 1.8; n = 30; SE ± 0.3).
The length of C. hatfieldi central projection and mesial process both extend to the margin of the shaft of the
gonopod and not beyond compared to C. sciotensis with the central projection extending beyond the margin of the
gonopod shaft (Fig. 4). Cambarus hatfieldi can be distinguished from Scioto River C. sciotensis easily by the
absence of a second row of mesial tubercles on the chelae palm, both New and Scioto River C. sciotensis also have
noticeably elongate chelae compared to C. hatfieldi’s squamous, truncated chelae. Finally, C. hatfieldi’s average
total body length is smaller ( x = 67.6 mm; n = 52; SE ± 7.3 mm) as an adult compared to C. sciotensis ( x = 80.5
mm; n = 30; SE ± 6.5 mm).
FIGURE 4. Form I gonopod’s of (A.) Cambarus hatfieldi, (B.) Cambarus sciotensis, and (C.) Cambarus angularis.
CAMBARUS HATFIELDI DESCRIPTION
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231
Cambarus angularis and C. hatfieldi share in common short, subrectriangular chelae with swollen palms and a
single row of mesial tubercles on the chelae. Cambarus hatfieldi can be differentiated from C. angularis by the
rostrum width/length ratio, areola length/carapace length ratio, shape of the cephalothorax, and length of the central
projection and mesial process in comparison to gonopod shaft in Form I-males (Fig. 4). Cambarus hatfieldi
rostrum width constitutes a smaller percentage of the rostrum length ( x = 59.4%; n = 52; SE ± 5.2%) compared to
C. angularis ( x = 67.2%; n = 22; SE ± 8.3%), and a smaller areola length/carapace length ratio ( x = 30.4%; n = 52;
SE ± 2.1%) compared to C. angularis ( x = 36.7%; n = 22; SE ± 1.7%).
Cambarus hatfieldi’s abdominal pleura are rounded ventrally compared to C. angularis’s subtruncate pleura.
Dorsally C. angularis cephalon is anteriorly swollen compared to C. hatfieldi cephalon, which begins to taper
anteriorly at the junction of the cephalon with the cervical groove. The length of C. hatfieldi central projection and
mesial process both extend to the caudal margin of the shaft of the gonopod but not beyond when compared to C.
angularis where both extending beyond the caudal margin of the gonopod shaft (Fig. 4). Finally, like C. sciotensis,
C. angularis’ total body length on average is larger ( x = 84.1 mm; n = 22; 9.8 mm) than C. hatfieldi ( x = 67.6
mm; n = 52; SE ± 7.3 mm).
Trees resulting from ML, MP and NJ analyses were identical in their placement of taxa within clades. The
phylogenetic tree based on ML analysis is shown in Fig. 5. Cambarus sciotensis sequences form two groups, New
River Basin, WV (Glade Creek, Meadow River, Cherry River, Bluestone River, Gauley River, and Whitewater
Branch) and Scioto River, OH, clustered separately and formed a large, well-supported clade. Both groups (i.e.,
New River Basin and Scioto River) are phylogenetically related (Fig. 5). Cambarus hatfieldi sequences (from
Browns Creek and Mate Creek, WV) are more distantly related to C. sciotensis and are placed in a well-supported
cluster outside of the above mentioned clade consisting of C. sciotensis taxa.
FIGURE 5. Maximum Likelihood tree of partial COI sequences from analysis of 17 unique sequences (599 bp) retrieved in
this study. The species and locations of Cambarus sp. are listed in Table 3. The COI gene of C. bartonii was used as the
outgroup. Accession numbers of sequences are indicated in parentheses. Bootstrap values from 1,000 replicates are indicated at
the nodes of branches (if >50). The scale bar represents the number of nucleotide changes.
232 · Zootaxa 3750 (3) © 2013 Magnolia Press
LOUGHMAN ET AL.
Pairwise nucleotide sequence divergence rates within the (i) C. sciotensis clade consisting of sequences from
New River Basin, WV, (ii) C. sciotensis cluster from Sciotto River, OH, (iii) C. hatfieldi clade (from Browns
Creek, Mate Creek, WV), and (iv) C. angularis from Sycamore Creek, TN, were 0.002–0.008, 0.002, 0.002–0.005
and 0.022–0.037, respectively. In contrast, C. sciotensis from New River Basin, WV, and C. sciotensis from Scioto
River, OH, diverged on average, 0.026 (approx. 3%) and 0.029 (approx. 3%), respectively, from C. hatfieldi (from
Browns Creek, Mate Creek), WV (Table 2). It is our hypothesis that divergence results indicate that C. hatfieldi is
an incipient species, and likely diverged from Teays River C. sciotensis populations sometime in the Pleistocene
epoch. GenBank accession numbers, specimen locations are provided in Table 3.
TABLE 2. Proportion of uncorrected base substitutions (from sequence comparisons of 599 bp) between all taxa (as
average values) from the identified clusters (above diagonal) and associated standard errors (below diagonal). Analyses
were performed using ClustalW2. New River sites include Glade Creek, Meadow River, Cherry River, Bluestone River,
Gauley River, Whitewater Branch; Tug Fork sites include Browns Creek and Mate Creek. Both the Scioto River and
Sycamore Creek represent type locations for their representative species.
C. sciotensis New River
Basin, WV
C. sciotensis New River
Basin, WV
C. sciotensis Sciotto C. hatfieldii Tug Fork C.angularis
River, OH
River, WV
Sycamore Cr, TN.
0.019
C. sciotensis Sciotto River, 0.001
OH
C. hatfieldii Tug Fork
River, WV
0.000
0.000
C.angularis Sycamore Cr,
TN.
0.001
0.002
0.026
0.057
0.029
0.061
0.055
0.001
Three additional Cambarus species occur in the Tug Fork River system that can easily be differentiated from
C. hatfieldi. Cambarus theepiensis replaces C. hatfieldi in the Big Sandy River system, but is syntopic with C.
hatfieldi in lower reaches of the Tug Fork River (ZJL personal observation). Cambarus bartonii cavatus occurs
throughout the headwaters of the Tug Fork River system, and is the dominant species in ephemeral streams
throughout the watershed. Both C. theepiensis and C. b. cavatus possess two rows of tubercles on the mesial
surface of the chelae’s palm, and 1–2 subpalmer tubercles on the ventral surface of the chelae. Cambarus hatfieldi
possesses a single row of adpressed tubercles on the mesial surface of the palm and lacks subpalmer tubercles.
Cambarus veteranus is syntopic with C. hatfieldi in mid to headwaters reaches of Tug Fork, though noticeably
rarer than the latter (ZJL and RFT, personal observation). Cambarus veteranus rostrum is lanceolate compared to
C. hatfieldi’s broad rostrum. In addition to rostrum morphology, C. veteranus possess strong cervical spines; C.
hatfieldi lacks both cervical spines and cervical tubercles.
Distribution of C. hatfieldi, C. theepiensis, and C. sciotensis in Kentucky and West Virginia. In light of C.
hatfieldi’s description herein, and the recent description of C. theepiensis, the distribution of C. sciotensis in both
Kentucky and West Virginia comes into question. Cambarus sciotensis previously was recorded in the Big and
Little Sandy River drainages of Kentucky, and hadn’t formerly been documented in the Tug Fork drainage in the
state (Taylor and Schuster 2004). Currently, populations previously recognized as C. sciotensis in the Big and Little
Sandy River watersheds of Kentucky are synonymized under C. theepiensis (Loughman et al. 2013). Records
for C. sciotensis do exist for Kentucky in Tygart Creek, a direct tributary to the Ohio River in the vicinity of the
Scioto River confluence with the Ohio River. At present Tygart Creek and streams in its associated watershed
possess the only populations of C. sciotensis known to occur in Kentucky. Thoma (2010) was the first to
document C. hatfieldi in Kentucky, and reported the species as C. angularis.
In West Virginia, C. sciotensis is distributed in the greater New River system upstream of Kanawha Falls,
Kanawha County as well as the Kanawha River mainstem and tributaries immediately downstream of Kanawha
Falls. Cambarus robustus replaces C. sciotensis downstream of Kanawha Falls, and is the dominant tertiary
burrowing Cambarus throughout the Coal, Upper, and Lower Kanawha drainages in West Virginia. What
previously was documented as C. sciotensis in the Guyandotte and Twelvepole systems of West Virginia (Jezerinac
et al. 1993; Loughman et al. 2009) is synonymized with C. theepiensis (Loughman et al. 2013).
CAMBARUS HATFIELDI DESCRIPTION
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233
TABLE 3. Genbank accession numbers and locations for Cambarus sciotensis, Cambarus hatfieldi, and Cambarus
angularis sequences used in this study.
Specimen#
GenBank Accession # Species
Location
Coordinates
CS1
KF437297
C. sciotensis
Meadow River, WV
37.446123 N 81.120730 W
CS2
KF437298
C. sciotensis
Meadow River, WV
37.446123 N 81.120730 W
CS3
KF437299
C. sciotensis
Meadow River, WV
37.446123 N 81.120730 W
CS4
KF437300
C. sciotensis
Meadow River, WV
37.446123 N 81.120730 W
CS5
KF437301
C. sciotensis
Meadow River, WV
37.446123 N 81.120730 W
CS6
KF437302
C. sciotensis
Gauley River, WV
38.290968 N 80.640797 W
CS7
KF437303
C. sciotensis
Whitewater Branch, WV
38.274921 N 80.9324328 W
CS8
KF437304
C. sciotensis
Whitewater Branch, WV
38.274921 N 80.9324328 W
CS9
KF437305
C. sciotensis
Cherry River, WV
38.254229 N 80.510992 W
CS10
KF437306
C. sciotensis
Glade Creek, WV
37.704200 N81.052423 W
CS11
KF437307
C. sciotensis
Glade Creek, WV
37.704200 N81.052423 W
CS12
KF437308
C. sciotensis
Glade Creek, WV
37.704200 N81.052423 W
CS13
KF437309
C. sciotensis
Bluestone River, WV
37.446120 N81.120730 W
CS14
KF437310
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CS15
KF437311
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CS16
KF437312
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CS17
KF437313
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CS18
KF437314
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CS19
KF437315
C. sciotensis
Scioto River, OH
40.141670 N 83.11968 W
CA1
KF437316
C.angularis
Little Sycamore Creek, TN
36.442270 N 83.51006 W
CA2
KF437317
C.angularis
Little Sycamore Creek, TN
36.442270 N 83.51006 W
CA3
KF437318
C.angularis
Little Sycamore Creek, TN
36.442270 N 83.51006 W
CA4
KF437319
C.angularis
Little Sycamore Creek, TN
36.442270 N 83.51006 W
CH1
KF437320
C.hatfieldii
Browns Creek, WV
37.443050 N 81.56886 W
CH2
KF437321
C.hatfieldii
Browns Creek, WV
37.443050 N 81.56886 W
CH3
KF437322
C.hatfieldii
Mate Creek, WV
37.648070 N 82.13524 W
CH4
KF437323
C.hatfieldii
Mate Creek, WV
37.648070 N 82.13524 W
TABLE 4. Pleopodal egg diameters and instar condition for Cambarus hatfieldi, n. sp. ED denotes average egg diameter
in mm; asterisk denotes incomplete complement of eggs or instars.
Stream
Date
Female
TCL
Condition of eggs/instars
Horse Creek, McDowell Co. WV
10 July 2009
26.1
13 attached eggs*; ED = 1.9
Horse Creek, McDowell Co. WV
10 July 2009
33.3
33 attached eggs; ED = 1.6
Horse Creek, McDowell Co. WV
10 July 2009
39.5
36 attached eggs; ED = 1.8
Barenske Creek, McDowell Co. WV
11July 2009
32.1
64 attached instars;
3 attached eggs* ED = 1.5
Resultant of the description of C. hatfieldi and C. theepiensis, the distribution of C. sciotensis, as it is currently
is understood, is disjunct with populations in the Scioto River basin of Central Ohio as well as direct Ohio River
tributaries in Kentucky near the Ohio /Scioto River confluence and in the New River system of West Virginia and
Virginia. Both populations were likely connected via the ancient Teays River, which is evident given the low COI
divergence values exhibited between both populations (Table 2). That being said, Scioto and New river
populations of C. sciotensis differ morphologically and are readily identifiable from each other using
234 · Zootaxa 3750 (3) © 2013 Magnolia Press
LOUGHMAN ET AL.
morphological characters (Z. J Loughman and R. F. Thoma personal observation). Given morphologic and
zoogeographic differences, further taxonomic investigation is warranted between the Scioto and New River
populations of C. sciotensis.
Etymology. Latinized form of Hatfield in honor of the Hatfield and McCoy feud which occurred in the Tug
Fork River Valley of Kentucky and West Virginia in the 1860s–1870s.
Common name. Tug Valley Crayfish.
Acknowledgments
We are grateful to two anonymous reviewers whose suggestions improved the overall quality of the manuscript and
to David Foltz, Nicole Garrison, Evan Hewitt, Kyle McGill, Matthew McKinney, and Christopher Vopal who
assisted with collection of the West Virginia material. Special thanks are extended to E. Joseph Nolan IV whose
creation and management of the West Liberty University Astacology database expedited the writing of this
manuscript. Financial support was provided by the West Liberty University Faculty Development grant program,
federal funding through the State Wildlife Grant program administered by the Kentucky Department of Fish and
Wildlife Resources, and a Cooperative Research, Education, and Management Grant from the West Virginia
Division of Natural Resources, Wildlife Resources Section, Wildlife Diversity Program. Any use of trade, firm, or
product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.
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