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XM- PROCAMBARUS, A NEW SUBGENUS OF THE
GENUS CAMBARUS.

By A. E. Ortmann, PH.D.

In a paper just published ( Proc. Amer. Phil. Soc., vol. 44, 1905,
p- 91), the present writer has divided the genus Caméaris into four
subgenera: Cambarus (sens. strict.), Cambarellus, Faxonius, and Bar-
tonius. The first of these (Caméarus) is framed to receive the first
and second group of this genus, as distinguished by Faxon (excluding
C. pellucidus, which I refer to Laxonius).

In the paper referred to (p. 99), the writer has indicated, that the
association of Faxon’s first and second group into one subgenus might
possibly not be entirely correct, but, that a division according to the
number of legs which possess hooks on the ischiopodite, as introduced
by Faxon, does not exactly correspond to the natural affinities. He
also pointed out, that possibly the shape of the copulatory organs of
the males of certain Mexican and Cuban species is more important in
this respect.

The Carnegie Museum has lately purchased from Professor J. S.
Hine, Columbus, O., a number of specimens of a new species of Cam-
barus from Guatemala (Acc. No. 2778; Cat. Nos. 74,560 and 74,561),
belonging in this group, which, in the paper referred to, has been
designated as the dgueti-group (of the section of C. dgweti), con-
taining hitherto three species (digueti, mexicanus, cubensis). An
examination of the copulatory organs of this new form, and their com-
parison with the figures of these organs of C. carinatus, = digueti and
cubensis, with the description of these organs in C. mexicanus, and
with a male cotype of C. digueti, kindly presented by Professor E.
Bouvier, of Paris, has led to the conclusion, that all these species, as
well as the new one to be described herein, possess a type of male
copulatory organs, which is quite peculiar, and differs considerably
from that seen in other species of the subgenus Camdbarus (type: C.
blanding? ).

The description of this organ, as given by Faxon for his first and
second groups, and by the present writer for the subgenus Caméarus,
does not apply at all to these Central American and Cuban species.

4355

43 ANNALS OF THE CARNEGIE MUSEUM.

An attempt has been made, in the paper referred to, to also include
these forms in Caméarus, but, as is now evident, this is impossible.
The material at hand when the paper was written was too poor (only
one male of the second form of C. digweti) to decide the question
and, moreover, the description of these organs in C. mexicanus was
not well understood, since no figure had been published.

However, Faxon’s description of the male organs of C. mexzcanus
suddenly became clear to me when I compared it with the new species,
and plainly these organs in these two species are very similar. Since
good figures of those of the other two species have been published, it
is now possible to give a correct account of these organs, and to point
out their chief peculiarities.

Faxon’s description of the male copulatory organs in his first and
second group is as follows: ‘‘ Outer part truncate at the tip, and fur-
nished with one to three small recurved teeth ; inner part terminated by
a short acute spine, which ts generally directed outward.”

For the subgenus Cambarus, I have given the following characters
of the male organs: ‘* Sexwal organs of male stout, more or less straight,
and comparatively short, truncated or blunt at the tip, the outer part
ending in one to three horny teeth, which are sometimes recurved, or
compressed, or plate-like, and are always sharply distinguishable by means
of the blunt end. Inner part terminated by a shorter or longer acute
spine, which 1s sometimes distinct from the tip of this part, so that it
appears two-pointed.”’

This latter description was framed with the desire to make it fit the
species of the dgwefi-group ; but it neglects the fact, that in these
species the outer part of the male organs does not possess any horny
teeth, as is generally the case in species of the type of d/andingz. In
the species of the d@gweti-group, it is the zzwer part that possesses, in
the male of the first form, a single horny spine, which has a variable
position in the different species, but is always highly characteristic for
one and the same species. It has taken me some time to ascertain
this fact; indeed, I admit that I may be mistaken in so far as that
there is a remote possibility that this horny spine belongs to the outer
part, since in the new species to be described here, its position is just
at the point where the two parts (inner and outer) become distinguish-
able at the anterior margin of the organ; but after examining the con-
‘dition rn second form males of the new species and of C. dguwe/7, I feel
rather sure that this accessory tip, which is not horny and spiniform

PrROCAMBARUS, A NEW SUBGENUS. 437

in this case, but tuberculiform, belongs to the inner part. (Compare
also Faxon’s figure of C. carinatus = digueti, Proc. U.S. Mus. 20,
T5q5; ple 63, f..2%)

Thus this organ is fundamentally different from that of the subgenus
Cambarus as represented by its type (C. d/andingz), and, moreover,
it possesses at its anterior margin, not far from the tip, a peculiar
angular projection, which I have called the ‘‘shoulder.’’ Such a
shoulder is found in a few cases in other species belonging to the sub-
genus Cambarus (C. clarki and troglodytes), and in numerous species
of the subgenus Faxonius (section of C. propinguus); Dut here it
always has a different position, being quite remote from the tip of this
organ.

Thus I think we are justified in emphasizing this peculiar feature of
the male organs by creating for it a separate subgenus, for which the
name Procambarus is proposed, and we consequently have now jive
subgenera within the genus. The diagnoses of the two first are as
follows.

Procambarus subgen. nov.

Sexual organs of male stout, more or less straight and comparatively

short, rather blunt at the end, but not truncate. The two parts in

[a close épposition up to the tips. The outer one ending bluntly, with-
out horny teeth ; the inner one being more or less similar to the outer
one, but possessing in various positions one horny spine. Anterior
margin with a shoulder near the tips. In the male the third pereiopods
have hooks.

Species: diguett, zeilliamsont, mexicanus, cubensis.

Cambarus (Ortmann sews. restrict).

Sexual organs of male stout, more or less straight and comparatively
short, very blunt, or truncated at the end. The two parts in close
[a épposition, with exception of the terminal spines. The outer part
ending in one to three horny teeth, which are often recurved, or com-
pressed, or plate-like, being sharply distinguishable from the blunt
end. Inner part terminated by a shorter, or longer, acute spine. A
shoulder on the anterior margin is rarely present, and it is, if present,
quite remote from the tip. In the male the third, or the third and
fourth, pereiopods have hooks.
The diagnoses of the other three subgenera: Cambarellus, Faxonius,
Bartonius, do not require any change.

438 ANNALS OF THE CARNEGIE MUSEUM.

Thus, from my subgenus Camébarus, certain species are removed as
the subgenus Procambarus, which would belong to Faxon’s second
group: but not the whole of Faxon’s second group belongs to Pro-
cambarus, since the other species, which I have designated (7. c.) as
the szmu/ans-group and the section of C. graczlis, possess sexual
organs that entirely correspond to those of the section of C. d/anding?.
In the subgenus Caméarus, consequently, three sections remain :

Section of C. simulans :

Male with hooks on third pereiopods. Areola rather narrow, but
never obliterated in the middle, slightly longer than half of the ante-
rior section of the carapace. Chelze elongated and subcylindrical.

Species: s¢mulans, gallinas.

Section of C. gracilis :

Male with hooks on third pereiopods. Areola obliterated in the
middle, considerably longer than half of the anterior section of cara-
pace. Chelz short, broad, ovate.

Species: gracilis, hagenianus, advena.

Section of C. blandingi :

Male with hooks on third and fourth pereiopods. Areola wide, or
narrow, rarely obliterated in the middle, shorter, or longer. Chelz
generally elongated, narrow and subcylindrical.

Divided into four groups, containing together about 15 species.

The species of the subgenus Procambarus are easily distinguished
by the following characters :

;- Inner face of male sexual organs flattened, but not broadly dilated.

6,. Sexual organs of male slightly curved backward at apex ; horny spine of
inner part straight, situated posterior to the tip of the outer part. Rostrum
carinated above. Carapace with one lateral spine on each side.

C. ( Procambarus) digueti Bouvier.

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1°

6,. Sexual organs of male straight; horny spine of inner part curved, situated
anterior to the tip of outer part. Rostrum not carinated above.

Rostrum with marginal spines. Carapace with two lateral spines on

each side. C. ( Procambarus) williamson? spec. nov.

Cy.

Rostrum without marginal spines. Carapace without marginal spines.
C. (Procambarus) mexicanus Erichson,

Co .

a,. Inner face of male sexual organs flattened and greatly dilated, forming a broad,
flat, setose plate. Rostrum not carinated, with marginal spines. Carapace
without lateral spines. C. ( Procambarus) cubensis Erichson.

PROCAMBARUS, A NEW SUBGENUS. 439

DESCRIPTION OF NEW SPECIES.

CAMBARUS (PROCAMBARUS) WILLIAMSONI spec. nov.

Diagnosis. — Male sexual organs with the outer part blunt, the
inner slightly longer than the outer, narrower, but blunt ; in the first
form with a horny, procurved spine at the anterior margin, which is
represented by a blunt tubercle in the male of the second form. The
inner face of inner part slightly dilated and flattened, with two rows
of bristles diverging from a low oblique rib. Rostrum with distinct
lateral spines, acumen short ; its upper surface subplane, without keel.
Sides of carapace granulated, with two spines on each side behind the
cervical groove. Areola very narrow, about half as long as the anterior
section of the carapace. Chel long, subcylindrical, thickly squamoso-
granulated, the granulations larger on inner side of the carpopodite
and hand, only in young specimens with an indistinct spine. Finger
shorter than the palm. ‘Third pair of pereiopods with hooks in the
male.

Description of Male of First Form. — Rostrum subplane, without
median keel, margins slightly elevated, convergent, with a distinct
lateral spine on each side a short distance from the tip. Acumen
short, triangular, with a fringe of stiff hairs. Postorbital ridges
divergent posteriorly, ending in a small spine anteriorly. Carapace
laterally compressed, suborbital angle rounded. Branchiostegal spine
distinct. Sides of carapace finely, but distinctly granulated; two
sharp lateral spines on each side behind the crevical groove. <Aveola
about half as long as the anterior section of carapace, including the
rostrum, very narrow in the middle, with a single irregular row of
punctations. Addomen as broad as, and slightly longer than, carapace ;
epimera rounded ; anterior section of telson with two spines on each
side. Anterior end of efzstoma broadly triangular. Antenna with the
flagellum longer than the carapace, but shorter than the whole body.
Antennal scale broad, broadest in the middle. Che/e subcylindrical,
long, thickly beset with squamiform granulations, which are largest
on the innerside. Fingers not gaping, shorter than palm, granulated,
and indistinctly longitudinally costate on upper side; slightly pubes-
cent at the cutting edges, and with a few longer hairs. Carpopodite
without sulcus above, granulated like hand, granules largest on inner
side, but not spiniform (with exception of young specimens, where
there is a small spine at the distal end of inner margin). Meropodite

440 ANNALS OF THE CARNEGIE MUSEUM.

granulated, almost smooth on proximal parts of outer and inner side,
with a small spine near distal end above. Lower side with two rows
of spiniform granulations, most distinct in young individuals. Ischi-
opodite of the ¢Azrd pair of peretopods with a strong hook. Cofulatory
org7ns (first pair of abdominal appendages) short and straight, with a
prominent shoulder on the anterior margin near
the tip. Outer and inner part in close apposition
to their tips ; outer part ending bluntly, the inner
slightly longer and thinner, but also blunt. ‘The
inner part carries, at its anterior side, not far from
the shoulder, a small, procurved, horny spine.
Inner face of inner part flattened, and slightly
dilated, with two rows of bristles, which diverge
from a low, oblique median rib.

The male of the second form differs in the less
distinctly granulated carapace; the chelz are

Gis weaker; the hooks of the third pereiopods smaller;
3 the copulatory organs are similar, but the horny,

Fic. 1. Male copu- procurved spine is replaced by a blunt tubercle
latory organ of right that is not horny. (This gives the whole organ

side seen from outside. three blunt tips. )
Fic. 2. Male copula-

tory organ of right side ; ; :
seen frominside. Fic, 47¢lus ventralis transversely rhomboid, forming
3. Annulus ventralis asimple tuberculiform elevation, with an S-shaped
of female. All figures ]ongitudinal fissure, but without any depressions.
enlarged twelve diam-
eters.

Female: Similar to the male of the second form.

Locality: Los Amates, Province of Izabal, Gua-
temala. In small streams and swales. January
20, 1905; collected by E. B. Williamson, of Bluffton, Ind. (About
4 to 5 miles due south of the town of Izabal, south of Rio Malagua.)

There are 27 specimens in the lot, 13 ¢', 14.9. Only one (the
largest) is a male of the first form. Most of the specimens are very
young.

The measurements of the largest J (1) and 9 are as follows:

d'. Total length 51.5 mm., carapace. 25 mm., anterior section of
carapace 16.5 mm., areola 8.5 mm., width of areola in the middle
o.5 mm., abdomen 26.5 mm., length of hand 20 mm., of palm 11
mm., of fingers 9 mm.

QO. Total length 49.5 mm., carapace 24.5 mm., anterior section of
carapace 16 mm., areola 8.5 mm., width of areola in the middle 0.5

PROCAMBARUS, A NEW SUBGENUS. 44]

mm., abdomen, 25 mm., length of hand 13 mm., of palm 7 mm., of
fingers 6 mm.

The specific name is given in honor of the discoverer, Mr. E. B.
Williamson. Before Mr. Williamson started for Guatemala, I called
his attention to the possible presence of a Cambarus in this country,
and his search was rewarded by the discovery of this species.

REMARKS ON THE GEOGRAPHICAL DISTRIBUTION.

The existence of a Camdbarus in Guatemala (Coban, Alta Vera
Paz) was first mentioned by Huxley (1878), but the species has
never been described. It remains doubtful whether the present
species is the same, since it comes from another part of the country,
although both localities belong to the Atlantic slope.

Cambarus williamsoni is sharply distinguished from all other species
of the genus, although it forms a natural group (subgenus Procamébarus )
with three other species, two of which are found in Mexico (dgwezz,
mextcanus), and the third in Cuba (cubensis). The most closely
allied form is C. mexicanus, but this is apparently more highly
specialized, as is shown by the lack of the marginal spines of the ros-
trum and the lateral spines of the carapace. Thus C. wzl/zamsonti is
to be regarded as a rather primitive form, and stands between C. dg-
uetti and C. mexicanus. Ihave regarded C. digwe/ as the most primi-
tive form of the whole genus; but second to it-is C. we/damsont. C.
mexicanus comes very Close to the latter, while C. cwdensis is more
remote from the rest on account of the broadly dilated inner face of
the male organs ; but the beginning of the development of this feature
is clearly indicated in the other three species.

Thus it is beyond doubt, that C. cwéenszs points in its affinities to
Guatemala and Mexico, and our Guatemalan species indicates the
way by which Cuba was populated. In my paper referred to (Prec.
Am. Phil. Soc., 49, 1905), I have already assumed this connection
on the map, without knowing anything of the existence of C. we/-
liamsont, and this assumption has been fully confirmed by the dis-
covery of this species.

The subgenus Procambarus appears to be the earliest type of the
genus, and its range is in the original home of the genus (Mexico and
Guatemala), with the exception of C. cuwdensi?s, which apparently is
an early emigrant.

The separation of the subgenus Procambarus from the rest of the

4492 ANNALS OF THE CARNEGIE MUSEUM.

genus, chiefly from the typical subgenus Camdarus, renders the Mexi-
can crayfish-fauna rather peculiar. In Mexico, two well known sub-
genera are represented: Procambarus and Cambarellus. The first
one has no representatives at all in the United States, while Cam-
barellus has only one (in Louisiana). There remains a Mexican
species, C. zvegmanni, which has been placed with species of the true
subgenus Cambarus, but its position is quite uncertain, the male
copulatory organs being unknown.

CARNEGIE MUSEUM,
June 10, 1905.

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