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Redescription and distribution of the crayfish, Procambarus

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Redescription and distribution of the crayfish, Procambarus (Ortmannicus) pearsei (Creaser, 1934) (Decapoda: Cambaridae), with notes on its biology John E. Cooper North Carolina State Museum of Natural Sciences Research Lab, 4301 Reedy Creek Road, Raleigh, North Carolina 27607, U.S.A., e-mail: [email protected]
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Page 1: Redescription and distribution of the crayfish, Procambarus

Redescription and distribution of the crayfish, Procambarus (Ortmannicus) pearsei (Creaser, 1934) (Decapoda: Cambaridae),

with notes on its biology

John E. Cooper

North Carolina State Museum of Natural Sciences Research Lab, 4301 Reedy Creek Road,

Raleigh, North Carolina 27607, U.S.A., e-mail: [email protected]

Page 2: Redescription and distribution of the crayfish, Procambarus
Page 3: Redescription and distribution of the crayfish, Procambarus

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 124(1):9–22. 2011.

Redescription and distribution of the crayfish, Procambarus (Ortmannicus) pearsei (Creaser, 1934) (Decapoda: Cambaridae),

with notes on its biology

John E. Cooper

North Carolina State Museum of Natural Sciences Research Lab, 4301 Reedy Creek Road,

Raleigh, North Carolina 27607, U.S.A., e-mail: [email protected]

Abstract.—Procambarus (O.) pearsei is endemic to several river basins in

southeastern North Carolina and northeastern South Carolina, with most of its range in the former state. It was described over 75 years ago, but the

original description lacked many salient taxonomic features and very little

else has been published about the species. Recent examination of the two

primary type specimens, and an additional 296 specimens from all three of

the river basins to which the species is restricted, has prompted a

redescription that diagnoses and illustrates the species, provides distribu­

tional data and color notes, describes the previously neglected form-II male,

assesses the location of the nebulous type locality, and provides some information on the biology of the species.

Keywords: Carolina crayfishes, crayfish, parapatric crayfishes

Based on their phenotypes, three appar- It is absent from the autonomous North­

ently parapatric crayfishes—Procambarus east Cape Fear River basin and the related

(Ortmannicus) pearsei (Creaser, 1934), several elements of the White Oak River

Procambarus (Ortmannicus) plumimanus drainage (New, Newport, etc.) in North

Hobbs & Walton, 1958, and Procambarus Carolina, where it is replaced by P. (O.)

(Ortmannicus) medialis Hobbs, 1975— plumimanus (Cooper & Braswell 1995:

comprise a disjunct enclave of the planir- 113, 122). The single locality reported

ostris group of the genus Procambarus for P. (O.) pearsei in the Neuse River

that inhabits the Coastal Plain of North basin in Johnston County (Hobbs 1975:

Carolina. The other members of this 14, 1989:70) has been shown to apply to

group occur in Alabama, Louisiana, and P. (O.) medialis (Cooper & Braswell

Mississippi (see Hobbs 1962, 1972a). 1995:110, 122), which is endemic to the

Procambarus (O.) pearsei is the widest Neuse and Tar-Pamlico River basins.

ranging member of the North Carolina Procambarus (O.) pearsei is relatively

trio, and the only one that is not endemic abundant in ponds, roadside ditches,

to North Carolina. Its range encompasses Carolina bays, and borrow pits, and is

the lower Cape Fear, Lumber-Little Pee sometimes found in burrows. Yet, al-

Dee, and Waccamaw River basins in the though the taxon was established over

Carolinas. In North Carolina it is known 75 years ago, the species has never been

from Bladen, Brunswick, Columbus, adequately described and illustrated.

Cumberland, Hoke, Robeson, Sampson, Creaser’s (1934) description of ‘‘Cam­

and Scotland counties. South Carolina barus’’ (Ortmannicus) pearsei, which was

records are available from Dillon, Horry, based on two form-I males and eight

and Marion counties, and the species females from the type locality, was quite

likely occurs in Marlboro County as well. minimal, did not include a morphotypic

Page 4: Redescription and distribution of the crayfish, Procambarus

10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(form II) male, lacked a diagnosis and

reliable color notes, and contained illus­

trations of only the antennal scale and

form-I male gonopod. Some basic outline

drawings were later provided by Hobbs

(1968:27, 1972b:27, 47; 1974:143, 1989:

190), but there is no indication that any of

them represented Creaser’s types, which

were transferred from UMMZ to USNM

in 1954 (Hobbs 1954:1). When Hobbs &

Walton (1958) described P. (O.) pearsei

plumimanus, they added a brief diagnosis

of the nominate subspecies. In support of

their decision to treat these taxa as sub-

specifically related, they included in their

discussion of variation what they inter­

preted as ‘‘intergrade’’ specimens from

the Neuse River basin. These ‘‘inter­

grades,’’ however, were later assigned to

the new species P. (O.) medialis, at which

time P. (O.) plumimanus and P. (O.)

pearsei were elevated to full species

(Hobbs 1975). Consequently, the diagno­

sis of P. (O.) p. pearsei in Hobbs &

Walton (1958:8) was compromised by

inclusion of a few characters that later

devolved to P. (O.) medialis.

This paper is based on an examination

of Creaser’s two primary type specimens,

both of which are in poor condition, and

an additional 296 specimens from all

three of the river basins to which P. (O.)

pearsei is restricted. It provides diagnoses,

redescribes and illustrates the species;

gives color notes; describes a form-II

male; assesses the location of its nebulous

type locality; and includes some informa­

tion on the biology of the species.

Methods and Abbreviations

Measurements were made to the near-

Museum of Natural Sciences, Raleigh;

PCL 5 postorbital carapace length; R 5 River; SR 5 state secondary (county)

road; TCL 5 total carapace length;

UMMZ 5 University of Michigan Mu­

seum of Zoology, Ann Arbor; US 5 United States highway; USGS 5 U.S.

Geological Survey; USNM 5 National

Museum of Natural History, Smithsonian

Institution, Washington, D.C.

Procambarus (Ortmannicus) pearsei

(Creaser, 1934)

Fig. 1, Table 1

Cambarus (Ortmannicus) pearsei Creaser,

1934:1.—Hobbs, 1975:1 (by implica­

tion).

Procambarus pearsei: Hobbs, 1942:343.—

Cooper & Braswell, 1995:110.—Taylor

et al., 1996:32; 2007:386.

Procambarus pearsei pearsei: Hobbs &

Walton, 1958:7.—Hobbs, 1962:288; 1968:

K10.

Procambarus (Ortmannicus) pearsei pear-

sei: Hobbs, 1972a:10; 1972b:59; 1974:59.

Procambarus (Ortmannicus) pearsei:

Hobbs,1975:15;1989:70.—Hobbs&Peters,

1977:6.—Fitzpatrick, 1983:213.—Cooper

& Braswell, 1995:108.—Cooper, 2002:178.

The above synonymy includes only

references that have appeared in the

published literature; no citations from

open-file and similar reports are included.

Diagnosis.—Body and eyes pigmented,

eye large (mean adult diameter 2.2 mm,

n 5 83). Rostrum acarinate, very broad;

margins narrow, slightly elevated, gently

curving to base of short acumen, then

slightly more converging to small apical

tubercle or spine; margins occasionally

with small tubercles at or slightly cephalic

{

{

{

length

caliper, using a stereomicroscope and comprising 15.2–29.6% (X 5 20.8%, n 5 85) of rostrum length, latter comprising

X 5 23.6%, n 5 86) of TCL.

X 5 5.1, n 5 87) times as

est 0.1 mm with a Fowler precision dial to base of acumen; of acumen

following the methods of Hobbs (1981:9,

10) unless otherwise noted. Abbreviations

used in the text are: Crk 5 Creek; j 5 21.5–26.8% (

Areola 3.2–7.1 (

juvenile; NC {

5 North Carolina State long as broad, constituting 29.8–41.0%

(X 5 32.6%, n 5 88) of TCL and 38.4– highway; NCSM 5 North Carolina State

Page 5: Redescription and distribution of the crayfish, Procambarus

11 VOLUME 124, NUMBER 1

Fig. 1. Procambarus (Ortmannicus) pearsei. A, D, H, L, M: holotypic male, form I (USNM 98336); B, C, G,

I: form-I male (NCSM 2663); E, F: male, form II (NCSM 24993); and J, K: allotypic female (USNM 98337). A,

lateral aspect of carapace; B, E, mesial aspect of gonopod (first pleopod); C, F, lateral aspect of gonopod; D,

dorsal aspect of carapace; G, caudal aspect of in situ gonopods; H, epistome; I, basis and ischium of third and

fourth pereopod; J, annulus ventralis and postannular sclerite; K, antennal scale; L, dorsal aspect of distal

podomeres of right cheliped; M, lateral aspect of fixed finger of propodus. Scale bar 5 2 mm.

Page 6: Redescription and distribution of the crayfish, Procambarus

12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Measurements (mm) of Procambarus (Ortmannicus) pearsei.

Holotypic male, form I Allotypic female Male form II

Carapace

Total length 28.8 27.3 24.4

Postorbital length 22.9 21.6 19.3

Width 13.7 12.1 12.0

Depth 13.2 14.0 10.6

Length rostrum 6.6 6.1 5.6

Length acumen 1.1 1.1 1.1

Length areola 9.1 8.7 8.2

Width areola 1.6 1.5 1.2

Antennal scale

Length – 4.9 5.2

Width – 2.6 2.6

Abdomen

Length 30.1 29.0 28.2

Width 11.5 11.8 10.5

Cheliped

Length lateral margin

propodus 26.8 16.1 18.3

Length mesial margin

palm 9.8 5.6 7.4

Width palm 7.9 5.8 5.8

Depth palm 5.4 3.5 4.1

Length dactyl 14.2 9.2* 9.5

Gonopod length 7.3 N/A 6.6

* Estimated.

{

44.3% (X 5 41.0%, n 5 88) of PCL, and {

with 3–6 (usually 3 4) punctations or

across narrowest part. Thoracic section

X 5{

{

{67.8–100.0% (X 5 81.2%, n 5 41) of length

in males, 81.0–120.0% (X 5 102.3%, n 5 45) of length in females; length of palm

constituting 33.5–42.4% (X 5 37.4%, n 5of carapace constituting 64.9–70.2% (

67.7%, n 5 88) of TCL; dorsally punctate,

dorsolaterally and laterally granulate; cephalic section laterally with granules or

very small tubercles, dorsally with scat­

tered punctations; hepatic region without

spines. Cervical spines reduced to small

tubercles; cervical groove interrupted.

Branchiostegal spine small, acute. Subor­

bital angle obtuse to obsolete, usually

without tubercle. Postorbital ridge long, {

83) of chela (propodus) length; mesial

margin of palm with several staggered, poorly defined rows of 9 to 13 tubercles;

dorsal surface of palm covered with strong

tubercles, those on median third smaller

than others; ventral surface of palm less

tuberculate, those tubercles on mesial half

larger than others; all tubercles on chela

with short, fanlike setae.

Fingers occluded; dactyl 1.2–1.6 (X 5 with shallow, continuous groove; cephalic

{margin tapered, occasionally with tuber­

cle. Antennal scale (Fig. 1K) 1.8–2.7 (X 5

{1.4, n 5 30) times length of mesial margin

of palm in form-I males, 1.3–1.8 (X 5 1.6,

n 5 53) in form-II males and females;

2.2, n 5 87) times as long as broad, widest proximal fourth or less of mesial surface

near midlength; lateral margin thickened, with serrate row of tubercles; lateral

{terminating in acute, usually short spine.

Palm of chela of cheliped 1.3–1.7 (X 5 surface of fixed finger of propodus with

deep, punctate longitudinal groove along

1.5, n 5 65) times as wide as deep; width entire length.

Page 7: Redescription and distribution of the crayfish, Procambarus

13 VOLUME 124, NUMBER 1

Hook on ischium of third and fourth

pereopods of males, that on third pereo­

pod largest of pair; neither hook opposed

by clearly defined tubercle on basis. In

situ gonopods (first pleopods) of form-I

male (Fig. 1B, G, based on NCSM 2663)

asymmetrical; proximomesial apophyses

produced, not overlapping, without cau­

dal spur; shaft straight, narrow; mesial

process tapered, acute, directed caudolat­

erally; in lateral aspect (Fig. 1C), distal

fourth of shaft inclined caudally at ca. 90u to plane of shaft, terminal elements

directed caudally; cephalic process curv­

ing, subacute, tip directed caudoproxi­

mally; mesial process originating mesial

to caudal process, tapering, acute, direct­

ed caudally; subdistal setae originating

along bases of cephalic process and

central projection, partly obscuring parts

of both; total length of gonopod 25.3% of

TCL, 31.9% of PCL.

Annulus ventralis (based on allotypic

female; Fig. 1J) spindle-shaped in ventral

outline, with elongated lateral corners, 2.5

times wider than long, width 30.6% of

carapace width; sinistral half or more very

prominent; most of broad cephalic trough

and excavation situated dextral to mid­

line; cephalic half depressed, strongly

sloping; sulcus very deep beneath moder­

ately thick dextral margin; transverse

tongue short, broad, plunging into reverse

C-shaped caudosinistral margin. Config­

uration of annulus apparently invariable;

no mirror image seen in 49 adult females

from throughout range.

Measurements of both primary types

and a form-II male provided in Table 1.

Description of holotypic male, form 1.—

Body and eyes pigmented, eye 2.2 mm

diam. Cephalothorax (Fig. 1A, D) subcy­

lindrical, relatively narrow, vaulted (max­

imum width and depth subequal); length of

cephalic section 2.2 times length of areola

and constituting 68.4% of TCL. Areola 5.7

times longer than wide, constituting 31.6%

of TCL (39.7% of PCL), with 3 puncta­

tions across narrowest part; width of areola

11.7% of greatest width of thorax; bran­

chiocardiac grooves well defined. Rostrum

broad, with narrow, moderately elevated

margins gently curving to base of short

acumen and devoid of tubercles or spines;

margins slightly more converging from

base of acumen to very weak apical

tubercle, tip of which reaching slightly

beyond midlength of penultimate segment

of antennular peduncle; acumen compris­

ing 16.7% of rostrum length, latter consti­

tuting 22.9% of TCL; inner margins of

dorsal ridge flanked by row of setiferous

punctations; floor (dorsal surface) of ros­

trum subplane, with large, scattered, seti­

ferous punctations; ventral keel of rostrum

very weak, shallow, lacking spines or

tubercles; subrostral ridge underslung, not

visible in dorsal aspect.

Postorbital ridge long, slightly curving,

groove shallow, mostly lateral; caudal

margin slightly inflated; cephalic margin

tapered, without tubercle. Suborbital an­

gle obtuse, without tubercle; orbital rim

with shallow concavity around base of

antennal peduncle; branchiostegal spine

small, acute. Cervical spine region with

several weak tubercles; cervical groove

interrupted just dorsal to tubercles, with

short sulcus cephalic to groove; area

ventral to anterior section of groove very

granulate. Carapace with dorsal surface

of thoracic section punctate, dorsolateral

and lateral surfaces granulate; lateral

surface of cephalic section granulate,

dorsal surface with large punctations,

gastric region punctate; caudal termini

of postorbital ridges continuing caudally,

then curving inward nearly to midline of

carapace, forming low, arcuate ridge.

Antennal peduncle without distolateral

spine or tubercle on basis, with small,

squamous tubercle on ventral surface of

ischium; antennular peduncle with small,

mesially situated spine at midlength of

ventral surface of basal segment; both

antennal scales missing (see description of

scale of allotypic female), antennal flagel­

lae damaged.

Page 8: Redescription and distribution of the crayfish, Procambarus

14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Abdomen longer than carapace, width

of abdomen 83.9% of greatest carapace

width; pleura of most abdominal seg­

ments rounded on all margins, surface

covered with large punctations; surfaces

of terga with small punctations. Proximal

podomere of uropod with small lateral

spine on lateral lobe, slightly larger spine

on mesial lobe; mesial ramus of uropod

with moderate caudolateral spine, and

strong submedian ridge terminating in

spine situated well cephalic to caudal

margin; lateral ramus with submedian

ridge on cephalic section; transverse

flexure of ramus with margin bearing

row of 19 fixed spines and large, articu­

lated sublateral spine. Telson with 2

strong spines in each caudolateral corner

of cephalic section, innermost spine artic­

ulated; transverse flexure strong; caudal

margin of telson subtruncated, with slight

median concavity.

Epistome (Fig. 1H) with broad, basi­

cally subcordiform cephalic lobe bearing

long, narrow cephalomedian projection;

margins of lobe moderately elevated

(ventrally), lateral apices slightly thick­

ened, rounded, not flared; floor (ventral

surface) of lobe broadly convex, punctate,

lacking setae; body of epistome with

broad central depression bearing slitlike

fovea; lamellae sparsely punctate, cepha­

lolateral margins strongly sloping, taper­

ing laterally to subtruncate margin with

weak caudal tubercle; zygoma strongly

arched, laterally expanded, flanked ceph­

alolaterally on each side by deep, elon­

gate pit.

Third maxilliped with distal segment

missing on left; tip of right endopodite

reaching midlength of ultimate segment

of antennal peduncle; exopodite not

notably setose, tip reaching slightly be­

yond distal margin of merus of endopo­

dite; distolateral corner of ischium slightly

produced, subacute; ventrolateral half of

ischium with punctuations bearing long

setae, and strong longitudinal ridge with

inner margin flanked by row of puncta­

tions with longer setae, all setae combined

obscuring most of ventrolateral surface;

ventromesial half of ischium, and ventral

surface of basis, with tufts of long, dense

bristles. Right mandible with incisor ridge

bearing 9 denticles.

Left cheliped regenerated; palm of right

chela (Fig. 1L) subovate in cross section,

elongate; length of chela (propodus) 3.4

times greatest width, 93.1% of TCL;

length of mesial margin of palm 1.8 times

greatest depth, margin with several, often

irregular rows of 8–10 tubercles each; all

surfaces of palm covered with prominent

tubercles of varying size, many of those

on median third of dorsum smaller than

others, most tubercles with fanlike setae;

dorsal tubercles encroaching on poorly

defined articular ridge, including its distal

margin. Fingers narrow, both slightly

curving distoventrally, opposable surfaces

occluded; dactyl bowed in dorsal aspect,

length 53.0% of chela length, 1.4 times

length of mesial margin of palm; dorsal

surface of dactyl with strong, narrow

longitudinal ridge, distal three-fourths of

which flanked mesially by broad surface

bearing setiferous punctations, proximal

fourth flanked by tubercles; ridge flanked

laterally by punctate groove on distal

three-fourths, by tubercles on proximal

fourth; mesial surface of finger with

several rows of strong tubercles on

proximal third, tubercles encroaching on

dorsal and ventral surfaces, basalmost

ones semierect, subacute; ventral surface

of finger with narrow keel, flanked

mesially by broad surface, proximal half

of which tuberculate, rest with large

punctations; keel flanked laterally by

large punctations; opposable surface of

finger with 18 tubercles dorsal to denti­

cles, 5 or 6 ventral to them, third from

base very large, protrusive; denticles in

dense pad arranged in 6–8 rows from tip

to about midlength, 2 or 3 rows from

there to base. Fixed finger with strong,

narrow dorsomedian ridge, flanked later­

ally by moderate, punctate groove, mesi­

Page 9: Redescription and distribution of the crayfish, Procambarus

15 VOLUME 124, NUMBER 1

ally by punctate surface; dorsolateral

margin strongly costate, proximolateral

fifth of dorsal surface with prominent

tubercles; lateral surface of finger with

deep, punctuate longitudinal groove

(Fig. 1M); ventral surface with strong,

rounded keel, flanked mesially by punc­

tations on distal two-thirds, by tubercles

on proximal third; keel flanked laterally

by punctations; opposable surface with

large subconical tubercle ventral to den­

ticles at base of distal two-fifths of finger

and 1 small tubercle slightly distal to it at

same level; 12 tubercles dorsal to denti­

cles, second from base massive, protru­

sive, overlapping third tubercle from

base of opposable surface of dactyl when

fingers closed; denticles in dense pad

arranged in 5 or 6 rows to level of sub-

conical tubercle, 2 or 3 rows from there to

base of finger.

Carpus of right cheliped (Fig. 1L) 1.5

times longer than broad, length 74.5% of

palm length; dorsal surface with broad,

shallow, oblique sulcus, surface mesial to

which with several admixed rows of

strong tubercles, including single strong,

subconical tubercle on distomesial corner

near articular condyl; surface lateral to

sulcus punctate; mesial surface with

several rows of strong, admixed tubercles,

some larger than others, distalmost larg­

est; ventral surface with weak distolateral

tubercle, strong conical distomedian tu­

bercle, and several strong conical tuber­

cles proximomesial to latter. Merus elon­

gate, 2.8 times longer than greatest depth,

dorsal surface with many strong subdistal

tubercles and row of prominent tubercles

along dorsal ridge; distolateral surface

and distal third of distomesial surface

with tubercles; ventrolateral ridge with 14

small, subacute tubercles and vestigial

distal tubercle, ventromesial ridge with

16 acute, larger tubercles and moderate

distal spine; 6 strong tubercles in diagonal

row between distal extremities of ridges,

and several small tubercles between ridg­

es; ischium with row of 5 strong tubercles

on ventral ridge, 4 smaller ones on dorsal

ridge.

Most podomeres of both fourth pereo­

pods missing; hook on ischium of third

pereopod (Fig. 1I) strong, tapering, obli­

que, overreaching basioischial articula­

tion by tip, not opposed by tubercle on

basis; hook on ischium of fourth pereo­

pod smaller, vertically disposed, tip not

reaching articulation; coxa of fourth

pereopod with strong, vertically disposed

caudomesial boss; coxa of fifth pereopod

with strong, flattened mesial boss. Left

gonopod missing; for description of right

gonopod see ‘‘Diagnosis.’’

Description of allotypic female.—Other

than secondary sexual characters, differ­

ing from holotypic male as follows: TCL

2.3 times maximum carapace width;

cephalic section of carapace 2.1 times

length of areola, latter 5.8 times longer

than wide. Rostrum with minute tubercle

on each side between base of acumen and

weak apical tubercle; acumen comprising

18.0% of rostrum length. Postorbital

ridge with minute cephalic tubercle; sub­

orbital angle with vestigial tubercle. An­

tennal peduncle with small distolateral

tubercle on basis and small tubercle on

ventral surface of ischium. Antennal scale

with lateral margin broadly and slightly

convex, tip of very short distolateral spine

reaching slightly beyond distal margin of

ultimate podomere of antennular pedun­

cle; width of mesial (lamellar) portion ca.

2.3 times width of thickened lateral

portion, distal margin slightly convex

before curving proximomesially to widest

part at midlength, then abruptly turning

proximolaterally and obliquely to base.

Width of abdomen 97.5% of greatest

carapace width.

Right cheliped missing, tip of left dactyl

off; palm of left chela 2.8 times longer

than wide, length 59.0% of TCL; length

of mesial margin 1.6 times depth of palm.

Length of dactyl ca. 57% of total chela

length, ca. 1.6 times length of mesial

margin of palm; opposable surface of

Page 10: Redescription and distribution of the crayfish, Procambarus

16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

finger with 10 tubercles, fourth from base

largest and situated ventral to denticles,

which in 1 or 2 rows. Opposable surface

of fixed finger with strong subconical

tubercle ventral to denticles at base of

distal third of finger, 10 tubercles dorsal

to denticles, fourth from base largest,

protrusive, not overlapping fourth tuber­

cle from base of opposable surface of

dactyl when fingers closed; denticles in 1

or 2 rows. Carpus and mesial surface of

palm subequal in length; ventral surface

of carpus with conical distolateral tuber­

cles. Merus 2.4 times longer than deep;

ventrolateral ridge with 12 tubercles and

minute distal spine, ventromesial ridge

with 14 tubercles and moderate distal

spine; ischium with 4 weak tubercles on

ventral ridge, 4 smaller ones on dorsal

ridge.

First pleopods moderately long; ovi­

duct apertures open, surrounded by long,

dense setae; preannular sternite with

broad, glabrous, U-shaped median floor

between steep walls (width of sternite

between ventral margins 33.9% of cara­

pace width). For description of annulus

ventralis see ‘‘Diagnosis.’’

Description of male, form II.—(Based

on NCSM 24993.) Differing from holo­

typic male as follows: areola 6.8 times

longer than wide, constituting 33.6% of

TCL (42.5% of PCL). Acumen compris­

ing 19.6% of rostrum length, latter

constituting 23.0% of TCL. Length of

cephalic section of carapace 2.0 times

areola length, comprising 66.4% of TCL.

Suborbital angle obsolete; antennal pe­

duncle with 2 small tubercles. Distolateral

corner of ischium of third maxilliped not

produced.

Both chelipeds normal; length of chela

(propodus) 75.0% of TCL, 3.2 times

longer than greatest width. Mesial margin

of palm with 3 irregular rows of 9 to 13

tubercles. Length of dactyl 51.9% of chela

length, 1.3 times length of mesial margin

of palm; opposable surface of dactyl with

12 tubercles dorsal to denticles, 1 large,

prominent tubercle ventral to denticles;

latter in 4 rows. Opposable surface of

fixed finger of propodus with 6 tubercles

dorsal to denticles, single large tubercle

ventral to denticles at base of distal third

of finger; denticles in 4 or 5 rows. Carpus

of right cheliped 1.6 times longer than

wide, length 87.8% of palm length. Merus

of cheliped 2.6 times longer than greatest

depth; ventrolateral ridge with 11 tuber­

cles (9 on left) and vestigial distal spine;

ventromesial ridge with 14 tubercles and

strong distal spine; ischium with 6 tuber­

cles (5 on left) on ventral ridge, none on

dorsal ridge. Width of abdomen 87.5% of

greatest carapace width.

Hooks on ischia of third and fourth

pereopods tuberclelike, no tubercles in

opposition on either basis; bosses on

coxae of fourth and fifth pereopods

moderately developed. In situ gonopods

slightly asymmetrical; proximomesial

apophyses weak, acute; cephalic process

curving caudolaterally; in lateral and

mesiao aspect (Fig. 1E, F), distal third

of gonopod curving at ca. 45 degrees to

plane of shaft; very short setae along base

of cephalic process; central projection

small, tuberclelike; juvenile suture faintly

detectable. Total length of gonopod

27.0% of TCL (34.2% of PCL).

Color notes.—Based on five live fe­

males and a live form-I male: ground

color of dorsal cephalothorax greenish-

tan, dark green-brown, or olivaceous,

dorsolaterally with black spots or dark

brown blotches; lateral surface of thoracic

section with dark and light mottling,

sometimes pinkish with pale blue ventral

margin; areola pale, bearing black spots;

pale longitudinal median stripe on ceph­

alothorax; tubercles on lateral surfaces

gray, those in cervical spine area white,

gray, or pale tan; orbital rim with white

horizontal band between branchiostegal

spine and anterior terminus of cervical

groove; antennal scale with dark lateral

margin, rest pale or olivaceous, often with

scattered black spots; antennal and an­

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17 VOLUME 124, NUMBER 1

tennular flagellae pale greenish or tan,

banded with paler interstices.

Dorsal surface of abdomen with pink­

ish or orangish-tan median stripe, flanked

each side by narrow dark stripe, which in

turn flanked dorsolaterally by broad,

light reddish or tannish-green stripe;

dorsolateral surface of each pleuron with

dark crescentric marking, series of mark­

ings forming narrow, irregular stripe;

ventrolateral pleura pinkish to orangish,

with scattered, fine dark markings. All

ventral surfaces of body caudal to man­

dibles blue-gray or translucent; mandibles

blue, with white margin along surface of

incisor ridge; epistome reddish or iodine;

circular margins of renal apertures white;

zygoma greenish-blue. Telson and uro­

pods blue-green or tannish, dorsal sur­

faces with pale orange, pink, or black

spots and flecks; dorsomedian ridge and

lateral margin of mesial ramus of uropod

dark; spines along transverse flexure of

lateral ramus black, producing narrow

transverse band.

Ventral surfaces of second pleopods of

male blue; caudal surfaces of gonopods

with blue markings. Walls of annulus

ventralis white or creamy, with some

blue flecks, and parts of caudal wall with

dark gray markings; postannular sclerite

(Fig. 1J) white or gray. Dorsal surface of

palm of cheliped ground color, most

tubercles black, some gray or oyster;

ventral surface of palm tan or ground

color, mesial half often darker than

lateral half, most tubercles light; tubercles

on mesial surface of palm sometimes very

dark. Fingers ground color; ventral sur­

face of dactyl frequently darker than

same surface of fixed finger; tips of both

fingers amber, or whitish with pale bluish

tinge, color not subtended by dark band;

base of dorsal surface of fixed finger often

with black spots; lateral surface of entire

propodus with greenish-brown longitudi­

nal stripe and occasionally black spots;

tubercles on opposable surfaces of fingers

white or tan. Dorsal surface of carpus,

merus, and ischium of cheliped, and all

podomeres of other pereopods, dark

greenish-gray, other surfaces varying

from white to light bluish-gray to dark;

tubercles on dorsal surface of carpus and

merus very dark, those on mesial and

ventral surfaces white or grayish.

Location of types.—The holotypic male,

form I, and the allotypic female are in

the USNM crayfish collection, catalog

numbers 98336 (originally UMMZ 53792)

and 98337 (originally UMMZ 53793),

respectively. The other eight of Creaser’s

specimens, all from the type locality, are at

USNM: 1 = I (69361); 1 R (69360); 6 R (98338; originally UMMZ 53794, all six

were designated paratypes by Creaser). The

male, form II, is in the NCSM crustacean

collection (24993).

Type locality.—The nebulous type lo­

cality was given as ‘‘Pond and ditch on

Highway No. 22, south of Fayetteville,

Cumberland County, North Carolina’’

(Creaser 1934:3). ‘‘Highway No. 22’’ is

now US 301, which is paralleled by I-95

and traverses elements of both the Cape

Fear and the Lumber-Little Pee Dee

River basins. Since the type locality is in

Cumberland County, though, it is within

the Cape Fear drainage. Obviously, in the

75+ years since the description of P. (O.)

pearsei there has been a great deal of

development in the probable area of the

type locality and considerable expansion

of the city of Fayetteville in all directions.

It is highly likely that habitat for the

species at what could have been the

original type locality no longer exists.

Range and specimens examined.—

Known from eight counties in the lower

Cape Fear, Lumber-Little Pee Dee, and

Waccamaw River basins in southeastern

North Carolina, and three counties in the

latter two basins in northeastern South

Carolina. Specimens have been collected

at the following localities (all are at

NCSM unless otherwise noted): NORTH

CAROLINA. Cape Fear River basin:

Bladen Co.—(1) pond along SR 1327,

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18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

2.6 rd km NW jct SR 1325, 9.4 air km W

of Ammon (34.80389N, 278.68166W); 1

= I, 1 j = (304), 17 Mar 1979, coll. A. L.

Braswell (ALB), R. E. Ashton, Jr. (REA),

P. S. Ashton (PSA); (2) baylike depres­

sion W of NC 210, S of Colly Crk,

4.5 air km NE of Kelly (34.49556N,

278.28722W); 2 = II, 1 R, 1 j ? (2692), 1

j =, 1 j R (2702), 30 Apr 1987, coll. ALB;

(3) Horseshoe Lake (Suggs Mill Pond),

1.1 km NE of SR 1327; 1 = I, 8 j =, 1 R, 6

j R (1602), 23 Feb 1980, coll. REA, PSA,

and others; (4) roadside ditch on SR 1730,

0.8 km NE jct NC 211 in subdrainage

Friar Swamp; 1 R (5266), 24 Jun 1999,

coll. W. C. Starnes (WCS), D. G. Cooper

(DGC), et al. Brunswick Co.—(5) depres­

sion pond 7.7 air km NW of Winnabow; 2

j R (7424), 13 Mar 2001, coll. ALB, R.

LeBlond (RL), J. T. Finnegan (JTF), N.

Murdock (NM); (6) sinkhole pond ca. 6.1

air km NW of Winnebow; 1 = I, 1 R, 1 j R (7425), 13 Mar 2001, coll. ALB, RL, JTF,

NM; (7) sinkhole pond ca. 14.2 air km

WSW of Leland; 1 = I, 1 R (7426), 13 Mar

2001, coll. ALB, RL, JTF, NM; (8)

sinkhole pond along Goose Pond Road,

ca. 15.2 air km WSW of Leland; 3 j =, 2 R,

2 j R (7427), 14 Mar 2001, coll. ALB, JTF;

(9) sinkhole pond ca. 8.0 air km NW of

Bolivia; 1 = I, 1 R (7428), 13 Mar 2001,

coll. ALB, RL, JTF, NM. Cumberland

Co.—(10) ‘‘pond and ditch on State

Route 22, south of Fayetteville’’; 2 = I

(USNM 69361, 98336), 1 R (USNM

98337), 6 R (USNM 98338), 25 Mar

1934; 1 R (USNM 69360), 4 Jul 1933;

coll. A. S. Pearse; (11) Black Ground Bay

east of jct NC 53 and SR 1228; 1 j =, 2 R (6012), 10 Jul 2000, coll. G. S. Grant

(GSG); (12) ‘‘Cape Fear Bluffs, Massen­

gale property’’; pair dried chelipeds,

probably = I (6010), 9 Jul 2000, coll.

GSG; (13) shallow grassy ditch along NE

side SR 1609, ca. 0.8 km NW jct SR 1705,

2.9 air km SE of Linden (35.195N,

278.72861W); 1 = II (2027), 26 Jan

1975, coll. M. R. Cooper (MRC), J. E.

Cooper (JEC); (14) roadside ditch on US

13, 9.6 km S of Sampson Co. line; 3 = I, 22

= II, 7 j =, 10 R, 11 j R (USNM 131773), 21

May 1971, coll. H. H. Hobbs, Jr. (HHH).

Sampson Co.—(15) roadside ditch along

NC 242, 4.3 rd km N of jct US 421, ca. 13

air km W of Newton Grove; 1 = II, 4 j =, 3

j R (2014), 24 Mar 1975, coll. MRC, JEC;

(16) alive on NC 242, 0.3 rd km N of jct

SR 1634, 2.9 air km NNW of Spiveys

Corner (35.22139N, 278.49528W), ca.

0922 hr EDT, light rain; 1 R (5053), 21

Mar 1999, coll. J. C. Beane (JCB), J. W.

Rowland, Jr.; (17) roadside ditch on US

13, 0.6 km SW of Wayne Co. line; 1 = I, 2 j

=, 4 R, 6 j R (USNM 131767), 21 May

1971, coll. HHH; (18) roadside ditch on

US 13, 10.7 km S of Newton Grove; 7 j =,

16 j R (USNM 131768), 21 May 1971, coll.

HHH.

Lumber-Little Pee Dee River basin:

Bladen Co.—(19) on SR 41, 6 km S of

Dublin (Hobbs & Peters 1977:52, 54).

Hoke Co.—(20) Antioch Church Bay on

E side NC 211, 1.0 rd km SSE of jct SR

1447, 3.2 air km SSE of Antioch

(34.86361N, 279.19945W); 4 = I, 3 R (2240), 1 Feb 1985; 1 = II, 2 R (2241), 23

Feb 1985; 1 R (2668), 7 Feb 1986, coll. D.

L. Stephan (DLS), R. W. Laney (RWL),

D. F. Lockwood (DFL); (21) Hamby’s

Bay at SR 1448, 1.9 rd km SE jct SR 1105,

2.4 air km E of Antioch (34.88445N,

279.1825W); 1 = I (2663), 21 Mar 1990,

coll. ALB; 2 R (4486), 14 Feb 1998; 1 = I, 1

= II (5011), 7 Feb 1999, coll. JCB, JTF, S.

J. Horton (SJH). Robeson Co.—(22)

Goose Pond Bay off SR 1704, 2.9 air km

WSW town Lumber Bridge (34.88194N,

279.10056W); 3 R with third instar young

(2246, 2248, 2249); 1 = I, 1 = II, 1 R, 1 R with young (2247), 5 = I, 2 = II, 11 R, 6

loose young (2250), 23 Feb 1985; 3 = I, 3 = II, 1 j =, 12 R (2676), 10 Jan 1987; 1 R (2315), 27 Apr 1985, coll. DLS, DFL; 1 R (4459), 17 Jan 1998, coll. JCB, R. A. Davis

(RAD); (23) dead on SR 1704 along

Goose Pond Bay; 1 R (24258), 24 Jan

1997, coll. JCB, RAD, J. Morgan-Davis;

(24) field near Goose Pond Bay; 1 dried

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VOLUME 124, NUMBER 1

chela & other parts (3719), 19 May 1997,

coll. DGC, D. A. Jackan, D. DeOliviera;

(25) borrow pit along US 74 near railroad

tracks, 0.5 rd km NW jct SR 1554, 2.9 air

km SSW of Pembroke (34.65639N,

279.20805W); 1 R (800), 13 May 1982,

coll. ALB, W. M. Palmer (WMP); 1 = I, 2

= II, 2 j =, 2 R, 4 j R (2664), 26 May 1987,

coll. ALB, DLS, DFL; (26) roadside ditch

on NC 130 (34.45406N, 278.97732W); 4 j

=, 1 R, 7 j R (4209), 16 Jul 1996, coll. J.

Alderman (JA), G. Motessi (GM), Kirk,

M. Savacool (MS); (27) Pretty Pond Bay S

of NC 20, 0.5 rd km E of jct SR 1924, 4.8

air km E of St Pauls (34.79417N,

278.92528W); 4 R (2245), 1 R with third

instar young (2243), 23 Feb 1985, coll.

DLS, RWL, DFL; 1 = I (2688), 26

Mar 1987, coll. ALB, DLS, DFL; (28)

Oak Savannah Bay II, along SR 1811,

0.5 rd km W SR 175 (34.80416N,

279.090228W); 3 = I, 1 R (2690), 8 May

1987, coll. ALB, H. M. Wilbur (HMW), J.

Fauth; (29) roadside ditch on NC 71w,

6.6 km S of Red Springs; 6 = I, 32 j =, 6 R,

34 j R (USNM 131779), 22 May 1971, coll.

HHH; (30) roadside ditch on NC 71,

8.8 km S of Wakulla; 3 = I, 2 = II, 1 j =,

14 R, 3 j R (USNM 131783), 22 May 1971,

coll. HHH; (31) Big Swamp at NC 211,

1.1 km W of Bladen Co. line; 1 = II

(USNM 132635), coll. P. D. Ross, Sr.,

HHH. Scotland Co.—(32) field pond on

Maxton-Laurinburg Air Base; 1 = I, 3 = II, 2 j =, 5 R, 1 j R (USNM 144791), 14

Oct 1973, coll. M. Odell; (33) borrow pit

along SR 1432 near jct SR 1400 & 1413,

5.6 air km NW of Wagram (34.9289N,

279.3949W); 1 = I, 1 j =, 2 j R (2011), 8

Oct 1974, coll. MRC, ALB, WMP, JEC; 1

R (494), 25 Jan 1978, coll. ALB, REA; 1 = I (2662), 18 Apr 1990, coll. ALB; 2 R (7812), 8 Jul 2001, coll. JCB, JTF, SJH;

(34) Muddy Crk on Sandhills Game Land

at SR 1328, 10.6 air km NNW of Silver

Hill (35.00333N, 279.53833W); 1 j R (4496), 23 Jul 1997, coll. GM, M. R.

Wood; (35) State Line Prairie Bay on The

Nature Conservancy property off S side

19

SR 1622 at jct SR 1625 & Marlboro Co.,

SC, line, 4.0 km SSW of Hasty; 1 = I, 1 = II, 1 R, 1 j R (4487), 28 Feb 1998, coll. JCB,

JTF, SJH.

Waccamaw River basin: Bladen Co.—

(36) roadside ditch on US 701, 5.5 km N

of Clarkton (Hobbs & Peters 1977:49);

(37) roadside ditch on SR 1730 & SR

1786, 0.8 km NE of jct NC 211

(34.42722N, 278.45639W); 2 j =, 1 j R (5144), 1 R & exuvium (5266), 24 Jun

1999, coll. WCS, DGC, R. T. Bryant et

al. Brunswick Co.—(38) Juniper Crk at

SR 1340, ca. 1.3 air km E of Makatoka,

16.8 air km NW of Supply (34.12505N,

278.39445W); 1 R (1390), 13 Feb 1984,

coll. F. C. Rohde; (39) along NC 211,

0.8 km SSE of Juniper (Driving?) Crk,

9.9 air km NNW of Supply (34.09889N,

278.30194W); 2 = I (1829), 2 Aug 1984,

coll. ALB, S. D. Smith, K. Green; (40)

‘‘9 mi NW of Supply’’; 1 j R (3757), 29

Aug 1975, coll. HMW; (41) roadside

drainage ditch on NC 211, ca. 11.4 km

N of Supply, probably near Juniper

Creek; 2 j = (4548), coll. E. E. Brown;

(42) on NC 211, 9.6 km N of US 17; 1 = II, 3 R (USNM 147810), 20 Feb 1976, coll.

J. R. Bailey (JRB). Columbus Co.—(43)

ditch on E side of NC 211, 12 8 km S of

Bolton; 1 R (USNM 147813), 21 Mar

1976, coll. JRB; (44) roadside borrow pit

along NC 211, ca. 1.8 rd km NW of crossing

Clear Branch (34.206N, 278.3899W); 4 = I, 2 = II, 1 j =, 4 R, 1 j R (24992), 1 = II (24993),

30 Apr 2005, coll. ALB, DLS; (45) roadside

borrow pit in mesic pine flatwoods along

NC 211, 1.9 rd km WNW of Brunswick Co.

line, 13.0 air km S of Bolton; 1 = I, 3 = II, 3 j

=, 6 R (3195), 14 May 1996, coll. ALB; (46)

on pavement of NC 211, 1.1 rd km S of

Bolton (34.30556N, 278/38695W), ca. 2045

hr EDT, wet night; 1 R (1951), 24 Mar 1975,

coll. MRC, ALB, WMP, JEC; (47) grassy

sloughs on NC 130, ca. 7.4 air km SE of Old

Dock; 5 = I, 6 = II, 6 j =, 5 R, 11 j R (2020), 1

= I, 1 = II, 1 j =, 1 R, 1 j R (uncat.), 24 Mar

1975, coll. MRC, ALB, WMP, JEC; (48)

1.0 km E of Hallsboro on US 76 (Hobbs &

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20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Peters 1977:54); (49) from stomach of red-

shouldered hawk from near Hallsboro; pair

of = I gonopods & 2 partial carapaces

(1277), 8 Apr 1983, coll. GSG; (50)

roadside ditch along SR 1117

(34.15803N, 278.75275W); 1 j =, 1 R, 5 j

R (4220), 6 Jul 1995, coll. JA, Tim Savidge

(TS), M. Hartman (MH), MS; (51)

roadside ditch on NC 130 (34.1297N,

278.58665W); 6 j =, 2 R, 14 j R (4225),

28 Sep 1995, coll. JA, GM, MS; (52) road­

side ditch on NC 904 (34.01067N,

278.60792W); 1 = II, 2 j =, 2 R, 3 j R (4208), 21 Sep 1995, coll. JA, GM, MS;

(53) water along railroad tracks, 1.0 km E

of Hallsboro; 1 = I, 1 R (4952), 16 Apr

1956, coll. HHH, E. T. Hall; (54) ‘‘Green

Swamp’’ (34.31472N, 278.44611W); 1 j = (2062), 23 Feb 1975, J. H. Gillespie, R.

Gordon; (55) roadside ditch on SR 1006

(34.13654N, 278.66315W); 1 j =, 5 j R (4221), coll. JA, TS, MH, MS; (56)

roadside ditch on SR 1157; 3 j R (4202),

6 Jul 1995, coll. JA, TS, MH, MS.

SOUTH CAROLINA. Lumber-Little

Pee Dee River basin: Dillon Co.—(57)

Little Reedy Crk at route 34, ca. 16 km

NW of center Dillon; 1 = (25118), 12 Mar

2001, coll. R. G. Arndt, E. W. Gaines, and

students. Marion Co.—(58) roadside ditch

on route 41A, 3.4 km SW of Dillon Co.

line, NE of Marion; 1 = I, 2 R (USNM

177310), 12 Apr 1981, coll. HHH; (59)

roadside ditch on route 41A, 5.0 km S of

US 501, Marion; 1 = I (USNM 177317),

12 Apr 1981, coll. HHH.

Waccamaw River basin: Horry Coun­

ty.—(60) swamp at S-32, 100 m SSE of

Waccamaw R; 1 = II, 2 j =, 3 j R (USNM

207820), 13 Nov 1982, coll. P. H. Carlson

(PHC); (61) roadside ditch on S-237,

0.16 km W of S-564 near Georgetown

Co. line; 5 = II, 2 j =, 5 R, 2 j R (USNM

207826), 14 Nov 1982, coll. PHC.

Size.—Form-I males ranged from 21.5– {

35.5 (X 5 27.4, n 5 33) mm TCL. The

largest specimen examined is the largest

form-I male, which is from the Lumber-

Little Pee Dee basin. The largest female

measured 32.8 mm TCL and is from the

same basin.

Life history and ecological notes.—

Form-I males were found from January

through May, and in August and Octo­

ber. Cooper (2002:178, 179) provided

data on six females carrying late-instar

young, collected at two sites in Robeson

County on 23 February 1985. One

additional female (2243), collected in

Pretty Pond Bay, Robeson Co., on the

same date, measures 32.1 mm TCL

(25.2 mm PCL) and has 22 third-instar

young measuring ca. 3.7 mm TCL.

Some of the ponds, bays and borrow

pits where P. (O.) pearsei occurs are

breeding sites of the ambystomatid sala­

manders, Ambystoma maculatum, A. opa­

cum, and A. tigrinum. The adult salaman­

ders are only seasonally present at the

sites, but their larvae, which may be quite

large and carnivorous, remain in the

water until metamorphosis occurs. Sever­

al of the sites contain anostracans of the

genera Streptocephalus (cf. S. seali) and

Eubranchipus, along with conchostracans

of the genera Lynceus (cf. L. gracilicor­

nis), Limnadia (cf. L. lenticularis), and

Eulimnadia.

Crayfish associates.—Procambarus (O.)

pearsei has been found with Procambarus

(Ortmannicus) acutus (Girard, 1852), Pro­

cambarus (Ortmannicus) ancylus Hobbs,

1958, Procambarus (Ortmannicus) blan­

dingii (Harlan, 1830), and Fallicambarus

(Creaserinus) fodiens (Cottle, 1863).

Affinities.—Morphologically, P. (O.)

pearsei is most similar to P. (O.) plumi­

manus and P. (O.) medialis, and the three

probably originated from the same com­

mon ancestor in the southeastern Coastal

Plain of North Carolina. It is clearly

distinguishable from both these species

by the strong caudolateral turn of the

mesial process on the gonopod of the

form-I male; the mesial processes of both

P. (O.) plumimanus and P. (O.) medialis

are strongly directed caudomesially. Also,

viewed in lateral or mesial aspect, the

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VOLUME 124, NUMBER 1

distal fourth of the shaft of the gonopod is

strongly curved caudally at about 90

degrees in P. (O.) pearsei but at about

45–50 degrees in the other two species. In

addition, females of P. (O.) pearsei can be

distinguished from those of P. (O.) plumi­

manus by the configuration of the annu­

lus. In the former species it is basically

spindle-shaped and has a long, broad

sulcus; in the latter it is more prominently

subovate and has a weak sulcus.

Etymology.—Creaser named the spe­

cies in honor of A. S. Pearse, a noted

aquatic biologist of the time at UMMZ,

who published a number of papers on

fishes, crayfishes, and other crustaceans,

and who collected all of Creaser’s type

specimens of P. pearsei.

Acknowledgments

My sincerest thanks go to those many

collectors who provided the specimens on

which this redescription is based; their

names appear in the section on range and

specimens examined. Special thanks go to

A. L. Braswell and D. L. Stephan, who

collected the form-II male and the live

specimens used for color notes, and to S.

A. Cooper, who provided technical assis­

tance at NCSM and assisted in prepara­

tion of the plate that appears as Figure 1.

Loan of the primary types from USNM

was arranged through the generosity of R.

Lemaitre and K. Reed. Two anonymous

reviewers provided helpful comments.

Literature Cited

Cooper, J. E. 2002. North Carolina crayfishes

(Decapoda: Cambaridae): notes on distribu­

tion, taxonomy, life history, and habitat.—

Journal of the North Carolina Academy of

Science 118(3):167–180.

———, & A. L. Braswell. 1995. Observations on

North Carolina crayfishes (Decapoda: Cam­

baridae).—Brimleyana 22:87–132.

Cottle, T. J. 1863. On the two species of Astacus

found in upper Canada.—Canadian Journal

of Industry, Science, and Art, new series

45:216–219.

21

Creaser, E. P. 1934. A new crayfish from North

Carolina.—Occasional Papers of the Museum

of Zoology, University of Michigan 285:1–4.

Fitzpatrick, J. F., Jr. 1983. How to know the

freshwater Crustacea. William C. Brown

Company Publishers, Dubuque, Iowa, 227

pp.

Girard, C. 1852. A revision of the North American

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geographical distribution.—Proceedings of

the Academy of Natural Sciences of Phila­

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Harlan, R. 1830. Description of a new species of the

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crayfishes of the subfamily Cambarinae

(Decapoda, Astacidae) with the description

of a new genus and species.—American

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Hobbs, H. H., Jr 1954. A redescription of Procam­

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———. 1968. Crustacea: Malacostraca. Pp. K1–

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———. 1974. A checklist of the North and Middle

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Smithsonian Contributions to Zoology 201:

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———. 1981. The crayfishes of Georgia.—Smithso­

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———. 1989. An illustrated checklist of the Amer­

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Associate Editor: Christopher B. Boyko.


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