Synopsis of the Families and

Genera of Crayfishes

(Crustacea: Decapoda)

HORTON H, HOBBS, JR.

m

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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 16 4

Synopsis of the Families and

Genera of Crayfishes

(Crustacea: Decapoda)

Horton H. Hobbs, Jr.

SMITHSONIAN INSTITUTION PRESS

City of Washington

1974
ABSTRACT

Hobbs, Horton H., Jr. Synopsis of the Families and Genera of Crayfishes (Crustacea: Decapoda). Smithsonian Contributions to Zoology, number IG4, 32 pages,

27 figures, 1974.—The subdivisions of the infraorder Astacidea, comprising the

lobsters (superfamily Nephropoidea) and crayfishes (superfamilies Astacoidea

and Parastacoidea) are briefly discussed and their affinities indicated. This is followed by a key to the superfamilies and of the families and genera of the Astacoidea and Parastacoidea. Two families, the Astacidae and Cambaridae, are recognized as constituting the holarctic Astacoidea, and the Parastacidae as the only

family of the Parastacoidea, which is confined to the Southern Hemisphere. Synonymies and diagnoses of these taxa and the genera composing them are presented; in addition, the range, number of species, references to summary treatments, and illustrations of a representative of each genus are included.

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 4969. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Hobbs, Horton Holcombe, Jr. 1914-

Synopsis of the families and genera of crayfishes (Crustacea: Decapoda)

(Smithsonian contributions to zoology, no. 164)

1. Crayfish. 2. Lobsters. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 164.

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Contents

Page

Introduction 1

Infraorder Astacidea 2

Acknowledgments 4

Key to the Families and Genera of the Infraorder Astacidea Latrielle, 1803,

Exclusive of the Superfamily Nephropoidea 4

Superfamily Asiacoidea 7

Family Astacidae 7

Genus Astacus 8

Genus Austropotamobius 9

Genus Pacifastacus 9

Family Cambaridae 10

Subfamily Cambarinae 10

Genus Barbicambarus 10

Genus Cambarus 11

Genus Fallicambarus 12

Genus Faxonella 12

Genus Hobbseus IS

Genus Orconectes 14

Genus Procambarus 15

Genus Troglocambarus 16

Subfamily Cambarellinae 17

Genus Cambarellus 17

Subfamily Cambaroidinae 17

Genus Cambaroides 17

Superfamily Parastacoidea 18

Family Parastacidae 18

Genus Astacoides 18

Genus Astacopsis 19

Genus Cherax 20

Genus Engaeus 20

Genus Engaewa 21

Genus Euastacoides 22

Genus Euastacus 22

Genus Geocharax 23

Genus Gramastacus 24

Genus Paranephrops 24

Genus Parastacoides 25

Genus Parastacus 25

Genus Samastacus 26

Genus Tenuibranchiurus 27

Literature Cited 27

i n

Synopsis of the Families and

Genera of Crayfishes

(Crustacea: Decapoda)

Horton H. Hobbs, Jr.

Introduction

For a number of years, it has become increasingly

apparent to me and to at least some of my colleagues that the assignment of the freshwater crayfishes to two families, the Astacidae restricted to the

Northern Hemisphere, and the Parastacidae, to the

Southern Hemisphere, does not adequately express

the diversity that exists in the holarctic assemblage.

The classification of crayfishes at the familial level

that seems generally to have been adopted is as follows:

Family Parastacidae

Family Astacidae

Subfamily Astacinae

Subfamily Cambaroidinae

Subfamily Cambarinae

Subfamily Cambarellinae.

The Parastacidae are distinguished from the

Astacidae primarily by the absence of the first pleopod and in the unmodified second pleopod in the

male, in lacking bilobed plaited laminae on any of

the podobranchiae, and the usual presence of

branchial filaments on the epipodite of the first

maxillipeds. Among the four subfamilies of the

Astacidae, the Astacinae differ from the other three

in that the male lacks clasping hooks on the ischia

of any of the pereiopods, by the loosely rolled or

Horton H. Hobbs, Jr., Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560

tubular distal portion of the first pleopod, and, more

fundamentally, in the absence of a cyclic dimorphism. The females differ from all except the few

species assigned to the Cambaroidinae in lacking a

well-developed, functional annulus ventralis. These

differences that distinguish the Astacinae from the

other crayfishes of the Northern Hemisphere seem

to me to be at least as remarkable as those that distinguish them from the Parastacidae, and the recognition of a third family of crayfishes is here proposed.

Considerably greater confidence exists in any

classification that is supported by a fossil record in

which implied dichotomies can be demonstrated.

Unfortunately, the fossil record of the freshwater

crayfishes is exceedingly poor. No pre-Pleistocene

crayfish fossil, to my knowledge, has been recorded

from the Southern Hemisphere. Not only are few

fossils from few localities available, but also many

features that are considered to be of importance in

assessing relationships are not discernible in those

that are at hand. The arrangement and structure of

the gills cannot be determined in any known fossil

assignable to a crayfish family, and the presence or

absence, to say nothing of the structure, of the first

pleopod of the male cannot be ascertained in those

specimens that have been examined by me. Furthermore, characters that would indicate a cyclic dimorphism in males are not evident in any fossil.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

Inasmuch as all male crayfishes that are known to

exhibit an alternation of a breeding form with a

nonbreeding one possess hooks on the ischia of one

or more of the second through fourth pereiopods,

the apparent absence of such hooks in the known

fossils suggests that none of them exhibited this

cyclic alternation of forms that in certain crayfishes

is unique among decapod crustaceans.

It seems unlikely that the physiological mechanisms that control this cyclic morphology have been

independently acquired in the eastern Asiatic (Cambaroidinae) and eastern North American (Cambarinae and Cambarellinae) stocks. Consequently,

the existence of a common ancestor more recent

than any shared with other crayfish lineages (Astacinae) seems certain (see page 3). In spite of the

fact that there is no evidence based on the fossil

record of crayfishes that underwent such a cyclic

change occurring in the Tertiary, the range of the

Cambarinae, one of the species groups in which the

cycle is typical, is such that one is almost forced to

conclude that a cyclic dimorphism had become established in this lineage before the end of the Miocene (Hobbs, 1967, 1971b).

The fact that the eastern Asiatic crayfishes are in

some respects intermediate between the Eurasianwestern North American species (possessing similar

arrangement of gills and a sclerite corresponding to

the annulus ventralis that lacks both a sinus and

fossa) and the eastern North American species

(bearing hooks on the ischia of the second and

third pereiopods in the male in which there is also

a cyclic dimorphism) does not necessitate the conclusion that the eastern Asiatic mien represents an

arrested morphology that at one time served as a

transitional form in the evolution of one from the

other. Equally likely is the possibility that from

some Mesozoic marine nephropoid ancestral stock

there developed two lines of evolution: one retaining the primitive absence of cyclic dimorphism and

another in which such a dimorphism became fixed

along with those characters that accompany it

(hooks on the ischia of certain pereiopods in the

male and an annular sclerite possessing a cavity or

fossa for sperm storage in the female).

The more conservative stock, in migrating into a

freshwater habitat, gave rise to that from which the

present-day Astacinae (Eurasia and western North

America) have descended. Those remaining in marine waters were presumably ancestral to modern

Nephropoidea. The other stock, that in which cyclic dimorphism became established, also migrated

into fresh water, giving rise to those crayfishes assigned to the Cambaroidinae, Cambarellinae, and

Cambarinae. Whether or not the Asiatic Cambaroidinae and the American subfamilies were derived

from a common freshwater-inhabiting ancestor, or

whether there were separate Asiatic and American

invasions of fresh water in post-Jurassic times may

never be known; certain possibilities, however, were

recently discussed by Holt (1968:85-86) and Hobbs

(1969:115-116). A possible origin of the Parastacidae has been discussed by Riek (1959:255-257).

A meaningful taxonomy of the holarctic assemblage must reflect the recognition of a fundamental

divergence in the physiology and morphology of

two stocks expressed in the presence or absence of

the cyclic production of sperm and provisions for its

conveyance to the female. The cyclic behavior of

the testis of a member of the Cambarinae was

shown by Word and Hobbs (1958) to be associated

with the alternating first and second forms of the

male in which there is also an alternation in the

relative size of the grasping hooks on the ischia of

one or more of the pereiopods and the shape and

texture of the terminal elements of the first pleopods. These characters are most assuredly as little,

and perhaps less, affected by environmental conditions as are the loss of one or more units of the

branchial series. They are also equally as important

in assessing relationships as are modifications of the

branchial laminae, and of no less importance than

the development and variations that occur in the

telson and presence or absence of first pleopods.

Taking cognizance of these characteristics that, in

my opinion, have been inadequately emphasized,

together with others cited in the diagnosis presented

below, the following restructuring of the Astacuran

Infraorder Astacidea is proposed.

Infraorder ASTACIDEA

Superfamily Nephropoidea *

Superfamily Astacoidea

Family Astacidae

Genus Astacus

Austropotamobitis

Pacifastacus

1The Atlantic Nephropoidea are being revised by L. B. Hoithuis, and included in his study will be a review of the generic and familial characteristics.
NUMBER 164

MAJOR CHARACTERISTICS INVOLVED IN THE EVOLUTION OF CRAYFISHES

FROM A PRO-NEPHROPOID ANCESTOR

CAMBAROIDINE ANCESTOR

FIRST PLEOPOD OF MALE WITH

SHALLOW SPERM GROOVE OPENING

DISTALLY AMONG TERMINAL

ELEMENTS

ANNULAR PLATE IN FEMALE ALWAYS

LACKING SINUS AND FOSSA

BRANCHIAL FORMULA 18 + 3R • EP

MALE WITH HOOKS ON ISCHIA OF

THIRD, FOURTH, OR THIRD AND

FOURTH PERE1OPODS

ANNULUS VENTRAL FIXED, OR, IF

MOVABLE, SELDOM THROUGH ARC

OF MORE THAN <»0 DEGREES

BRANCHIAL FORMULA 17 + EP

CAMBARELLINE ANCES

MALE WITH HOOKS ON ISCHIA OF

SECOND AND THIRD PEREIOPODS

ANNULUS VENTRALIS MOVABLE

THROUGH ARC OF AT LEAST 75

DEGREES

BRANCHIAL FORMULA 16 + EP

ASTACID AN(

CYCLIC DIMORPHISM ABSENT IN

MALE

FIRST PLEOPOD OF MALE SUBTU8ULAR

DISTALLY AND LACKING

ORNAMENTATION OTHER THAN SPOONLIKE LOBES

ISCHIA OF ALL PEREIOPODS OF MALE

LACKING HOOKS

ANNULAR PLATE IN FEMALE WITHOUT

SINUS OR FOSSA

PRO-CAMBAR INE ANCESTOR

FIRST PLEOPOD OF MALE WITH

DEEP SPERM GROOVE OPENING

DISTALLY ON CENTRAL PROJECTION

ANNULAR PLATE IN FEMALE ALWAYS

WITH SINUS AND FOSSA

BRANCHIAL FORMULA 17 + EP,

OR 16 + EP

CAMBARID ANCESTOR

CYCLIC DIMORPHISM IN MALE

FIRST PLEOPOD OF MALE WITH

SHALLOW OR DEEP SPERM GROOVE

AND WITH TERMINAL ORNAMENTATION

ISCHIA OF ONE OR MORE PEREIOPODS

OF MALE WITH HOOKS

ANNULAR PLATE IN FEMALE WITH OR

WITHOUT SINUS AND FOSSA

ASTACOIDEAN ANCESTOR

FIRST PLEOPODS OF MALE EACH

WITH SPERM CONDUIT

SECOND PLEOPOD OF MALE WITH

SPIRAL ELEMENT ON ENDOPODITE

PODOBRANCHIAE WITH FUSED BRANCHIAL

AND EPI PODITIC PORTIONS AND

WITH BILOBED PLAITED LAMINAE

STERNUM BETWEEN FIFTH PEREIOPODS

NOT FUSED WITH ANTERIOR STERNAL

PLATE

EGGS HATCHING AS MINIATURES OF

ADULT

NEPHROPOIDEAN ANCESTOR

FIRST PLEOPODS OF MALE

SERVING TOGETHER TO FORM

SINGLE SPERM CONDUIT

SECOND PLEOPOD OF MALE WITHOUT SPIRAL ELEMENT ON

ENDOPODITE

PODOBRANCHIAE WITH DISCRETE

BRANCHIAL AND EPIPODITIC

PORTIONS

STERNUM BETWEEN FIFTH

PEREIOPODS FUSED WITH

ANTERIOR STERNAL PLATE

EGGS HATCHING AS LARVAE

PARASTACOIDEAN ANCESTOR

FIRST PLEOPODS ABSENT IN

MALE AND FEMALE

SECOND PLEOPOD OF MALE

WITHOUT SPIRAL ELEMENT ON

ENDOPODITE

PODOBRANCHIAE WITH FUSED

BRANCHIAL AND EPIPODITIC

PORTIONS BUT WITHOUT

BILOBED PLAITED LAMINAE

STERNUM BETWEEN FIFTH

PEREIOPODS NOT FUSED WITH

ANTERIOR STERNAL PLATE

EGGS HATCHING AS MINIATURES

OF ADULT

PRO-NEPHROPOID ANCESTOR
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Family Cambaridae

Subfamily Cambarinae

Genus Barbicambarus

Cambarus

Fallicambarus

Faxonella

Hobbseus

Orconectes

Procambarus

Troglocambarus

Subfamily Cambarellinae

Genus Cambarellus

Subfamily Cambaroidinae

Genus Cambaroides

Superfamily Parastacoidea

Family Parastacidae

Genus Astacoides

Astacopsis

Cherax

Engaeus

Engaewa

Euastacoides

Euastacus

Geocharax

Gramastacus

Paranephrops

Parastacoides

Parastacus

Samastacus

Tenuibranchiurus

Because of the scattered literature treating the

three families of crayfishes—no composite summaries of them have been attempted since those of

Faxon (1885a, 1914)—a key to the taxa comprising

crayfishes, together with synonymies and diagnosis

of each, are presented here. In addition to the diagnoses of the genera, the following are cited: range

of each, genus, number of described species and

nonnominate subspecies currently recognized, and

references to monographic or summary treatments

and keys.

The generic structure of the Astacidae utilized is

that of Bott (1950), and that of the Parastacidae

has been proposed by Riek (1969, 1971, 1972). I

have drawn freely from these works in composing

the diagnoses. The organization of the Cambaridae

is based primarily on studies of Hagen (1870),

Faxon (1885a), Ortmann (1905a, 1906), and Hobbs

(1942a, 1969, 1972a).

The illustrations included are depicted as though

all of the species were of approximately the same

size. They were so scaled as to allow an appreciation

of the differences in proportions of the structures

included.

Shortly after this manuscript had been completed,

Riek's study (1972) on the phylogeny of the Parastacidae appeared—in which not only are representatives of each of the parastacid genera illustrated but also a new genus, Gramastacus, is described. The present manuscript has been revised to

include the new genus, the treatment of which is

based on Riek's diagnosis and illustrations. It is unfortunate that neither of us was aware of the endeavors of the other.

ACKNOWLEDGMENTS.—Among those who have

assisted me in this undertaking are Lipke B. Hoithuis, who not only placed at my disposal his

synonymies of a number of the taxa included

herein, but who also critically read an early draft

of the manuscript. Fenner A. Chace, Jr., was most

helpful in discussing a number of problems that

arose during the preparation of the manuscript and

in reading the final draft. Without the cooperation

of D. J. G. Griffin in lending me specimens in the

collection of the Australian Museum, the illustrations of Astacopsis, Engaewa, and Tenuibranchiurus

would be incomplete or lacking. Martha R. Cooper,

Joseph F. Fitzpatrick, Jr., and Raymond B. Manning offered considerable encouragement and advice, and read a preliminary draft of the paper. To

all of these persons, I am most grateful. I also wish

to acknowledge, with appreciation, the assistance of

Carolyn B. Gast in inking the illustrations, that of

Carolyn S. Hahn in obtaining a number of the

early publications that were not readily available

in the Smithsonian Library, and that of Margaret

A. Daniel in all aspects of the preparation of the

manuscript.

Key to the Families and Genera of the Infraorder Astacidea Latrielle,

1803, Exclusive of the Superfamily Nephropoidea

Carapace with dorsomedian longitudinal suture or with simple or spinous ridge extending from caudal margin of carapace at least to base of rostrum; sternal plate

between fifth pereiopods fused with that between fourth pair; podobranchiae of first

3 pereiopods with discrete branchial and epipoditic parts; first pleopod of male without individual sperm conduit Superfamily NEPHROPOIDEA Dana, 1852
NUMBER 164

Carapace never with longitudinal suture nor with ridge extending from caudal margin

of carapace lo base of rostrum: sternal plate between fifth pereiopods never fused

with that between fourth pair; podobranchiae of first 3 pereiopods with branchial and

epipoditic elements united; first pleopod of male each with complete sperm conduit

or first pkopod of male absent 2

2(1). First pleopod of male well developed and serving as organ of sperm transfer, reduced

or absent in females; podobranchiae of second and third maxillipeds and first 3

perciopods with bilobed. plaited laminae; epipodite of first maxilliped without branchial filaments; coxopoditic setae acute apically; telson usually (except in Cambaroides,

Hacifaitacus and rarely in Fallicambarus) divided by transverse suture

Sup* ifamily ASTACOIDEA 3

First pleopod lacking in both sexes; podobranchiae lacking bilobed, plaited laminae

although stem may be alate; epipodite of first maxilliped usually with branchial filaments; coxopoditic setae with apical hooks; telson never completely divided by transverse suture Superfamily PARASTACOIDEA, Family PARASTACIDAE 17

S (2). Male never exhibiting cyclic dimorphism; first pleopod with distal portion rolled to

form cylinder; ischia of all pereiopods without hooks; female lacking annulus ventralis.

Family ASTACIDAE 4

Male always exhibiting cyclic dimorphism; first pleopod with distal portion either bearing shallow tperm groove crphalomesially or complexly and tightly folded with sperm

<<in.Inn opening on one of 2 or more terminal elements; male with hooks on one or

more of second through fourth pereiopods; female with or without annulus ventralis

Family CAMRARIDAE 6

4 (S). Distal part of first pleopod of male contracted to form narrow tube; exopodite of second

pleopod shorter than endopodite; merus of third maxilliped with uniform row of

spines on mesial margin and strong spine at distal end of row; epistome without spines

or ridges posterior to renal papilla Pacifastacus

Distal part of first pleopod of male not contracted but bearing 2 apical lobes; exopodite

of second pleopod as long as, or shorter than, endopodite; merus of third maxilliped

with mesial margin bearing row of spines or with 1 or 2 distally; epistome with spine

or ridges posterior to renal papilla 5

5 (4). Distal lobes of first pleopod conspicuously unequal in length; exopodite of second pleopod as long or almost as long as endopodite; merus of third maxilliped with single

large mesiodistal spine, latter occasionally accompanied by single smaller spine distally Astacus

Distal lobes of first pleopod subequal in length; exopodite of second pleopod distinctly

shorter than endopodite; merus of third maxilliped usually with row of spines along

mesial margin, rarely with only 1 distal spine Austropotamobhis

6(3). Branchial formula 17 + ep; male with hooks on ischia of third, fourth, or third and

fourth pereiopods; first pleopod with distal portion complexly and tightly folded and

sperm conduit opening on one of 2 or more terminal elements; annulus ventralis fixed

or movable, if latter, seldom through arc of as much as 40 degrees

Subfamily CAMBARINAE 8

Branchial formula 16 + ep or 18 + 3r + ep; male with hooks on ischia of second and

third pereiopods; first pleopod with distal part bearing shallow sperm groove or complexly folded as in Cambarinae but always bearing at least 3 terminal elements; annulus ventralis present (moving through arc of at least 75 degrees) or absent (plate lacking sinus and fossa) 7

7 (6). Branchial formula 16 + ep: first pleopod of male with distal portion complexly and

tightly folded and sperm conduit opening on one of 3 terminal elements; annulus ventralis present Subfamily CAMBARELLINAE Cambarellus

Branchial formula 18 +3r + ep; first pleopod of male with shallow sperm conduit not

opening on terminal element, annulus ventralis absent, plate lacking sinus and fossa

Subfamily CAMBAROIDINAE Cambaroides

8 (6). Third maxillipeds much enlarged; ischium without teeth on opposable margin

Troglocambarus

Third maxillipeds not conspicuously large; ischium with teeth on opposable margin 9

9 (8). First pleopod of male terminating in 2 elements 10

First pleopod of male terminating in more than 2 elements 15

10(9). Mesial process of first pleopod of male no more than one-half length of elongate central
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

projection; central projections of paired pleopods of first form male nearly straight and

overlapping Faxonella

Mesial process of first pleopod of male greater than one-half length of central projection;

central projections of paired pleopods overlapping only in occasional individuals of

Fallicambarus dissitus Penn in which strongly recurved 11

11(10). Distal one-third of first pleopod of male with prominent shoulder on cephalic surface.

or central projection forming distally projecting triangular plate, or central projection

arising from enlarged subterminal or terminal area Procambarui

Distal one-third of first pleopod of male lacking shoulder on cephalic surface; central

projection never forming distally projecting triangular plate nor arising from enlarged

subterminal or terminal area 12

12(11). Coxa of fourth pereiopod in male lacking caudomesial boss 13

Coxa of fourth pereiopod in male with caudomesial boss 14

13(12) First pleopods, of male, in resting position, deeply withdrawn between bases of pereiopods

and largely concealed by dense setiferous mat extending from ventrolateral margins of

sternum Hobbmu

First pleopods of male, in resting position, never deeply withdrawn between bates of

pereiopods and never concealed by dense sctifcrou* mat extending from ventrolateral

margins of sternum OrconecUt

14(12). Opposable margin of dactyl of chela with abrupt excision in proximal one-half

FaltUambanu

Opposable margin of dactyl of chela without abrupt excision in proximal om- half 15

15(14). Central projection of first pleopod bladelike, always directed caudally or caudodistally,

and frequently bearing subterminal notch Cambanu

Central projection of first pleopod seldom bladelike, if so, directed laterodistally or lacking subterminal notch 16

16(15). First pleopod with both central projection and mesial process curved caudally at least

90 degrees to principal axis of shaft of appendage FallUambmui

First pleopod never with both central projection and mesial process curved caudally at

angle of so much as 90 degrees to principal axis of shaft of appendage . Ptocambanu

17 (2). Branchiocardiac groove without conspicuous anterolateral extension; chela with dactyl

moving in subhorizontal plane; branchial formula 21 + epr 18

Branchiocardiac groove with anterolateral extension situated subparallel to, or converging on, cervical groove for some distance; chela with dactyl moving in subvertical or

subhorizontal plane; branchial formula variable 21

18(17). Telson entirely calcified, lacking trace of transverse suture; stems of podobranchiae without winglike expansion Aitacoprii

Telson with posterior part membranous, with or without trace of transverse suture;

stems of podobranchiae with winglike expansion 19

19(18). Distal portion of inner ramus of uropod membranous; cephalothorax rarely with small

tubercles; abdominal pleura without spines or tubercles; ventral surface of third

maxilliped with setae covering most of mesial half; male genitalia consisting of large

papilla arising from mesial surface of coxa C.herax

Distal portion of inner ramus of uropod calcified; cephalothorax with enlarged spines or

tubercles; abdominal pleura with or without spines or tubercles; ventral surface of

ischium of third maxilliped with median row of large setiferous punctations; male

genitalia consisting of short papilla with calcified ring situated on ventral surface of

coxa 20

20(19). Abdomen with spines or tubercles, sometimes limited to pleura; carpus of cheliped with

enlarged submedian spine ventrodistally; ventrolateral margin of chela with one row

of spines and usually with additional row or low carina Euaslacus

Abdomen without spines or tubercles but with many setiferous punctations on pleura;

carpus of cheliped with irregularly arranged small spine or tubercles on or near median ventrodistal margin; ventrolateral margin of chela with single row of weak

sPine s

 Euastacoides

21 (17). Chela with dactyl moving in subvertical plane 22

Chela with dactyl moving in subhorizontal plane 26

22 (21). Pleural lobe of first abdominal segment not overlapped by anterior lobe of pleuron of

second segment; palp of third maxilliped not reaching distal end of ischium Engaetu

Pleural lobe of first abdominal segment partially overlapped by anterior lobe of pleuron
NUMBER 164

of second segment; palp of third maxilliped overreaching distal end of ischium 23

23 (22). Anterolateral lobe of branchiocardiac groove (ubcontiguous with cervical groove; branchial formula 20 + epr Fmrmstocut

Anterolateral lobe of branchiocardiac groove widely separated from cervical groove;

branchial formula 21 + epr or 18 + epr 24

24 (23). Postorbital ridge* well developed; eyestalks long; palm of chela depressed; carpus of

chelipcd produced diitomesially Geochmrmx

Postorbital ridges very weak or absent; eyestalks short and bulbous; palm of chela inSated; carpus of cheliped rounded distomesially 25

25 (24). Lateral rostral carina present; lateral process of sternum between fourth pereiopods not

separated by median longitudinal carina; branchial formula 21 + ep Engmemm

Lateral rostral carina absent; lateral processes of sternum between fourth pereiopods

separated by median longitudinal carina; branchial formula 18 + ep Temribrmmchhtnu

26(21). Ventrolateral surface of chela with 2 rows of spines, and additional rows of spines or

enlarged tubercles on dorsal and ventral surfaces; branchial formula 20 + epr

Pmrmttephrpps

Surface of chela never with longitudinal rows of large'spines or tubercles; branchial

formula other than 20 + epr 27

27 (26). Carapace and pleuron of second abdominal segment with large spines or tubercles; anterolateral extension of branchiocardiac groove almost contiguous with cervical groove; telson with trace of transverse suture laterally; branchial formula 12 + epr + 5r

Carapace and pleuron of second abdominal segment never with large spines or tubercles;

anterolateral extension of branchiocardiac groove with wide to moderately narrow gap

between it and cervical groove; telson without trace of transverse suture; branchial

formula other than 12 + epr + 5r 28

28 (27). Junction or approximation of branchiocardiac and cervical grooves situated no farther

laterally than level of caudolateral extremity of postorbital ridge; branchial formula

17 + epr Pmrmrtacotdet

Junction or approximation of branchiocardiac and cervical grooves situated distinctly

lateral to level of caudolateral extremity of postorbital ridge; branchial formula 20 +

epr or 21 -I- epr 29

29 (28). Anterolateral part of branchiocardiac groove subparallel to cervical groove with comparatively little space between them; branchial formula 20 + epr Sammitmcus

Anterolateral part of branchiocardiac groove widely separated from, but converging

throughout its length with, cervical groove; branchial formula 21 + epr SO

30(29). Male genitalia large, longer than coxa; pleuron of second abdominal segment enlarged

in female Grmnuutmou

Male genitalia small, much shorter than coxa; pleuron of second abdominal segment not

enlarged in female Gtthmmx

Superfamily ASTACOIDEA

AsTACorocA De Haan, 1841:148, 160.

ASTAQNA Huxley, 1880a:254. 255.

DIAGNOSIS.—Carapace lacking dorsomedian longitudinal suture or ridge in cardiac and posterior

gastric regions. Sternal plate between fifth pereiopods not fused with sternal complex anteriorly.

Telson almost always partially, and usually completely, divided by transverse suture. Podobranchiae

of first three pereiopods not differentiated into

branchial and epipoditic portions but with bilobed

plaited laminae. Branchial formula 16 + ep; 17 +

ep; 18 + 2r + ep; or 18 + 3r + ep. Articles of

lateral ramus of antennule, in region of asthetascs,

usually bearing two dusters, only one in Cambaroidinae. First pleopod present in male and possessing individual sperm groove, present or absent in

female; second pleopod in male with spiral element,

frequently borne on subtriangular lobe. Eggs large

and hatching as miniatures of adult. All members

occurring in fresh water, some migrating into salt

water for part of year.

Family ASTACIDAE

astacini Latreille, 1802-1803:32.

AsUkoiden Dumeril, 1806:172.

Astacoidea Dumeril, 1806:332.—De Haan, 1841:148, 160.

Astacia Rafinesque, 1815:98.

Samouelle, 1819:94.
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Astaddes Billberg, 1820:134, 135.

ASTACINAE Latreille, 1825:279.

Astacida Vogt, 1851:459.

Astacina Gerstaeker, 1863:373.

POTAMOBIIDAE Huxley, 1879:776, 781, 785.

POTAMOBIINAE Ortmann, 1891:10.

PATAMOBIIDAE Spandl, 1926:141 [erroneous spelling].

DIAGNOSIS.—Male never exhibiting cyclic dimorphism; first pleopod with distal portion rolled

to form cylinder, and distalmost part contracted to

form tube or produced into 2 simple spoonlike

lobes. Ischia of pereiopods of male without hooks.

Female lacking annulus ventralis (sclerite present

but lacking sinus and fossa). Branchial formula 18

-f 2r + ep; or 18 + 3r + ep. Some articles of lateral ramus of antennule bearing 2 clusters of asthetascs.

Genus Astacus

FIGURE 1

Astacus Fabricius, 1775:413. [Type-species, by Direction 12 of

the International Commission on Zoological Nomenclature,

Cancer Astacus Linnaeus, 1758:631. Gender: masculine.

Junior homonym of Astacus Pallas, 1772:81, latter suppressed by Commission in same direction; Astacus Fabricius, 1775:413, also placed on Official List of Generic Names

in Zoology.]

astaca Dum£ril, 1806:183 [erroneous spelling].

Potamobie Leach, 1818:75 [vernacular name and nomen

nudum].

Potamobius Samouelle, 1819:95. [Type-species, by monotypy.

Cancer Astacus Linnaeus, 1758:631. Gender: masculine.]

Potamobia Desmarest, 1823:246, 424 [latinization of Potamobie Leach, 1818].

Patamobius Gray, 1845:410 [erroneous spelling].

Carabis Marchand, Lamy, and de Boisvillette, 1874:55. [Typespecies, by monotypy, Cancer Astacus Linnaeus, 1758:631.

Gender: masculine.]

Actacus Philippi, 1894:378 [erroneous spelling].

Astachus Ninni, 1923:13 [erroneous spelling].

Subgenus Pontastacus Bott, 1950:12. [Type-species, by original

designation, Astacus Icptodactylus Eschscholtz, 1823:109.

Gender: masculine.]

DIAGNOSIS.—Carapace with at least rudiments of

both anterior and posterior postorbital ridges;

merus of third maxilliped with single large mesiodistal spine, sometimes accompanied by single

smaller spine distally; epistome with spine or ridge

(often bearing at least one spine) posterior to renal

papilla; first pleopod of male terminating in two

FIGURE 1.—Astacus (A.) astacus (Linnaeus, 1758) from Ljumgan River, Alby, Sweden: a, dorsal view of carapace and

chelipcds of male; b, ventral view of merus of third maxilliped; c, lateral view of first pleopod of male; d, basal

podomere* of second through fifth pereiopods; e, caudal

thoracic sternal area of female; /, epistome and proximal

podomerc of antennae.

lobes, one reaching distinctly farther distally than

other; exopodite of second pleopod extending almost or quite as far distally as endopodite.

RANGE.—Middle and eastern Europe.

NUMBER OF SPECIES.—Bott (1950) recognized

four species and four subspecies assigned to two subgenera, and two additional species of uncertain relationships. Karaman (1962) treated four species

and six subspecies and "natio" which were assigned

to two subgenera.

REFERENCES.—Bott, 1950, 1972; Karaman, 1962;

Curra, 1967.
NUMBER 164

FIGURE 2.—Austropotamobius torren tins danubicus Karaman,

1962, from Bucharest, Rumania. (For explanation, see Figure

Genus Austropotamobius

FIGURE 2

Austropotomobius Skorikov, 1908:116. [Type-species, by subsequent designation (Bott, 1950:18), Cancer torrentium

Schrank, 1803:247. Gender: masculine. Originally proposed

as a subgenus of Potamobius; elevated to generic rank by

Bott, 1950:18.]

Subgenus Atlantoastacus Bott, 1950:21. [Type-species, by original designation, Astacus pallipes Lereboullet, 1858:7. Gender: masculine.]

DIAGNOSIS.—Carapace without trace of posterior

postorbital ridge; merus of third maxilliped with

row of spines along mesial border; epistome with

spine or ridge, latter often bearing at least one

spine, posterior to renal papilla; first pleopod of

male terminating in 2 lobes reaching approximately

same level distally; exopodite of second pleopod

never extending so far distally as endopodite.

FIGURE S.—Pacifastacus leniusculus klamathensis (Stimpson)

from Klamath River, Klamath County, Oregon, U.S.A. (For

explanation, see Figure 1).

RANGE.—West and middle Europe.

NUMBER OF SPECIES.—Bott (1950, 1972) recognized three species and five subspecies assigned to

two subgenera. Karaman (1962) treated three species and eight subspecies [one of his subspecies, A.

pallipes bispinosus Karaman, was declared a synonym of Cambaroides schrenckii (Kessler, 1874:361)

by Holthuis, 1964b: 46]; no subgenera were recognized by him.

REFERENCES.—Bott, 1950, 1972; Karaman, 1962;

Curra, 1967.

Genus Pacifastacus

FIGURE 3

Pacifastacus Bott, 1950:3, 24. [Type-species, by original designation, Astacus klamathensis Stimpson, 1857:87. Gender:

masculine.]
10

DIAGNOSIS.—Carapace usually with at least rudiments of both anterior and posterior postorbital

ridges; merus of third maxilliped with row of spines

along mesial margin; epistome without spine or

ridge posterior to renal papilla; first pleopod of

male with distal portion contracted to form narrow tube, lacking lobes apically; exopodite of second pleopod not reaching distal end of endopodite.

RANGE.—Pacific drainage of North America and

headwaters of the Missouri River in Wyoming. Introduced into Sweden and Japan.

NUMBER OF SPECIES.—Five species, one of which

comprises three subspecies.

REFERENCES.—Faxon, 1885a, 1898; Bott, 1950;

Hobbs, 1972b.

Family CAMBARIDAE

CAMRARINAE Hobbs. 1942a: 338.

GAMBARINAE Penn, 1957:93 [erroneous spelling].

DIAGNOSIS.—Male exhibiting cyclic dimorphism

(first and second form males); first pleopod either

bearing shallow sperm groove mesially or distal portion tightly folded with distal end of sperm groove

opening on one (central projection) of 2-4 terminal elements. Male with hooks present on ischium of one or more of second through fourth

pereiopods. Female with or without annulus ventralis and first pleopod. Branchial formula 18 +

3r 4- ep; 17 + ep; or 16 + ep. Some articles of lateral ramus of antennule bearing either 1 or 2 clusters of asthetascs.

Subfamily CAMBARINAE

DIAGNOSIS.—First pleopod of male with distal

portion tightly folded, and distal end of sperm

groove opening on one (central projection) of 2-4

terminal elements; hooks present on ischia of third,

fourth, or third and fourth pereiopods. Female with

fixed or movable annulus ventralis, if latter seldom

through arc of as much as 40 degrees; with or without first pleopod. Branchial formula 17 + ep or

16 4- ep. Some articles of lateral ramus of antennule

bearing 2 clusters of asthetascs.

Genus Barbicambanu

FIGURE 4

Barbicambarus Hobbs, 1969:95, 98. [Type-species, by original

designation, Cambarus cornutus Faxon, 1884:120. Gender:

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

masculine. Proposed as subgenus of Cambarus; elevated to

generic rank by Bouchard, 1972:56.]

DIAGNOSIS.—Antenna heavily fringed on mesial

border. Third maxilliped with teeth on mesial margin of ischium. Mesial margin of palm of chela and

lateral margin of fixed finger with single row of

spiniform tubercles, no more than 12 tubercles in

former; opposable margin of dactyl never with

prominent excision. Areola broad at midlength.

Ischium of third pereiopod with hook. Coxa of

fourth pereiopod with caudomesial boss bearing

deep setiferous pit. First pleopods of first form male

symmetrical, widely separated at base, and with

distal portion of shaft inclined caudally; terminal

elements consisting of very short, distally rounded

FIGURE 4.—Barbicambarus cornutus (Faxon, 1884) from Green

River, 3.4 miles northeast of Donansburg, Green County,

Kentucky, U.S.A.: a, dorsal view of carapace and chelipeds

of male, form I; b, lateral view of first pleopod of male,

form II; c, lateral view of first pleopod of male, form I; d,

proximal podomeres of second through fifth pereiopods; e,

annulus ventralis of female.
NUMBER 164 11

central projection curved caudally at much more

than right angle, and truncate, partially cornified

mesial process bearing fingerlike projection reaching caudally beyond tip of central projection. Female with annulus ventralis immovable; first pleopod rudimentary. Branchial formula 17 -f ep.

RANGE.—North America: Barren and Green river

systems in Kentucky and Tennessee.

NUMBER OF SPECIES.—Monotypic.

REFERENCES.—Faxon. 1884, 1885a; Hobbs, 1969,

1972b.

Genus Cambarus

FlCUME 5

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with

teeth on mesial margin of ischium. Mesial margin

of palm of chela with row of fewer than 12 tubercles except in albinistic species in which more

present; lateral margin of fixed finger never bearing spiniform tubercles; opposable margin of dactyl almost never with prominent excision. Areola

broad to obliterated or linear at midlength. Ischium

of third pereiopod with hook. Coxa of fourth

pereiopod of male with caudomesial boss. First

pleopods of first form male symmetrical, contiguous

basally, with distal portion of shaft never inclined

caudally, and terminating in 2 or 3 distinct parts

(mesial process, central projection, and occasionally

Cambarus Erichson. 1846:97. [Type ipeciei, by subsequent

designation (Faxon, 1898:644), Astacus Bartonii Fabricius,

1798:407. Gender: masculine. Proposed as subgenus of

Attacus; elevated to generic rank by Girard, 1852:88.]

Gambarus Huxley. 1880b:81 [erroneous spelling].

Camburus Faxon, 1885b: 358 [erroneous spelling].

Camberus Miller, 1895:336 [erroneous spelling].

Bartonius Ortmann, 1905a:97. [Type-spedes, by original designation, "C. bartoni" Fabricius, 1798:407. Gender: masculine. Proposed as subgenus of Cambarus, treated unintentionally as generic name by Williamson, 1907:749, and declared a synonym of Cambarus by Fowler, 1912:341.]

Canbarus Thompson, 1967:47 [erroneous spelling].

Cambaroides U nest am, 1969:204 [lapsus for Cambarus].

cambarus Padgett, 1970:19.

Cambaras Bouchard, 1973:106 [erroneous spelling].

Subgenus Aviticambarus Hobbs, 1969:96, 99. [Type-species, by

original designation, Orconectes hamulatus Cope, 1881:881.

Gender: masculine.]

Subgenus Depressicambarus Hobbs, 1969:96, 102. [Typespecies, by original designation, Astacus latimanus LeConte,

1856:402. Gender: masculine.]

Subgenus Erebicambarus Hobbs, 1969:95, 99. [Type-spedes,

by original designation, Cambarus bartoni tenebrosus Hay,

1902:232. Gender: masculine.]

Subgenus Hiaticambarus Hobbs, 1969:95, 105. [Type-spedes,

by original designation, Cambarus longulus Girard, 1852:90.

Gender: masculine.]

Subgenus Jugicambarus Hobbs, 1969:95, 106. [Type-spedes.

by original designation, Cambarus bartonii asperimanus

Faxon, 1914:391. Gender: masculine.]

Subgenus Lacunicambarus Hobbs. 1969:96, 110. [Type-spedes,

by original designation, Cambarus diogenes Girard, 1852:88.

Gender: masculine.]

Subgenus Puncticambarus Hobbs, 1969:96, 101. [Type-spedes,

by original designation, Cambarus extraneus Hagen, 1870:

73. Gender: masculine.]

Subgenus Veticambarus Hobbs, 1969:95, 96. [Type-spedes, by

original designation, Cambarus pristinus Hobbs, 1965: 268.

Gender: masculine.]

FIGURE 5.—Cambarus (C.) bartonii bartonii (Fabricius, 1798)

from 8.6 miles south of Waynesboro, Augusta County, Virginia, U.S.A. (For explanation, see Figure 4.)
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

caudal knob; cephalic process always absent), 2

prominent ones bent caudally or caudolaterally

between 45 and 100 degrees or with central projection forming arc approaching 180 degrees; central

projection bladelike or tapering from base, with or

without subapical notch; mesial process subconical,

bulbiform, or conspicuously inflated at base, seldom

corneous, never appearing twisted or subspatulate

distally, and lacking eminence on cephalic (morphological) border; caudal element seldom present,

but occasionally represented by knoblike prominence at caudolateral base of central projection.

Female with annulus ventralis immovable or with

caudal half slightly movable; first pleopod present,

rudimentary, or absent. Branchial formula 17 -f ep.

RANGE.—North America: From the coastal region of New Brunswick, Canada, southward to the

panhandle of Florida, westward to Texas, and

northward to Minnesota and southern Ontario.

FIGURE 6.—Fallicambarus fodiens (Cottle, 1863) from 5 miles

west of Ann Arbor, Washtcnaw County, Michigan, U.S.A.

(For explanation, see Figure 4.)

NUMBER OF SPECIES.—Fifty-five species and three

subspecies.

REFERENCES.—Ortmann, 1931; Hobbs, 1969,

1972b.

Genus Fallicambarus

FIGURE 6

Fallicambarus Hobbs, 1969:111. [Type-species, by original

designation, Cambarus sirawni Reimer, 1966:11. Gender:

masculine.]

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with

teeth on mesial margin of ischium. Mesial margin

of palm of depressed chela with row of fewer than

12 tubercles; lateral margin of fixed finger never

witli row of spines or tubercles; opposable margin

of dactyl usually with prominent excision. Areola

linear or obliterated at midlength. Ischium of third

pereiopod, and sometimes that of fourth, of male

with hook. Coxa of fourth pereiopod of male with

prominent caudomesial boss. First pleopods of first

form male symmetrical, contiguous basally, and

terminating in 2 or S elements (mesial process, central projection, and, occasionally, cephalic process)

bent caudally or caudomesially at angle greater

than 95 degrees to main shaft or forming broad arc

of almost 180 degrees; central projection bladelike

or tapering, and usually lacking subapical notch;

mesial process never bulbiform but appearing

twisted and often with eminence on cephalic (morphological) border; cephalic process at least partially corneous, situated at mesial base of mesial

process, and directed caudad or caudodistad. Female with annulus ventralis only slightly movable;

first pleopod present. Branchial formula 17 + ep.

RANGE.—North America: From Texas to Ontario

and southward to the panhandle of Florida.

NUMBER OF SPECIES.—Nine.

REFERENCES.—Hobbs, 1969, 1972b.

Genus Faxonella

FIGURE 7

Faxonella Creaser, 1933:21. [Type-species, by monotypy, Cambarus clypeatus Hay, 1899:122. Gender: feminine. Proposed

as a subgenus of Faxonius; elevated to generic rank by

Fitzpatrick, 1963:61, and questionably by Creaser, 1962:3.]
NUMBER 164 IS

FIGURE l.—Faxonella clypeata (Hay, 1899) from 6.2 miles east

of Midway, Barbour County, Alabama, U.S.A. (For explanation, see Figure 4; except b, caudal view of first pleopod of

male, form I.)

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with

teeth on mesial margin of ischium. Mesial margin

of palm of chela with or without much reduced

tubercles; lateral margin of fixed finger never with

row of spines or tubercles; opposable margin of dactyl never with prominent excision. Areola moderately broad at midlength. Ischium of third pereiopod of male with hook. Coxa of fourth pereiopod

of male without caudomesial boss. First pleopods

of first form male symmetrical, contiguous basally,

and terminating in one long (central projection)

and one short (mesial process) element, latter no

more than half length of former; central projection

of paired appendages always overlapping. Female

with annulus ventralis freely movable; first pleopod

rudimentary to virtually absent. Branchial formula

17 + ep.

FIGURE 8.—Hobbseus cristatus (Hobbs, 1955) from Scooba

Creek, 2 miles south of Scooba, Kemper County, Mississippi,

U.S.A. (For explanation, see Figure 4.)

RANGE.—North America: Oklahoma and Texas

east to Gadsden County, Florida, and Richland

County, South Carolina.

NUMBER OF SPECIES.—Three.

REFERENCES.—Fitzpatrick, 1963; Hobbs, 1972b.

Genus Hobbseus

FIGURE 8

Hobbseus Fitzpatrick and Payne, 1968:15. [Type-species, by

original designation, Cnmbarus cristatus Hobbs, 1955:95.

Gender: masculine.]

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with

teeth on mesial margin of ischium. Mesial margin

of palm of depressed chela with row of more than

12 tubercles; lateral margin of fixed finger never

with row of spiniform tubercles; opposable margin

of dactyl never with prominent excision. Areola
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

moderately broad at midlength. Ischium of third

pereiopod with hook. Coxa of fourth pereiopod

never with prominent caudomesial boss. First pleopod of first form male symmetrical or slightly asymmetrical, deeply withdrawn between bases of

pereiopods and largely hidden by dense setal mat

extending from ventrolateral margin of sternum,

and terminating in two distinct parts (mesial process and central projection) almost straight, bent

caudally as much as 159 degrees, or forming broad

arc of as much as 180 degrees; central projection

tapering to point distally and lacking subterminal

notch; mesial process never bulbiform, with slender

distal portion, and lacking eminence on cephalic

(morphological) border. Female with annulus ventralis freely movable; first pleopod present. Branchial formula 17 + ep.

RANGE.—North America: Eastern Mississippi and

western Alabama.

NUMBER OF SPECIES.—Five.

REFERENCES.—Fitzpatrick and Payne, 1968;

Hobbs, 1969, 1972b.

Genus Orconectes

FIGURE 9

Orconectes Cope, 1872:419. [Type-species, by subsequent designation (Fowler, 1912:339), Orconectes inermis Cope,

1872:419. Gender: masculine.]

Camtarus Packard, 1888:156 [erroneous spelling].

Cambrus Price, 1900:155 [erroneous spelling].

Orconectis Harris, 1903:11 [erroneous spelling].

Camborus Clawson, 1905:103 [erroneous spelling].

Faxonius Ortmann, 1905a:97. [Type-spedes, by original designation, Astacus limosus Rafinesque, 1817:42. Gender: masculine. Proposed as subgenus of Cambarus, elevated to

generic rank by Creaser, 1933:1, and declared synonym of

Orconectes by Hobbs, 1942a:352.]

G[ambarus] Ortmann, 1905a: 112 [incomplete spelling].

Cityphlobius Joleaud, 1939:14 [error for C. typhlobius].

Cambarrus Fleming, 1939:305 [erroneous spelling].

Orconecctes Williams, 1952:330 [erroneous spelling].

Gambarus Croizat, 1958:908 [erroneous spelling].

Camburus Bacescu, 1967:218 [erroneous spelling].

Oronectes Bacescu, 1967:218 [erroneous spelling].

Orchonectes Dimond et al., 1968:760 [erroneous spelling].

Orconectus Ray and Stevens, 1970:58 [erroneous spelling].

Orconnectes Molley and Prins, 1973:70 [erroneous spelling].

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with

teeth on mesial margin of ischium. Mesial margin

of palm of chela usually with row of less than 12

V

FIGURE 9.—Orconectes limosus (Rafinesque, 1817) from Cattail

Branch at Route 97, Howard County, Maryland, U.S.A. (For

explanation, see Figure 4.)

tubercles; lateral margin of fixed finger never bearing row of spiniforra tubercles; opposable margin

of dactyl seldom with prominent excision. Areola

broad to obliterated at midlength. Ischium of third,

rarely third and fourth, pereiopod with hook. Coxa

of fourth pereiopod of male lacking caudomesial

boss except in troglobitic members. First pleopod of

first form male almost always symmetrical, never

deeply withdrawn between bases of pereiopods nor

concealed by dense setal mat extending from ventrolateral margins of sternum, and contiguous basally;

terminal elements (usually 2, ocassionally S in troglobitic members) highly variable in length and

disposition—divergent, straight, or curved caudodistally or caudally; central projection never

abruptly curved caudally at base nor forming arc of

more than 90 degrees. Female with annulus ventralis immovable or slightly movable in troglobitic
NUMBER 164 15

species; first pleopod usually present. Branchial formula 17 + ep.

RANGE.—North America: Arizona and Alberta

eastward to the Atlantic Ocean except eastern seaboard from South Carolina to Florida; also in

Rogue River in Oregon. Introduced into western

Europe.

NUMBER OF SPECIES.—Sixty-three species and

seven subspecies.

REFERENCES.—Ortmann, 1931; Rhoades, 1944;

Williams, 1954; Fitzpatrick, 1967; Crocker and

Barr, 1968; Hobbs, 1972b.

Genus Procambartu

FIGURE 10

Cambarus Ortmann. 1905a:96 (not Erichion, 1846). [Typespecies, by original designation. Astacus Mandingii Harlan,

1830:464. Gender: masculine. Proposed as typical subgenus,

but type-species previously selected by Faxon, 1898:644,

Astacus Bartonii Fabriciui, 1798:407.]

Procambarus Ortmann. l9O5c:435, 437. [Type-species, by subsequent designation (Fowler, 1912:340), Cambarus Digueti

Bouvier, 1897:225. Gender: masculine. Proposed as subgenus of Cambarus, elevated to generic rank by Hobbs,

1942a:34l.]

Paracambarus Ortmann, 1906:1. [Type-species, by monotypy,

Cambarus (Paracambarus) paradox us Ortmann, 1906:3.

Gender: masculine. Proposed as subgenus of Cambarus,

elevated to generic rank by Hobbs, 1942a:S44, and reduced

to subgenus of Procambarus by Hobbs, 1972a: 10.]

Ortmannicus Fowler, 1912:34, 341. [Type-species, by original

designation, Astacus Blandingii Harlan, 1830:464. Gender:

masculine. Replacement name for Ortmann's subgenus

Cambarus, utilized as generic name by Rhoades, 1944:114,

and as a subgenus of Procambarus by Hobbs, 1972a:2-5, 9.]

Cambarcllu* Creaser, 1933:21 [lapsus for Cajnbarus].

procambarus Padget. 1970:19.

Subgenus Gitardiella Lyle, 1938:76. [Type-species, by monotypy, Cambarus Hagenianus Faxon, 1884:141. Gender:

feminine.]

Subgenus Acucauda Hobbs, 1972a:S-5. [Type-species, by

original designation, Procambarus fttzpatricki Hobbs, 1971a:

461. Gender: feminine.]

Subgenus Austrocambarus Hobbs, 1972a:2-5. [Type-species.

by original designation, Procambarus vazquezae Villolobos,

1954:328. Gender: masculine.]

Subgenus Capillicambarus Hobbs, 1972a:3-4, 6. [Type-spedes,

by original designation, Cambarus (Cambarus) hind Ortmann, 1905b:401. Gender: masculine.]

Subgenus Hagenides Hobbs, 1972a:2-~4. 7. [Type-spedes, by

original designation, Astacus advena LeConte, 1856:402.

Gender: masculine.]

Subgenus Leconticambarus Hobbs, 1972a:2-5. 7. [Type-spedes,

by original designation, Cambarus barbatus Faxon, 1890:

621. Gender: masculine.]

FIGURE 10.—Procambarus (Ortmannicus) blandingii (Harlan,

1830) from Town Creek, 3 miles south of Camden, Kershaw

County, South Carolina. U.SA. (For explanation, see Figure

*)•

Subgenus Lonnbergius Hobbs, 1972a:2-4, 8. [Type-spedes, by

original designation, Cambarus acherontis Lonnberg, 1895:

6. Gender: masculine.]

Subgenus Mexicambarus Hobbs. 1972a:3. 4, 8. [Type-spedes,

by original designation, Cambarus (Cambarus) bouvieri

Ortmann, 1909:159. Gender: masculine.]

Subgenus Pennides Hobbs, 1972a:2-4, 10. [Type-spedes, by

original designation, Procambarus natchitochae Penn, 1953:

5. Gender: masculine.]

Subgenus Remoticambarus Hobbs, 19?2a:3, 4, 11. [Typespedes, by original designation, Procambarus pecki Hobbs,

1967:2. Gender: masculine.]

Subgenus Scapulicambarus Hobbs, 1972a:2, 3, 5. [Type-spedes,

by original designation, Cambarus clarkii paeninsulanus

Faxon, 1914:369. Gender: masculine.]

Subgenus Tenuicambarus Hobbs, 1972a:S. 5, 12. [Type-spedes,

by original designation, Procambarus tenuis Hobbs, 1950:

194. Gender: masculine.]

Subgenus Villalobosus Hobbs, 1972a:3, 5, 12. [Type-spedes, by

original designation, Paracambarus riojae Villalobos, 1944:

161. Gender: masculine.]

DIAGNOSIS.—Antenna never with conspicuous

fringe on mesial border. Third maxilliped with
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

teeth on mesial margin of ischium. Mesial margin

of palm of chela with none to many tubercles; lateral margin of fixed finger never bearing spiniform

tubercles; opposable margin of dactyl occasionally

with prominent excision. Areola broad to obliterated at midlength. Ischia of third, fourth, or third

and fourth pereiopods of male with hook. Coxa of

fourth pereiopod of male with or without caudomesial boss. First pleopods of first form male

symmetrical or asymmetrical, sometimes deeply

withdrawn between bases of pereiopods and at least

partially concealed by setae extending from ventrolateral margin of sternum; subcontiguous, contiguous, or partially overlapping basally, and terminating in 2 or more, usually 3 or 4 elements; presence

of subterminal setae in many members of genus

unique; terminal elements highly variable in form

and disposition; if only 2 elements present (mesial

process and central projection), shoulder present

on distal third of cephalic surface of appendage, or

central projection forming distally projecting triangular plate, or central projection arising from caudal margin of enlarged terminal region; if more

than 2 elements present, central projection seldom

bladelike, if so, directed laterodistally or lacking

subterminal notch, and elements never bent caudally at angle of so much as 90 degrees. Female with

annulus ventralis freely movable, although sometimes partially covered ventrally by caudally projecting prominences from sternal plate immediately

cephalic to it; first pleopod usually present. Branchial formula 17 + ep.

RANGE.—North America: Guatemala and Cuba

to Minnesota and southern New England, except in

mountains. Introduced into California, Hawaii, and

Japan.

NUMBER OF SPECIES.—One hundred seventeen

species and six subspecies.

REFERENCES.—Hobbs, 1942a, 1942b, 1962, 1972a,

1972b; Villalobos, 1955; Penn, 1959.

Genus Troglocambarus

FIGURE 11

Troglocambarus Hobbs, 1942a:354, 339, 341, 345. [Typespecies, by original designation, Troglocambarus maclanei

Hobbs, 1942a:345. Gender: masculine.]

DIAGNOSIS.—Albinistic, eyes reduced and without

pigment. Antenna never with conspicuous fringe

FICURE 11.—Troglocambarus maclanei Hobbs, 1942, from

Squirrel Chimney (Sec. 21, Twp. 9S, R. 18E), Alachua County,

Florida, U.S.A. (For explanation, see Figure 4.)

on mesial border. Third maxilliped greatly enlarged

and without teeth on mesial margin of ischium.

Mesial margin of palm of chela weakly tuberculate

or smooth; lateral margin of fixed finger lacking

row of spines; opposable margin of dactyl without

prominent excision. Areola broad at midlength.

Ischium of third and fourth pereiopods with hooks.

Coxa of fourth pereiopod of male with caudomesial

boss. First pleopod of first form male asymmetrical,

not conspicuously withdrawn between bases of

pereiopods, partially overlapping basally, and terminating in 4 distinct elements: mesial process

slender, acute, and directed caudolaterally; cephalic

process acute and partially hooding cephalic base

of central projection; central projection large, some-
NUMBER 164 17

what beaklike, with apex directed caudally; and

caudal element consisting of rudimentary caudal

knob and bladelike caudal process situated along

caudolateral base of central projection. Female with

annulus ventralis freely movable; first pleopod rudimentary. Branchial formula 16 + ep.

RANGE.—North America: Peninsular Florida.

NUMBER OF SPECIES.—Monotypic.

REFERENCES.—Hobbs, 1942a, 1972b.

Subfamily CAMBARELLINAE

CAMBARFM INAE Laguarda. 1961:69. [Originally proposed as a

iubfamily of the Astacidae.]

DIAGNOSIS.—Male with hooks on ischia of second

and third pereiopods; first pleopod with distal portion tightly folded, and distal end of sperm groove

opening on one (central projection) of S terminal

elements (mesial and caudal processes and central

projection). Female with annulus ventralis freely

FIGURE 12.— Carnbarellus montezumae (Saussure, 1858) from

near Pachuca, Hidalgo, Mexico. (For explanation, see Figure

movable through arc of at least 75 degrees; first

pleopod lacking. Branchial formula 16+ ep (posterior arthrobranch absent at base of fourth pereiopod). Some articles of lateral ramus of antennule

bearing 2 clusters of asthetascs.

Genus Cambarellus

FicuftE 12

Cambarellus Ortmann, 1905a:97, 106. [Type-species, by original designation, Cambarus Montezumae Saussure, 1857:102.

Gender: masculine. Proposed as subgenus of Cambarus;

elevated to generic rank by Hobbs, 1942a:S50.]

Cambarella Carlisle and Knowles, 1959:50, 51 [erroneous

spelling].

DIAGNOSIS.—Same as that of subfamily.

RANGE.—North America: Pacific slope and central plateau of Mexico, coastal area along the Gulf

of Mexico from Texas to the Suwannee River in

Florida, and northward in the Mississippi Basin to

southern Illinois.

NUMBER OF SPECIES.—Twelve.

REFERENCES.—Villalobos, 1955; Penn, 1959;

Hobbs, 1972b.

Subfamily CAMBAROIDINAE

CAMBAROIMNAE Villalobos, 1955:7 [originally proposed as a

subfamily of the Astaddae].—Laguarda, 1961:17. 69, 70.

DIAGNOSIS.—Male with hooks on ischia of second

and third pereiopods; first pleopod with distal portion not complexly folded, instead bearing shallow

sperm groove with distal extremity situated cephalomesially or cephalically and not on any of 3 or more

terminal elements. Female lacking annulus ventralis (sclerite present but without sinus and fossa);

first pleopods lacking. Branchial formula 18 + 3r

+ ep. Articles of lateral ramus of antennule bearing

only one cluster of asthetascs.

Genus Cambaroides

FIGURE IS

Astacus Pallas, 1772:81. [Type-species, by monotypy, Astacus

Dauuricus Pallas, 1772:81. Gender: masculine. Name suppressed by International Commission on Zoological Nomenclature in Direction 12 (1955).]

Cambaroides Faxon, 1884:149, 150. [Type-species, by subsequent designation (Faxon, 1898:665), Astacus japonicus De
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Cambaroides similis (Koelbel, 1892) bom near

Pusan, Korea. (For explanation, see Figure 4.)

Haan, 1842:164. Gendei: masculine. Originally proposed as

a subgenus of Astacus; elevated to generic rank by Skorikov,

1908:116.]

Gambaroides Parisi, 1917:17 [erroneous spelling].

Catnberoides Sowerby, 1922:58 [erroneous spelling].

DIAGNOSIS.—Same as that of subfamily.

RANGE.—Eastern Asia: Amur Basin, Korea, and

Japan.

NUMBER OF SPECIES.—Four species. Birstein and

Vinogradov (1934) recognized, in addition to the

nominate subspecies, two subspecies of C. dauricus

(Pallas) and one of C. schrenckii (Kessler, 1874:

363).

REFERENCES.—Faxon, 1884, 1885a, 1898; Skorikov, 1908; Birstein and Vinogradov, 1934.

Superfamily PARASTACOIDEA

PARASTACIDAE Huxley, 1879:771, 775.

PARASTACINAE Ortmann, 1891:5, 7.

AUSTROASTACIDAE Clark, 1936:8. [Synonymued. by implication.

with Parastacidae by Rick, 1959:255.]

EUASTACIDAE Rick, 1959:255. [Synonymized with Parastacidae

by Riek. 1969:861.]

DIAGNOSIS.—Carapace lacking dorsomedian, longitudinal suture or ridge in cardiac and posterior

gastric regions. Sternal plate between fifth pereiopods not fused with sternal complex anteriorly. Telson never completely divided by transverse suture.

Podobranchiae of first three pereiopods not differentiated into branchial and epipoditic portions and

lacking bilobed plaited laminae; epipodite of first

maxilliped usually with branchial filaments. Branchial formula ranging from 12 + epr + 5r to 21 +

ep. Articles of lateral ramus of antennule never

with more than one cluster of asthetascs. First pleo»

pod absent m male and female; second pleopod

similar to third. Eggs large and hatching as miniatures of adult. All members occurring in fresh

water.

Family PARASTACIDAE

DIAGNOSIS.—Same as that of superfamily.

Genus Astacoides

FIGURE 14

Astacoides Guerin-Mlneville, 1839:109. [Type-spedes, by

monotypy, Astacoides Goudotii GuerinMeneville, 1839:109,

a subjective junior synonym of Astacus madagascarensis H.

Milne Edwards and Audouin, 1839:152. Gender: matculinr]

DIAGNOSIS.—Carapace with conspicuous spines or

tubercles, in addition to rostral, postorbital, branchiosiegal, and cervical; anterolateral portion of

branchiocardiac groove extremely narrowly separated from, and parallel to, cervical groove dorsolaterally; viewed dorsally, cervical groove broadly

U-shaped and deeply impressed; postorbital ridges

weak to almost obsolete. Abdomen with spines or

tubercles on pleura of one or more segments; pleuron of first abdominal segment distinct and partially overlapped by that of second. Telson sometimes with spines dorsally, with or without faint

transverse suture laterally, and entirely calcified.

Third maxilliped with ventral surface bearing scattered setiferous punctations; distolateral portion

produced in heavy spine; exopodite reduced, not

reaching distolateral extremity of ischium. Chela
NUMBER 164 19

with ventrolateral margin tuberculate or costate

but never with 2 rows of heavy spines or tubercles;

when upper surface of carpus of cheliped held in

horizontal plane, dactyl moving subhorizontally;

carpus of cheliped with enlarged spines or tubercles

mesially and ventrally. Male genitalia, situated on

ventral surface of coxa, consisting of slender articulated, partially sclerotized cylinder with subapical

setae. Branchial formula 12 + epr + 5r; pleurobranch on segment XIV only, and anterior

arthrobranch on segment VIII and posterior

arthrobranchiae on segments X through XIII rudimentary; stem of podobranchiae without winglike

expansion.

RANGE.—Madagascar.

NUMBER OP SPECIES.—One species comprising

four subspecies.

REFERENCES.—Monod and Petit, 1929; Holthuis,

1964a.

FIGURE 14.—Astacoides madagascarensis betsileoensis Petit,

1923, purchased in market place in Tananarive, Madagascar:

a, dorsal view of carapace and chelipeds of male; b, lateral

view of first three abdominal segments; c, dorsal view of

telson; d, ventral view of ischium and exopodite of third

maxilliped; e, basal podomeres of fifth pereiopod illustrating

male genitalia on mesial surface of coxa.

FICURE 15.—Astacopsis flvviatilis Riek, 1969, from Hobart,

Tasmania, Australia. (For explanation, see Figure 14.)

Genus Astacopsis

FIGURE 15

Astacopsis Huxley, 1879:760. 764. [Type-species, by monotypy.

Astacus franklinii Huxley, 1879:764 (not Gray, 1845:409),

("Astacopsis gouldi Clark, 1936:35). Gender feminine.]

DIAGNOSIS.—Carapace with spines or tubercles,

other than rostral, postorbital, branchiostegal, and

cervical; anterolateral extremity of branchiocardiac

groove fusing with cervical groove rather high on

carapace. Abdomen with spines on pleura of only

first and second abdominal segments; pleuron of

first abdominal segment distinct and partially overlapped by that of second. Telson with no trace of

transverse suture and entirely calcified. Third

maxilliped with ventral surface of ischium provided

with one row of large punctations bearing clusters

of stiff setae, and often produced distolaterally in

acute prominence; exopodite reduced, never reaching distal end of ischium. Chela with ventrolateral

margin usually bearing 2 rows of spines, occasionally

only one; when upper surface of carpus of cheliped

held in horizontal plane, dactyl moving subhorizontally; carpus of cheliped with 2-4 large spines

mesially and 1 or 2 on, or adjacent to, ventrodistal

margin. Male genitalia consisting of short papilla
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

bearing calcified ring. Branchial formula 21 + epr;

stem of podobranch continuous to apex of gill but

without winglike expansion; 6 posterior arthrobranchiae very reduced.

RANGE.—Australia: Northwestern and central

southern part of Tasmania.

NUMBER OF SPECIES.—Four.

REFERENCES.—Clark, 1936, 1939; Riek, 1969.

Genus Cherax

FIGURE 16

Cherax Erichson, 1846:88. [Type-species, by monotypy, Astacus (Cheraps) Preissii Erichson, 1846:101. Gender: masculine. Proposed as subgenus of Astacus with two spellings;

elevated to generic rank with third spelling by Huxley,

1879:769; placed on official list of generic names by International Commission on Zoological Nomenclature, Opinion

519 (Hemming, 1958).]

Cheraps Erichson, 1846:101 [erroneous spelling].

Cherops Von Martens, 1869:516.—Bate, 1888:195 [erroneous

spelling].

Chaeraps Huxley, 1879:752, 755, 764, 767-771, 775, 776. 786

[erroneous spelling].

Chaerops Huxley, 1880b:79 [erroneous spelling].

Astaconephrops Nobili, 1899:244. [Type-species, by monotypy,

Astaconephrops Albertisii Nobili, 1899:244. Gender: masculine.]

FIGURE 16.—Cherax albidus Clark, 1951, from Hawthorne,

Victoria, Australia. (For explanation, see Figure 14.)

Parachaerapi Smith, 1912:145. 148. 150-153. 161. [Type-species.

by monotypy. Aslacus bicarinatus Gray, 1845:410. Gender:

masculine.]

Paracheraps Baer, 1945:505. 509. 510 [erroneous spelling].

DIAGNOSIS.—Carapace smooth, punctate, or finely

granulate, rarely with spines or tubercles other than

rostral, postorbital, branchiostegal, and cervical;

anterolateral extremity of branchiocardiac groove

fused with cervical groove rather high on carapace.

Abdomen rarely with spines or tubercles on pleura;

pleuron of first abdominal segment distinct and partially overlapped by that of second. Telson usually

without indication of transverse suture and with

posterior part membranous. Third maxilliped with

mesial half of ventral surface of ischium completely

covered with tufts of long setae; distolateral extremity of ischium subangular, only slightly, if at all,

produced; exopodite usually reaching beyond distal end of ischium. Chela with ventrolateral margin

lacking spines or large tubercles, sometimes with

noncalcified area; when upper surface of carpus of

cheliped held in horizontal plane, dactyl moving

subhorizontally; carpus of cheliped with 0-4 large

spines mesially and 1 to several on or adjacent to

ventrodistal margin. Male genitalia, situated on

mesial surface of coxa, consisting of complex uncalcified papilla. Branchial formula 21 + epr; stem

of podobranchiae produced in broad winglike expansion.

RANGE.—Australia and New Guinea: Eastern half

and extreme southwestern part of Australia; New

Guinea, Aru Islands, and Misool.

NUMBER OF SPECIES.—Thirty-nine species and

three subspecies.

REFERENCES.—Clark, 1936; Holthuis, 1949; Riek,

1969.

Genus Engaeus

FIGURE 17

Engaeus Erichson, 1846:88, 89, 102. [Type-species, by subsequent designation (Clark, 1936:37), Astacus (Engaeus)

fossor Erichson, 1846:102. Gender: masculine. Proposed as

subgenus of Astacus; elevated to generic rank by Huxley,

1879:769.]

Eugaeus Huxley, 1880b:79 [erroneous spelling].

Pseudengaeus Clark, 1936:47. [Type-species, by original designation, Pseudengaeus strictifrons Clark, 1936:47. Gender:

masculine. Synonymized with Engaeus by Riek, 1969:863.]

Austroastacus Clark, 1936:51. [Type-species, by original designation, Engaeus hemicirratulus Smith and Schuster, 1913:
NUMBER 164 21

FIGURE 17.—Engaeus orientalis Clark, 1941, from 4 miles north

of Bonong, Victoria, Australia. (For explanation, see Figure

14.)

12S. Gender: masculine. Synonymized with Engaeus by

Riek, 1969:863.]

DIAGNOSIS.—Carapace generally smooth, rarely

with granules or tubercles; anterolateral portion of

branchiocardiac groove widely separated from, and

parallel to, cervical groove almost to midheight of

carapace; viewed dorsally, cervical groove tending

toward V-shape and not deeply impressed; postorbital ridges obsolete. Abdomen smooth or punctate and conspicuously narrow; pleuron of first abdominal segment greatly reduced and not partially

covered by anterior lobe of pleuron of second. Telson with no trace of transverse suture and entirely

calcified. Third maxilliped with mesial half of ventral surface of ischium bearing dense tufts of stiff

and plumose setae; distolateral extremity of ischium

angular or rounded, only slightly produced; exopodite reduced, vestigial, or absent, never reaching

distal end of ischium. Chela with ventrolateral margin rounded, lacking spines or large tubercles;

when upper surface of carpus of cheliped held in

horizontal plane, dactyl moving subvertically; carpus of cheliped without enlarged spines or tubercles

mesially or ventrally, but often with row of small

tubercles dorsomesially or mesiodistally. Male genitalia consisting of small simple papilla. Branchial

formula 21 + epr; stem of podobranchiae not produced in winglike expansion.

RANGE.—Australia: Victoria, New South Wales,

and Tasmania.

NUMBER OF SPECIES.—Twenty-three.

REFERENCES.—Clark, 1936, 19S9; Riek, 1969.

Genus Engaetua

FIGURE 18

Engaewa Riek, 1967a: 106. [Type-species, by original designation, Engaewa subcoerulea Riek, 1967a: 107. Gender: feminine.]

DIAGNOSIS.—Carapace smooth; anterolateral porFICURE -18.—Engaewa subcoerulea Riek, 1967, from Walpole,

Western Australia, Australia. (For explanation, see Figure 14.)
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tion of branchiocardiac groove widely separated

from, and parallel to, cervical groove almost to midheight of carapace; viewed dorsally, cervical groove

broadly U-shaped and deeply impressed; postorbital

ridges absent. Abdomen smooth; pleuron of first

abdominal segment partially overlapped by anterior

lobe of pleuron of second. Telson without transverse suture and entirely calcified. Third maxilliped

with mesial portion of ventral surface of ischium

bearing long stiff setae, otherwise without setae except for submarginal lateral row; exopodite well

developed, extending beyond distal extremity of

ischium. Chela with ventrolateral margin smooth;

when upper surface of carpus of cheliped held in

horizontal plane, dactyl moving subvertically; carpus of cheliped without enlarged spines or tubercles

mesially or ventrally, feebly tuberculate dorsomesially. Male genitalia consisting of simple small papilla. Branchial formula 21 + ep, all four pleurobranchiae present; stem of podobranchiae with

small winglike expansion.

RANGE.—Australia: Western Australia.

NUMBER OF SPECIES.—Three.

REFERENCES.—Riek, 1967a, 1969.

FIGURE 19.—Euastacoides setosus Riek, 1956, from unknown

locality. (For explanation, see Figure 14.)

Genus Euastacoides

FlCURE 19

Euastacoides Riek. 1956:4. [Type species, by original designation. Euastacoides setosus Riek. 1956:4. Gender: masculine.]

DIAGNOSIS.—Carapace with dense setiferous punctations and very small tubercles; anterolateral extremity of branchiocardiac groove fused with cervical groove rather high on carapace. Abdomen without spines but pleura provided with dense setiferous punctations; pleuron of first abdominal segment

distinct and partially overlapped by that of second.

Telson divided laterally by transverse suture; posterior portion membranous. Third maxilliped with

ventral surface of ischium provided with one row

of large punctations bearing clusters of stiff setae,

and produced distolaterally in acute prominence;

exopodite reduced, never reaching distal end of

ischium. Chela with ventrolateral margin bearing

single row of spines; when upper surface of carpus

of cheliped held in horizontal plane, dactyl moving

subhorizontally; carpus of cheliped with 2-4 large

spines mesially and irregularly arranged small

spines or tubercles on or adjacent to ventrodistal

margin. Male genitalia consisting of short papilla

bearing calcified ring. Branchial formula 21 + epr;

stem of podobranchiae, except on fourth pereiopod,

produced in winglike expansion.

RANGE.—Australia: Queensland.

NUMBER OF SPECIES.—Two.

REFERENCES.—Riek, 1956,1969.

Genus Euastacus

FIGURE 20

Euastacus Clark, 1939:10. [Type-species, by original designation, Cancer serratus Shaw, 1794:21 (see Riek, 1969:869,

893, 894). Gender: masculine.]

Euastaeus Riek, 1956:6 [erroneous spelling].

DIAGNOSIS.—Carapace with spines or tubercles

other than rostral, postorbital, branchiostegal, and

cervical; anterolateral extremity of branchiocardiac

groove fusing with cervical groove rather high on

carapace. Abdomen often with 3 or more rows of

spines or tubercles, sometimes confined to margins

of one or more pleura; pleuron of first abdominal

segment distinct and partially overlapped by that

of second. Telson, frequently with spines dorsally,

divided laterally by transverse suture; posterior por-
NUMBER 164 23

FIGURE 20.—Euastacus bispinosus Clark, 1941, from Otways,

Beauchamp Falls. Victoria, Australia. (For explanation, see

Figure 14.)

don membranous. Third maxilliped with ventral

surface of ischium provided with one row of large

punctations bearing clusters of stiff setae, and produced distolaterally in acute prominence; exopodite

much reduced, but flagellum often reaching distal

end of ischium. Chela with ventrolateral margin

bearing one row of spines and usually with additional row or subparallel low carina; when upper

surface of carpus of cheliped held in horizontal

plane, dactyl moving subhorizontally; carpus of

cheliped with 2-4 large spines raesially and 1 or 2

on, or adjacent to, ventrodistal margin. Male genitalia, situated on ventral surface of coxa, consisting

of short papilla bearing calcified ring. Branchial

formula 21 4- epr; stem of podobranchiae, except

on fourth pereiopod, produced in winglike expansion.

RANGE.—Australia: eastern states.

NUMBER OF SPECIES.—Twenty-seven.

REFERENCES.—Clark, 1936, 1941; Riek, 1969.

Genus Geocharax

FIGURE 21

Geocharax Clark, 1936:31. [Type-species, by original designation, Geocharax graciUs Clark, 1936:31. Gender: masculine.]

DIAGNOSIS.—Carapace punctate or tuberculate;

anterolateral portion of branchiocardiac groove

widely separated from, and parallel to, cervical

groove almost to midheight of carapace; viewed

dorsally, cervical groove broadly U-shaped and

deeply impressed; postorbital ridges well developed.

Abdomen smooth, punctate, or setose; pleuron of

first abdominal segment distinct and partially overlapped by that of second. Telson without transverse

suture and entirely calcified. Third maxilliped with

mesial half of ventral surface of ischium bearing

setae; exopodite well developed, extending beyond

distal extremity of ischium. Chela with ventrolateral margin bearing small tubercles; when upper

surface of carpus of cheliped held in horizontal

plane, dactyl moving subvertically but not so clearly

as in Engaens; carpus of cheliped without enlarged

spines or tubercles, but with row of tubercles dorsomesially. Male genitalia consisting of small simple

papilla. Branchial formula 21 + epr; stem of podobranchiae with small winglike expansion.

FICURE 21.—Geocharax falcata Clark, 1941, from Wannon

River, Western Australia, Australia. (For explanation, see

Figure 14.)
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

RANGE.—Australia: Victoria to King Island, and

northwestern Tasmania.

NUMBER OF SPECIES.—Two.

REFERENCES.—Clark, 1936; Riek, 1969.

Genus Gramastacus

FICUKE 22

Gramastacus Riek, 1972:385. [Type-species, by original designation, Gramastacus insolitus Riek, 1972:386. Gender: masculine.]

DIAGNOSIS.—Carapace punctate; anterolateral

portion of branchiocardiac groove widely separated

from, and parallel to, cervical groove almost to

midheight of carapace; viewed dorsally, cervical

groove U-shaped; postorbital ridges well developed.

Abdomen without prominent tubercles; pleuron of

first abdominal segment distinct and partially overlapped by that of second. Telson entirely calcified.

Third maxilliped with "dense hairs over all lower

surface" (Riek, 1972:385) of merus. Chela with

ventrolateral margin lacking large tubercles; when

upper surface of carpus of cheliped held in horizontal plane, dactyl moving subhorizontally; carpus

of cheliped with "slightly enlarged tooth on mesial

side towards apex." Male genitalia consisting of

"very large, uncalcified (only slightly sclerotized)

process, distinctly longer than wide."

RANGE.—Australia: "Grampians area of western

Victoria" (Riek, 1972:385).

NUMBER OF SPECIES.—Two.

REFERENCES.—Riek, 1972.

Genus Paranephrops

FIGURE 23

Paranrphropi While, 1842:79. (Type ipccirs. by monotypy,

Paranephrops planifrom White, 1842:79. Gender: masculine.]

Paranephros Andre, 1937:32. 34 [erroneous spelling].

Parancnphrops Cunningham et al. 1953:373 [erroneous spelling).

DIAGNOSIS.—Carapace with spines or tubercles,

other than rostral, postorbital, branchiostegal, and

cervical; anterolateral portion of branchiocardiac

groove widely separated from and parallel to cervical groove; postcervical groove prominent and

fusing with anterolateral arms of branchiocardiac

FIGURE 22.—Gramastacus insolitus Riek, 1972 (after Riek",

1972). (For explanation, see Figure 14; except d, basal podomere of fifth pereiopod illustrating male genitalia on mesial

surface of coxa.)

FICUKE 23.—Paranephrops planifrom White, 1842, from Rotorua District, North Island, New Zealand. (For explanation,

see Figure 14.)
NUMBER 164

grooves; viewed dorsally, cervical groove broadly

U-shaped and deeply impressed; post orbital ridges

well developed. Abdomen with setiferous punctations, lacking spines on pleura; pleuron of first

abdominal segment distinct and partially overlapped by that of second. Telson without transverse

suture and entirely calcified. Third maxilliped

usually with submedian ventral row of spiniform

tubercles; mesial half of ventral surface bearing

stiff setae; distolateral portion produced in spine;

exopodite well developed, extending beyond distal

extremity of ischium. Chela with 2 ventrolateral

rows of large spines or tubercles, and additional

rows both dorsally and ventrally; when upper surface of carpus of cheliped held in horizontal plane,

dactyl moving more horizontally than vertically;

carpus of cheliped with large spines mesially and

ventrally. Male genitalia, situated on mesial surface

of coxa, consisting of large, articulated, partially

sclerotized lobe. Gills 20 -f epr + r; pleurobranchiae on somites XI to XIV, posterior arthrobranch

on XIII rudimentary; stem without winglike expansion.

RANGE.—New Zealand.

NUMBER OF SPECIES.—Two.

REFERENCES.—Hopkins, 1970.

Genus Parastacoides

FIGURE 24

Parastacoides Clark, 1936:10, 48. [Type-specie*, by original

designation, Astacus (Astacus) Tasmanicus Erichson, 1846:

94. Gender: masculine.]

DIAGNOSIS.—Carapace setose, punctate, or granulate; anterolateral portion of branchiocardiac

groove extremely narrowly separated from, and

parallel to, cervical groove dorsolaterally; viewed

dorsally, cervical groove broadly U-shaped and

deeply impressed; postorbital ridges weakly developed. Abdomen punctate to smooth; pleuron of

first abdominal segment distinct and partially overlapped by that of second. Telson without transverse

suture and entirely calcified. Third maxilliped with

ventral surface bearing scattered setiferous punctations; exopodite well developed, extending to or

beyond distal extremity of ischium. Chela with

ventrolateral margin smooth; when upper surface

of carpus of cheliped held in horizontal plane, dactyl moving subhorizontally; carpus of cheliped with

25

FIGURE 24.—Parastacoides tasmanicus (Erichson, 1846) from

Derwent Bridge, Tasmania, Australia. (For explanation, see

Figure 14.)

2 to several large tubercles mesially, rare on ventrodistal margin. Male genitalia consisting of large,

nonlobed, fleshy, semicylindrical outgrowth on

mesial side of coxa. Branchial formula 17 + epr;

pleurobranchiae absent; posterior arthrobranchiae

reduced; stem of podobranchiae without winglike

expansion.

RANGE.—Australia: Tasmania.

NUMBER OF SPECIES.—Six.

REFERENCES.—Clark, 1939; Riek, 1967b, 1969.

Genus Parastacus

FIGURE 25

Parastacus Huxley. 1879:759, 771. [Type-species, by subsequent designation (Faxon, 1898:683), Astacus pilimanus von

Martens, 1869:15. Gender: masculine.]

DIAGNOSIS.—Carapace punctate or tuberculate;

anterolateral portion of branchiocardiac groove

often hardly distinguishable because closely approximating deeply impressed cervical groove, two

merging dorsolaterally; viewed dorsally, cervical

groove somewhat V-shaped; postorbital ridges rang-
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Parastacus pugnax (Pocppig, 18S5) from Concepcion, Chile. (For explanation, see Figure 14.)

FIGURE 26.—Samastaetu spinifrom (Philippi. 1882) from

Talcahuano, Chile. (For explanation, tee Figure 14.)

ing from well developed to virtually obsolete. Abdomen without tubercles or spines; pleuron of first

abdominal segment distinct and partially overlapped by that of second. Telson without transverse

suture and entirely, sometimes weakly, calcified.

Third maxilliped with mesial half of ventral surface of ischium bearing setiferous punctations; exopodite reaching to, or slightly beyond, distal extremity of ischium. Chela with ventrolateral margin smooth or bearing small tubercles; when upper

surface of carpus of cheliped held in horizontal

plane, dactyl moving subvertically; carpus with or

without enlarged spiniform tubercles. Male genitalia consisting of fixed, slightly elevated ventromesial ridge bearing n on calcified papilla; all members with male and female genital apertures. Branchial formula 20 + epr + r or 20 + ep + r;

pleurobranchiae on segments XI to XIV; posterior

arthrobranch on XIII rudimentary; stem without

winglike expansion.

RANGE.—South America: Chile, Argentina, Uruguay, and southern Brazil.

NUMBER OF SPECIES.—Six.

REFERENCES.—Faxon, 1885a, 1898, 1914; Riek,

1971.

Genus Sanuutacus

Ficuu 26

Samastaetu Riek, 1971:134. [Type-species, by original designation, Astacus spinifrom Philippi, 1882:627- Gender: masculine.]

DIAGNOSIS.—Carapace punctate or smooth; anterolateral portion of branchiocardiac groove extending subparallel to cervical groove, two merging laterally; viewed dorsally, cervical groove

broadly U-shaped; postorbital ridges well developed

with anterior spine or tubercle. Abdomen without

tubercles or spines; pleuron of first abdominal segment distinct and partially overlapped by that of

second. Telson without transverse suture and entirely calcified. Third maxilliped with setiferous

punctations over entire ventral surface; exopodite

reaching to or slightly beyond distal extremity of

ischium. Chela with ventrolateral margin smooth;
NUMBER 164 27

when upper surface of carpus of cheliped held in

horizontal plane, dactyl moving subhorizontally;

carpus without enlarged spiniform tubercles mesially or ventrally. Male genitalia consisting of articulated, slender, tubular, calcified projection; members never with both male and female apertures.

Branchial formula 20 + epr + r; stem without

winglike expansion.

RANGE.—South America: Chile.

NUMBER OF SPECIES.—Two.

REFERENCES.—Riek, 1971.

Genus Tenuibranchhtnu

FicinuE 27

Tenuibranchiurus Riek. 1951:368, 909. 881. [Type-spede*. by

original designation. Tenuibranchiurus gtyptictu Riek.

1951 :S82. Gender: masculine.]

DIAGNOSIS.—Carapace punctate to finely granulate; anterolateral portion of branchiocardiac

groove widely separated from, and parallel to, cervical groove almost to midheight of carapace;

viewed dorsally, cervical groove broadly U-shaped

and deeply impressed; postorbital ridges greatly reduced. Abdomen smooth; pleuron of first abdominal segment not rudimentary and partially covered

by anterior lobe of pleuron of second. Telson without transverse suture and entirely calcified. Third

maxilliped with ventral surface of ischium studded

with both plumose and simple stiff setae; exopodite

well developed, extending beyond distal extremity

of ischium. Chela with ventrolateral margin

smooth; when upper surface of carpus of cheliped

held in horizontal plane, dactyl moving subvertically; carpus of cheliped without enlarged spines or

FIGURE 27.—Tenuibranchiurus glypticus Riek, 1951, from

Caloundra, Queensland, Australia. (For explanation, see Figure 14).

tubercles mesially or ventrally, feebly tuberculate

dorsomesially. Male genitalia consisting of "flattened disc-like flange" (Riek, 1969:867). Branchial

formula 18 + ep; some females, however, with 2

additional pleurobranchiae (on segments XII and

XIII) = 20 + ep; stem of podobranchiae not produced in winglike expansion.

RANGE.—Australia: Queensland.

NUMBER OF SPECIES.—Monotypic.

REFERENCES.—Riek, 1951,1969.

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