SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Institution, Joseph Henry articulated a program that included the following statement:·“It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series:

Smithsonian Annals of Flight

Smithsonian Contributions to Anthropology

Smithsonian Contributions to Astrophysics

Smithsonian Contributions to Botany

Smithsonian Contributions to the Earth Sciences

Smithsonian Contributions to Paleobiology

Smithsonian Contributions to Zoology

Smithsonian Studies in History and Technology

In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These publications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the world. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available.

S. DILLON RIPLEY

Secretary

Smithsonian Institution

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 117

The Subgenera of the Crayfish Genus Procambarus (Decapoda: Astacidae)

Horton H. Hobbs, Jr.

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972

ERRATA FOR SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY, NUMBER 117

Page 4, couplet 3(2′): Reference to figure should read “… (Figure 13c) …”, not 19c.

Page 4, couplet 3′: Reference to figure should read “… (Figure 19c) …”, not 13c.

ABSTRACT

Hobbs, Horton H., Jr. The Subgenera of the Crayfish Genus Procambarus (Decapoda: Astacidae). Smithsonian Contributions to Zoology, number 117, 22 pages, 20 figures, 1 table. 1972.—A brief nomenclatural history of the genus Procambarus is followed by a key to the proposed 16 subgenera comprising 121 species and subspecies. Four of the subgenera, Girardiella (5 species and subspecies), Paracambarus (2 species), Procambarus (monotypic), and Ortmannicus (42 species and subspecies) have been previously recognized, and since 1942, Paracambarus has been accorded generic rank. The following subgenera are recognized as new: Acucauda (monotypic), Austrocambarus (16 species and subspecies), Capillicambarus (2 species), Hagenides (8 species and subspecies), Leconticambarus (13 species), Lonnbergius (monotypic), Mexicambarus (monotypic), Pennides (15 species), Remoticambarus (monotypic), Scapulicambarus (4 species), Tenuicambarus (monotypic), and Villalobosus (8 species). A diagnosis is provided for each subgenus together with illustrations of the type-species. Maps depicting the range of each subgenus are also included.

Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1972

For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402-Price 35 cents (paper cover)

Contents

A Brief History of the Genus Procambarus

Key to Subgenera of Genus Procambarus

Acucauda, new subgenus

Austrocambarus, new subgenus

Capillicambarus, new subgenus

Subgenus Girardiella Lyle

Hagenides, new subgenus

Leconticambarus, new subgenus

Lonnbergius, new subgenus

Mexicambarus, new subgenus

Subgenus Ortmannicus Fowler

Subgenus Paracambarus Ortmann

Pennides, new subgenus

Subgenus Procambarus Ortmann

Remoticambarus, new subgenus

Scapulicambarus, new subgenus

Tenuicambarus, new subgenus

Villalobosus, new subgenus

Literature Cited

Plates

Publication in Smithsonian Contributions to Zoology

The Subgenera of the Crayfish Genus Procambarus (Decapoda: Astacidae)

Horton H. Hobbs, Jr.

__________

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

The genus Procambarus, the largest of the crayfish genera, comprises 121 species and subspecies and, excluding introductions, is confined to North America, the northern part of Central America, and Cuba. With comparatively few exceptions, its members can be distinguished from those of other Cambarine genera by their possession of three or more terminal elements on the first pleopod of the first form male and a movable annulus ventralis in the female—both characters shared with only a few of the more primitive members of the genera Orconectes, Hobbseus, Cambarus, and Fallicambarus. (For discussions of the intergeneric relationships, see Hobbs 1967:8–15 and 1969:119–126.)

Table 1, modified from Hobbs (1962:278), summarizes the currently recognized species-groups of the genus in relation to the modifications that are proposed here. Intrageneric affinities are discussed by Hobbs (1962) and will be expanded in a monograph of the genus now in progress, in which more detailed synonymies for the infrageneric taxa are included.

Labeled illustrations depict the terminology utilized in the key to the 16 subgenera and in their diagnoses, both based solely on the first form male. Comparable figures of the type-species of each of the subgenera, including the first pleopod of the second form male and the annulus ventralis of the female, are also presented.

A few new terms have been introduced into the key and diagnoses. To clarify them, the following explanations are offered, the first three applying to the first pleopod of the male: The lamellae comprise the three layers of the doubly folded distal portion of the appendage (Figure 1a, b). The proximomedian lobe (Figure 2d, pml) extends proximally from the caudomedian base of the appendage; it is well developed in Figure 2a, d, e, f, g, o, and p. The proximomesial spur (Figure 2l, pms) is situated immediately proximal to the mesial base of the appendage; it is also conspicuous in Figure 2c, d, h, l, and o. The infraorbital spines (Figure 1c, ios) are those which are situated along the anterior margin of the carapace between the anterior extremity of the cervical groove and the suborbital angle.

I am grateful to Fenner A. Chace, Jr., Martha R. Cooper, Margaret A. Daniel, and Joseph F. Fitzpatrick, Jr., for their critical reading of the manuscript, and to Carolyn B. Gast for the rendition of Figure 3.

A Brief History of the Genus Procambarus

Thirty-two years after the first crayfish had been described from North America, R. Harlan (1830:464) assigned the name Astacus Blandingii to a crayfish that “inhabits the southern states, where it is common in the marshes and rivulets.” This was the first species to be described that is now assigned to the genus Procambarus. Erichson (1846), in his study of the genus Astacus, assigned nine species (four subsequently placed in the genus Procambarus: A. blandingii, A. cubensis, A. mexicanus, and A. wiegmanni) to his new subgenus Cambarus. Six years later, Girard (1852) elevated Erichson’s subgenus Cambarus to generic rank, assigning to his “Third Groups” C. Blandingii, C. acutissimus, C. acutus, and C. Clarkii, the latter three new species. LeConte (1856) was the last person who relegated a member of this species-group (Procambarus) to the genus Astacus, describing six new members from Georgia: A. advena, A. angustatus, A. fossarum, A. maniculatus, A. penicillatus, and A. spiculifer. Two new species of the group were added by Saussure (1857), C. aztecus and C. consobrinus.

In Hagen’s (1870) monograph of the North American Astacidae, he included 15 members of this species group, introducing three new species. In his “Group I” he placed C. acutus, C. angustatus, C. Blandingii, C. Clarkii, C. Fallax, C. Lecontei, C. maniculatus, C. penicillatus, C. spiculifer, C. troglodytes, C. versutus, and C. Wiegmanni. (C. acutissimus, C. aztecus, and C. consobrinus were considered by Hagen to be synonyms of C. acutus, C. wiegmanni, and C. cubensis, respectively.) In his “Group III” he placed C. advena, C. Cubensis, and C. Mexicanus.

Between 1870 and 1900, only 11 names were added to this species group: Cambarus gracilis Bundy, 1876; C. alleni Faxon, 1884; C. hagenianus Faxon, 1884; C. hayi Faxon, 1884; C. pubescens Faxon, 1884; C. simulans Faxon, 1884; C. acherontis Lönnberg, 1895; C. evermanni Faxon, 1890; C. digueti Bouvier, 1897; C. barbatus Faxon, 1898 to replace LeConte’s (1856) Astacus penicillatus; and C. gallinus Cockrell and Porter, 1900.

In Faxon’s (1885) revision of the Astacidae, the groupings are essentially similar to those of Hagen’s except the latter’s “Group III” corresponds to Faxon’s “Group II.” Only three species were added to Faxon’s Group II: Cambarus gracilis, C. simulans, and C. carolinus Hagen (not Erichson) (=C. hagenianus). The remaining species mentioned above were assigned to “Group I” with synonymies almost identical to those recognized by Hagen.

Within the decade 1901–1910, Ortmann (1905a–c, 1906, 1909) was the sole contributor to our knowledge of members of the current genus Procambarus. In addition to describing four new members, he recognized (1905a) four subgenera of the genus Cambarus: Bartonius, Cambarus, Faxonius, and Cambarellus, the first three of which were subdivided into “Sections” and “Groups.” His subgenus Cambarus (now recognized as the genus Procambarus) was subdivided as follows:

In later papers (1905c and 1906) Ortmann elevated his Section of C. digueti to subgeneric rank, designating it Procambarus, and proposed the monotypic subgenus Paracambarus to receive a new species, Cambarus (Paracambarus) paradoxus.

From 1911 to 1920, four additional species were described by Faxon, Pearse, and Ortmann. Fowler (1912), in treating the crayfishes of New Jersey, pointed out an error that Ortmann (1905a) had made in his proposal of the subgenera Cambarus and Bartonius: he had selected Cambarus blandingii as the type of Cambarus and C. bartonii as that of Bartonius. Faxon’s (1898:644) earlier designation of C. bartonii as the type-species of the genus Cambarus, however, invalidated Ortmann’s subsequent selection of C. blandingii as the type of the nominate subgenus, and inasmuch as Faxon’s type-species had been chosen by Ortmann as that of Bartonius, the latter became a still-born synonym of Cambarus, thus leaving Ortmann’s species-group, designated by him as Cambarus, without a name. For this assemblage, Fowler proposed the name Ortmannicus.

During the years 1921 to 1930, no new taxa were introduced, nor were there any alterations in the supraspecific categories affecting the assemblage of species now assigned to the genus Procambarus. In the following 10 years, however, Creaser and Hobbs added 11 new species to the group, and Lyle (1938) proposed that those species Faxon (1885) had referred to his Group II be assigned to a new subgenus, Girardiella.

Between 1941 and 1960, Hobbs, Hobbs and Walton, Penn, and Villalobos described 64 additional members of this species-group. Hobbs (1942a) proposed a revision of the genus Cambarus, involving chiefly the elevation of Ortmann’s subgenera to generic rank, combining under the older name, Procambarus, however, Ortmann’s Procambarus, Fowler’s Ortmannicus (=Ortmann’s Cambarus), and Lyle’s Girardiella.

Since 1960, only 11 species have been added to the genus, and Hobbs (1962:278) summarized and defined the supraspecific categories that were being utilized at that time. This summary, together with necessary emendations and equivalents of the proposals made herein, are outlined in Table 1.

At the time Hobbs (1942a) elevated Ortmann’s subgenera to generic rank, Paracambarus was a monotypic subgenus comprising the single, very peculiar C. paradoxus, possessing among other unique characters, hooks on the ischia of only the fourth pereiopods of the male. With the discovery by Villalobos (1949) of Paracambarus ortmanni in Puebla and related species of Procambarus (recorded from 1944 to 1951 and here referred to the subgenus Villalobosus) in Puebla and Veracruz, the characteristics which distinguish Paracambarus became much less well defined, and, in my opinion, hardly worthy of generic distinction. Therefore, this taxon is here reduced to subgeneric status in the genus Procambarus.

From 1942 until the present, no subgenera have been recognized within the genus; however, with the treatment proposed here, Fowler’s Ortmannicus and Lyle’s Girardiella are available for the groups that are indicated in Table 1.

Two monotypic subgenera that have no earlier counterparts are proposed to encompass the disjunct Procambarus pecki Hobbs, 1967, and P. fitzpatricki Hobbs, 1971.

TABLE 1.—Infrageneric Categories of the Genus Procambarus

Key to Subgenera of Genus Procambarus

Acucauda, new subgenus

(FIGURES 1l, 2a, 4,20d)

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum without marginal spines, tubercles, or median carina. Carapace without cervical spines. Areola 3.6 to 5.6 times longer than broad and constituting 29.3 to 34.2 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of plumose setae extending from basis to merus. Simple hooks on ischia of fourth pereiopods, absent on third. Coxa of fourth pereiopod with caudally subacute caudomesial boss directed in longitudinal plane of body. First pleopod reaching, or almost reaching, coxa of second pereiopod, slightly asymmetrical, with prominent proximomedian lobe, lacking proximomesial spur; cephalic surface with subangular shoulder at base of cephalic process; subapical setae present. Terminal elements of first pleopod consisting of subacute mesial process directed distolaterally beyond other terminal elements; cephalically situated, distally rounded cephalic process; central projection consisting of broad subrectangular, almost bladelike, centrocaudal process and narrower centrocephalic process; and caudal element represented by ridge around caudal base of centrocephalic process. Mesial ramus of uropod with median spine extending distinctly beyond distal margin of ramus.

TYPE-SPECIES.—Procambarus fitzpatricki Hobbs, 1971:461.

GENDER.—Feminine.

LIST OF SPECIES.—Monotypic, Procambarus (Acucauda) fitzpatricki Hobbs, 1971.

ETYMOLOGY.—Acus (L. = needle) + Cauda (L. = tail), signifying the long median spine on the distal margin of the mesial ramus of the uropod.

Austrocambarus, new subgenus

(FIGURES 2b, 5, 20a)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Procambarus Ortmann, 1905c:437 (part).

DIAGNOSIS.—Body pigmented or albinistic; eyes well developed or with pigment much reduced. Rostrum with or without marginal spines or tubercles, always lacking median carina. Carapace with two, one, or no cervical spines. Areola obliterated at midlength or as wide as 2.1 times longer than broad and constituting 24 to 37 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending distally from basis (merus sometimes with weak brush). Simple hooks on ischia of third pereiopods, absent on fourth. Coxa of fourth pereiopod without caudomesial boss. First pleopod reaching coxa of third pereiopod, symmetrical, with or without weak proximomedian lobe but sometimes with small proximomesial spur; cephalic surface with angular or rounded shoulder proximal to bases of terminal elements; subapical setae absent. Terminal elements of first pleopod consisting of distally or laterodistally directed simple-acute or lanceolate mesial process; cephalically or cephalodistally inclined acute or bladelike central projection; vestigial cephalic process rarely present at cephalomesial base of central projection; and caudal element, although sometimes absent or vestigial, usually represented by variously shaped caudal knob sometimes supporting small dentiform or tuberculiform caudal process. Mesial ramus of uropod with median spine acute or reduced to tubercle but never extending beyond distal margin of ramus.

TYPE-SPFCIES.—Procambarus vazquezae Villalobos, 1954:328.

GENDER.—Masculine.

LIST OF SPECIES.—Mexicanus Group (those species lacking or possessing one cervical spine, having no infraorbital spines, and with first pleopod lacking prominent bulbous enlargement between level of shoulder and base of central projection, and its mesial lamella not protruding caudally): Procambarus (Austrocambarus) aztecus (Saussure, 1857); P. (A.) mexicanus (Erichson, 1846); P. (A.) mirandai Villalobos, 1954; P. (A.) rodiguezi Hobbs, 1943; P. (A.) ruthveni ruthveni (Pearse, 1911); P. (A.) ruthveni zapoapensis Villalobos, 1954; P. (A.) vazquezae Villalobos, 1954; P. (A.) veracruzanus Villalobos, 1954.

Pilosimanus Group (those species possessing two or more cervical spines and at least one infraorbital spine): Procambarus (Austrocambarus) acanthophorus Villalobos, 1948; P. (A.) llamasi Villalobos, 1954; P. (A.) pilosimanus (Ortmann, 1906); P. (A.) williamsoni (Ortmann, 1905c).

Cubensis Group (those species lacking or possessing only one cervical spine, having no infraorbital spines, and with first pleopod bearing prominent bulbous enlargement between level of shoulder and base of central projection and its mesial lamella protruding caudally beyond principal shaft of appendage): Procambarus (Austrocambarus) atkinsoni (Ortmann, 1913); P. (A.) cubensis cubensis (Erichson, 1846); P. (A.) cubensis rivalis (Faxon, 1912); P. (A) niveus Hobbs and Villalobos, 1964.

ETYMOLOGY.—Auster (L. = south) in combination with the generic name Cambarus, referring to their occupying the southernmost area of the range of the members of the family Astacidae.

Capillicambarus, new subgenus

(FIGURES 1n, 2c, 6, 20d)

Subgenus Cambarus Ortmann, 1905b:401 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum with or without small marginal tubercles, lacking median carina. Carapace with or without single cervical spine. Areola 2.5 to 4.0 times longer than broad and constituting 29 to 35 percent of entire length of carapace. First three pairs of pereiopods with conspicuous brush of plumose setae extending from basis to at least proximal part of merus. Simple hooks on ischia of third and fourth pereiopods. Coxa of fourth pereiopod with caudomesial boss. First pleopod reaching coxa of third pereiopod, symmetrical, without proximomedian lobe, with prominent proximomesial spur; cephalic surface without shoulder; subapical setae absent. Terminal elements of first pleopod comprising caudally or caudodistally directed mesial process; small cephalic process present or not, if so, situated on mesial surface caudal to central projection; cephalically located central projection directed distally or caudally; caudal element lacking. Mesial ramus of uropod with median spine acute or reduced to tubercle but never extending beyond distal margin of ramus.

TYPE-SPECIES.—Cambarus (Cambarus) hinei Ortmann, 1905b:401.

GENDER.—Masculine.

LIST OF SPECIES.—Procambarus (Capillicambarus) hinei (Ortmann, 1905); P. (C.) incilis Penn, 1962.

ETYMOLOGY.—Capillus (L. = hair) in combination with the generic name, Cambarus, alluding to the abundant setation of the gnathal appendages and anterior pereiopods.

Subgenus Girardiella Lyle

(FIGURES 1k, 2d, 7, 20b)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

Subgenus Girardiella Lyle, 1938:76.

DIAGNOSIS.—Body and eyes pigmented, latter often small but well developed. Rostrum rarely with small marginal tubercles, lacking spines and median carina. Carapace without cervical spines, occasionally represented by small tubercles. Areola obliterated at midlength or as wide as 7.5 times longer than broad and constituting 34 to 42 percent (usually more than 36) of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third pereiopods, absent on fourth. Coxa of fourth pereiopod without caudomesial boss. First pleopods reaching coxa of third pereiopod, symmetrical, with prominent proximomedian lobe, usually with proximomesial spur; prominent shoulder on cephalic surface at cephalic base of terminal elements; subapical setae absent. Terminal elements of first pleopod consisting of distally or caudodistally directed, subspiculiform mesial process usually extending beyond other terminal elements; cephalomesially situated acute or subacute cephalic process (absent in Procambarus hagenianus); caudolaterally placed, usually somewhat compressed, caudal process; and prominent, centrally located, bladelike central projection. Mesial ramus of uropod with median spine extending beyond distal margin of ramus only in P. hagenianus.

TYPE-SPECIES.—Cambarus hagenianus Faxon, 1884:141; by monotypy. (The subspecies listed by Lyle (1938:76) are nomena nuda.)

LIST OF SPECIES.—Procambarus (Girardiella) gracilis (Bundy, 1876); P. (G.) hagenianus (Faxon, 1884); P. (G.) simulans simulans (Faxon, 1884); P. (G.) simulans regiomontanus Villalobos, 1954; P. (G.) tulanei Penn, 1953.

Hagenides, new subgenus

(FIGURES 2e, 8, 20b)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body and eyes pigmented, latter usually small but well developed. Rostrum without marginal spines, tubercles, or median carina. Carapace without cervical spines. Areola very narrow or obliterated at midlength, or as wide as 32 times longer than broad, and constituting 33 to 42 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third or third and fourth pereiopods, latter only in Procambarus geodytes. Coxa of fourth pereiopod with caudomesial boss. First pleopod reaching coxa of second pereiopod, asymmetrical, with proximomedian lobe, without proximomesial spur; cephalic surface without shoulder; subapical setae absent. Terminal elements of first pleopod represented by distally to caudally directed, acute, usually long mesial process; prominent beaklike or broad platelike central projection; cephalic process rarely well developed, often vestigial or lacking; and caudal elements usually consisting of prominent caudal knob although much reduced in Procambarus pygmaeus. Mesial ramus of uropod with median spine acute or reduced to tubercle but never extending beyond distal margin of ramus.

TYPE-SPECIES.—Astacus advena LeConte, 1856:402.

GENDER.—Masculine.

LIST OF SPECIES.—Advena Group (those species with first pleopod bearing acute central projection directed caudodistally or caudally, and with caudal knob, if evident, never fingerlike): Procambarus (Hagenides) advena (LeConte, 1856); P. (H.) geodytes Hobbs, 1942; P. (H.) pygmaeus Hobbs, 1942; P. (H.) truculentus Hobbs, 1954.

Rogersi Group (those species with first pleopod bearing platelike central projection directed distolaterally or obliquely, and with prominent finger- or thumblike caudal knob): Procambarus (Hagenides) rogersi rogersi (Hobbs, 1938); P. (H.) rogersi campestris Hobbs, 1942: P. (H.) rogersi expletus Hobbs and Hart, 1959; P. (H.) rogersi ochlocknensis Hobbs, 1942.

ETYMOLOGY.—Named in honor of Hermann A. Hagen whose monograph of the North American Astacidae (1870) constituted the first adequate summary of our knowledge of the crayfishes of North America.

Leconticambarus, new subgenus

(FIGURES 1m, 2f, 9, 20d)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body with or without pigment; eyes usually well developed, pigmented area much reduced in P. milleri. Rostrum with or without marginal spines or tubercles, lacking median carina. Carapace with, or more often without, cervical spines. Areola 2.5 to 13 times longer than broad and constituting 27 to 36 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Usually simple, occasionally bituberculate, hooks present on ischia of third or third and fourth pereiopods. Coxa with well developed, occasionally weakly developed, caudomesial boss. First pleopod reaching coxa of third, rarely second, pereiopod, asymmetrical, with or without proximomedian lobe, sometimes with proximomesial spur; setiferous shoulder, rounded knob, or slope present on cephalic surface at base of terminal elements; subapical setae disposed as just indicated. Terminal elements of first pleopod consisting of caudodistally to cephalodistally directed, slender or massive mesial process; cephalomesially situated, usually small, rounded or acute cephalic process; prominent caudal knob or process; and usually small dentiform central projection. Mesial ramus of uropod with median spine acute or reduced to tubercle but never extending beyond distal margin of ramus.

TYPE-SPECIES.—Cambarus barbatus Faxon, 1890:621.

GENDER.—Masculine.

LIST OF SPECIES.—Barbatus Group (those species possessing massive, or slender, curved or straight mesial process on first pleopod): Procambarus (Leconticambarus) apalachicolae Hobbs, 1942; P. (L.) barbatus Faxon, 1890); P. (L.) capillatus Hobbs, 1971; P. (L.) econfinae Hobbs, 1942; P. (L.) escambiensis Hobbs, 1942; P. (L.) hubbelli (Hobbs, 1940); P. (L.) kilbyi (Hobbs, 1940); P. (L.) latipleurum Hobbs, 1942; P. (L.) pubischelae Hobbs, 1942; P. (L.) rathbunae (Hobbs, 1940); P. (L.) shermani Hobbs, 1942.

Alleni Group (those species possessing spiculiform, sinuous mesial process on first pleopod): Procambarus (Leconticambarus) alleni (Faxon, 1884); P. (L.) milleri Hobbs, 1971.

ETYMOLOGY.—Named in honor of John LeConte, who, in 1856, described the first crayfishes from the area frequented by members of this subgenus.

Lonnbergius, new subgenus

(FIGURES 2g, 10, 20e)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Albinistic; eyes reduced and without pigment. Rostrum with marginal spines or tubercles, lacking median carina. Carapace without cervical spines. Areola 14 to 16 times longer than broad and constituting 41 to 43 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Bituberculate hooks on ischia of third and fourth pereiopods. Coxa of fourth pereiopod with prominent, vertically disposed caudomesial boss. First pleopod reaching coxa of third pereiopod, symmetrical, with proximomedian lobe and without proximomesial spur; no shoulder on cephalic surface; subapical setae lacking. Terminal elements of first pleopod comprising mesial process (not reaching distally nearly so far as other terminal elements); cephalic process rudimentary or absent; caudal process prominent, flanked by protuberances on ridge of caudal knob; and central projection very prominent, subacute, and directed distally. (Mesial lamella, joining mesial process and vestigial cephalic process, much shorter than in other members of genus, exposing much of median lamella between caudal process and centrocaudal proccess.) Mesial ramus of uropod with median spine acute but never reaching distal margin of ramus.

TYPE-SPECIES.—Cambarus acherontis Lönnberg, 1895:6.

GENDER.—Masculine.

LIST OF SPECIES.—Monotypic, Procambarus (Lonnbergius) acherontis (Lönnberg, 1895).

ETYMOLOGY.—Named in honor of Einar Lönnberg, who described the only known member of the subgenus.

Mexicambarus, new subgenus

(FIGURES 2h, 11, 20a)

Subgenus Cambarus Ortmann, 1909:159 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum without marginal spines, tubercles, or median carina. Carapace without cervical spines. Areola 6 to 11 times longer than broad and constituting 29 to 32 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third pereiopods, absent on fourth. Coxa of fourth pereiopod without caudomesial boss. First pleopod reaching coxa of second pereiopod, symmetrical, with proximomedian lobe somewhat laterally situated, proximomesial spur tuberculiform; shoulder absent from cephalic surface; subapical setae lacking. Terminal elements of first pleopod consisting of small, acute caudodistally directed mesial process; very prominent, corneous, curved, lamelliform cephalic process; caudolaterally situated caudal knob at level proximal to other terminal elements; and small central projection at caudolateral base of cephalic process. Mesial ramus of uropod with median spine never extending beyond margin of ramus.

TYPE-SPECIES.—Cambarus (Cambarus) bouvieri Ortmann, 1909:159.

GENDER.—Masculine.

LIST OF SPECIES.—Monotypic, Procambarus (Mexicambarus) bouvieri (Ortmann, 1909).

ETYMOLOGY.—Prefix of Mexico combined with the generic name, Cambarus; of the four peculiarly Mexican subgenera, Mexicambarus, Paracambarus, Procambarus, and Villalobosus, its affinities with members of the other subgenera seem most remote.

Subgenus Ortmannicus Fowler

(FIGURE 2i, 12, 20a)

Subgenus Cambarus Ortmann, 1905a:97 (part) [not Cambarus Erichson, 1846:95].

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body pigmented or albinistic, eyes with or without pigment, reduced or well developed. Rostrum with or without marginal spines, tubercles, and median carina. Carapace with or without one cervical spine. Areola obliterated or as wide as 2.9 times longer than broad and constituting 25 to 43 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third and fourth pereiopods, those on fourth sometimes bituberculate. Coxa of fourth pereiopod with bulbous or subangular caudomesial boss. First pleopod reaching coxa of third pereiopod, almost always asymmetrical, lacking or with weak proximomedian lobe and seldom bearing proximomesial spur; shoulder on cephalic surface usually absent, if present, shoulder on right pleopod never folded caudally against mesial surface of mesial lamella of pleopod; subapical setae usually present (absent in Procambarus bivittatus, P. hybus, P. jaculus, and P. lewisi). Terminal elements of first pleopod including subspiculiform to sublanceolate mesial process directed caudally, caudodistally or distally and usually somewhat laterally; setiform, hoodlike, or bladelike cephalic process (absent in P. mancus and in some populations of P. viaeviridis) situated cephalic, mesial, or lateral to central projection, and directed caudally, caudodistally, or distally; caudal process and/or caudal knob usually present (both absent in P. toltecae), sometimes vestigial (caudal process bladelike to knoblike, always situated caudally, caudomesially, or caudolaterally; caudal knob highly variable in size and form, varying in position from caudal surface laterally to cephalic surface); and central projection massive to minute, beaklike or bladelike to dentiform, and directed caudally, caudodistally, or distally. Mesial ramus of uropod with median spine often reduced and never extending beyond margin of ramus.

TYPE-SPECIES.—Astacus blandingii Harlan, 1830:464; designated by Ortmann, 1905a:97.

LIST OF SPECIES.—Blandingii Group (for diagnosis, see Hobbs, 1962:286.): Procambarus (Ortmannicus) acutissimus (Girard, 1852); P. (O.) acutus acutus (Girard, 1852); P. (O). acutus cuevachicae (Hobbs, 1941); P. (O.) blandingii (Harlan, 1830); P. (O.) caballeroi Villalobos, 1944; P. (O.) hayi (Faxon, 1884); P. (O.) lecontei (Hagen, 1870); P. (O.) lophotus Hobbs and Walton, 1960; P. (O.) texanus Hobbs, 1971; P. (O.) verrucosus Hobbs, 1952; P. (O.) viaeviridis (Faxon, 1914).

Pictus Group (for diagnosis, see Hobbs, 1962:283): Procambarus (Ortmannicus) angustatus (LeConte, 1856); P. (O.) chacei Hobbs, 1958; P. (O.) enoplosternum Hobbs, 1947; P. (O.) epicyrtus Hobbs, 1958; P. (O.) hirsutus Hobbs, 1958; P. (O.) lepidodactylus Hobbs, 1947; P. (O.) litosternum Hobbs, 1947; P. (O.) pictus (Hobbs, 1940); P. (O.) pubescens (Faxon, 1884); P. (O.) youngi Hobbs, 1942.

Seminolae Group (for diagnosis, see Hobbs, 1962:284): Procambarus (Ortmannicus) ancylus Hobbs, 1958; P. (O.) fallax (Hagen, 1870); P. (O.) leonensis Hobbs, 1942; P. (O.) lunzi (Hobbs, 1940); P. (O.) pycnogonopodus Hobbs, 1942; P. (O.) seminolae Hobbs, 1942.

Lucifugus Group (for diagnosis, see Hobbs, 1962:285): Procambarus (Ortmannicus) lucifugus lucifugus (Hobbs, 1940); P. (O.) lucifugus alachua (Hobbs, 1940); P. (O.) pallidus (Hobbs, 1940).

Planirostris Group (for diagnosis, see Hobbs, 1962:288): Procambarus (Ortmannicus) evermanni (Faxon, 1890); P. (O.) hybus Hobbs and Walton, 1957:P. (O.) jaculus Hobbs and Walton, 1957; P. (O.) mancus Hobbs and Walton, 1957; P. (O). pearsei pearsei (Creaser, 1934); P. (O.) pearsei plumimanus Hobbs and Walton, 1958; P. (O.) planirostris Penn, 1953.

Disjunct Species: Procambarus (Ortmannicus) bivittatus Hobbs, 1942; P. (O.) gonopodocristatus Villalobos, 1958; P. (O.) lewisi Hobbs and Walton, 1959; P. (O.) toltecae Hobbs, 1943; P. (O.) villalobosi Hobbs, 1969.

Subgenus Paracambarus Ortmann

(FIGURES 2j, 13, 20a)

Subgenus Paracambarus Ortmann, 1906:1.

Genus Paracambarus.—Hobbs, 1942a:344.

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum without marginal spines, tubercles, or median carina. Areola 3 to 4 times longer than broad and constituting 31 to 36 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of fourth pereiopods, absent on third. Coxa of fourth pereiopod with prominent simple or bituberculate caudomesial boss. First pleopod reaching coxa of second pereiopod, asymmetrical, with proximomedian lobe situated somewhat laterally, without well-defined proximomesial spur; shoulder on cephalic surface poorly developed or lacking; subapical setae absent. Terminal elements of first pleopod consisting of large caudodistally or smaller caudolaterally disposed mesial processes; cephalic process very small or absent; central projection beaklike; caudal element obsolete. Mesial ramus of uropod with median spine never extending beyond margin of ramus.

TYPE-SPECIES.—Cambarus (Paracambarus) paradoxus Ortmann, 1906:3; designated by Ortmann, 1906:1.

LIST OF SPECIES.—Procambarus (Paracambarus) ortmanni (Villalobos, 1949); P. (P.) paradoxus (Ortmann, 1906).

Pennides, new subgenus

(FIGURES 2k, 14, 20e)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum with marginal spines, seldom with median carina. Carapace with two or more cervical spines. Areola 3 to 5 times longer than broad and constituting 25 to 29 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third and fourth pereiopods. Coxa of fourth pereiopod with bulbous or subangular caudomesial boss. First pleopod reaching coxa of third pereiopod, asymmetrical, with or without broad, somewhat laterally situated proximomedian lobe, without proximomesial spur; seldom with shoulder on cephalic surface; subapical setae present. Terminal elements of first pleopod with caudally to distally directed, slender, usually acute mesial process; mesially to cephalolaterally situated cephalic process (absent in Procambarus gibbus, P. raneyi, P. spiculifer, and occasionally in P. ouachitae); caudal element variable, typically with caudal knob and process, but sometimes reduced, and occasionally lacking process; and central projection subdentiform to almost bladelike. Mesial ramus of uropod with median spine never extending markedly beyond margin of ramus, usually not reaching margin.

TYPE-SPECIES.—Procambarus natchitochae Penn, 1953c:5.

GENDER.—Masculine.

LIST OF SPECIES.—Procambarus (Pennides) ablusus Penn, 1963; P. (P.) dupratzi Penn, 1953; P. (P.) echinatus Hobbs, 1956; P. (P.) elegans Hobbs, 1969; P. (P.) gibbus Hobbs, 1969; P. (P.) lagniappe Black, 1968; P. (P.) lylei Fitzpatrick and Hobbs, 1971; P. (P.) natchitochae Penn, 1953; P. (P.) ouachitae Penn, 1956; P. (P.) penni Hobbs, 1951; P. (P.) raneyi Hobbs, 1953; P. (P.) spiculifer (LeConte, 1856); P. (P.) suttkusi Hobbs, 1953; P. (P.) versutus (Hagen, 1870); P. (P.) vioscai Penn, 1946.

ETYMOLOGY.—Named in honor of my good friend, the late George H. Penn, Jr., who contributed much to our knowledge of the crayfishes of the southern part of the United States and who was in the process of monographing this species-group at the time of his death.

Subgenus Procambarus Ortmann

(FIGURES 2l, 3a, 15, 20a)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Procambarus Ortmann, 1905c:437 (part).

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum with marginal spines or tubercles and median carina. Carapace with one cervical spine. Areola 4 to 6 times longer than broad and constituting 30 to 32 percent of entire length of carapace. Second pair of pereiopods with moderately conspicuous brush of setae extending from basis to merus, first and third with brush much reduced. Simple hooks on ischia of third pereiopods, absent on fourth. Coxa of fourth pereiopod without caudomesial boss. First pleopod reaching coxa of third pereiopod, symmetrical, without proximomedian lobe, with proximomesial spur; angular shoulder present on cephalic surface at base of terminal elements; subapical setae lacking. Terminal elements of first pleopod consisting of setiform, distally directed mesial process; lamelliform, mesially situated cephalic process; and very small caudal process contiguous with caudal base of prominent, bladelike central projection. Mesial ramus of uropod with median spine never extending beyond margin of ramus.

TYPE-SPECIES.—Cambarus digueti Bouvier, 1897:225; designated by Hobbs, 1942a:341.

LIST OF SPECIES.—Monotypic, Procambarus (Procambarus) digueti (Bouvier, 1897).

Remoticambarus, new subgenus

(FIGURES 2m, 16, 20b)

DIAGNOSIS.—Albinistic; eyes without pigment and reduced. Rostrum with marginal spines, lacking median carina. Carapace with one to three cervical spines. Areola 5 to 6.5 times longer than broad and constituting 38 to 43 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third pereiopods, absent on fourth. Coxa of fourth pereiopod with caudomesial bulbous boss. First pleopod reaching coxa of third pereiopod, symmetrical, without proximomedian lobe or proximomesial spur; shoulder present on cephalic surface proximal to terminal elements; subapical setae lacking. Terminal elements of first pleopod consisting of heavy, cephalically grooved, distolaterally directed mesial process obscuring, in caudal aspect, acute, cephalodistally directed central projection; cephalic and caudal processes lacking. Mesial ramus of uropod without median spine.

TYPE-SPECIES.—Procambarus pecki Hobbs, 1967:2.

GENDER.—Masculine.

LIST OF SPECIES.—Monotypic, Procambarus (Remoticambarus) pecki Hobbs, 1967.

ETYMOLOGY.—Remotus (L.=distant) in combination with the generic name Cambarus, alluding to the apparent distant relationships with other members of the genus.

Scapulicambarus, new subgenus

(FIGURES 2n, 3b, 17, 20c)

Subgenus Cambarus Ortmann, 1905a:97 (part).

Subgenus Ortmannicus Fowler, 1912:341 (part).

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum with or without marginal spines or tubercles, lacking median carina. Carapace with or without one cervical spine or tubercle. Areola obliterated or as wide as 7.5 times as long as broad, and constituting 22 to 38 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third and fourth pereiopods. Coxa of fourth pereiopod with bulbous caudomesial boss. First pleopod reaching coxa of third pereiopod, symmetrical or asymmetrical, with broad, short proximomedian lobe, without proximomesial spur; prominent shoulder on cephalic surface of left pleopod, that on right always folded caudomesially to lie against mesial face of mesial lamella, thus making shoulders asymmetrical; subapical setae present but often sparse. First pleopod with terminal elements consisting of caudodistally directed acute mesial process; cephalically situated acute or lamelliform cephalic process; caudolateral subspatulate caudal process (distinct caudal knob always absent); and small to prominent central projection. Mesial ramus of uropod with median spine never extending beyond margin of ramus.

TYPE-SPECIES.—Cambarus paeninsulanus Faxon, 1914:369.

GENDER.—Masculine.

LIST OF SPECIES.—Procambarus (Scapulicambarus) clarkii (Girard, 1852); P. (S.) howellae Hobbs, 1952; P. (S.) okaloosae Hobbs, 1942; P. (S.) paeninsulanus (Faxon, 1914); P. (S.) troglodytes (LeConte, 1856).

ETYMOLOGY.—Scapula (L.=shoulder-blade) in combination with the generic name Cambarus, referring to the peculiar folded shoulder on the right first pleopod of the first form male.

Tenuicambarus, new subgenus

(FIGURES 2o, 3d, 18, 20d)

DIAGNOSIS.—Body strongly compressed laterally and pigmented; eyes with pigment and well developed. Rostrum without marginal spines, tubercles or median carina. Carapace without cervical spines. Areola 15 to 18 times longer than broad and constituting 35 to 37 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple hooks on ischia of third and fourth pereiopods. Coxa of fourth pereiopod with very massive caudomesial boss, somewhat compressed in proximodistal axis of podomere and flared. First pleopod of male reaching coxa of third pereiopod, symmetrical, with long proximomesial lobe and with small proximomesial spur; shoulder on cephalic surface absent; subapical setae lacking. First pleopod with terminal elements comprising caudally directed, slender, acute mesial process; broad, acute, mesially situated cephalic process; curved scythelike central projection; caudal element lacking. Mesial ramus of uropod with median spine never extending beyond margin of ramus.

TYPE-SPECIES.—Procambarus tenuis Hobbs, 1950:194.

GENDER.—Masculine.

LIST OF SPECIES.—Monotypic, Procambarus (Tenuicambarus) tenuis Hobbs, 1950.

ETYMOLOGY.—Tenuis (L.=thin) in combination with the generic name Cambarus, in token of the compressed body of the only member of the subgenus.

Villalobosus, new subgenus

(FIGURES 2p, 3c, 19, 20a)

DIAGNOSIS.—Body and eyes pigmented, latter well developed. Rostrum without marginal spines or tubercles; median carina present or absent. Carapace without cervical spines. Areola 3 to 37 times longer than broad and constituting 29 to 38 percent of entire length of carapace. First three pairs of pereiopods without conspicuous brush of setae extending from basis to merus. Simple or bituberculate hooks on ischia of fourth pereiopods, occasionally vestigial ones on ischia of third. Coxa of fourth pereiopod with subcylindrical, subacute, caudomesial boss directed distally. First pleopod of first form male reaching coxa of second pereiopod, asymmetrical with proximomedian lobe (sometimes displaced laterally); proximomesial spur and shoulder on cephalic surface absent; subapical setae usually absent, present on Procambarus teziutlanensis, P. tlapacoyanensis, and sparsely in P. zihuateutlensis. Terminal elements of first pleopod consisting of subspiculiform to massive, short or long mesial process; spiniform, platelike, or vestigial cephalic or cephalomesially situated cephalic process; toothlike to prominent, flared caudal process; and tuberculiform to acute centrally located central projection. Mesial ramus of uropod with median spine sometimes almost obsolete, never extending beyond margin of ramus.

TYPE-SPECIES.—Paracambarus riojae [sic] Villalobos, 1944:161.

GENDER.—Masculine.

LIST OF SPECIES.—Erichsoni Group (those species in which caudal process of first pleopod consists of broad curved or flared corneous plate); Procambarus (Villalobosus) contrerasi (Creaser, 1931); P. (V.) erichsoni Villalobos, 1950; P. (V.) zihuateutlensis Villalobos, 1950.

Riojai Group (those species in which caudal process of first pleopod never consists of broad curved or flared corneous plate): Procambarus (Villalobosus) hoffmanni (Villalobos, 1944); P. (V.) hortonhobbsi Villalobos, 1950; P. (V.) riojai (Villalobos, 1944); P. (V.) teziutlanensis (Villalobos, 1947); P. (V.) tlapacoyanensis (Villalobos, 1947).

ETYMOLOGY.—Named in honor of Alejandro Villalobos F., whose detailed study of the crayfishes of Mexico will remain a classic for that faunal region.

Literature Cited

Bouvier, E. L.

1897. Sur les Cambarus recueillis au Mexique par M. Diguet. Bulletin du Muséum d’Histoire Naturelle, 3(6):224–228.

Erichson, W. F.

1846. Ubersicht der Arten der Gattung Astacus. Archives für Naturgeschichte, 1:86–103, 375–377.

Faxon, Walter

1884. Descriptions of New Species of Cambarus, to which is Added a Synonymical List of the Known Species of Cambarus and Astacus. Proceedings of the American Academy of Arts and Sciences, 20:107–158.

1885. A Revision of the Astacidae; Part I: The Genera Cambarus and Astacus. Memoirs of the Museum of Comparative Zoology, Harvard College, 10:vi + 186 pages, 10 plates.

1890. Notes on North American Crayfishes, Family Astacidae. Proceedings of the United States National Museum, 12(785):619–634.

1898. Observations on the Astacidae in the United States National Museum and in the Museum of Comparative Zoology, with Descriptions of New Species. Proceedings of the United States National Museum, 20(1136):643–694, 8 plates.

1914. Notes on the Crayfishes in the United States National Museum and the Museum of Comparative Zoology with Descriptions of New Species and Subspecies to which is Appended a Catalogue of the Known Species and Subspecies. Memoirs of the Museum of Comparative Zoology, Harvard College, 40(8):351–427, 13 plates.

Fowler, Henry W.

1912. The Crustacea of New Jersey. Annual Report of the New Jersey State Museum for 1911, pages 31–650, 150 plates.

Girard, Charles

1852. A Revision of the North American Astaci with Observations on their Habits and Geographical Distribution. Proceedings of the Academy of Natural Sciences of Philadelphia, 6:87–91.

Hagen, Hermann A.

1870. Monograph of the North American Astacidae. Illustrated Catalogue of the Museum of Comparative Zoology, Harvard College, number 3, 109 pages, 11 plates.

Harlan, R.

1830. Description of a New Species of the Genus Astacus. Transactions of the American Philosophical Society, 3(15):464–465.

Hobbs, Horton H., Jr.

1942a. A Generic Revision of the Crayfishes of the Subfamily Cambarinae (Decapoda, Astacidae) with the Description of a New Genus and Species. The American Midland Naturalist, 28(2):334–357, 23 figures.

1942b. The Crayfishes of Florida. University of Florida Publication, Biological Science Series, 3(2):1–279, 346 figures.

1950. A New Crayfish of the Genus Procambarus from Oklahoma and Arkansas (Decapoda, Astacidae). Journal of the Washington Academy of Sciences, 40(6):194–198, 12 figures.

1962. Notes on the Affinities of the Members of the Blandingii Section of the Crayfish Genus Procambarus (Decapoda, Astacidae). Tulane Studies in Zoology, 9(5):273–293, 72 figures.

1967. A New Crayfish from Alabama Caves with Notes on the Origin of the Genera Orconectes and Cambarus (Decapoda: Astacidae). Proceedings of the United States National Museum, 123(3621):1–17, 21 figures.

1969. On the Distribution and Phylogeny of the Crayfish Genus Cambarus. In Perry C. Holt, Richard L. Hoffman, and C. Willard Hart, Jr., The Distributional History of the Biota of the Southern Appalachians, Part I: Invertebrates. Virginia Polytechnic Institute, Research Division Monograph, 1:93–178, 20 figures.

1971. A New Crayfish of the Genus Procambarus from Mississippi (Decapoda: Astacidae). Proceedings of the Biological Society of Washington, 83(40):459–468, 1 figure.

LeConte, John

1856. Descriptions of New Species of Astacus from Georgia. Proceedings of the Academy of Natural Sciences of Philadelphia, 7:400–402.

Lönnberg, Einar

1895. Cambarids from Florida, a New Blind Species. Bihang till Koniglische Svenska Ventenskaps-Akademiens Handlingar, 22(4):3–14, 6 figures.

Lyle, Clay

1938. The Crawfishes of Mississippi, with Special Reference to the Biology and Control of Destructive Species. Iowa State College Journal of Science, 13:75–77.

Ortmann, Arnold E.

1905a. The Mutual Affinities of the Species of the Genus Cambarus, and their Dispersal over the United States. Proceedings of the American Philosophical Society, 44(180):91–136, 1 plate.

1905b. A New Species of Cambarus from Louisiana. Ohio Naturalist, 6(2):401–403, 1 figure.

1905c. Procambarus, a New Subgenus of the Genus Cambarus. Annals of the Carnegie Museum, 3(3):435–442, 3 figures.

1906. Mexican, Central American, and Cuban Cambari. Proceedings of the Washington Academy of Sciences, 8:1–24, 4 figures.

1909. Une écrevisse nouvelle du Mexique. Annales des Sciences Naturelles, Neuvième Série, 7:159–166, 4 figures.

Penn, George Henry, Jr.

1953a. A Redescription of the Crawfish Procambarus hinei (Ortmann) (Decapoda, Astacidae). Tulane Studies in Zoology, 1(5):63–68, 15 figures.

1953b. A New Burrowing Crawfish of the Genus Procambarus from Louisiana and Mississippi (Decapoda, Astacidae). Tulane Studies in Zoology, 1(6):71–76, 12 figures.

1953c. Two New Crawfishes of the Genus Procambarus from Texas, Louisiana, and Arkansas (Decapoda, Astacidae). American Museum Novitates, number 1636:1–10, 19 figures.

1956. The Genus Procambarus in Louisiana (Decapoda, Astacidae). The American Midland Naturalist, 6(2):406–422.

Saussure, H. de

1857. Diagnoses de quel Crustacés nouveaux de l’Amérique tropicale. Revue et Magasin Zoologie Pure et Appliquée, series 2, 9:501–510.

Villalobos, Alejandro

1944. Estudios de los cambarinos mexicanos, II: Dos especies nuevas del genero Paracambarus. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, 15(1):161–174, plates 1–2.

1949. Estudios de los cambarinos mexicanos, VIII: Descripción de una nueva especie del género Paracambarus. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, 19(1):175–182, plates 1–2.

1954.Estudios de los cambarinos mexicanos, XII: Revisión de las especies afines a Procambarus mexicanus (Erichson), con descripción de nuevas formas. Anales del Instituto de Biología, Universidad Nacional Autónoma de México, 25(1 y 2):299–379, plates 1–19.

1955.Cambarinos de la fauna mexicana. Tesis, Facultad de Ciencias. Universidad Nacional Autónoma de México, pages 1–290, plates 1–62.

FIGURE 1.—Characters utilized in distinguishing between the subgenera of the genus Procambarus: a, mesial view of hypothetical first pleopod of first form male; b, lateral view of same; c, lateral view of carapace showing two cervical spines [ios, infraorbital spines; bs, branchiostegal spine; cs, cervical spines]; d, lateral view of carapace showing no cervical spines; e, caudomesial view of first right pleopod of first form male with shoulder folded caudally against surface of mesial lamella [P. (Scapulicambarus) paeninsulanus]; f, caudomesial view of first left pleopod of first form male with shoulder extending cephalically [same]; g–j, ventral view of basal podomeres of pereiopods showing variation in number and form of hooks on ischia (stippled) of third and fourth; k, l, dorsal view of sixth abdominal segment, telson, and uropods [k, P. (Girardiella) gracilis; l, P. (Acucauda) fitzpatricki]; m, n, ventral view of third maxillipeds and first three pereiopods [m, P. (Leconticambarus) barbatus; n, P. (Capillicambarus) hinei].

FIGURE 2.—Caudal views of first pleopods of first form males of the type-species of the subgenera of the genus Procambarus: a, P. (Acucauda) fitzpatricki; b, P. (Austrocambarus) vazquezae; c, P. (Capillicambarus) hinei; d, P. (Girardiella) hagenianus [pml, proximomedian lobe]; e, P. (Hagenides) advena; f, P. (Leconticambarus) barbatus; g, P. (Lonnbergius) acherontis; h, P. (Mexicambarus) bouvieri; i, P. (Ortmannicus) blandingii; j, P. (Paracambarus) paradoxus; k, P. (Pennides) natchitochae; l, P. (Procambarus) digueti [pms, proximomesial spur]; m, P. (Remoticambarus) pecki; n, P. (Scapulicambarus) paeninsulanus; o, P. (Tenuicambarus) tenuis; p, P. (Villalobosus) riojai.

FIGURE 3.—Ventral views of basal podomeres of third, fourth, and fifth pereiopods of Procambarus showing certain variations in the coxa (shaded) of the fourth: a, P. (Procambarus) digueti, lacking a caudomesial boss; b, P. (Scapulicambarus) paeninsulanus, with bulbiform caudomesial boss; c, P. (Villalobosus) riojai, with acute caudomesial boss directed distally; d, P. (Tenuicambarus) tenuis, with compressed caudomesial boss directed in longitudinal axis of body. (Broad articular membranes in flat tone.)

FIGURES 4–7.—Type-species of the subgenera of Procambarus (4, Procambarus (Acucauda) fitzpatricki Hobbs; 5, Procambarus (Austrocambarus) vazquezae Villalobos; 6, Procambarus (Capillicambarus) hinei (Ortmann); 7, Procambarus (Girardiella) hagenianus (Faxon)): a, dorsal view of carapace; b, lateral view of first left pleopod of first form male; c, lateral view of distal portion of first left pleopod of first form male (subapical setae, if present, removed); d, mesial view of distal portion of first left pleopod of first form male; e, annulus ventralis of female; f, lateral view of first left pleopod of second form male; g, dorsal view of chela of first form male.

FIGURES 8–11.—Type-species of the subgenera of Procambarus (8, Procambarus (Hagenides) advena (LeConte); 9, Procambarus (Leconticambarus) barbatus (Faxon); 10, Procambarus (Lonnbergius) acherontis (Lönnberg); 11, Procambarus (Mexicambarus) bouvieri (Ortmann)): a, dorsal view of carapace; b, lateral view of first left pleopod of first form male; c, lateral view of distal portion of first left pleopod of first form male (subapical setae, if present, removed); d, mesial view of distal portion of first left pleopod of first form male; e, annulus ventralis of female; f, lateral view of first left pleopod of second form male; g, dorsal view of chela of first form male.

FIGURES 12–15.–Type-species of the subgenera of Procambarus (12, Procambarus (Ortmannicus) blandingii (Harlan)l 13, Procambarus (Paracambarus) pardoxus (Ortmann); 14, Procambarus (Pennides) natchitochae penn; 15, Procambarus (Procambarus) digueti (Bouvier)): a, dorsal view of carapace; b, lateral view of first left pleopod of first form male; c, lateral view of distal portion of first left pleopod of first form male (subapical setae, if present, removed); d, mesial view of distal portion of first left pleopod of firwst form male; e, annulus ventralis of female; f, lateral view of first left pleopod of second form make; g, dorsal view of chela of first form male.

FIGURES 16–19.—Type-species of the subgenera of Procambarus (Remoticambarus) pecki Hobbs; 17, Procambarus (Scapulicambarus) paeninsulanus (Faxon); 18, Procambarus (Tenuicambarus) tenuis Hobbs; 19, Procambarus (Villalobosus) roijai (Villalobos)): a, dorsal view of carapace; b, lateral view of first left pleopod of first form male; c, lateral view of distal portion of first left pleopod of first form male (subapical setae, if present, removed); d, mesial view of distal portion of first left pleopod of first form male; e, annulus ventralis of female; f, lateral view of first left pleopod of second form male; g, dorsal view of chela of first form male.

FIGURE 20.—Ranges of the subgenera of the genus Procambarus: a, 1: Ortmannicus; 2: Procambarus; 3: Mexicambarus; 4: Paracambarus and Villalobosus; 5: Austrocambarus; b, 1: Girardiella; 2: Remoticambarus; 3: Hagenides; c, Scapulicambarus; d, 1: Tenuicambarus; 2: Capillicambarus; 3: Acucauda; 4: Leconticambarus; e, 1: Pennides; 2: Lonnbergius.

Publication in Smithsonian Contributions to Zoology

Manuscripts for serial publications are accepted by the Smithsonian Institution Press, subject to substantive review, only through departments of the various Smithsonian museums. Non-Smithsonian authors should address inquiries to the appropriate department. If submission is invited, the following format requirements of the Press will govern the preparation of copy.

Copy must be typewritten, double-spaced, on one side of standard white bond paper, with 1½″ top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompanied by the original artwork. Duplicate copies of all material, including illustrations, should be retained by the author. There may be several paragraphs to a page, but each page should begin with a new paragraph. Number consecutively all pages, including title page, abstract, text, literature cited, legends, and tables. The minimum length is 30 pages, including typescript and illustrations.

The title should be complete and clear for easy indexing by abstracting services. Taxonomic titles will carry a final line indicating the higher categories to which the taxon is referable: “(Hymenoptera: Sphecidae).” Include an abstract as an introductory part of the text. Identify the author on the first page of text with an unnumbered footnote that includes his professional mailing address. A table of contents is optional. An index, if required, may be supplied by the author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet, omit the number. For easy reference, number also the taxa and their corresponding headings throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date:page) with a full reference at the end of the paper under “Literature Cited.” Begin each taxon at the left margin with subsequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate each reference. Enclose with square brackets any annotation in, or at the end of, the entry. For references within the text, use the author-date system: “(Jones 1910)” and “Jones (1910).” If the reference is expanded, abbreviate the data: “Jones (1910:122, pl. 20: fig. 1).”

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they should not contain rules. Formal tables must be submitted as pages separate from the text, and each table, no matter how large, should be pasted up as a single sheet of copy.

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

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be indicated by lowercase italic letters on the illustration, in the legend, and in text references: “Figure 9b.” If illustrations (usually tone photographs) are printed separately from the text as full pages on a different stock of paper, they will be termed Plates, and individual components should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the numbering system of text illustrations with that of plate illustrations. Submit all legends on pages separate from the text and not attached to the artwork. An instruction booklet for the preparation of illustrations is available from the Press on request.

In the bibliography (usually called “Literature Cited’), spell out book, journal, and article titles, using initial caps with all words except minor terms such as “and, of, the.’ For capitalization of titles in foreign languages, follow the national practice of each language. Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume, number, and page citations: “10(2):5-9.” Spell out such words as “figures,” “plates,” “pages.”

For free copies of his own paper, a Smithsonian author should indicate his requirements on “Form 36” (submitted to the Press with the manuscript). A non-Smithsonian author will receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.