A New Subgenus and

Two New Species of Crayfishes of

the Genus Cambarus

(Decapoda: Cambaridae) from the

Southeastern United States

RAYMOND W. BOUCHARD

and

HORTON H. HOBBS, JR.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 224
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 22 4

A New Subgenus and

Two New Species of Crayfishes of

the Genus Cambarus

(Decapoda: Cambaridae) from the

Southeastern United States

Raymond W. Bouchard

and Horton H. Hobbs, Jr.

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976
ABSTRAC T

Bouchard, Raymond W., and Horton H. Hobbs, Jr. A New Subgenus and Two

New Species of Crayfishes of the Genus Cambarus (Decapoda: Cambaridae)

from the Southeastern United States. Smithsonian Contributions to Zoology,

number 224, 15 pages, 3 figures, 1976.—A new subgenus Exilicambarus is proposed to receive a single species, Cambarus (Exilicambarus) cracens, described

from southeastern tributaries of Guntersville Lake (impounded Tennessee

River) in DeKalb and Marshall counties, Alabama. A second previously undescribed species, Cambarus (Jugicambariis) nodosus, is a burrowing crayfish closely

allied to Cambarus (/.) carolinus (Erichson), and is reported from the Hiwassee,

Savannah, and Chattahoochee drainage systems in Tennessee, North Carolina,

and Georgia.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6163. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Calaloging in Publication Data

Bouchard, Raymond VV.

A new subgenus and two new species of crayfishes of the genus Cambarus (Decapoda, Cambaridae) from the southeastern United States.

(Smithsonian contributions to zoology ; no. 224)

Bibliography: p.

Supt. of Docs, no.: SI 1.27:224

1. Cambarus cracens. 2. Cambarus nodosus. 3. Cambarus. 4. Crustacea—Southern States.

I. Hobbs, Horton Holcombe, 1914- II. Title: A new subgenus and two new species of crayfishes of the genus Cambarus ... . III. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 224.

QL1.S54 no. 224 [QL444.M33] 591'.08s [595'.3841] 75-619424
A New Subgenus and

Two New Species of Crayfishes of

the Genus Cambarus

(Decapoda: Cambaridae) from the

Southeastern United States

Raymond W, Bouchard

and Horton H. Hobbs, Jr.

Introduction

The recognition of 10 subgenera within the

genus Cambarus (Hobbs, 1969) has received general acceptance among crayfish systematists. Only

the monotypic subgenus Barbicambarus has been

elevated to generic rank (Bouchard, 1972), suggesting a greater divergence from a common ancestry

in it than in any other member or assemblage

within the genus. Cambarus (Exilicambarus) cracens, described herein, provides evidence of a

previously unknown evolutionary line within the

genus; nevertheless, it bears morphological resemblances to members of three of the currently recognized subgenera. Certain characteristics of the

areola, rostrum, cervical spines, and chela are similar to those in members of the subgenus Puncticambarus. Other characters of the chela and of the

first form male gonopod are suggestive of crayfishes assigned to the subgenus Jugicambarus,

while in some respects the chela and areola exhibit

features reminiscent of those occurring in the

monotypic subgenus Veticambarus. Details of

Raymond W. Bouchard and Horton H. Hobbs, Jr., Department of Invertebrate Zoology, National Museum of Natural

History, Smithsonian Institution, Washington, D.C. 20560.

these characters are more completely discussed in

the section on "Relationships." The array of

unique and shared characters of this new species

provides the rationale prompting the proposal

of the recognition of the new subgenus Exilicambarus.

The second species described herein represents

one of a growing number of burrowing crayfishes

to be reported. The difficulty in obtaining specimens and the variation exhibited by numerous

populations of burrowers compound the problem

of their recognition as distinct taxa. Specimens of

this species have been available since 1958, but its

close kinship to another poorly known species,

C. carolinus (Erichson, 1846), suggested that it

was prudent not to describe it until additional

specimens from other localities became available.

A previous study by us (Hobbs and Bouchard,

1973) presented a better understanding of C.

carolinus and C. dubius, which was previously

synonymized with the former. Improved concepts

of the latter two species and the accumulation of

additional data on the range and variations of the

undescribed species encourage a description of

the latter.

ACKNOWLEDGMENTS.—We wish to express our

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gratitude to the many persons who either contributed specimens or assisted us in the field: J. W.

Bouchard, T. A. English, Jr., P. L. Holcomb, B. E.

Oakberg, F. L. Oakberg, F. K. Parrish, D. J. Peters,

J. E. Pugh, K. W. Simonds, T. Unestam, and C. H.

Wharton. Our appreciation is also extended to

Judith W. Bouchard and Fenner A. Chace, Jr., for

reviewing the manuscript and to Carolyn B. Gast

for the rendition of Figures 2 and 3.

Exilicambarus, new subgenus

DIAGNOSIS.—Eyes pigmented and of moderate

size. Rostrum with margins slightly thickened and

almost always bearing marginal tubercles or spines.

Postorbital ridge with spine or tubercle. Cervical

spine well developed, often flanked dorsally by

secondary small acute tubercle. Suborbital angle

rounded or obsolete. Hepatic spines absent, branchiostegal spine well developed or vestigial.

Areola broad (about 3 or 4 times as long as wide),

constituting 31.1 to 34.8 percent of entire length

of carapace (41.5 to 44.6 percent of postorbital

carapace length) with 6 to 10 punctations across

narrowest part. Antennal scale 2 to 2.5 times as

long as broad, broadest at or distal to midlength.

Chela comparatively slender with mesial surface

of elongate palm bearing 2, usually irregular, rows

of tubercles (mesialmost consisting of 7 to 9); at

least mesial half of dorsal surface, as well as ventral surface of palm, studded with squamous tubercles; lateral margin of fixed finger costate. Hook

on ischium of third pereiopod of male overreaching basioischial articulation, not opposed by tubercle on basis. First pleopod of first form male with

central projection corneous, bladelike, somewhat

tapering from base, bearing distinct subapical

notch, bent caudoproximally at angle of about

135 degrees (distal half almost straight), and, if

straightened, constituting approximately one-fifth

total length of appendage; mesial process swollen,

tapering distally, directed at 90 to 110 degrees to

main shaft of appendage, and usually bearing 1

or 2 acute apical prominences; caudal element

represented by small rounded lobe at caudolateral base of central projection. Annulus ventralis

shallowly embedded in sternum, subspindle

shaped. First pleopod reaching or almost reaching midlength of annulus when abdomen flexed.

TYPE-SPECIES.—Cambarus (Exilicambarus) cracens, here described.

LIST OF SPECIES.—Monotypic.

ETYMOLOGY.—Exilis (L., = slender) in combination with Cambarus; in reference to the comparatively elongate, slender, chela.

Cambarus (Exilicambarus) cracens, new species

FIGURES 1, 2

DIAGNOSIS.—Identical to diagnosis of subgenus.

HOLOTYPE MALE, FORM I.—Cephalothorax

(Figure \e, see also Figure la, j) subcylindrical

in cross section. Abdomen narrower than thorax

(18.4 and 19.9 mm). Greatest width of carapace

greater than height at caudodorsal margin of cervical groove (17.6 and 14.8 mm). Areola 3.4

times as long as broad with 6 widely spaced punctations across narrowest part; length of areola

31.9 percent of entire length of carapace (41.9 percent of postorbital carapace length). Rostrum

with slightly thickened, convergent margins bearing prominent acute corneous tubercles at base of

acumen; latter almost reaching distal end of

ultimate podomere of antennular peduncle and

terminating in corneous, subspiculiform tip, acumen slightly upturned apically; dorsal surface of

rostrum concave and bearing small setiferous punctations, latter more numerous in depression at

level of caudal margin of orbit. Subrostral ridge

weak and evident in dorsal aspect almost to base

of marginal tubercle of rostrum. Postorbital ridge

moderately strong with prominent knob at caudal

extremity, grooved dorsolaterally, and with spiniform corneous tubercle cephalically. Suborbital

angle broadly rounded, virtually nonexistent;

branchiostegal spine moderately large, acute, and

curved cephalodorsally. Cervical spine well developed and flanked by several small tubercles, at

least one of which acute; hepatic area with scattered tubercles; cephaloventral part of branchiostegite with crowded tubercles and few adjacent

and caudoventral to cervical spine, remaining lateral portion of branchiostegite granulate and dorsolateral portion finely, but densely, punctate.

Abdomen (Figure la) shorter than carapace

(35.0 and 35.4 mm); pleura of moderate length,

broad with caudoventral extremity of second

through fifth rounded. Cephalic section of telson
NUMBER 224

FIGURE 1.—Catnbarus (Exilicambarus) cracens, new species (holotypic male, form I): a, lateral

view of abdomen; b, dorsal view of caudal extremity of abdomen; c, mesial view of first pleopod;

d, lateral view of first pleopod; e, dorsal view of carapace; /, dorsal view of distal podomeres

of cheliped.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figure 16) with 1 spine in caudodextral corner

and 2 in caudosinistral, more mesial one movable.

Proximal podomere of uropod with small distal

spine on lateral lobe and larger one on mesial

lobe; both rami with distinct median keel, that on

mesial ramus with well-developed premarginal

spine; lateral margin of lateral ramus subserrate

with fixed and movable spines at extremity of proximal section; corresponding margin of mesial ramus with spine at distolateral angle.

Cephalic lobe of epistome (see Figure 2t of paratypic male, form I) apparently injured, rounded

but with asymmetrical emargination cephalodextrally, margins only slightly thickened and marginal areas somewhat elevated, cephalomedian

projection rather weak; main body of epistome

with fovea represented by shallow depression not

nearly so distinct as that illustrated, and arched

epistomal zygoma flanked cephalolaterally by

usual pair of slitlike fossae. Proximal segment of

antennule with prominent spine on ventral surface at base of distal two-fifths. Antennal peduncle

with lateral spine on basis and distal spine on

ischium, flagellum extending caudally to third abdominal tergum. Antennal scale (see Figure 2g

of paratypic male) rather broad, broadest distal

to midlength, distomesial portion of lamellar area

evenly rounded; distolateral spine strong, with

long corneous tip, and reaching slightly beyond

tip of rostrum. Postaxial surface of third maxilliped with stiff setae studding mesial half, row of

submarginal shorter setae laterally, and with

scattered setiferous punctations between submarginal row and clusters of stiff setae; distolateral

extremity angular but not produced in spiniform

projection.

Right chela (Figure If; see also Figure 2/) 3.7

times as long as broad (28.1 and 10.7 mm), subovate, slightly depressed; mesial margin of palm

with irregular row of 8 tubercles, 2 immediately

ventral to row and irregular row of 8 immediately

dorsolaterally; remainder of palm studded with

squamous tubercles, although few present ventromesially; distoventral surface of palm with usual

prominent tubercle at base of dactyl and several

somewhat larger than neighboring ones extending

proximomesially from submedian ridge on fixed

finger. Lateral surface of propodus costate with

row of punctations causing irregular appearance

of proximolateral margin on fixed finger and distolateral part of palm. Both fingers with well-defined

submedian ridges flanked by setiferous punctations dorsally and ventrally; opposable surface of

fixed finger with row of 5 rounded tubercles,

fourth from base largest, along proximal two-fifths

of fingers, and broad band of minute denticles

(between, ventral to, and beyond tubercles) reaching base of corneous tip of finger; usual large,

single, more ventral tubercle lacking. Opposable

margin of dactyl with row of 5 tubercles along

proximal three-fifths; proximal 3 tubercles subequal in size and much larger than distal 2, distalmost somewhat removed from more proximal 4,

and additional large tubercle present more ventrally between distal 2 tubercles of row; minute

denticles dispersed very similarly to those on fixed

finger; lateral surface of dactyl tuberculate along

proximal half and punctate distally.

Carpus longer than broad (10.3 and 6.9 mm)

with moderately deep oblique furrow dorsally;

mesial surface with prominent procurved spine

near midlength and very small tubercle proximally (carpus of left chela with two procurved

spines); distoventral margin with strong submedian spiniform tubercle, another on ventrolateral

articular knob, and 2 smaller tubercles proximomesial to submedian one; podomere otherwise

sparsely punctate.

Dorsal surface of merus with irregular row of

small tubercles and 2 subdistal spines; mesial and

lateral surfaces with scattered punctations; ventral

surface with mesial row of 10 (8 on left cheliped),

mostly spiniform, tubercles, and 2 (4 on left)

strong ones representing lateral row; also spine

present on ventral margin of distolateral articular

knob. Ischium with row of 5 small tubercles ventromesially.

Hook on ischium of third pereiopod only (see

Figure 2h of paratypic male, form I); hook simple,

overreaching basioischial articulation and not opposed by tubercle on basis. Coxa of fourth pereiopod with moderately prominent caudomesial boss,

that of fifth well developed but much less conspicuous than that on fourth. (See "Measurements.")

First pleopods (Figure \c,d; see also Figure 1b,

f) reaching coxae of third pereiopods when abdomen flexed. See "Diagnosis" for description.

(The almost straight distal half of the pleopod

directed at about 135 degrees to the main shaft
NUMBER 224

of the appendage is a unique feature of this

crayfish.)

ALLOTYPIC FEMALE.—Excluding secondary sexual features, differing from holotype in following

respects: epistome symmetrical, more nearly subtriangular and fovea slightly deeper; cephalic section of telson with 2 spines in each caudolateral

corner; more lateral row of tubercles on dorsomesial surface of palm of chela consisting of 6 or

7 tubercles; ventral surface of palm with fewer

and smaller tubercles, row extending proximomesially from submedian ridge on fixed finger in

holotype not recognizable; opposable margin of

fixed finger with row of 10 tubercles, that of dactyl

with 9 on right and 11 on left chela; carpus of

both chelipeds with 2 procurved tubercles mesially

but lacking small tubercle on proximomesial surface; only 1 small tubercle proximomesial to median ventromarginal tubercle; merus with 3 subdistal acute tubercles dorsally on left cheliped,

ventral surface with 10 tubercles in mesial row and

4 in lateral, right cheliped with 12 and 3 respectively; ischium with 4 tubercles on right and 3 on

left cheliped. (See "Measurements.")

Annulus ventralis (Figure 2k) distinctly asymmetrical and cephalic region not nearly so strongly

calcified as caudal; cephalic half with shallow, submedian, longitudinal trough; sinus originating at

sinistrocaudal end of latter and extending sinistrally, making hairpin turn, returning almost to

median line before forming broad arc over caudal

wall, there crossing median line and ending very

slightly dextral to it; tongue directed sinistrally

into fossa. Postannular sclerite 2.5 times as broad

as long and half as wide as annulus. First pleopod

almost reaching midlength of annulus when abdomen flexed.

MORPHOTYPIC MALE, FORM II.—Differing from

holotype in following respects: epistome resembling that of allotype; tubercles flanking cervical

spine on holotype and allotype not evident; cephalic section of telson with 2 spines in each caudolateral corner; opposable margin of fixed finger

of right cheliped with 3 (4 on left) tubercles

(third from base largest) along proximal third of

finger, and large, more ventrally placed tubercle

at base of distal two-fifths (absent on left); opposable margin of dactyl of right chela with 3 (5 on

left) tubercles along proximal fourth and more

ventrally placed one (absent on left) at end of

proximal third; mesial surface of carpus of left

cheliped with 2 procurved spines and smaller

proximal tubercle; 3 subdistal spines dorsally on

merus of right cheliped, ventral surface with mesial

row of 9 and lateral one of 3 tubercles on both

chelipeds; ischia of both with row of 3 tubercles;

hook on ischium of third pereiopod much reduced,

not overreaching basioischial articulation; bosses

on coxae of fourth and fifth pereiopods also

greatly reduced. (See "Measurements".) First

pleopods (Figure 2c,e) with mesial process lacking

acute extension; central projection noncorneous,

more massive, not so strongly recurved, with subacute apex directed caudally at approximately 90

degrees; only trace of caudal element evident.

COLOR NOTES.—Carapace olive tan with irregular cream tan mottlings dorsally and dorsolaterally,

fading ventrally; rostral margins, suffused with

red, flanking olive brown dorsum; hepatic region

largely mustard yellow with prominent olive

brown spot. Area over and between attachments

of mandibular muscles dark olive, and median

part of cervical groove flanked cephalically by

broad dark brown band paralleling contour of

groove. First through sixth abdominal terga olive

tan with paired, pale to dark brown markings

dorsolaterally (larger, although less intensely

colored on first and second segments); terga with

or without narrow, almost linear, scarlet band on

caudal margin; if present, band continuing ventrally along cephalic and cephaloventral margins

of pleuron of next segment, becoming reddish

brown on pleuron and fading before reaching caudoventral extremity; first abdominal pleuron

cream to white, succeeding ones with conspicuous

similarly colored spot; bases of second through

fifth pleura with oblique reddish brown bar subparallel to ventral margin. Margins of telson, as

well as keels, lateral margins, and proximal borders of transverse articulations of uropods reddish

brown. Remaining pereiopods largely cream to

white with olive suffusion dorsally on distal part

of merus, carpus, and proximal part of propodus.

Peduncle of antennule, antenna, and antennal

scale brown mottled with olive cream; flagella

olive brown. Dorsal surface of cheliped distal to

midlength of merus olive tan, fingers fading to

base of distal third where becoming vermilion to

scarlet; tubercles and articular knobs olive cream

to almost white.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Cambarus (Exilicambarus) cracens, new species (all depicting paratypic male, form I,

except c and e of morphotypic male, form II, and k of allotype): a, lateral view of carapace;

bjc, mesial view of first pleopod; d, caudal view of first pleopods; e,f, lateral view of first

pleopod; g, antennal scale; h, basal podomeres of third, fourth, and fifth pereiopods; i, epistome;

/, dorsal view of carapace; k, annulus ventralis; I, dorsal view of distal podomeres of cheliped.
NUMBER 224

MEASUREMENTS

Characters

Carapace:

Total length

of carapace

Postorbital

carapace length

Width

Height

Areola:

Width

Length

Rostrum:

Width

Length

Chela:

Length, niesial

margin of palm

Width, palm

Length, lateral

margin

Length, dactyl

Abdomen:

Width

Length

(in mm).—

Holotype

35.4

27.0

17.6

13.6

3.3

11.3

4.9

9.7

10.4

10.7

28.1

15.5

14.8

35.0

Allotype

39.7

30.7

19.9

17.1

4.1

13.4

6.0

11.1

10.3

10.9

28.1

15.6

18.4

425

Morpho

24.8

19.2

12.0

10.0

2.3

8.1

4.0

7.2

52

6.5

15.1

9.0

10.9

27.0

TYPE-LOCALITY.—Short Creek at State Route

75, 1.1 miles southwest of the junction with State

Route 68 (T.8S, R.4E, Sec. 36), Marshall County,

Alabama. There the clear stream, some 35 to 40

feet wide and mostly less than one foot deep, flows

rather sluggishly over a bed-rock and sandy bottom littered with large rocks. Shading the shore

line are Platanus occidentalis, Liriodendron tulipifera, Quercus sp., and Pinus sp. Only two other

crayfishes, Cambarus (Depressicambarus) striatus

Hay (1902:437) and C. (Jugicambarus) distans

Rhoades (1944:136), share this locality with C.

(E.) cracens.

DISPOSITION OF TYPES.—The holotypic male,

form I (no. 146082), the allotype (no. 146083),

and the morphotypic male, form II (no. 146084),

are deposited in the National Museum of Natural

History (NMNH), Smithsonian Institution. Paratypes consisting of 10 a I, 1 $ II, 9 $, 28j $ , and

19j2 are also in NMNH, and 9^1, 1 $ II, 2$ ,

2j $ , and 6j $ are in the collection of the senior

author.

RANGE AND SPECIMENS EXAMINED.—This crayfish is known only from the localities cited here,

all in the southeastern tributaries of Guntersville

Lake (impounded Tennessee River) in DeKalb

and Marshall counties, Alabama.

DEKALB COUNTY: Scarham Creek at State Route 20, 1 mile

west of Skirum (T.2S, R.6E, Sec. 23), 1$I, l^II, 1$ ,

1V/29/70, P. L. Holcomb and H. H. H., Jr., coll.; 2^1, 39 ,

9j<5» 7j9, XI/11/73, T. A. English, Jr., and H. H. H., Jr.,

coll.; 8 $ I, 1 $ II, 2j $, 2j $, IH/26/74, J. W. Bouchard and

R. W. B., coll. Scarham Creek at State Route 277, 1.4 miles

north of Crossville (T.8S, R.6E, Sec. 27), A$l, 2$ , 10j£,

7j $, XI/12/73, T. A. E., Jr. and H. H. H., Jr., coll. Bengis

Creek at County Road 116 (T.5S, R.8E, Sec. 25), l^II

(morphotype), X/31/74, J. W. B. and R. W. B., coll. MARSHALL COUNTY: Type-locality, 4£I, 4$ , 9j $, 5j 9 , XI/12/73,

T. A. E., Jr., and H. H. H., Jr., coll. Shoal Creek at unpaved

road off County Road 60 (T.8S, R.5E, Sec. 4), 1$I, 29 ,

4j 9, VIII/21/73, J. W. B. and R. W. B., coll.

VARIATIONS.—Among the variations noted, only

one seems to be restricted to some degree to a part

of the range. The spination of the carapace (marginal spines of the rostrum, postorbital tubercles

or spines, and cervical spines) is more pronounced

in most of the specimens from the type-locality

than from elsewhere. In many of the crayfish from

the other localities, the marginal spines on the

rostrum are reduced to small tubercles and in

one or two are reduced to the extent that while

the rostral margins at the base of the acumen are

angular they are not produced in spines or tubercles. The postorbital tubercles are frequently

markedly reduced or they may be well developed

and spiniform; the cervical spines, always present,

may be long and very sharp, or somewhat tuberculiform, although always bearing an acute tip.

Frequently, one (sometimes two) spiniform tubercle flanking the cervical spine dorsally is somewhat

larger than others in the vicinity. The chelipeds

are rather uniform in general contour, and the

numbers of tubercles pointed out on the cheliped

vary little from the ranges indicated in the descriptions of the three specimens above. The first

pleopod of the male, form I, exhibits few variations, the only conspicuous one being in the mesial

process—in two specimens the distal extremity is

rounded instead of possessing a short, apical,

spiniform process on the margin adjacent to the

central projection. The concavity of that margin,

illustrated in Figure 2b and /, is present in only

one other specimen, and both are probable artifacts of preservation.

SIZE.—The largest specimen available is a female having a carapace length of 39.7 mm (postorbital carapace length 30.7 mm). The corresponding measurements of the largest first form
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

male are 37.3 and 28.6 mm, and of the smallest

first form male, 24.7 and 18.7 mm.

LIFE HISTORY NOTES.—First form males have

been collected in March, August, and November,

and the exoskeletons of most are encrusted. No

large second form males are available, so there is

no direct evidence as to the probable molting seasons in the adult male population. Neither ovigerous females nor those carrying young have been

found, although well-developed cement glands

were observed on two females collected in November. Judging by the sizes of the limited number of

specimens at hand, three generations are present

in the populations sampled. The smallest individual is a juvenile male, collected on 11 November

1974, having a carapace length of 12.2 mm.

ECOLOGICAL NOTES.—All of our collections were

made in rock-littered areas of the streams. The

first lot was obtained from Scarham Creek in April

1970 when the stream was at flood stage, some four

feet above usual water level, and so swift that one

could not stand in it except in lee and shallow

marginal areas.

In November 1973, the stream was quite sluggish. At that season, the rock-littered areas were

so choked with leaves that it is possible the crayfish were among the latter rather than having

sought cover beneath the rocks. All of the specimens collected by the senior author were taken

from under rocks. In none of the localities was the

water swift except during the period of flood mentioned above.

RELATIONSHIPS.—Our assessment of the affinities

of this crayfish is to some degree based on negative

conclusions. Most of the characteristics of the

carapace, except the comparatively small number

of punctations in the areola in most of the specimens, are those usually associated with members

of the subgenus Puncticambarus. The rather long,

strongly recurved, central projection of the first

pleopod of the first form male, the few punctations in the areola of most of the specimens, and

the slightly downcurved fingers of the cheliped are

suggestive of crayfishes comprising the subgenus

Jugicambarus. The elongate chela resembles that

of the monotypic subgenus Veticambarus, and to

a lesser degree that of C. (Puncticambarus) reburrus Prins (1968:458). In few species of the genus

is the palm of the chela so nearly completely covered with tubercles; only in some members of the

subgenus Depressicambarus is such a tuberculate

condition approximated.

The tapering central projection of the first pleopod of the first form male, although strongly recurved, is unique in that the morphologically cephalodistal margin is almost straight.

For the following reasons, it does not seem appropriate to assign this crayfish to any of these

subgenera. The usually limited number of punctations in the areola, the strongly recurved central

projection of the first pleopod of the male, and

the absence of a tubercle on the basis of the third

pereiopod opposing the hook on the ischium constitute a combination of features that does not

occur elsewhere in members of the subgenus Puncticambarus. The long rostrum with marginal tubercles or spines, the strongly developed cervical

spine, and the elongate chela studded with tubercles, distinguish it from all known epigean

members of the subgenus Jugicambarus. From the

monotypic Cambarus (Veticambarus) pristinus

Hobbs (1965:268) it differs in possessing marginal

spines on the rostrum, lacking an acute suborbital

angle, a tubercle on the basis of the third pereiopod that opposes the hook on the ischium, and a

cephalic excavation on the boss on the coxa of the

fourth pereiopod. In addition, the following differences exist in the first pleopod of the first form

male: the central projection is more strongly recurved; the mesial process is bent caudally at at

least 90 degrees to the main shaft of appendage;

and the caudal process is not nearly so well developed. The shape and armature of the chela

and the nature of the first pleopod of the first

form male isolate this crayfish from members of

the subgenus Depressicambarus. Accordingly, it

has been assigned to the new monotypic subgenus

Exilicambarus.

ETYMOLOGY.—The name cracens (L., = slender)

was chosen because of the elongate, comparatively

slender chela.

Cambarus (Jugicambarus) nodosus, new species

FIGURE 3a-/

Cambarus carolinus.—Holt, 1968:32.

DIAGNOSIS.—Body and eyes with pigment, latter

small. Rostrum without marginal spines or tubercles. Areola 5.0 to 13.6 times as long as broad and
NUMBER 224

constituting 33.9 to 41.4 percent of total length

of carapace (38.2 to 46.5 percent of postorbital

carapace length) with 2 to 4 punctations across

narrowest part. Cervical spine lacking, although

small rounded tubercle present. Hepatic spines

absent; branchiostegal tubercle small and rounded;

suborbital angle moderately to well developed.

Postorbital ridge rather well developed without

cephalic tubercle. Antennal scale longer than

broad, broadest slightly distal to midlength. Chela

with 2 rows of tubercles along mesial margin of

palm, mesialmost subserrate, secondary row uneven; scattered tubercles over dorsomesial half of

palm; well-defined longitudinal ridges on dorsal

surface of fingers, less obvious ones on ventral side;

lateral surface of fixed finger weakly costate. Hook

on ischium of third pereiopod of first form male

overreaching basioischial articulation and not o p

posed by tubercle on basis. First pleopod of first

form male with central projection corneous, bladelike, tapering, bearing distinct, small subapical

notch, and recurved at angle of approximately 90

degrees to shaft of appendage; mesial process tumescent, tapering to acute tip and directed caudally at angle of approximately 90 degrees. Both

elements of first pleopod appreciably longer than

cephalocaudal diameter of shaft of appendage. Annulus ventralis asymmetrical, subcylindrical and

movable; cephalic half bearing longitudinal median trough between anteriorly converging ridges,

and caudal half with sinuate dextral or sinistral

sinus. Caudal wall of annulus elevated. First pleopod of female uniramous and reaching at least

midlength of annulus when abdomen flexed.

HOLOTYPIC MALE, FORM I.—Body subovate, depressed (Figure 3a,;). Abdomen narrower than

thorax (12.9 and 14.9 mm). Greatest width of carapace greater than height at caudodorsal margin of

cervical groove (14.9 and 12.0 mm). Areola 7.1

times as long as wide with 3 punctations across

narrowest part; length of areola 37 percent of entire length of carapace (42.3 percent of postorbital carapace length). Rostrum with thickened,

slightly concave, subparallel margins devoid of

marginal spines or tubercles. Acumen set off from

basal portion of rostrum, with concave, oblique,

slender margins, and terminating in small u p

turned tubercle. Rostrum excavate dorsally, especially deep anteriorly, with submarginal punctations and few scattered ones basally. Ventrolateral

margin of rostrum with long, thick setae except

at apex of acumen. Postorbital ridges moderately

well developed, shallowly grooved dorsolaterally

and lacking cephalic tubercle. Suborbital angle

moderately strong, lacking corneous tip; branchiostegal spine represented by small tubercle. Cervical spine absent although represented by very

small rounded tubercle. Hepatic area and lateral

portion of branchiostegite tuberculate; dorsal surface of carapace punctate.

Abdomen longer than carapace (29.3 and 28.9

mm); pleura of moderate length with caudoventral extremities subangular. Cephalic section of

telson with marginal spine and submarginal movable spine in each caudolateral corner and partially delimited from caudal section by paired

oblique incisions. Basal podomere of uropod with

spine extending over mesial ramus; lateral ramus

with median and submedian ridges, former terminating in acute spine at transverse flexure. Proximal portion of lateral ramus with row of small

spines distally and large, submarginal movable

spine at caudolateral corner. Mesial ramus of uropod with median ridge terminating distally in premarginal acute spine; lateral margin with distolateral acute spine. Dorsal surface of telson and

uropod lightly setiferous and fringed caudally.

Cephalic lobe of epistome (Figure 3g) rounded

with small cephalomedian projection and slightly

irregular, thickened cephalolateral margins. Main

body of epistome with very shallow median fovea

and pair of obliquely disposed slitlike fossae immediately cephalic and subparallel to thickened,

slightly arched epistomal zygoma; lateral extremities with several small tubercles. Proximal segment of antennule with small spine on ventral surface near midlength. Antennae broken. Antennal

scale (Figure Si) broadly declivous, broadest distal

to midlength; thickened lateral portion terminating in prominent, acute, corneous-tipped spine

projecting anteriorly beyond tip of rostrum.

Right chela (Figure 3/) about 2 times as long as

broad (21.9 and 11.0 mm), inflated, especially

proximolaterally; mesial margin of palm with

primary subserrate row of 7 tubercles and uneven

secondary row of slightly smaller tubercles dorsolateral to primary row; scattered tubercles over

dorsomesial half of palm; distoventral surface of

palm with 2 large, swollen tubercles at base of

dactyl, mesial one with small, corneous, rounded
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Cambarus (Jugicambarus) nodosus, new species (all depicting holotypic male,

form I, except b and / of paratypic male, form I (Union County, Georgia), c and e of morphotypic male, form II, and k of allotype): a, lateral view of carapace; b,c, mesial view of first

pleopod; d, caudal view of first pleopods; e,f, lateral view of first pleopod; g, epistome; h, basal

podomeres of third, fourth, and fifth pereiopods; i, antennal scale; /, dorsal view of carapace;

k, annulus ventralis; I, dorsal view of distal podomeres of cheliped.
NUMBER 224 11

tip. Lateral surface of propodus weakly costate

with row of punctations rendering proximolateral

base slightly impressed dorsally, less so ventrally;

dorsal and ventral surfaces of fixed finger with

distinct submedian ridges flanked by setiferous

punctations; opposable surface with row of 5

tubercles along proximal half, third from base

largest; distal tubercle moderately acute, others

knoblike; additional acute tubercle present on

lower level near base of distal fourth; opposable

surface also with row of minute denticles extending distally from fourth tubercle from base, row

interrupted only by fifth. Dorsal and ventral surfaces of dactyl with median longitudinal ridges

flanked by setiferous punctations; opposable margin with row of 7 tubercles situated in slight excavation, proximal 3 rounded (basalmost largest),

followed by 2 moderately acute and 2 acute distal

ones; row of denticles, broken by tubercles, extending from third tubercle to distal end of finger.

Fingers terminating in large, acute, corneous

spines.

Carpus longer than broad (9.2 and 6.0 mm)

with deep oblique furrow dorsally; mesial surface

with large spine near midlength and small tubercle near proximal base of latter; 3 rounded tubercles on dorsomesial surface; distoventral margin

with 3 rounded tubercles; podomere otherwise

punctate.

Upper surface of merus with 3 rounded subdistal tubercles and 11 along crest of podomere; ventral surface with lateral row of 5 subacute spines,

and mesial row of 11 subacute ones, both rows decreasing in size proximally. Ischium with row of

3 small tubercles on ventromesial margin.

Hook on ischium of third pereiopod only (Figure Sh); hook simple, overreaching basioischial

articulation and not opposed by tubercle on basis.

Coxa of fourth pereiopod with prominent caudomesial boss, that of fifth without prominence. (See

"Measurements".)

First pleopods (Figure Sb,d,f) reaching caudal

portion of coxae of third pereiopods when abdomen flexed. (See "Diagnosis" for description.)

ALLOTYPIC FEMALE.—Differing from holotype in

following respects: areola constituting 37.7 percent

of entire length of carapace (43.1 percent of postorbital carapace length) and 5.8 times as long as

broad. Cephalic lobe of epistome triangular. Palm

of right chela with primary row of 8 tubercles.

Mesial surface of carpus with horizontal row of 3

tubercles proximal to large spine; dorsomesial surface with 3 very small tubercles (6 on left cheliped); distoventral surface with cluster of 5 tubercles. Upper surface of merus of right cheliped

with 2 subdistal rounded tubercles and 10 (8

on left) along crest of podomere; ventrolateral

margin with row of 7 spines on both chelipeds;

ventral surface with more acute spines. (See

"Measurements".)

Sternum between third and fourth pereiopods

broadly V-shaped and moderately deep. Annulus

ventralis (Figure 5k) with cephalic half bearing

median longitudinal trough flanked by ridges converging anteriad; caudal half with sinistral sinus.

Caudal wall of annulus elevated, especially caudolaterally. First pleopod uniramous and reaching

at least midlength of annulus when abdomen

flexed.

MORPHOTYPIC MALE, FORM II.—Differing from

holotype in following respects: rostrum less excavate anteriorly; ventral margin with setae sparse

on sides of rostrum and lacking on most of acumen; cephalic tubercle more spiniform. Areola

constituting 37.9 percent of entire length of carapace (43.8 percent of postorbital carapace length)

and 7.1 times as long as broad with 2 punctations

across narrowest part. Palm on left chela with

primary row of 6 tubercles. Mesial surface of carpus of both left and right chelipeds with horizontal row of 3 tubercles proximal to large spine;

dorsomesial surface of both with 2 tubercles. Upper

surface of merus of right cheliped with 2 rounded

subdistal tubercles and 11 along crest of both right

and left, latter with additional laterally disposed

tubercle; ventrolateral margin of merus of left

cheliped with row of 6 spines, and ventromesial

margin of left and right with row of 10 and 9, respectively. Caudolateral corners of epistome especially elevated. Propodus of chela with third and

fourth tubercles from proximal end more acute.

Hook on ischium of third pereiopod much reduced, not reaching basioischial articulation. Boss

on coxa of fourth pereiopod somewhat smaller

and less sharply defined. (See "Measurements".)

First pleopods (Figure Sc,e) of uniform texture,

neither element corneous, central projection

heavier, rounded caudally, and recurved at angle

of approximately 90 degrees to shaft of appendage; mesial process tumescent, terminating ce-
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

phalically in acute tip and directed caudally at

angle of approximately 90 degrees. Both elements

of first pleopod appreciably longer than cephalocaudal diameter of shaft of appendage.

COLOR NOTES.—Based on specimens from the

type-locality. Cephalothorax and abdomen concolorous blue gray, blue green to brown dorsally

with pair of submedian broken, dark brown stripes

on abdomen. Stripes evident in younger individuals and less apparent, or faded, in adults.

Branchiostegites and hepatic region light brown.

Chela blue gray, blue green to brown dorsally

with lighter proximolateral area. Distal half or

less of fingers orange to yellow. Pereiopods brown

dorsolaterally; distal podomeres darker dorsally

than proximal ones.

Rostral margins and postorbital ridges light

brown. Lateral margin of antennal scale dark

brown. Color of antenna and antennule like that

of dorsal surface of body. Tubercles on dorsal

surface of chela, carpus, merus, and lateral surface

of cephalothorax (especially cephalic region of

carapace) cream to yellow. Large swollen tubercle

at base of dactyl and articular knob on dorsal

surface at base of palm yellow.

The following color notes are based on specimens from Towns and Union counties, Georgia.

Dorsum of carapace olive tan fading ventrally to

pale tan. Rostral margins, postorbital ridges, and

cephalolateral margins of carapace orange tan.

Abdomen, telson, and uropods olive tan, slightly

darker than dorsum of carapace. Antennular and

antennal peduncles olive tan with pale orange

tan ring on distal extremity of each podomere;

flagella olive tan. Cheliped olive tan dorsally from

midlength of merus over dorsal surface of palm

but with vermilion tubercles; distal portion (occasionally almost all) of both fingers, ventral,

mesial, and lateral surfaces from ischium distally

also vermilion; ventral surface of cheliped orange

basally becoming vermilion on fingers; articular

membranes pinkish cream. Remaining pereiopods

orange tan proximally, suffused with olive on

distal half of merus and becoming pale green over

propodus and dactyl. Ventral surface of third

maxilliped, sternum, and ventral surface of proximal podomeres of pereiopods pinkish cream.

The orange coloration on the ventral surface of

C. nodosus is shared by certain other close relatives

belonging to the subgenus Jugicambarus: C. asperimanus Faxon (1914:391) (Blue Ridge province of North Carolina, Tennessee, Georgia, and

South Carolina), C. conasaugaensis Hobbs and

Hobbs (1962:41) (Blue Ridge province of Tennessee and Georgia), and C. parvoculus Hobbs

and Shoup (1947:142) (Cumberland Plateau and

Cumberland Mountains, descending to western

edge of Ridge and Valley province in Virginia,

Kentucky, Tennessee, and probably Georgia).

(See Bouchard, in press.)

MEASUREMENTS (in mm).—

Characters

Carapace:

Total length

of carapace

Postorbital

carapace length

Width

Height

Areola:

Width

Length

Rostrum:

Width

Length

Chela:

Length, mesial

margin of palm

Width, palm

Length, lateral

margin

Length, dactyl

Abdomen:

Width

Length

Holotype

28.9

25.3

14.9

12.0

1.5

10.7

3.8

3.6

7.8

11.0

21.9

13.1

12.9

29.3

Allotype

30.8

26.9

15.1

12.8

2.0

11.6

42

3.9

7.4

10.4

212

12.2

14.6

29.6

Morpho

31.7

27.4

16.4

12.6

1.7

12.0

4.2

4.3

82

10.5

23.0

135

14.0

31.6

TYPE-LOCALITY.—A small unnamed tributary of

North Potato Creek (Hiwassee River system via

Ocoee River) between 0.7 and 0.9 mile west of the

Tennessee-North Carolina border on U.S. Highway 64, Polk County, Tennessee. During the autumn, the water level was low, and the depth

varied from several inches in the riffles to less than

15 inches in the pools. The stream is approximately 2 to 6 feet wide and flows over a substrate

of sand and mica with stretches of bedrock and

scattered small to large rocks. The banks consist

of sand with scattered embedded rocks. At this

locality the stream is bordered with grasses and a

dense growth of Bidens sp.

DISPOSITION OF TYPES.—The holotypic male,

form I (no. 146756), the allotype (no. 146757),

and the morphotypic male, form II (no. 146758)

are deposited in the National Museum of Natural
NUMBER 224 13

History, Smithsonian Institution. Paratypes consisting of 5 $ I, 8 $ II, 20 $ , 11 j $ , 7j 9 , 4 9 with

eggs and 2 5 with young are also in NMNH, and

4 $ II, 4 $, lj $ and 1 j $ are in the collection of

the senior author.

RANGE AND SPECIMENS EXAMINED.—The crayfish is known from the Hiwassee (Ocoee, Nottely,

and Toccoa river basins), Savannah (Chatooga

and Tugaloo river basins) and Chattahoochee

river systems in Tennessee, North Carolina, and

Georgia. It seems possible that it also may be

found in South Carolina. The presence of an additional member of the subgenus Jugicambarus

in the Blue Ridge province lends countenance to

the hypothesis that the subgenus used this province as a path of migration and radiation.

GEORGIA. LUMPKIN COUNTY: (1) 3.6 miles south of

Union County line on U.S. Highway 19 and 0.4 mile west

of Highway in DeSota Falls camping area, 1 $ II, 3j $, 2j 9,

VI/17/75, D. J. Peters, J. E. Pugh, and H. H. H., Jr., coll.

RABUN COUNTY: (2) County Road just south of U.S. Highway 76, 2 miles east of Clayton, 19, IV/25/67, Torgny

Unestam and H. H. H., Jr., coll. TOWNS COUNTY: (3) Bog in

headwaters of Tallulah River north of Tate City, 2 $ I, 2 9,

IV/11/67, C. H. Wharton and F. K. Parrish, coll.; 1£I,

VI/7/67, C. H. W., coll.; 19, VI/23/67, C. H. W., coll.;

1 311, 1 9 , VII/6/67, C. H. W., coll. (4) Along roadside seepage area on Forest Road 70 at edge of Tate City, 2 9, lj $,

2 ovigerous 9,1 9 with young, VI/16/75; D. J. P., J. E. P.,

and H. H. H., Jr., coll. (5) Roadside ditch 5 miles south of

junction of U.S. Highway 76 on State Route 17, 2 9, 1 ovigerous 9,1 9 with young, VI/17/75, D. J. P., J. E. P., and

H. H. H., Jr., coll. UNION COUNTY: (6) Seepage area 3.6 miles

west of U.S. Highway 19 on State Route 180 (top of mountain west of Vogel State Park), lj 9, IV/27/67, T. U. and

H. H. H., Jr., coll.; 1 $ II, 3 9 , 1 j $, VI/17/75, D. J. P., J. E. P.,

H. H. H., Jr., coll. (7) Stream 0.5 mile north of Vogel State

Park on U.S. Highway 19, 1$I, XI/5/58, K. W. Simonds,

coll. WHITE COUNTY: (8) Seepage area across road from Spoil

Cane Creek, 9.1 miles south of Towns County line on State

Route 17, 23II, 39 , 2j£, 2j 9, 1 ovigerous 9, VI/17/75,

D. J. P., J. E. P., and H. H. H., Jr., coll.

NORTH CAROLINA. CHEROKEE COUNTY: (1) 1.6 miles

west of junction U.S. Highway 64 and North Carolina State

Route 60 on latter, 19, VI/12/60, K. W. S., coll. (2) 6.9

miles east of Tennessee-North Carolina state line on U.S.

Highway 64, 1 9, IV/15/62, K. W. S., coll.

TENNESSEE. POLK COUNTY: (1) Type-locality, 4$11, 49 ,

l\$, Ij9, XI/5/72, F. L. Oakberg, B. E. Oakberg, J. W.

Bouchard and R. W. B., coll.; \ $1, 1$II, 19, X/23/71,

J. W. B. and R. W. B., coll.; 1 $1, 1 $ II, XII/9/59, K. W. S.,

coll.; 1$H, 19, lj$, VI/8/59, K. W. S., coll. (2) Tennessee

Route 68 in Harbuck, 1 $ II, VI/8/59, K. W. S., coll. (3) Public picnic area 0.9 mile west of Turtletown on Tennessee

Route 68, 29 , 3}$, 2j9, VI/12/59, K. W. S., coll.

VARIATIONS.—The most evident variations in

this species are found in the chela, suborbital

angle, rostrum, and areola. The tubercles on the

dorsal surface of the chela vary in size and area of

dispersal. The suborbital angle varies from subacute to broadly rounded, while the rostrum on

the Georgia material often possesses a distinct carina at the tip. The areola of specimens from the

headwaters of the Hiwassee and Savannah rivers

in Georgia is narrower, ranging from 8.5 to 13.6

times as long as broad. None of the other specimens available possess such a narrow areola.

Other than an expected range in meristic

and mensural characters, as well as in density of

setae, there exist few notable variations. (See

"Diagnosis.")

LIFE HISTORY NOTES.—First form males have

been collected in April, June, and from October

to December. Females carrying eggs or young were

collected during the month of June.

ECOLOGICAL NOTES.—Cambarus (/.) nodosus has

been collected from burrows in the stream bank

and in seepage areas, under rocks on shore or at

the shoreline, and under rocks in midstream of

pools and slow runs. Despite this broad array of

habitats, the reduced size of the eyes and the relatively narrow areola indicate that C. (J.) nodosus

is primarily a burrowing species. Most of our specimens were dug from complex burrows consisting

of branching galleries with two or more openings

to the surface. Several were taken from a sphagnum bog in pit traps set to catch small mammals.

Collected with Cambarus (J.) nodosus in one

or more localities were Cambarus (C.) bartonii

(Fabricius, 1798:407), C. (Hiaticambarus) longirostris Faxon (1885:64), and two undescribed

congeners, one each of the subgenera Puncticambarus and Lacunicambarus. Cambarus (Lacunicambarus) sp. was collected only from burrows in

the stream bank, and C. (C.) bartonii was taken

in the stream as well as under rocks at or on the

shore. Cambarus (H.) longirostris was found

primarily in runs and C. (Puncticambarus) sp.

in pools and riffle areas of streams.

RELATIONSHIPS.—Cambarus (J.) nodosus has its

closest affinities with C. (/.) carolinus, sharing

with it a similar first form male gonopod and general mien. It differs from C. (/.) carolinus primarily in exhibiting the following characters: the

chela of C. (/.) nodosus possesses two well-defined
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rows of tubercles along the mesial surface of the

palm with additional smaller ones dispersed over

the dorsal surface; the central projection of the

first form male gonopod bears a distinct, small

subapical notch; and the suborbital angle is moderately to well developed. An additional character

that will serve to separate C. (/.) nodosus from

C. (/.) carolinus is exhibited in the areola width,

the former possessing an areola 5.0 to 13.6 times

as long as broad compared to a range of 13.7 to

39.3 in the latter (data from Hobbs and Bouchard,

1973).

The more subdued coloration of C. (/.) nodosus is in sharp contrast to the bright red to red

orange colors exhibited by C. (/.) carolinus. Only

a single individual of the latter is known to have

exhibited a different color after molting—from red

to blue gray in captivity.

SIZE.—The largest specimen available is a female from Polk County, Tennessee, with a carapace length of 36.5 mm (postorbital carapace

length 32.4 mm). The smallest first form male is

from Towns County, Georgia, and has corresponding lengths of 26.8 and 23.9 mm. The smallest female with eggs or young is also from Towns

County and has comparable lengths of 26.5 and

23.4 mm.

ETYMOLOGY.—The name nodosus (L., = knotty)

was chosen because of the large number of welldeveloped tubercles on the dorsomesial half of the

palm of the chela and on the hepatic region of

the carapace.
Literature Cited

Bouchard, Raymond W.

1972. A New Genus of Crayfish from Kentucky and Tennessee (Crustacea, Decapoda, Astacidae). [Abstract.]

ASB Bulletin, 19(2):56.

1973. A New Crayfish of the Subgenus Jugicambarus

from Tennessee with an Emended Definition of the

Subgenus (Astacidae, Decapoda). American Midland

Naturalist, 89 (l):103-lll, 1 figure.

in press. Geography and Ecology of Crayfishes of the

Cumberland Plateau and Cumberland Mountains,

Kentucky, Virginia, Tennessee, Georgia, and Alabama, Part II: The Genera Fallicambarus and Cambarus. In James W. Avault, Jr., editor, Freshwater

Crayfish. [Papers from the Second International

Symposium on Freshwater Crayfish, Baton Rouge,

Louisiana, U.S.A., Division of Continuing Education, Louisiana State University.]

Erichson, W. F.

1846. Ubersicht der Arten der Gattung Astacus. Archiv

fur Naturgeschichte, Berlin, 12(l):86-103.

Fabricius, Johan C.

1798. Supplementum Entomologiae Systematicae. 572

pages. Hafniae: Proft et Storch.

Faxon, Walter

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):

vi + 186 pages, 10 plates.

1914. Notes on the Crayfishes in the United States

National Museum and the Museum of Comparative

Zoology, with Descriptions of New Species and Subspecies to Which Is Appended a Catalogue of the

Known Species and Subspecies. Memoirs of the

Museum of Comparative Zoology at Harvard College, 40(8):351-427, 13 plates.

Hay, William P.

1902. Observations on the Crustacean Fauna of Nickajack Cave, Tennessee, and Vicinity. Proceedings of

the United States National Museum, 25(1292):

417^39, 8 figures.

Hobbs, Horton H., Jr.

1965. A New Crayfish of the Genus Cambarus from

Tennessee with an Emended Definition of the

Genus (Decapoda, Astacidae). Proceedings of the

Biological Society of Washington, 78 (33):265-273,

12 figures.

1969. On the Distribution and Phylogeny of the Crayfish Genus Cambarus. In Perry C. Holt, Richard L.

Hoffman, and C. Willard Hart, Jr., The Distributional History of the Biota of the Southern Appalachians, Part I: Invertebrates. Virginia Polytechnic

Institute, Research Division Monograph, 1:93-178,

20 figures.

Hobbs, Horton H., Jr., and Raymond W. Bouchard

1973. A New Crayfish from the Cumberland River System

with Notes on Cambarus carolinus (Erichson).

Proceedings of the Biological Society of Washington, 86 (5):41-68, 4 figures.

Hobbs, Horton H., Jr., and H. H. Hobbs III

1962. A New Crayfish of the Genus Cambarus from

Georgia (Decapoda, Astaddae). Proceedings of the

Biological Society of Washington, 75(5):41-45, 10

figures.

Hobbs, Horton H., Jr., and Charles S. Shoup

1947. Two New Crayfishes (Decapoda, Astacidae) from

the Obey River Drainage in Tennessee. Journal of

the Tennessee Academy of Science, 22(2):138-145,

22 figures.

Holt, Perry C.

1968. The Genus Pterodrilus (Annelida: Branchiobdellida). Proceedings of the United States National

Museum, 125 (3668): 1-44, 12 figures.

Prins, Rudolph

1968. A New Crayfish of the Genus Cambarus from

North Carolina (Decapoda, Astacidae). Journal of

the Elisha Mitchell Scientific Society, 84(4):458-461,

11 figures.

Rhoades, Rendell

1944. The Crayfishes of Kentucky, with Notes on Variation, Distribution, and Descriptions of New Species

and Subspecies. American Midland Naturalist, 31(1):

111-149, 10 figures.

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15



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retained by the author. There may be several paragraphs to a page, but each page should begin

with a new paragraph. Number consecutively all pages, including title page, abstract, text, literature cited, legends, and tables. The minimum length is 30 pages, including typescript and illustrations.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic

titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further

headings may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date:page) with a full reference at the

end of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate

each reference. Enclose with square brackets any annotation in, or at the end of, the entry.

For references within the text, use the author-date system: "(Jones, 1910)" and "Jones

(1910)." If the reference is expanded, abbreviate the data: "Jones (1910:122, pi. 20: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Use the metric system instead of, or in addition to, the English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on

pages separate from the text and not attached to the artwork. An instruction booklet for the

preparation of illustrations is available from the Press on request.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and

article titles, using initial caps with all words except minor terms such as "and, of, the." For

capitalization of titles in foreign languages, follow the national practice of each language.

Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume,

number, and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," "pages."

For free copies of his own paper, a Smithsonian author should indicate his requirements

on "Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will

receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.