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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 164

Synopsis of the Families and Genera of Crayfishes (Crustacea: Decapoda)

Horton H. Hobbs, Jr.

SMITHSONIAN INSTITUTION PRESS

City of Washington

1974

A B S T R A C T

Hobbs, Horton H., Jr. Synopsis of the Families and Genera of Crayfishes (Crustacea: Decapoda). Smithsonian Contributions to Zoology, number 164, 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 arc 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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Synopsis of the Families and Genera of Crayfishes (Crustacea: Decapoda)

Horton H. Hobbs, Jr.

____________

Horton H. Hobbs, Jr., Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560

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 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. 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 Eurasian-western 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 (1969:113–116). 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 spcrm 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 Nephropoidea¹

Superfamily Astacoidea

Family Astacidae

Genus Astacus

Austropotamobius

Pacifastacus

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

Euaslacus

Geocharax

Gramastacus

Paranephrops

Parastacoides

Parastacus

Samastacus

Tenuibranchiurus

MAJOR CHARACTERISTICS INVOLVED IN THE EVOLUTION OF CRAYFISHES FROM A PRO-NEPHROPOID ANCESTOR

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. Holthuis, 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

Superfamily ASTACOIDEA

ASTACODEA De Haan, 1841:148, 160.

Astaona 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 + Sr + ep. Articles of lateral ramus of antennule, in region of asthetascs, usually bearing two clusters, 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.

Astakoiden Duméril, 1806:172.

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

Astacia Rafinesque, 1815:98.

ASTACIDAE Samouelle, 1819:94.

Astacides 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 + 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 Duméril, 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. [Type-species, 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 lcptodactylus 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 lobes, one reaching distinctly farther distally than other; exopodite of second pleopod extending almost or quite as far distally as endopodite.

FIGURE 1.—Astacus (A.) astacus (Linnaeus, 1758) from Ljumgan River, Alby, Sweden: a, dorsal view of carapace and chelipeds of male; b, ventral view of merus of third maxilliped; c, lateral view of first pleopod of male; d, basal podomeres of second through fifth pereiopods; e, caudal thoracic sternal area of female; f, epistome and proximal podomerc of antennae.

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.

FIGURE 2.—Austropotamobius torrentius danubicus Karaman, 1962, from Bucharest, Rumania. (For explanation, see Figure 1.)

Genus Austropotamobius

FIGURE 2

Austropotomobius Skorikov, 1908:116. [Type-species, by subsequent designation /Bou, 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 3.—Pacifastacus leniusculus klamathensis (Stimpson) (rom 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.]

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

CAMBARINAE 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 + 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 + ep. Some articles of lateral ramus of antennule bearing 2 clusters of asthetascs.

Genus Barbicambarus

FIGURE 4

Barbicambarus Hobbs, 1969:95, 98. [Type-species, by original designation, Cambarus cornutus Faxon, 1884:120. Gender: 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 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 + ep.

FIGURE 4.—Barbicambarus comutus (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.

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

FIGURE 5

Cambarus Erichson. 1846:97. [Type-species. by subsequent designation (Faxon, 1898:644), Astacus Bartonii Fabricius, 1798:407. Gender: masculine. Proposed as subgenus of Astacus; elevated to generic rank by Girard, 1852:88.]

Cambarus Huxley, 1880b:81 [erroneous spelling].

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

Camberus Miller, 1895:336 [erroneous spelling].

Bartonius Ortmann, I905a:97. [Type-species, 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 Unestam, 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. [Type-species, by original designation, Astacus latimanus LeConte, 1856:402. Gender: masculine.]

Subgenus Erebicambarus Hobbs, 1969:95, 99. [Type-species, by original designation, Cambarus bartoni tenebrosus Hay, 1902:232. Gender: masculine.]

Subgenus Hiaticambarus Hobbs, 1969:95, 105. [Type-species, by original designation, Cambarus longulus Girard, 1852:90. Gender: masculine.]

Subgenus Jugicambarus Hobbs, 1969:95, 106. [Type-species, by original designation, Cambarus bartonii asperimanus Faxon, 1914:391. Gender: masculine.]

Subgenus Lacunicambarus Hobbs, 1969:96, 110. [Type-species, by original designation, Cambarus diogenes Girard, 1852:88. Gender: masculine.]

Subgenus Puncticambarus Hobbs, 1969:96, 101. [Type-species, by original designation, Cambarus extraneus Hagen, 1870: 73. Gender: masculine.]

Subgenus Veticambarus Hobbs, 1969:95, 96. [Type-species, by original designation, Cambarus pristinus Hobbs, 1965: 268. 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 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 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 + ep.

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.)

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, Washtenaw County, Michigan, U.S.A. (For explanation, sec 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 strawni 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 with 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 3 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.]

FIGURE 7.—Faxonella clypteata (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, Cambarus 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 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-species, 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 tubercles; lateral margin of fixed finger never bearing row of spiniform 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 3 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 species; first pleopod usually present. Branchial formula 17 + ep.

FIGURE 9.—Orconectes limosus (Rafinesque, 1817) from Cattail Branch at Route 97, Howard County. Maryland. U.S.A. (For explanation, see Figure 4.)

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 Procambarus

FIGURE 10

Cambarus Ortmann. 1905a:96 (not Erichson. 1846). [Type-species. by original designation. Ailacus Blandingii Harlan, 1830:464. Gender: masculine. Proposed as typical subgenus, but type-species previously selected by Faxon, 1898:644, Astacus Bartonii Fabricius, 1798:407.]

Procambarus Ortmann, 1905c: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:341.]

Paracambarus Ortmann. 1906:1. [Type-species, by monotypy, Cambarus (Paracambarus) paradoxus Ortmann, 1906:3. Gender: masculine. Proposed as subgenus of Cambarus, elevated to generic rank by Hobbs, 1942a:344, 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.]

Cambarellus Creaser, 1933:21 [lapsus for Cambarus].

procambarus Padget, 1970:19.

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

Subgenus Acucauda Hobbs, 1972a:3–5. [Type-species, by original designation, Procambarus fitzpatricki 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-species, by original designation, Cambarus (Cambarus) hinei Ortmann, 1905b:401. Gender: masculine.]

Subgenus Hagenides Hobbs, 1972a:2–4, 7. [Type-species, by original designation, Astacus advena LeConte, 1856:402. Gender: masculine.]

Subgenus Leconticambarus Hobbs, 1972a:2–5, 7. [Type-species, 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.S.A. (For explanation, see Figure 4).

Subgenus Lonnbergius Hobbs, 1972a:2–4, 8. [Type-species, by original designation, Cambarus acherontis Lonnberg, 1895: 6. Gender: masculine.]

Subgenus Mexicambarus Hobos, 1972a:3, 4, 8. (Type-species, by original designation, Cambarus (Cambarus) bouvieri Ortmann, 1909:159. Gender: masculine.]

Subgenus Pennides Hobbs, 1972a:2–4, 10. [Type-species, by original designation, Procambarus natchitochae Penn, 1953: 5. Gender: masculine.]

Subgenus Remoticambarus Hobbs, 1972a:3, 4, 11. [Type-species, by original designation, Procambarus pecki Hobbs, 1967:2. Gender: masculine.]

Subgenus Scapulicambarus Hobbs, 1972a:2, 3, 5. [Type-species, by original designation, Cambarus clarkii paeninsulanus Faxon, 1914:369. Gender: masculine.]

Subgenus Tenuicambarus Hobbs, 1972a:3, 5, 12. [Type-species, by original designation, Procambarus tenuis Hobbs, 1950: 194. Gender: masculine.]

Subgenus Villalobosus Hobbs, 1972a:3, 5, 12. [Type-species, by original designation, Paracambarus riojae Villalobos, 1944: 161. 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 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. [Type-species, by original designation, Troglocambarus maclanei Hobbs, 1942a:345. Gender: masculine.]

DIAGNOSIS.—Albinistic, eyes reduced and without pigment. Antenna never with conspicuous fringe 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, somewhat 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.

FIGURE 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.)

RANGE.—North America: Peninsular Florida.

NUMBER OF SPECIES.—Monotypic.

REFERENCES—Hobbs, 1942a, 1972b.

Subfamily CAMBARELLINAE

CAMBARELLINAE Laguarda. 1961:69. [Originally proposed as a lubfamily of the Astacidac.]

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 3 terminal elements (mesial and caudal processes and central projection). Female with annulus ventralis freely 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.

FIGURE 12.— Cambarellus montezumae (Saussure, 1858) from near Pachuca, Hidalgo, Mexico. (For explanation, see Figure 4).

Genus Cambarellus

FIGURE 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:550.]

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

CAMBAROIDINAE Villalobos, 1955:7 [originally proposed as a subfamily of the Astacidae].—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 cephalo-mesially 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 13

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 Haan, 1842:164. Gender: masculine. Originally proposed as a subgenus of Astacus; elevated to generic rank by Skorikov, 1908:116.]

FIGURE 13.—Cambaroides similis (Koelbel, 1892) from near Pusan, Korea. (For explanation, see Figure 4.)

Gambaroides Parisi, 1917:17 [erroneous spelling].

Camberoides 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 die 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. [Synonymized, by implication. with Parastacidae by Riek, 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. Fint pleo pod absent in 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 Guérin-Méneville, 1839:109. [Type-species. by monotypy, Astacoides Goudotii Guérin-Méneville, 1839:109, a subjective junior synonym of Astacus madagascarensis H. Milne Edwards and Audouin, 1839:152. Gender: masculine.]

DIAGNOSIS.—Carapace with conspicuous spines or tubercles, in addition to rostral, postorbital, branchiostegal, 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 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; pleuro-branch 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 OF 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.

FIGURE 15.—Astacopsis fluviatilis 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 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, 1888:193 [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.)

Parachaeraps Smith, 1912:145, 148, 150–153. 161. [Type-species, by monotypy. Astacus 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, 1956:47. [Type-species, by original designation, Pseudengaeus strictifrons Clark, 1956:47. Gender: masculine. Synonymized with Engaeus by Riek, 1969:863.]

Austroastacus Clark, 1936:51. [Type-species, by original designation, Engaeus hemicirratulus Smith and Schuster, 1913: 123. Gender: masculine. Synonymized with Engaeus by Riek, 1969:863.]

FIGURE 17.—Engaeus orientalis Clark. 1941. from 4 miles north of Bonong, Victoria, Australia. (For explanation, see Figure 14.)

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, 1939; Riek, 1969.

Genus Engaewa

FIGURE 18

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

DIAGNOSIS.—Carapace smooth; 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 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.

FIGURE 18.—Engaewa subcoerulea Riek, 1967, from Walpole, Western Australia, Australia. (For explanation, see Figure 14.)

RANGE.—Australia: Western Australia.

NUMBER of SPECIES.—Three.

REFERENCES.—Riek, 1967a, 1969.

FIGURE 19.—Euastacoides selosus Riek, 1956, from unknown locality. (For explanation, tee Figure 14.)

Genus Euastacus

FIGURE 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.

RANCE.—Australia: Queensland.

NUMBER OF SPECIES.—Two.

REFERENCES.—Riek, 1956, 1969.

Genus Euastacus

FIGURE 20

Euastacus Clark, 1939:10. [Type-species, by original designa tion. Cancer serratus Shaw. 1794:21 (sec Riek, 1969:869, 893, 894). Gender: masculine.)

Euastacus Riek, 1956:6 [erroneous spelling].

DIAGNOSIS.—Carapace with spines or tubercles other than rostral, postorbital, branchiostegal, and cervical; anterolateral extremity of branchiocardiac groove fusing widi 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 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 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 chèliped with 2–4 large spines mesially and 1 or 2 on, or adjacent to, ventrodistal margin. Male genitalia, situated on ventral surface of coxa, consisting of short papilla bearing calcined ring. Branchial formula 21 + epr; stem of podobranchiae, except on fourth pereiopod, produced in winglike expansion.

FIGURE 20.—euaslncui bispinosus Clark, 1941, from Otways,. Beauchamp Falls, Victoria, Australia. (For explanation, see Figure 14.)

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 gracilis 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 Engaeus; 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.

FIGURE 21.—Geocharax falcata Clark, 1941, from Wannon River, Western Australia, Australia. (For explanation, see Figure 14.)

RANGE.—Australia: Victoria to King Island, and northwestern Tasmania.

NUMBER OF SPECIES.—Two.

REFERENCES.—Clark, 1936; Riek, 1969.

Genus Gramastacus

FIGURE 22

Gramastacus Rick, 1972:385. [Type-species, by original designation, Gramastacus insolitus Rick, 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.”

FIGURE 22.—Gramastacus insolitus Riek, 1972 (after Riek, 1972). (For explanation, see Figure 14; except d, basal podo-mere of fifth pereiopod illustrating male genitalia on mesial surface of coxa.)

RANGE.—Australia: “Grampians area of western Victoria” (Riek, 1972:385).

NUMBER OF SPECIES.—Two.

REFERENCES.—Riek, 1972.

Genus Paranephrops

FIGURE 23

Paranephrops White. 1842:79. [Type-species, by monotypy, Paranephrops planifrons White, 1842:79. Cender: masculine.]

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

Paranenphrops 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 grooves; viewed dorsally, cervical groove broadly U-shaped and deeply impressed; postorbital ridges well developed. Abdomen with setiferous punctations, lacking spines on pleura; pleuron of first abdominal segment distinct and partially over lapped by that of second. Telson without transverse suture and entirely calcified. Third maxilliped usually with submedian ventral row of spinifom 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 + epr + r; pleurobranchiae on somites XI to XIV, posterior arthrobranch on XIII rudimentary; stem without winglike expansion.

FIGURE 23.—Paranephrops planifrons White, 1842, from Ro-torua District, North Island, New Zealand. (For explanation, see Figure 14.)

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 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.

FIGURE 24.—Parastacoides tasmanicus (Erichson, 1846) from Derwent Bridge, Tasmania, Australia. (For explanation, see Figure 14.)

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 ranging 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 ventro-mesial ridge bearing noncalcified 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.

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

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

NUMBER OF SPECIES.—Six.

FIGURE 26.—Samastacus spinifrons (Philippi, 1882) from Talcahuano, Chile. (For explanation, see Figure 14.)

REFERENCES.—Faxon. 1885a, 1898, 1914; Riek, 1971.

Genus Samastacus

FIGURE 26

Samastacus Riek, 1971:134. [Type-specie, by original designation, Aslacus spinifrons 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; 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 or SPECIES.—Two.

REFERENCES.—Rick, 1971.

Genus Tenuibranchiurus

FIGURE 27

Tenuibranchiurus Rick. 1951:368. 969, 581. [Type-species. by original designation, Tenuibranchiurus glypticus Rick, 1951:582. 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 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.

FIGURE 27.—Tenuibranchiurus glypticus Riek, 1951, from Caloundra, Queensland, Australia. (For explanation, see Figure 14).

RANGE.—Australia: Queensland.

NUMBER of SPECIES.—Monotypic.

REFERENCES.—Riek, 1951, 1969.

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1970. Using Baytex to Control Crayfish in Ponds. Progressive Fish-Culturist, 33:58–60.

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1966. Two New Species of the Genus Cambarus from Arkansas (Decapoda, Astacidae). Tulane Studies in Zoology, 13(1):9–15, 18 figures.

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1944. The Crayfishes of Kentucky with Notes on Variation, Distribution, and Descriptions of New Species and Subspecies. American Midland Naturalist, 31(1): 111–149, 10 figures.

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1956. Additions to the Australian Freshwater Crayfish. Records of The Australian Museum, 24(1):1–6, 1 plate.

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

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