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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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 2 2 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
A B S T R A C 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 pro-
posed 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 unde-
scribed 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, Cam-
baridae) 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 cray-
fishes of the genus Cambarus . . . . III. Series: Smithsonian Institution. Smithsonian contri-
butions 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 gen-
eral acceptance among crayfish systematists. Only
the monotypic subgenus Barbicambarus has been
elevated to generic rank (Bouchard, 1972), suggest-
ing a greater divergence from a common ancestry
in it than in any other member or assemblage
within the genus. Cambarus (Exilicambarus) cra-
cens, described herein, provides evidence of a
previously unknown evolutionary line within the
genus; nevertheless, it bears morphological resem-
blances to members of three of the currently rec-
ognized subgenera. Certain characteristics of the
areola, rostrum, cervical spines, and chela are simi-
lar to those in members of the subgenus Puncti-
cambarus. Other characters of the chela and of the
first form male gonopod are suggestive of cray-
fishes 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., Depart-
ment 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 Exilicam-
barus.
The second species described herein represents
one of a growing number of burrowing crayfishes
to be reported. The difficulty in obtaining speci-
mens 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 contrib-
uted 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, bran-
chiostegal 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 ven-
tral surface of palm, studded with squamous tu-
bercles; lateral margin of fixed finger costate. Hook
on ischium of third pereiopod of male overreach-
ing basioischial articulation, not opposed by tuber-
cle 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 caudolat-
eral base of central projection. Annulus ventralis
shallowly embedded in sternum, subspindle
shaped. First pleopod reaching or almost reach-
ing midlength of annulus when abdomen flexed.
TYPE-SPECIES.?Cambarus (Exilicambarus) cra-
cens, here described.
LIST OF SPECIES.?Monotypic.
ETYMOLOGY.?Exilis (L., = slender) in combi-
nation with Cambarus; in reference to the com-
paratively 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 cer-
vical groove (17.6 and 14.8 mm). Areola 3.4
times as long as broad with 6 widely spaced punc-
tations across narrowest part; length of areola
31.9 percent of entire length of carapace (41.9 per-
cent of postorbital carapace length). Rostrum
with slightly thickened, convergent margins bear-
ing 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, acu-
men slightly upturned apically; dorsal surface of
rostrum concave and bearing small setiferous punc-
tations, 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 spini-
form corneous tubercle cephalically. Suborbital
angle broadly rounded, virtually nonexistent;
branchiostegal spine moderately large, acute, and
curved cephalodorsally. Cervical spine well devel-
oped and flanked by several small tubercles, at
least one of which acute; hepatic area with scat-
tered tubercles; cephaloventral part of branchio-
stegite with crowded tubercles and few adjacent
and caudoventral to cervical spine, remaining lat-
eral portion of branchiostegite granulate and dor-
solateral 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 prox-
imal section; corresponding margin of mesial ra-
mus with spine at distolateral angle.
Cephalic lobe of epistome (see Figure 2t of para-
typic male, form I) apparently injured, rounded
but with asymmetrical emargination cephalodex-
trally, margins only slightly thickened and mar-
ginal 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 sur-
face at base of distal two-fifths. Antennal peduncle
with lateral spine on basis and distal spine on
ischium, flagellum extending caudally to third ab-
dominal 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 maxil-
liped with stiff setae studding mesial half, row of
submarginal shorter setae laterally, and with
scattered setiferous punctations between submar-
ginal 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), sub-
ovate, 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 ventro-
mesially; 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 disto-
lateral part of palm. Both fingers with well-defined
submedian ridges flanked by setiferous puncta-
tions 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) reach-
ing 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 sub-
equal in size and much larger than distal 2, distal-
most somewhat removed from more proximal 4,
and additional large tubercle present more ven-
trally 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 proxi-
mally (carpus of left chela with two procurved
spines); distoventral margin with strong subme-
dian spiniform tubercle, another on ventrolateral
articular knob, and 2 smaller tubercles proximo-
mesial 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 ven-
tromesially.
Hook on ischium of third pereiopod only (see
Figure 2h of paratypic male, form I); hook simple,
overreaching basioischial articulation and not op-
posed by tubercle on basis. Coxa of fourth pereio-
pod with moderately prominent caudomesial boss,
that of fifth well developed but much less con-
spicuous than that on fourth. (See "Measure-
ments.")
First pleopods (Figure \c,d; see also Figure 1b,
f) reaching coxae of third pereiopods when ab-
domen 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 sex-
ual features, differing from holotype in following
respects: epistome symmetrical, more nearly sub-
triangular and fovea slightly deeper; cephalic sec-
tion of telson with 2 spines in each caudolateral
corner; more lateral row of tubercles on dorso-
mesial surface of palm of chela consisting of 6 or
7 tubercles; ventral surface of palm with fewer
and smaller tubercles, row extending proximo-
mesially 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 sur-
face; only 1 small tubercle proximomesial to me-
dian ventromarginal tubercle; merus with 3 sub-
distal acute tubercles dorsally on left cheliped,
ventral surface with 10 tubercles in mesial row and
4 in lateral, right cheliped with 12 and 3 respec-
tively; ischium with 4 tubercles on right and 3 on
left cheliped. (See "Measurements.")
Annulus ventralis (Figure 2k) distinctly asym-
metrical and cephalic region not nearly so strongly
calcified as caudal; cephalic half with shallow, sub-
median, longitudinal trough; sinus originating at
sinistrocaudal end of latter and extending sinis-
trally, 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 ab-
domen flexed.
MORPHOTYPIC MALE, FORM II.?Differing from
holotype in following respects: epistome resem-
bling that of allotype; tubercles flanking cervical
spine on holotype and allotype not evident; ceph-
alic section of telson with 2 spines in each cau-
dolateral 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); oppos-
able 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 sub-
acute apex directed caudally at approximately 90
degrees; only trace of caudal element evident.
COLOR NOTES.?Carapace olive tan with irregu-
lar 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 ven-
trally along cephalic and cephaloventral margins
of pleuron of next segment, becoming reddish
brown on pleuron and fading before reaching cau-
doventral 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 sub-
parallel to ventral margin. Margins of telson, as
well as keels, lateral margins, and proximal bor-
ders 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 bot-
tom littered with large rocks. Shading the shore
line are Platanus occidentalis, Liriodendron tulipi-
fera, 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. Par-
atypes 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 cray-
fish 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 , 3 9 ,
9j<5? 7 j9 , 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^I I
(morphotype), X/31/74, J. W. B. and R. W. B., coll. MAR-
SHALL 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 , 2 9 ,
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 (mar-
ginal 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 tub-
ercles. 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 tuber-
culiform, although always bearing an acute tip.
Frequently, one (sometimes two) spiniform tuber-
cle 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 de-
scriptions of the three specimens above. The first
pleopod of the male, form I, exhibits few varia-
tions, 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 arti-
facts of preservation.
SIZE.?The largest specimen available is a fe-
male having a carapace length of 39.7 mm (post-
orbital carapace length 30.7 mm). The corre-
sponding 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 sea-
sons in the adult male population. Neither oviger-
ous females nor those carrying young have been
found, although well-developed cement glands
were observed on two females collected in Novem-
ber. Judging by the sizes of the limited number of
specimens at hand, three generations are present
in the populations sampled. The smallest individ-
ual 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 slug-
gish. At that season, the rock-littered areas were
so choked with leaves that it is possible the cray-
fish were among the latter rather than having
sought cover beneath the rocks. All of the speci-
mens 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 men-
tioned 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 speci-
mens, 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 puncta-
tions 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) rebur-
rus Prins (1968:458). In few species of the genus
is the palm of the chela so nearly completely cov-
ered with tubercles; only in some members of the
subgenus Depressicambarus is such a tuberculate
condition approximated.
The tapering central projection of the first pleo-
pod of the first form male, although strongly re-
curved, is unique in that the morphologically ceph-
alodistal margin is almost straight.
For the following reasons, it does not seem ap-
propriate to assign this crayfish to any of these
subgenera. The usually limited number of punc-
tations 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 con-
stitute a combination of features that does not
occur elsewhere in members of the subgenus Punc-
ticambarus. The long rostrum with marginal tu-
bercles or spines, the strongly developed cervical
spine, and the elongate chela studded with tu-
bercles, 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 pereio-
pod 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 dif-
ferences exist in the first pleopod of the first form
male: the central projection is more strongly re-
curved; 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 de-
veloped. 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 tuber-
cles. 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 un-
even; 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, blade-
like, tapering, bearing distinct, small subapical
notch, and recurved at angle of approximately 90
degrees to shaft of appendage; mesial process tu-
mescent, tapering to acute tip and directed cau-
dally at angle of approximately 90 degrees. Both
elements of first pleopod appreciably longer than
cephalocaudal diameter of shaft of appendage. An-
nulus ventralis asymmetrical, subcylindrical and
movable; cephalic half bearing longitudinal me-
dian trough between anteriorly converging ridges,
and caudal half with sinuate dextral or sinistral
sinus. Caudal wall of annulus elevated. First pleo-
pod of female uniramous and reaching at least
midlength of annulus when abdomen flexed.
HOLOTYPIC MALE, FORM I.?Body subovate, de-
pressed (Figure 3a,;). Abdomen narrower than
thorax (12.9 and 14.9 mm). Greatest width of cara-
pace 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 en-
tire length of carapace (42.3 percent of postor-
bital 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, espe-
cially deep anteriorly, with submarginal puncta-
tions 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; branchio-
stegal spine represented by small tubercle. Cervi-
cal spine absent although represented by very
small rounded tubercle. Hepatic area and lateral
portion of branchiostegite tuberculate; dorsal sur-
face of carapace punctate.
Abdomen longer than carapace (29.3 and 28.9
mm); pleura of moderate length with caudoven-
tral extremities subangular. Cephalic section of
telson with marginal spine and submarginal mov-
able spine in each caudolateral corner and par-
tially 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 termi-
nating in acute spine at transverse flexure. Proxi-
mal portion of lateral ramus with row of small
spines distally and large, submarginal movable
spine at caudolateral corner. Mesial ramus of uro-
pod with median ridge terminating distally in pre-
marginal acute spine; lateral margin with disto-
lateral 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 im-
mediately cephalic and subparallel to thickened,
slightly arched epistomal zygoma; lateral extremi-
ties with several small tubercles. Proximal seg-
ment of antennule with small spine on ventral sur-
face near midlength. Antennae broken. Antennal
scale (Figure Si) broadly declivous, broadest distal
to midlength; thickened lateral portion terminat-
ing 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 dorso-
lateral 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 morpho-
typic 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 extend-
ing distally from fourth tubercle from base, row
interrupted only by fifth. Dorsal and ventral sur-
faces of dactyl with median longitudinal ridges
flanked by setiferous punctations; opposable mar-
gin with row of 7 tubercles situated in slight exca-
vation, proximal 3 rounded (basalmost largest),
followed by 2 moderately acute and 2 acute distal
ones; row of denticles, broken by tubercles, ex-
tending 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 tuber-
cle near proximal base of latter; 3 rounded tuber-
cles on dorsomesial surface; distoventral margin
with 3 rounded tubercles; podomere otherwise
punctate.
Upper surface of merus with 3 rounded subdis-
tal tubercles and 11 along crest of podomere; ven-
tral surface with lateral row of 5 subacute spines,
and mesial row of 11 subacute ones, both rows de-
creasing in size proximally. Ischium with row of
3 small tubercles on ventromesial margin.
Hook on ischium of third pereiopod only (Fig-
ure Sh); hook simple, overreaching basioischial
articulation and not opposed by tubercle on basis.
Coxa of fourth pereiopod with prominent caudo-
mesial boss, that of fifth without prominence. (See
"Measurements".)
First pleopods (Figure Sb,d,f) reaching caudal
portion of coxae of third pereiopods when abdo-
men 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 post-
orbital 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 sur-
face with 3 very small tubercles (6 on left cheli-
ped); distoventral surface with cluster of 5 tuber-
cles. 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 con-
verging anteriad; caudal half with sinistral sinus.
Caudal wall of annulus elevated, especially caudo-
laterally. 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 exca-
vate anteriorly; ventral margin with setae sparse
on sides of rostrum and lacking on most of acu-
men; cephalic tubercle more spiniform. Areola
constituting 37.9 percent of entire length of cara-
pace (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 car-
pus of both left and right chelipeds with hori-
zontal 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, re-
spectively. Caudolateral corners of epistome espe-
cially elevated. Propodus of chela with third and
fourth tubercles from proximal end more acute.
Hook on ischium of third pereiopod much re-
duced, 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 append-
age; 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 cephalo-
caudal diameter of shaft of appendage.
COLOR NOTES.?Based on specimens from the
type-locality. Cephalothorax and abdomen con-
colorous blue gray, blue green to brown dorsally
with pair of submedian broken, dark brown stripes
on abdomen. Stripes evident in younger indi-
viduals 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 speci-
mens 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 (oc-
casionally 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 proxi-
mal 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. as-
perimanus Faxon (1914:391) (Blue Ridge prov-
ince of North Carolina, Tennessee, Georgia, and
South Carolina), C. conasaugaensis Hobbs and
Hobbs (1962:41) (Blue Ridge province of Ten-
nessee 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. High-
way 64, Polk County, Tennessee. During the au-
tumn, 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 approxi-
mately 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 con-
sisting 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 cray-
fish 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 ad-
ditional member of the subgenus Jugicambarus
in the Blue Ridge province lends countenance to
the hypothesis that the subgenus used this prov-
ince 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. High-
way 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 seep-
age 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 oviger-
ous 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 moun-
tain 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, 3 9 , 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, 4 9 ,
l\$, I j9 , 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) Pub-
lic picnic area 0.9 mile west of Turtletown on Tennessee
Route 68, 2 9 , 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 sub-
acute to broadly rounded, while the rostrum on
the Georgia material often possesses a distinct ca-
rina 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 speci-
mens 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 rela-
tively narrow areola indicate that C. (J.) nodosus
is primarily a burrowing species. Most of our speci-
mens were dug from complex burrows consisting
of branching galleries with two or more openings
to the surface. Several were taken from a sphag-
num 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) longi-
rostris Faxon (1885:64), and two undescribed
congeners, one each of the subgenera Puncticam-
barus and Lacunicambarus. Cambarus (Lacuni-
cambarus) 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 gen-
eral mien. It differs from C. (/.) carolinus primar-
ily 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 mod-
erately 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. (/.) nodo-
sus 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 fe-
male from Polk County, Tennessee, with a cara-
pace length of 36.5 mm (postorbital carapace
length 32.4 mm). The smallest first form male is
from Towns County, Georgia, and has correspond-
ing lengths of 26.8 and 23.9 mm. The smallest fe-
male 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 well-
developed 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 Ten-
nessee (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 Ala-
bama, Part II: The Genera Fallicambarus and Cam-
barus. 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 Educa-
tion, Louisiana State University.]
Erichson, W. F.
1846. Ubersicht der Arten der Gattung Astacus. Archiv
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