Cambarus (Lacunicambarus) erythrodactylus, a new species of crayfish (Decapoda: Cambaridae) of the Cambarus diogenes complex from Alabama and Mississippi, U.S.A. Author(s): Thomas P. Simon and Charles C. Morris Source: Proceedings of the Biological Society of Washington, 127(4):572-584. 2015. Published By: Biological Society of Washington DOI: http://dx.doi.org/10.2988/0006-324X-127.4.572 URL: http://www.bioone.org/doi/full/10.2988/0006-324X-127.4.572
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 127(4):572–584. 2014.
Cambarus (Lacunicambarus) erythrodactylus, a new species of crayfish (Decapoda: Cambaridae) of the Cambarus diogenes complex from Alabama and Mississippi, U.S.A. Thomas P. Simon* and Charles C. Morris (TPS) Indiana Biological Survey, Aquatic Research Center, 6400 Fairfax Road, Bloomington, Indiana 47401, U.S.A., e-mail:
[email protected]; (CCM) Troy University, Department of Environmental and Biological Sciences, Troy, Alabama 36081, U.S.A.; Current Address (TPS & CCM): Indiana State University, Department of Biology, Terre Haute, Indiana 47809, U.S.A.
Abstract.—A new species of crayfish, Cambarus (Lacunicambarus) erythrodactylus, is separated from the Cambarus (L.) diogenes complex. The new species is morphologically similar to C. (L.) diogenes but differs from it in the presence of three or four spines along the cervical groove, greatest depth anterior mid-horizontal beneath the antennal scale, the presence of two rows of tubercles along the distal medial palm, with an additional two or three partial or full rows of tubercles on the palm, subpalm of the chela with one to four (mode ¼ three) tubercles, and a gonopod extending greater than 908 and bent cephalodistally 208 at the shoulder. The species occurs in the Pea, Choctawhatchee, Sepulga, and Conecuh River systems of southeastern Alabama, in the Black Warrior and Tombigbee River systems in western Alabama and eastern Mississippi, in the Alabama River in central Alabama, and Bear Creek drainage of the Tennessee River, and Yazoo River drainages and occurring throughout the central and eastern portions of Mississippi. Keywords: biogeography, chelae characters, gonopod, morphometrics, warpaint mudbug
The Cambarus diogenes complex has been recognized for over a century as a combination of multiple species (Faxon 1885, Marlow 1960, Hobbs 1969, 1974, 1989). The subgenus Lacunicambarus was originally diagnosed by Hobbs (1989), but Jezerinac (1993) amended and split the subgenus, forming two subgenera. The subgenera Lacunicambarus and Tubericambarus can be separated based on chelae characteristics, including the placement and arrangement of palmar tubercles (Jezerinac 1993). Three described species and at least two other forms that are awaiting description or revision are referred to the subgenus * Corresponding author.
Lacunicambarus (T. P. Simon pers. obs.), including Cambarus diogenes Girard, 1852 and related forms. As a result of the 1871 Chicago Fire, the type material was lost and limited further study, which has clouded the definition of this group. Faxon (1884) initially described C. ludovicianus as a subspecies or variation of C. diogenes from New Orleans, Louisiana (Hay 1899, Marlow 1960). Hobbs (1989) ultimately elevated it to full species status. Cambarus (L.) miltus Fitzpatrick, 1978 is the third described species. Cambarus miltus was reportedly limited to the area around Spanish Fort, east of Mobile, Alabama. However, this species was recently reported from southern Alabama and north central Florida (Taylor & Shuster 2010).
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We present a new Lacunicambarus that has previously been referred to as C. (L.) species ‘‘B’’ (Jezerinac 1993, Heath et al. 2010) and as an undescribed species of the Cambarus (Lacunicambarus) diogenes complex (Thoma et al. 2005). Systematics Cambarus (Lacunicambarus) erythrodactylus, new species Figs. 1, 3, 4, Table 1 Cambarus diogenes.—Girard, 1852:88 (in part).—Hay, 1896:489 (in part).—Williamson, 1899:48 (in part).—Ortmann, 1905:123 (in part).—Marlow, 1960:231 (in part).—Lukhaup, 2003:123 (in part). Cambarus diogenes diogenes.—Hay, 1899:959 (in part).—Williamson, 1907:762 (in part).—Marlow, 1960:233. Cambarus (Bartonius) diogenes.—Ortmann, 1906:402 (in part). Cambarus (Lacunicambarus) diogenes diogenes.—Hobbs, 1969:110 (in part), 1974:20 (in part).—Bouchard, 1972:56 (in part), 1974:595 (in part).—Jezerinac, 1985:7 (in part). Cambarus (Lacunicambarus) diogenes.— Bouchard, 1976:13–21 (in part).—Jezerinac, 1985:7 (in part).—Hobbs, 1989:24 (in part).—Buchanan, 1992:1–185 (in part).—Schuster & Taylor, 2004:9, 13, 44, 48 (in part).— Thoma et al., 2005:332.—Miller et al. 2014:15–24. Cambarus (Lacunicambarus) sp. B.—Jezerinac, 1986:178 (in part).—Heath et al., 2010:139, 143, 147, 150. Diagnosis.—The gonopod is diagnostic. Form I male with coxa of fourth pereiopod lacking large ventral setiferous pit on caudomesial boss; first pleopods contiguous at base, with convexity near midlength of cephalic surface; terminal elements consisting of 1) short, nontapering, distally truncate central projection bent cephalodistally 208 angle to
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shoulder; 2) conically shaped mesial process, both directed caudally at angle slightly greater than 908 to shaft; and 3) central projection apical notch present. Hooks on ischium of pereiopods absent. Female annulus ventralis elliptical, slightly longer than broad, and rather deeply embedded in V-shaped sternum. Body pigmented; eyes small (X¯ ¼ 5.3% CL), pigmented. Antennae not heavily fringed on mesial border. Carapace subcylindrical, slightly dorsoventrally flattened. Rostrum margins moderately thickened, straight, without marginal spines or tubercles, abruptly curved cephalically. Postorbital and cervical spines absent, three or four spines along cervical groove. Suborbital angle prominent and acute. Areola obliterated, comprising 38.2– 44.8% (X¯ ¼ 40.9%, n ¼ 59) of total length of carapace, without punctuations in narrowest part. Antennal scale width 37.7% of antennal length (range: 26.8– 44.6%), broadest anterior midlength. Chela palm mesial surface with two distinct rows of adpressed tubercles and additional two to three partial to full rows on surface of palm. Chela (Fig. 1A) moderately robust with dactyl length/palm length 30.4% on first form males; chela palm dorsomesial surface with two welldeveloped basiodactyl tubercle rows modal number six (range: 3–7) tubercles each, dorsomesial surface mostly punctuate laterally; propodus lateral margin subcostate, with punctuations but never bearing rows of spines; fingers gaping and with moderately well-defined dorsomedian longitudinal ridges; proximal opposable margin of dactyl distinctly concave; generally without tuft of setae present at mesial base of fixed fingers, dorsolateral base slightly impressed; modal number of subpalmar tubercles two (range: 0–3). Dactyl medial surface with distal 18.3% margined with small denticles, with five small tubercles (range: 4–6), a single large tubercle, and two (rarely three) small proximal tubercles. Propodus
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Fig. 1. Cambarus (Lacunicambarus) erythrodactylus; all from holotype male, form I (OSUMC 2556), except B, C, from morphotype male, form II (OSUMC 2556), and F from allotype female (OSUMC 2555). A, B, lateral aspect of gonopod (first pleopod); C, D, mesial aspect of gonopod; E, dorsal aspect of antennal scale; F, caudal aspect of annulus ventralis; G, dorsal aspect of carapace; H, epistome; I, dorsal aspect of distal podomeres of left cheliped.
medial surface with distal 23.5% marginated with small denticles and with single canine-shaped tooth-like denticle, five small (range: 4–6) tubercles, single large tubercle offset from large tubercle on dactyl, and two (rarely three) moderately-sized proximal tubercles. Dorsolateral impression at base of propodus moderate to strong. Carpus without spiniform tubercles on ventral surface along distal articular rim. Mesial ramus uropod me-
dial spine never reaching caudal margin of ramus. Holotype male, Form I.—Body subovate (Fig. 1G), laterally compressed. Abdomen narrower than cephalothorax (22.6 and 28.6 mm); carapace maximum width less than depth at caudodorsal margin of cervical groove (28.6 and 30.2 mm). Areola closed, without punctuations in narrowest part; length comprising 41.2% of total length of carapace. Rostrum straight, with
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Fig. 2. Distribution of Cambarus (Lacunicambarus) erythrodactylus. Solid dots ¼ sites from which C. (L.) erythrodactylus was collected; dark gray shading represents the hypothesized range. Abbreviations: AL, Alabama; FL, Florida; GA, Georgia; MS, Mississippi. Insert: contiguous 48 states of United States of America; darkened area indicates Alabama, Florida, Georgia, and Mississippi.
slightly thickened margins; acumen basally not distinctly delimited, anterior tip upturned and reaching base of ultimate podomere of antennular peduncle; rostrum upper surface concave without punctuations other than usual submarginal ones. Subrostral ridge weak but evident in dorsal aspect along basal two-thirds of rostrum. Postorbital ridge weak, grooved dorsolaterally, ending cephalically without spine or corneous tubercle. Acute suborbital angle very prominent; branchiostegal spine represented by small tubercle. Cervical groove with three spines on each side of areola above antennal scale. Hepatic and branchiostegal region with granules. Remainder of carapace punctuate dorsally and granular laterally. Abdomen subequal in
length to carapace, pleura short, subtruncate, rounded caudoventrally. Telson cephalic section with two spines on caudolateral corner. Proximal uropod podomeres with weak distal spine on mesial lobe; mesial uropod ramus with prominent median rib ending distally in strong distomedian spine, not overreaching margin of ramus; laterodistal ramus spine strong. Cephalomedian epistome lobe (Fig. 1H) short and subtriangular with uniform margins, ventral surface rather flat; main body with shallow fovea; epistomal zygoma weakly arched. Ventral surface of antennular peduncle proximal podomere with small acute spine at base of distal third. Antennal peduncle without spines;
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Fig. 3. Cambarus (Lacunicambarus) erythrodactylus, male form I. A, coloration representative of Mississippi form (photo U.S. Forest Service, S. B. Adams); B, typical Alabama form coloration (photo TPS).
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Fig. 4. Cambarus (Lacunicambarus) erythrodactylus, male form I. A, anterior view; B, coloration variation based on molt stage (photo TPS).
antennal scale (Fig. 1E) 2.89 times as long as broad, broadest slightly distal to midlength, mesial border forming strong arc; distal spine strong, reaching distal extremTable 1.—Measurements (mm) of Cambarus (Lacunicambarus) erythrodactylus. Character
Carapace Height Width Length Areola Length Rostrum Width at eyes Length Postorbital ridge Width Chelae–right Length of lateral margin of chelae Length of mesial margin of palm Width of palm Length of dactyl Thickness of palm Abdomen Length Width Gonopod Length Antennal scale Length Width
Holotype Allotype Morphotype
30.2 28.6 58.2
28.1 25.7 53.0
21.8 19.8 42.2
24.0
22.7
18.9
5.5 10.5
4.9 8.9
4.6 6.7
9.0
8.4
6.5
50.9
38.9
28.7
14.3 19.1 31.5 12.2
11.6 16.1 25.3 10.1
8.6 12.5 18.2 7.7
54.4 22.6
54.6 21.5
42.8 15.9
13.6
N/A
10.3
8.1 2.8
8.2 3.3
6.6 2.8
ity of antennular peduncle. Mesial half of ventral surface of ischium of third maxilliped studded with irregular rows of long, stiff setae; submarginal lateral rows on podomere consisting of much smaller flexible ones; distolateral angle not acute. Right chelae length 87.5% that of carapace; width 38.9% of length; palm length 28.1% of chela length; dactyl length 2.2 times palm length. Dorsomesial palm quarter surface with two or three rows of tubercles, mesial-most row composed of six tubercles, secondary row with six tubercles, tertiary row with three tubercles, and basiodactyl row with six tubercles; dorsolateral half punctuate, punctuations deep and large in vicinity of dorsolateral base of propodus; palm and propodus lateral surface subcostate; palm ventral surface punctuate, with small corneous tubercle on articular rim opposite base of dactyl; subpalmar tubercle modal one (range: 0–3) (Fig. 1D). Both chela fingers with well-defined submedian ridges dorsally and ventrally; distal 20.9% of propodus without denticles; single amber canine-like tearing tooth, followed by serrate opposable margin of propodus with row of four tubercles at distal edge, with single large, pale yellow tubercle near mid-length of propodus; with two to four moderately-
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sized basal tubercles along proximal third of finger. Dactyl opposable margin distal 22.1% without denticles; possessing five tubercles above mid-length; single large tubercle offset and inter-digitizing with propodus (large mid-length tubercle), dactyl with slightly incised margin with three moderately sized tubercles along proximal half; single row of seven adpressed tubercles extending distally from mid-denticles; dactyl mesial surface with row of six adpressed tubercles basally giving way to punctuations distally. Dorsomedian fingers with well-developed longitudinal ridges. Dactyl and propodus distal edges with amber, canine-like hooked tooth. Base of fixed finger with moderate dorsolateral impression. Cheliped carpus (Fig. 1I) with distinct furrow dorsally; dorsomesial surface with row of 6 tubercles; dorsolateral surface with sparse punctuations; mesial surface with six large spiniform tubercles and three additional small spiniform tubercles; ventral surface with single spiniform tubercle on distal articular rim. Merus dorsally with three premarginal tubercles, ventrolateral row of six tubercles, and ventromesial row of six; podomere otherwise smooth. Ventral ischium ridge without small tubercles. Third pereiopod ischium with simple hook extending proximally over basioischial articulation, not opposed by tubercles on basis. Fourth pereiopod coxa with vertically disposed caudomesial boss; fifth pereiopod lacking boss, ventral membrane bearing oblique row of small sclerites armed with stiff setae. First pleopods contiguous at base, reaching coxa of third pereiopod; central projections short, hooked but not tapering, possessing subapial notch, rounded apically, and not extending beyond mesial process; mesial process conical, tapering, and directed essentially caudolaterally. Both terminal elements bent caudally at angle slightly greater than 908, with entire structure bent cephalodistally from shoulder at angle of 208; caudal knob absent.
Allotype female.—Excluding secondary sexual characteristics, differing from holotype in following respects: areola length 42.8% of total length of carapace; epistome cephalomedian lobe with thickened margins; antennal scale 2.48 times as long as broad; right chela 73.4% of carapace length; dactyl opposable margin with row of eight tubercles, only first tubercle enlarged; dactyl mesial surface with five squamous tubercles; merus dorsally with three premarginal tubercles, ventrolateral row of six tubercles, and ventromesial row of six; and four cervical groove spines beneath antennal scale. Annulus ventralis (Fig. 1F) deeply embedded in V-shaped sternum, subcircular in outline, with narrow median longitudinal furrow in cephalic half, ending in central depression; tongue extending caudosinistrally across caudal side of depression, disappearing beneath thickened caudosinistral wall; reverse S-shaped sinus and tilted sinistrally at almost 458, ending under caudal wall slightly dextral to median line. Postannular sclerite oval. First pleopod reaching mid-length of annulus when abdomen flexed. Morphotype male, Form II.—Differing from holotype in following respects: areola length 44.8% of carapace length; antennal scale 2.36 times as long as broad; right chela 68.0% of carapace length; palm length 30.0% of chela length; dactyl distal margin 13.4% of length without denticles followed by five denticles distal to midlength. Propodus distal 20.8% of margin free of denticles; single canine-like amber tooth, and four serrate denticles distal to mid-length. Opposable margin of right fixed finger with single, enlarged tubercle at mid-length; dactyl opposable margin with single large and two moderately sized proximal enlarged tubercles; two subpalmar tubercles; merus dorsally with three premarginal tubercles, ventrolateral row of six tubercles and ventromesial row of six; first pleopod central projection length 24.4% of CL, non-corneous and blunt,
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directed cephalodistally at 20% angle caudally at shoulder. Type locality.—U.S.A.: Alabama: Fayette County: Section 18, Range 12W (3.2 air km S of Fayette; 8.9 air km ENE of Belk); a roadside ditch at the intersection of State Route (St. Rte.) 171, 0.5 mi S of St. Rte. 159 [33840 0 8.7 00 N, 87850 0 7.5 00 W]. The specimens were dug from burrows with chimneys in a ditch having permanently flowing water from a spring. The surrounding vegetation was grass (old field) and pine forest. The collection was made by Raymond Jezerinac, R. F. Thoma, G. W. Stocker, D. Chrisman, P. Donaker, C. Pickerel, Z. Thoma on 21 March 1989 and consisted of 5 ?I, 4 ?II, and 10 /. Disposition of the types.—The holotype, allotype, and morphotype are in the collection of the Ohio State University Crustacean Collection (OSUMC 2656, 4 ?I, 3 ?II, and 10 /; OSUMC 2655, 7 ?I, and 7 /). Paratypes are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM 2071511) and in the Indiana Biological Survey Aquatic Research Center Astacology Collection (INBS 101222, 1 ?I, 1 ?II, and 1 /). Range.—The new species occurs in the Pea, Choctawhatchee, Sepulga, and Conecuh River systems of southeastern Alabama and possibly northcentral Florida, in the Black Warrior and Tombigbee River systems in western Alabama and Mississippi, in the Alabama River in central Alabama, and the Bear Creek drainage of eastern Mississippi in the Tennessee River drainage (Fig. 2). The species has also been collected from central and southwestern Mississippi including the Yazoo River watershed. A small portion of the species’ range may include northwestern Georgia; however, this has not been confirmed. Material examined.—We measured 59 specimens from 28 localities in Alabama and Mississippi (see Appendix). Eighteen specimens were Form I males, 10 were
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Form II males, 16 were Form I females, and 15 were Form II females. This material is curated in the University of Alabama Decapod Collection, Indiana Biological Survey Astacology Collection, and The Ohio State University Crustacean Collection. An additional 143 specimens were examined to determine the species range and are deposited in these same museum collections. These additional specimens comprise juveniles, females, and form II males. Many of the collections were made from roadside ditches. Mississippi distribution records are based on material curated at Tulane University as uncatalogued material. Conservation status.—Cambarus (L.) erythrodactylus is widely distributed and frequently found in headwater streams, disturbed environments such as partially drained wetlands, ditched streams surrounded by anthropogenic activity, as well as undisturbed and more natural habitats, including wooded wetlands. It appears to be negatively affected by silviculture. It should be considered Currently Stable following Taylor et al. (2007) and Least Concern (LC) using IUCN criteria (IUCN 2001). Color notes.—Two color patterns exist and represent a west-to-east variation cline that is based on molt stage (Figs. 3, 4). The color pattern at the western periphery of the range is more variable. In western Mississippi (Issaquena County), the carapace tends to be more tan to brown with the abdomen being darker brown than the thoracic portion of the carapace (Figs. 3A, 4A). The color pattern of individuals from the central portion of the species range is rather uniform consisting of dark olive to brownish-burgundy carapace and abdomen (Figs. 3B, 4B). Both coloration patterns have the distal two-thirds of the tips of the propodus and dactyl being cherry red. The lateral margin of the chela and dorsal knob on the chela (at the base of the dactyl) is cherry to orangish-red. The lateral edge of the palm can become
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Table 2.—Seasonal data of Cambarus (Lacunicambarus) erythrodactylus. # First form males
January February March April May June July August November December
4 16 1 3
4 1
# Second form males
4 1 10 6 2 8 6 2
# Females1
Sex ratio (M:F)
Ovigerous females
4 2 24 2 5 2 8 7 2 3
(1:0.5) (1:2) (1:0.92) (1:2.0) (1:0.6) (1:1) (1:1) (1:0.7) (1:2.0) (1:1.3)
0 0 16 1 0 0 0 0 0 0
1
Numbers include ovigerous females and Form II females.
lighter orange to tan-orange, whereas the rostral margins are usually brown-burgundy. The ventrum is brown to cream. Red bands outline every articulation joint and the caudal margin of the abdominal tergites. Red bands occur along the margins and mid-laterally along the uropods and telson. Variations.—The material was examined for clinal variation and characters that might be restricted to local populations. All characters were normally distributed with data distributions found to be within a single standard deviation from the mean. As expected in a species that occupies a narrow geographic range, little variation occurs in most body structures and body proportions. Sexual dimorphism is exhibited in two characters. Form I male chelae (X¯ ¼ 79.7% chelae/CL) are significantly longer (ANOVA, F ¼ 14.13, P , 0.001) than form II male, female, and ovigerous female individuals. Likewise, ovigerous female abdominal widths were significantly wider (X¯ ¼ 46.7% abdomen width/CL) than both form I and II male and female individuals. Some color variation is present in specimens collected in Mississippi, which tend to possess brown-olive carapaces, rather than the typical burgundycolored Alabama specimens. There appears to be more variation in color than
in body morphometric or meristic structures. Additional meristic and morphometric data (simple descriptive statistics, ratios, and regression analysis) are available from the senior author (TPS) or from the library at the Indiana Biological Survey. Size.—The largest specimen examined was a male with a carapace length of 58.2 mm (47.7 mm Posterior Orbit Ridge [POR] carapace length) from Washington County, Alabama. The largest Form I male measured 58.2 mm and the smallest 33.7 mm (POR carapace length 28.6 mm). The largest Form I female measured 49.2 mm (POR carapace length 40.8 mm) and the smallest 34.3 (POR carapace length 29.5 mm). For measurements see Table 1. Life-history notes.—First and second form males are most likely present at all times of the year (Table 2). Ovigerous females are only observed in the spring months. Ovigerous females were collected in mid-March to late April. Sex ratio is 1:1.15 male to female, whereas Miller et al. (2014) reported 1:1.3. Egg size was not correlated with female length. Females 39.5 mm CL had 6–31 ova attached to their abdomen. Miller et al. (2014) reported females matured at a larger mean length 41.2 mm CL and two females (length not reported) taken with 154–255 ova. Ova are brown with diameters averaging 2.3 mm (range: 1.6–2.83 mm). Miller et al. (2014) reported females with glair in November thru January. We found that females collected in early March had sperm plugs, whereas ova were attached to abdominal swimmerets by mid-March. This suggests an early spring mating period. Females (n ¼ 7) with mean 61 (range: 48–93) neonate young attached to their abdomen were captured in late April; while Miller et al. (2014) reported data for a single female taken with 255 instars. Additional seasonal reproductive data are presented in Table 2, which validates hypotheses reported by Miller et al. (2014). Cambarus (L.) erythrodactylus consumes both plant material
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and animal foods, including algae, dipterans and other aquatic insects, and annelids, based on laboratory diet experiments. Similar to other burrowing species, such as Cambarus (T.) thomai and C. (T.) polychromatus, this species is nocturnal and is most active at night. Individuals can be observed perched at the mouth of their burrow with their chelae and head exposed. Response to approaching predators or threats involves individual escape by free-falling or dropping down the burrow shaft. Individuals are photophobic. Aggressive behavior was observed, with the species responding to prey by lunging out of its burrow and attempting to grab the organism with its chelae. Aggression was not biased toward either sex, but neither male nor females ventured away from the burrow entrance. Adults could be lured no more than 150–300 mm from the burrow entrance with food (i.e., red worms). Burrows typically possess a single primary shaft and usually two secondary escape shafts. Burrow depths are usually less than one meter with mean depths of 0.7 m to the central chamber. Chimney heights range between 200 and 1225 mm. Crayfish associates.—Collected at one or more locations in Alabama and Mississippi with Procambarus (Pennides) suttkusi Hobbs, 1953, P. (Pennides) spiculifer (Le Conte, 1856), P. (Pennides) versutus (Hagen, 1870), P. (Scapulicambarus) okaloosae Hobbs, 1942, C. (Depressicambarus) striatus Hay, 1902, C. (Depressicambarus) latimanus (Le Conte, 1856), and C. (Tubericambarus) sp. B. Relationships.—Cambarus (L.) erythrodactylus is most similar to C. (L.) diogenes but differs from the latter in possessing many more spines and tubercles on the palm, cervical groove, mesial process, and the distal and proximal podomeres. Cambarus (L.) diogenes does not possess more than two rows of mesial-palm tubercles, whereas C. (L.) erythrodactylus has two rows of mesial palm tubercles, as well as an additional two to three partial to full rows
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on the palm. The gonopod shape is diagnostic. Cambarus (L.) erythrodactylus possesses the central and mesial projections equal in length (Fig. 1B, F), and the first pleopods are contiguous at the base, with convexity near midlength of cephalic surface. The distally truncated Form I male central projection and mesial projection are both directed caudally at an angle greater than 908 and are bent cephalodistally at a 208 angle from the shoulder. Cambarus diogenes has a caudally projected central projection at a 908 angle to the shoulder and shaft. The gonopod and antennal scale of Cambarus diogenes are wider than those of C. erythrodactylus. Etymology.—The prominent red distal edges of the propodus and dactyl, which comprise over two-thirds of the chela, account for the scientific name erythro(red) and -dactylus (claw). The vernacular name chosen for the species is the warpaint mudbug, based on the extensive red pigment outlining the rostral margins, joints of the pleopods, uropod margins, and red coloration along the distal twothirds of the dactyl and propodus, resembling warpaint markings. Acknowledgments Thanks are extended to J. F. Fitzpatrick, Jr. (deceased), Tulane University; R. F. Thoma, Midwest Biodiversity Institute; and P. M. Stewart, Troy University, for their insight and assistance on the project. R. F. Thoma and C. A. Taylor provided assistance on selecting diagnostic characters and other professional courtesies. We thank all individuals past and present that assisted in the field collection of specimens used in this study. Individuals include a virtual who’s who of current astacologists, including D. Chrisman, P. Donaker, T. Jones, J. Miller, C. Pickeral, P. M. Stewart, G. W. Stocker, R. F. Thoma, and Z. Thoma. Julia Ferguson prepared Figure 1. Figure 2 was prepared by the coauthor (CCM). Figure 3A is from U.S.
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Forest Service (S. B. Adams), and Figures 3B, 4 were taken by the senior author. This project was completed through the support of Troy University, Department of Environmental and Biological Sciences, Troy, Alabama, and the Indiana Biological Survey Aquatic Research Center, Bloomington, Indiana. Literature Cited Adams, S. B., & G. Henderson. 2009. Mississippi crayfishes database v. 3.1. United States Department of Agriculture Forest Service, Oxford, Mississippi. ,http://maps.fs.fed.us/ crayfish/speciesmap.jsp; (Last accessed 27 December 2009; no longer available as of 6 November 2014.) Bouchard, R. W. 1972. A contribution to the knowledge of Tennessee crayfish. Unpublished Ph.D. dissertation, University of Tennessee, Knoxville, 113 pp. Bouchard, R. W. 1974. Crayfishes of the Nashville Basin, Tennessee, Alabama, and Kentucky (Decapoda, Astacidae). [Abstract.] The Association of Southeastern Biologists Bulletin 21(2):41. Bouchard, R. W. 1976. Crayfishes and shrimps. Pp. 13–21 in H. Boschung, ed., Endangered and threatened plants and animals of Alabama. Bulletin of the Alabama Museum of Natural History 2, 93 pp. Buchanan, J. V. 1992. The burrowing crayfishes (Decapoda: Cambaridae) of the Cahaba River system. Unpublished M.S. thesis, University of South Alabama, Mobile, Alabama, 185 pp. Faxon, W. 1884. Descriptions of new species of Cambarus; to which is added a synonymical list of the known species of Cambarus and Astacus. Proceedings of the American Academy of Arts and Sciences 20:107–158. Faxon, W. 1885. A revision of the Astacidae. Part I. The genera Cambarus and Astacus. Memoirs of the Museum of Comparative Zoology at Harvard College 10(4):1–186. Fitzpatrick, J. F., Jr. 1978. A new burrowing crawfish of the genus Cambarus from southwest Alabama (Decapoda, Cambaridae). Proceedings of the Biological Society of Washington 91:748–755. Girard, C. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution. Proceedings of the Academy of Natural Sciences of Philadelphia 6:87–91. Hagen, H. A. 1870. Monograph of the North American Astacidae. Illustrated Catalogue of
the Museum of Comparative Zoology, at Harvard College 3:1–109. Hay, W. P. 1896. The Crawfishes of the State of Indiana. Pp. 476–507 in Indiana Department of Geology and Natural Resources, Twentieth Annual Report for 1895, 520 pp. Hay, W. P. 1899. Synopses of North American invertebrates. VI. The Astacidae of North America. American Naturalist 33:957–966. Hay, W. P. 1902. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and vicinity. Proceedings of the United States National Museum 25:417–439. Heath, W. H., P. M. Stewart, T. P. Simon, & J. M. Miller. 2010. Distributional survey of crayfish (Crustacea: Decapoda) in wadeable streams in the coastal plains of southeastern Alabama. Pp. 139–154 in Z. Loughman, S. Welsh, T. P. Simon, eds., Conservation, biology, and natural history of crayfishes from the southern United States: proceedings of a 2008 symposium of the Southern Division of the American Fisheries Society. Southeastern Naturalist 9, Special Issue 3. Hobbs, H. H., Jr. 1942. The Crayfishes of Florida. University of Florida Publication, Biological Science Series 3(2):1–179. Hobbs, H. H., Jr. 1953. A new crayfish of the genus Procambarus from Alabama and Florida (Decapoda, Astacidae). Proceedings of the Biological Society of Washington 66:173–178. Hobbs, H. H., Jr. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Pp. 93–178 in P. C. Holt, R. L. Hoffman, & C. W. Hart, Jr., eds., The distributional history of the biota of the southern Appalachians. Part I: Invertebrates. Research Division Monograph 1, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, 295 pp. Hobbs, H. H., Jr. 1974. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambaridae). Smithsonian Contributions to Zoology 166:1–161. Hobbs, H. H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Smithsonian Contributions to Zoology 488:1–236. IUCN. 2001. IUCN Red List Categories and Criteria: Version 3.1. Second edition. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, United Kingdom, 30 pp. Jezerinac, R. F. 1985. On the two forms of Cambarus (L.) diogenes (Decapoda: Cambaridae) in Ohio. April program abstracts, 94th annual meeting, The Ohio Academy of Science, April 19–21, 1985. The Ohio Journal of Science 85(2):7.
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Jezerinac, R. F. 1986. Endangered and threatened crayfishes (Decapoda: Cambaridae) of Ohio. Ohio Journal of Science 86(4):177–180. Jezerinac, R. F. 1993. A new subgenus and species of crayfish (Decapoda: Cambaridae) of the genus Cambarus, with an amended description of the subgenus Lacunicambarus. Proceedings of the Biological Society of Washington 106:532– 544. Le Conte, J. 1856 [1855]. Descriptions of new species of Astacus from Georgia. Proceedings of the Academy of Natural Sciences of Philadelphia 7:400–402. Lukhaup, C. 2003. S¨uwasserkrebse aus aller Welt. D¨ahne Verlag GmbH, Ettlingen, Germany, 292 pp. Marlow, G. 1960. The subspecies of Cambarus diogenes. American Midland Naturalist 64:229–250. Miller, J. M., B. B. Niraula, E. G. Rea´tegui-Zirena, & P. M. Stewart. 2014. Life history and physical observations of primary burrowing crayfish (Decapoda: Cambaridae) Cambarus (Lacunicambarus) diogenes and Cambarus (Tubericambarus) polychromatus. Journal of Crustacean Biology 34:15–24. Ortmann, A. E. 1905. The mutual affinities of the species of the genus Cambarus, and their dispersal over the United States. Proceedings of the American Philosophical Society 44(180):91–136. Ortmann, A. E. 1906. The crawfishes of the state of Pennsylvania. Memoirs of the Carnegie Museum 2(10):343–524 þ pls. A, B, 39–43. Schuster, G. A., & C. A. Taylor. 2004. Report on the crayfishes of Alabama: literature and museum database review, species list with abbreviated annotations and proposed conservation statuses. Illinois Natural History Survey, Center for Biodiversity Technical Report 2004 (12), 53 pp. Taylor, C. A., & G. A. Schuster. 2010. Status and distribution of the rusty gravedigger, Cambarus miltus: a case of recovery or neglect? [Abstract.] P. 6 in Z. J. Loughman, T. P. Simon, & S. A. Welsh, Forward; pp. 1–10 in Z. Loughman, S. Welsh, T. P. Simon, eds., Conservation, biology, and natural history of crayfishes from the southern United States: proceedings of a 2008 symposium of the Southern Division of the American Fisheries Society. Southeastern Naturalist 9, Special Issue 3. Taylor, C. A., G. A. Schuster, J. E. Cooper, R. J. DiStefano, A. G. Eversole, P. Hamr, H. H. Hobbs, III, H. W. Robison, C. E. Skelton, & R. F. Thoma. 2007. A reassessment of the conservation status of crayfishes of the United
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States and Canada after 10þ years of increased awareness. Fisheries 32(8):372–389. Thoma, R. F., R. F. Jezerinac, & T. P. Simon. 2005. Cambarus (Tubericambarus) polychromatus (Decapoda: Cambaridae), a new species of crayfish from the United States. Proceedings of the Biological Society of Washington 118:326–336. Williamson, E. B. 1899. Notes on Ohio Astacidae. Annual Report of the Ohio Academy of Science 7:47–48. Williamson, E. B. 1907. Notes on the crayfish of Wells County, Indiana, with description of a new species. Pp. 749–763 in 31st Annual Report of the Indiana Department of Geology and Natural Resources for 1906, 763 pp. Associate Editor: Christopher B. Boyko.
Appendix Collection groups Coll. 1: T. P. Simon (TPS) & C. C. Morris (CCM); Coll. 2: TPS & P. M. Stewart (PMS); Coll. 3: J. Miller (JM) & A. McCall (AM); Coll. 4: R. F. Jezerinac (RFJ), R. F. Thoma (RFT), G. W. Stocker (GWS), D. Chrisman (DC), P. Donaker (PD), C. Pickerel (CP), & Z. Thoma (ZT); Coll. 5: RFJ, GWS, DC, & T. Jones (TJ); Coll. 6: RFJ, GWS, & TJ; and Coll. 7: J. V. Buchanan (JVB) & L. L. White (LLW). Institutional abbreviations INBS, Indiana Biological Survey Aquatic Research Center Astacology Collection; OSUMC, Ohio State University Crustacean Collection; UADC, University of Alabama Decapod Collection; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C. Material examined ALABAMA. Bullock Co: INBS 229, 1 /, tributary Conecuh River, Co Rd 28 bridge, 0.25 mi SW Inverness, 20 Apr 2001, Coll. 1; Butler Co: OSUMC 2661, 1 /, Persimmon Creek, St Rd 106 bridge, 5.5 mi E Georgiana, J-88-14, 22 Mar 1988, Coll. 6; Cherokee Co: OSUMC 2657, 2 //, Terrapin Creek, St Rd 8, 2 mi SE US HWY 411 and St Rd 25 intersection, 3.7 mi SE Centre and 1.9 mi WSW Moshat, J-89-6, 14 Apr 1989, Coll. 4; Chilton Co: UADC 452.01, 1 /, Mahan Creek, Co Rd. 42 bridge, T 23N R13E S 21, JVB-02-2392-7, 23 Feb 1992, Coll. 7; Crenshaw Co: OSUMC 2660, 1 ?I, 1 ?II, 3 //, Patsaliga Creek, St Rd 106, 8.4 mi W Brantley (7.9 mi SE of Pigeon Creek), J-88-15, 23 Mar 1988, Coll. 4; Dale Co: INBS 101190, 1 /,
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Little Choctawhatchee River, site 7, JMM-08-07, 15 May 2008, Coll. 3; INBS 101191, 3 ?I, 2 ?II, 3 //, Little Choctawhatchee River, site 12, JMM-08-12, 12 May 2008, Coll. 3; INBS 101192, 4 ?I, 4 ?II, 4 //, Little Choctawhatchee River, site 11, JMM-07-11, 16 Jan 2007, Coll. 3; INBS 101193, 1 ?II, 1 /, Little Choctawhatchee River, site 9, JMM-07-09, 11 Feb 2007, Coll. 3; INBS 101194, 1 ?I, Little Choctawhatchee River, site 5, JMM-07-05, 31 Aug 2007, Coll. 3; INBS 101198, 2 ?II, 3 //, Little Choctawhatchee River, site 5, JMM-07-05, 7 Dec 2007, Coll. 3; INBS 101199, 1 ?I, 2 //, Little Choctawhatchee River, site 9, JMM-07-09, 28 Nov 2007, Coll. 3; Dallas Co: INBS 100899, 1 /, Soapstone Creek, US Hwy 80 bridge, 5 mi SE Selma, TPS-0216, 21 Jul 2002, Coll. 1; Elmore Co: INBS 100789, 1 /, unnamed tributary Tallapoosa River, Co Rd 4 (Rifle Range Rd), Old Bingham, TPS-03-27, 11 Apr 2003, Coll. 1; Fayette Co: OSUMC 2655, 7 ?I, 7 // , ditch, St Rd 171 just S of St Rd 159, 2.0 mi S Fayette, 5.5 mi ENE Belk, J-88-5, 20 Mar 1988, Coll. 5; OSUMC 2656, 5 ?I, 3 ?II, 10 //, ditch, St Rd 171 just S of St Rd 159, 2.0 mi S Fayette, 5.5 mi ENE Belk, J-87-117, 21 Mar 1989, Coll. 4; Greene Co: INBS 100838, 1 ?I, 2 //, Tubbs Creek, Co Rd 125 bridge, 0.3 mi N West Greene, TPS-02-14, 21 Jul 2002, Coll. 1; Jefferson Co: INBS 100922, 1 ?I, Rock Creek, Taylor Ferry Rd, 1.5 mi W Hopkins, TPS-02-07, 20 Jul 2002, Coll. 1; INBS 100954, 1 ?I,
Valley Creek, Warrior River Rd, 1 mi E Ezra, TPS02-08, 20 Jul 2002, Coll. 1; INBS 100956, 2 ?I, 2 // , unnamed tributary Mud Creek, Lock 12 Rd bridge and Griffis Rd intersection, TPS 02-09, 20 Mar 2002, Coll. 1; Montgomery Co: INBS 100950, 3 ?I, 4 //, Catoma Creek, St Rd 22 bridge, TPS-02-06, Coll. 1; Perry Co: UADC 450.01, 2 //, Little Creek, St. Rd. 183, 7.7 mi N St Rd. 14, T 20N, R 7E, S 17 SE1/4, JVB-09-0691-10, 6 Sep 1991, Coll. 7; OSUMC J-89-14, 1 ?II, unnamed ditch of Oakmulgee Creek, St Rd 183, ca. 8 mi W of US Rt. 82, 1.0 mi SW Oakmulgee, 11.5 mi SE Centreville, J-89-14, 20 Mar 1989, Coll. 5; Pickens Co: INBS 100968, 1 ?I, 1 /, Beaver Creek, St Rd 14 bridge, 5 mi W Aliceville, TPS-02-13, 21 Jul 2002, Coll. 1; Pike Co: INBS 228, 1 ?I, 1 /, unnamed tributary Walnut Creek, 1 mi NE Troy, Co Rd 26 bridge, 18 Apr 2002, Coll. 1; Washington Co: OSUMC 2659, 1 ?I, 4 ?II, 1 /, ditch, US HWY 43 and St Rd 13 intersection, 3.3 mi SW of Jackson, RFJ- J88-10, 21 Mar 1988, Coll. 5. MISSISSIPPI. Issaquena Co: OSUMC 3144, 1 ?I, 2 ?II, 4 //, ditch, US HWY 61, 2 mi N St Rd 3, 10 mi N Redwood, 20 mi S Valley Park, J-90-19, 20 Mar 1990, Coll. 4. Additional distribution records include specimens curated at Tulane University from Clarke, Covington, Lafayette, Lamar, Lee, Lowndes, Monroe, Pearl River, Rankin, Scott, Union, and Webster counties (Adams & Henderson 2009).
