HORTON H. HOBBS, The Subgenera

of the Crayfish

Genus Procambarus

(Decapoda: Astacidae)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 11 7

Horton H. Hobbs, jr. Th e Subgenera

of the Crayfish

Genus Procambarus

(Decapoda: Astacidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
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NUMBER 117

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ABSTRACT

Hobbs, Horton H.3 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.

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Horton H. Hobbs, Jr. The Subgenera

of the Crayfish

Genus Procambarus

(Decapoda: Astacidae)

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

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

planations 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 \a, 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 21, pms) is situated immediately proximal to

the mesial base of the appendage; it is also conspicuous in Figure 2c, d, h, I, and o. The infraorbital

spines (Figure \c, 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
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(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 speciesgroup {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 Lonnberg, 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:

Section of C. digueti

Subgeneric Equivalents Here

Proposed

Procambarus and Aust'rocambarm

Section of C. gracilis Girardiella and Hagenides

Section of C. blandingi

Group of C. spiculifer = Pennides and Ortmannicus

(part)

Group of C. blandingi = Lonnbergius and Ortmannicus

(part)

Group of C. clarki — Scapulicambarus

Group of C. alleni = Leconticambarus and Ortmannicus (part)

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
NUMBER 1 1 7

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

Table I.— Infrageneric Categories of the Genus Procambarus

Previously Recognized

(Modified from Hobbs, 1962)

Proposed

Sections Groups Subgenera Groups

Blandingii (Ortmann, 1905a:98)

Tenuis (Hobbs, 1962:278)

Riojae (Villalobos, 1955:94)

Digueti (Ortmann, 1905a:98)

Advena (hobbs, 1942b:73)

Acherontis (Hobbs, 1942b:91)

Hinei (Penn, 1953a:67)

Barbatus (Hobbs, 1942b:33)

Gracilis (Ortmann, 1905a:98)

Mexicanus (Villalobos, 1955:159)

Members not known in 1962

Relegated to separate

genus in 1962

Spiculifer (Ortmann, 1905a:100)

Clarkii (Ortmann, 1905a:100)

Blandingii (Ortmann, 1905a:100)

Pictus (Hobbs, 1942b:129)

Seminolae (Hobbs, 1942b:142)

Lucifugus (Hobbs, 1942b:134)

Planirostris (Penn, 1953b:75)

Disjunct members

Riojae (Villalobos, 1955:96)

Erichsoni (Villalobos, 1955:130)

Advena (Hobbs, 1942b:73)

Rogersi (Hobbs, 1942b:88)

Barbatus (Hobbs, 1942b:35)

Alleni (Ortmann, 1905a:100)

Disjunct members

Simulans (Penn, 1956:418)

Mexicanus (Villalobos, 1955:160) j

Pilosimanus (Villalobos, 1955:160),

Cubensis (Hobbs, 1962:278) ]

Pennides *(15)

Scapulicambarus (4)

Ortmannicus (Fowler, 1912:341) (42)

Tenuicambarus (1)

Villalobosus (8)

Procambarus (Ortmann, 1905c:437) (1)

Mexicambarus (1)

Lonnbergius (1)

Capillicambarus (2)

Leconticambarus (13)

Girardiella (Lyle, 1938:76) (5)

Austrocambarus (16)

Acucauda (1)

Remoticambarus (1)

Paracambarus (Ortmann, 1906:1) (2)

Blandingii

Pictus

Seminolae

Lucifugus

Planirostris

Disjunct members

Riojai

Erichsoni

Advena

Rogersi

Barbatus

Alleni

Mexicanus

Pilosimanus

Cubensis

•Numbers in parentheses following subgeneric names indicate numbers of species and subspecies assigned to them.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Subgenera of Genus Procambarus

1 Hooks on ischia of fourth pereiopods only (Fgure l/i) 2

V Hooks on ischia of third or third and fourth pereiopods (Figure \g, i, j) 4

2(1) Median spine on mesial ramus of uropod extending beyond distal margin of ramus

(Figure U) Acucauda

2' Median spine on mesial ramus of uropod not extending beyond distal margin of

ramus (Figure Ik), reduced to premarginal tubercle, or absent 3

3(2') First pleopod reaching coxa of third pereiopod, central projection beaklike (Figure

19C) Paracambarus

3' First pleopod reaching coxa of second pereiopod, central projection never beaklike

(Figure 13c) Villalobosus

4(1') Hooks on ischia of third pereiopods only (Figure \g) 5

4' Hooks on ischia of third and fourth pereiopods (Figure \i, j) 11

5(4) First pleopod reaching coxa of second pereiopod 6

5' First pleopod reaching coxa of third pereiopod 7

6(5) First pleopods asymmetrical (Figure 2e) ; coxa of fourth pereiopod with caudomesial

boss (Figure 3b) Hagenides (part)

6' First pleopods symmetrical (Figure 2h); coxa of fourth pereiopod without caudomesial boss (Figure 3a) Mexicambarus

7(5') Coxa of fourth pereiopod with caudomesial boss (Figure 3b) 8

7' Coxa of fourth pereiopod without caudomesial boss (Figure 3a) 9

8(7) Albinistic; first pleopod with only two terminal elements; subapical setae absent

(Figure 16b-d) Remoticambarus

8' Pigmented; first pleopod with three or four terminal elements; subapical setae present

(Figure 9b—d) Leconticambarus (part)

9(7') First pleopod with platelike cephalic process. (Figure 15fe—d); rostrum with median

carina (Figure 15a) Procambarus

9' First pleopod with cephalic process tapering, rudimentary, or absent (Figures 5b—d;

7b—d); rostrum without median carina (Figures 5a, 7a) 10

10(9') First pleopod with prominent proximomedian lobe (Figure 2d); cephalic process

well developed, absent only in Procambarus hagenianus (latter with median spine

on mesial ramus of uropod extending beyond distal margin of ramus (Figure

1/) ; dorsal surface of palm of chela with widely scattered tubercles (Figure

7g) Girardielta

10' First pleopod with reduced proximomedian lobe (Figure 2b); cephalic process

rudimentary or absent; median spine on mesial ramus of uropod never extending beyond distal margin of ramus (Figure 1*) ; dorsal surface of palm of chela

with crowded squamous tubercles (Figure 5g) Austrocambarus

11(4') First three pairs of pereiopods with conspicuous brush of plumose setae extending

from basis to at least proximal part of merus (Figure In) Capillicambarus

11' First three pairs of pereiopods never with conspicuous brush of plumose setae extending from basis to merus (Figure \m) 12

12(11') Carapace with two cervical spine on each side (Figure Ic) Pennides

12' Carapace with one or no cervical spine on each side (Figure Id) 13

13(12') First pleopod usually reaching coxa of second pereiopod, never with sinuous mesial

process nor with Made like caudal process (Figure 8b-d); chela strongly depressed

with cristiform row of tubercles on mesial surface of palm (Figure 8g).

Hagenides (part)

13' First pleopod reaching coxa of third pereiopod, if reaching second, mesial process long

and sinuous, or caudal process bladelike and cornerous (Figures 9, 10, 12, 17,

\8b-d) ; chela not depressed and lacking cristiform row of tubercles on mesial

surface of palm (Figures 9, 10, 12, 17, 18g) 14

14(13') Bituberculate hooks on ischia of third and fourth pereiopods (Figure 1;).

Lonnbergius

14' Bituberculate hooks never present on ischia of both third and fourth pereiopods

(Figure 1») 15
NUMBER 1 1 7

Key to Subgenera of Genus Procambarus—Continued

15(14') Carapace strongly compressed laterally (Figure 18a); boss on coxa of fourth pereiopod greatly expanded ventrally, and directed in longitudinal axis of body (Figure

3d) Tenuicambarus

15' Carapace never strongly compressed laterally (Figures 9a, 12a, 17a); boss on coxa

of fourth pereiopod sometimes massive but never greatly expanded ventrally

(Figure 3b) 16

16(15') Shoulder on cephalic surface of right first pleopod folded caudally against mesial

face of mesial lamella (Figure le), that on left projecting cephalically.

Scapulicambarus

16' Shoulder present or absent on cephalic surface of first pleopods, if present, similarly

disposed on both members of pair, that of right member never folded caudomesially

(like Figure 1/) 17

17(16') Mesial surface of palm of chela barbate (Figure 9g) and/or cephalic process of first

pleopod arising from mesial surface of appendage and never hooding central projection (Figure 9a") ; cephalic surface of appendage never with fingerlike extension Leconticambarus (part)

17' Mesial surface of palm of chela never barbate (Figure I2g); cephalic process of

first pleopod arising from cephalic (Figure 12c) or lateral surface of appendage

(often hooding central projection) except in Procambarus viaeviridis in which

distally directed fingerlike extension, bearing apical tuft of setae, present at cephalic

base of central projection Ortmannicus

Acucauda, new subgenus

(FIGURES 1/, 2a, 4, 20rf)

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 26, 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
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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-SPECIES.—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 In, 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 Ik, 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.
NUMBER 1 1 7

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, 206)

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. (//.) 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 lm, 2/, 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.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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, 20*)

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 Lonnberg,

1895:6.

GENDER.—Masculine.

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

ETYMOLOGY.—Named in honor of Einar Lonnberg,

who described the only known member of the subgenus.

Mexicantbarus, 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 caudo-
NUMBER 1 1 7

mesial 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 2», 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.) enofilosternum 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 luci-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fugus (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 2;\ 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
NUMBER 1 1 7 11

of monographing this species-group at the time of his

death.

Subgenus Procambarus Ortmann

(FIGURES 21, 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, 206)

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
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with median spine never extending beyond margin of

ramus.

TYPE-SPECIES.—Cambarus paeninsulanus Faxon,

1914:369.

GENDER.—Masculine.

LIS T 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.
NUMBER 1 1 7 13

Literature Cited

Bouvier, E. L.

1897. Sur les Cambarus recueillis au Mexique par M.

Diguet. Bulletin du Museum d'Histoire Naturelle,

3(6):224-228.

Erichson, W. F.

1846. Ubersicht der Arten der Gattung Astacus. Archives

fur 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.

Lonnberg, Einar

1895. Cambarids from Florida, a New Blind Species.

Bihang till Koniglische Svenska VentenskapsAkademiens 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 Gcrevisse nouvelle du Mexique. Annales des

Sciences Naturelles, Neuviime Sirie, 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.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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.

1933c. 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 Crust aces nouveaux de l'Ainerique tropicale. Revue et Magasin Zoologie Pure

et Appliquee, series 2, 9:501-510.

Villalobos, Alejandro

1944. Estudios de los cambarinos mexicanos, II: Dos

especies nuevas del genero Paracambarus. Anales

del Instituto de Biologia, Uniiersidad Nacional

Autonoma de Mexico, 15( 1): 161-174, plates 1-2.

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Descripcion de una nueva especie del g6nero

Paracambarus. Anales del Instituto de Biologia,

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379, plates 1-19.

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de Ciencias. Universidad Nacional Autonoma de

Mexico, pages 1-290, plates 1-62.
NUMBER 1 1 7 15

cephalic process

-central projection

-caudal process

-mesial process

\—caudal knob—.

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: t, 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, I, dorsal view of sixth abdominal segment, telson, and uropods [k,

P. (Girardiella) gracilis; I, P. (Acucauda) fitzpatricki]; m, n, ventral view of third maxillipeds

and first three pereiopods [m, P. (Leconticambarus) barbatus; n, P. (Capillicambarus) hinei].
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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; I, P. (Procambarus) digueti [pms, proximomesial

spur]; m, P. (Remoticambarus) pecki; n, P. (Scapulicambarus) paeninsulanus; o, P. (Tenuicambarus) tennis; p, P. (Villalobosus) riojai.
NUMBER 1 1 7 17

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 d is tally: d,

P. (Tenuicambarus) tenuis, with compressed caudomesial boss directed in longitudinal axis of

body. (Broad articular membranes in flat tone.)
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

b d

FIGURES 4-7.—Type-species of the subgenera of Procambarus (4, Procambarus (Acucauda)

fttzpatricki Hobbs; 5, Procambarus (Austrocambarus) vazquezae Villalohos; 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; /, lateral view of first left pleopod of second form male; g, dorsal view of chela of first

form male.
NUMBER 1 1 7 19

FIGURES 8-11.—Type-species of the subgenera of Procambarus (8, Procambarus (Hagenides)

advena (LeConte) ; 9, Procambarus (Leconticambarus) barbatus (Faxon); 10, Procambarus

(LonnbergiusJ acherontis (Lonnberg) ; 1 1, 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; /, lateral view of first left pleopod of second form male; g, dorsal view of chela of first

form male.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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; 1:

Capillicambarus; 3: Acucauda; 4: Leconticambarus; e, 1: Pennides; 2: Lonnbergius.

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