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dALIFdRNIA ACADEMY OF SCIENCES.
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Third Series.
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Zoology.
Vol. I," No. 9.
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IX.
BY .
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THOMAS H. MONtaOMERY,. JRm PH.D.,
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David S. Jordan, " V^ii*uam E- Ritter.
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PROOBBDINOS
CALIFORNIA ACADEMY OF SCIENCES.
Third Series.
Zoology.
Vol. I, No. 9.
The Gordiacea of Certain American
Collections, with Particular Reference
to the North American Fauna.
II.
BY
THOMAS H. MONTGOMERY, JR., PH.D.,
Lttturtr in Zatlagy, Viiiotrtify tf Ptmuyhmnia.
With Two Platbs.
Issued October t», 1898.
SAN FRANCISCO:
PUBLUHBD BY THB ACADBHV.
189&
V.
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THE GORDIACEA OF CERTAIN AMERICAN
COLLECTIONS, WITH PARTICULAR REF-
ERENCE TO THE NORTH AMERICAN
FAUNA.— II. *
BY THOMAS H. MONTGOMERY, JR., PH. D.,
Ltclmrtr im Zt«U g r> Univmitf »f Puuuylvanm,
Plates XIX and XX.
The present paper deals with Gordiacea, principally of
California, and with a few specimens from neighboring
regions (Baja California, Arizona, Nevada, and Mexico).
The fauna of California has been heretofore but little
known, so that the comparatively large number of speci-
mens from that locality which have been offered to me for
study have a special interest. In a previous contribution *
Gordiacea were described by me from other portions of
North America.
The following collections are herein described : that of
the California Academy of Sciences, San Francisco; of
the University of California; and of Stanford Uni-
versity, California. For the opportunity to study these col-
lections, my thanks are due to Dr. Gustav Eisen of the
California Academy of Sciences, and to Prof. Henry B.
Ward of the University of Nebraska.
Appended to this paper is a Key for the determination of
the species of Gordiacea on the North American continent,
north of Mexico. While this key may serve to determine
with rapidity certain species, it must be noted that some of
them, and particularly the females of Gordius and Chor-
dodes, can only be satisfactorily determined by means of
sections and of portions of the cuticle cleared in glycerine
or some clearing oil (I am accustomed to clear them in
cedar oil, after dehydration, and to study them mounted in
Canada bakam). For descriptions of species which are
I BvU. Mm. Conpar. ZooL, Uarrard Univenity, April, i8g8.
L333J
Oct. 1, 1898.
ifi^
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334
CALIFORNIA ACADEMY Of SCIENCES. [Proc. 3D Sbh.
here mentioned only briefly, the reader is referred to my
previous paper, 1. c. In this Key the females of Gordius
aquaticus difficilis, G. longareolatus, n. sp., Chordodes puer-
ilis, and the male of C. morgani find a place, though I have
seen only the males of the first three mentioned, and the
female of the last; this was done with the assumption that
the cuticular characters of the two sexes are similar in
these forms.
It is here shown that Chordodes gordioides Montg. is
identical with C. occidentalis Montg.
z. Gordius longareolatus, sp. nov.
Type.—Qo^. Cal. Acad. Sci.: i <S , San Francisco, Cal, Sept. 10, 1896.
/bTM.— Of equal diameter in its whole length, nearly cylindrical, with faint
median grooves. Head end (fig. i) conically rounded, the tip slightly con-
stricted off, mouth ventral (on the ventral side of the white tip of the head).
Tail lobes (figs, i, 3) short, thick, nearly parallel, terminally rounded. Their
point of bifurcation (on the ventral surface) is at a distance from the oval
cloacal aperture about equal to the length of the lobes. Each lobe is approx-
imately cylindrical, except on its median surface which is concave ; this con-
cavity is deepest nearest the posterior end of the lobes and is bounded by a
sharp line. The cloacal aperture is situated upon a slightly elevated papilla ;
firom this papilla to the point of bifurcation of the tail lobes extends a deep
groove( deepest caudad). The area around the cloacal aperture, and extend-
ing caudad to about the point of bifurcation of the tail lobes, is covered with
numerous, minute, conical spines.
C»/tV^.— With elevated areoles of elongate form, their longitudinal axes
parallel to that of the body (fig. 4). The areoles do not produce particular
longitudinal rows, though everywhere the spaces which separate them are
about equal in length. They are darker than the intermediate portions of
the cuticle. Most of them are several times longer than broad, with nearly
parallel margins ; a smaller number are shorter and thicker, but even these
are somewhat elongate. On cross section of the body (fig. 5) the areoles are
seen to be hyaline and homogeneous, their darker, denser bases sunk into
the surface of the fibrous cuticle. On section they show a rounded-conical
form, without any median groove. No hairs lie between the areoles, but
wklely separated from one another are found a few hyaline, finger-shaped
processes, of slightly greater elevation than the areoles ; such processes have
heretofore been noticed by me only in the genus Chordodes.
Coftw.— Deep olive brown, slightly iridescent ; the tip of the head white.
iMfNtfiwibfU.— Length, 115 mm.; greatest diameter, 0.5 mm.
«^
'«^
,^»«».
ZooL.— Vol. i.]
MONTGOMER Y—COKPIACEA.
335
Comparison. — The only other Gordiacea with longitudi-
nally arranged elongate areoles are G. raphaelis, alfredi,
and latastei oi Camerano, and G, abbreviatus ViWoV, but
all of these differ from G. longareolatus in having the
areoles longitudinally grooved.
a. Gordius aquaticns Linn,
CoU. Cat. Acad. Sci.: 2 i i , City of Mexico; 2 S 6 , Berkeley, Cal., winter
of 1891. Coll. Univ. of Cal.: i , Berkeley, Cal., January, 1895.
These are the only specimens as yet seen by me of the
true aguaticus from North America ; in all of them the white
spots (figs. 13, 14) are clearly marked on the surface of
the cuticle and may be seen with a low-power hand lens.
3. Gordius tquatictts robustus {Leidy.)
Coll. Cal. Acad. Sci. : $ , San Francisco, Cal. ; $ , Berkeley, Cal. Coll. Univ.
Cal.: 9, Berkeley, Cal., January, 1895.
The largest specimen measured 590 mm.
This is the form, corresponding to G. robustus Leidy,
which has been regarded by me (1. c.) as a mere variety
of the preceding, from which it differs merely in the absence
of the white cuticular spots. In the true European aguati-
cus the white spots are present in most individuals, accord-
ing to the description of Rdmer'; while in specimens of
robustus from the eastern United States (from Kansas east-
ward) I have never found these spots, though I have studied
one or two hundred specimens. Westward of California
occurs the true aguaticus, eastward the aguaticus robustus,
while in California the two appear to overlap. If white-
spotted specimens be subsequently found in any number
from the eastern United States, then aguaticus robustus can
no longer be retained as a separate subspecies ; but with
the facts at hand we are justified in assigning it that
position.
> Beitrag car SyttematUc dcr Gordiiden Abh. ScckcnberK. Om., Bd; XXin, 1896.
i !ifij).i| l !l | l l Wtt* H«ll>W«W! aW«lll»'!H. I
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336
CALIFORNIA ACADEMY OF SCIENCES. [P«OC. 3D Sbk.
4. Gorditts densareolatns Montg.
Coll. Cal. Acad. Sci. : i , Lake, San Francisco, Cal.
The posterior end of this specimen was as in the type
(cf. Montgomery, 1. c), except that there was only a slight
precloacal groove.
In my first description of this species (1. c.) the cuticle
of the female only was described. I have since re-examined
the type of the male (Montana, Leidy coll., No. 5063) and
find the cuticle in general like that of the female, except
that the areoles show less tendency to confluence and are
more rounded in outline.
>»>^
5. Paragordins yarius {Leidy.)
Coll. Cal. Acad. Sci.: 3 J J , 3 v $, San Jos^ del Cabo, Baja Cal., July,
1896; $, St. Helena, Napa Co., Cal. Coll. Stanford Univ., collected by
C. H. Gilbert: 5 $ $, Napa Creek, Calistoga, Napa Co., Cal., May 31,
1897.
This appears to have the broadest range of any North
American species, extending completely across the conti-
nent, and, according to Camerano, into South America.
In the females the tail lobes may be either parallel or
divergent, so that they are probably opened and closed by
muscular action ; while in the males of this genus and of
Gordius these lobes appear to be to great extent independ-
ent of such action.
6. Chordodes occidentalis Montg.
Coll. Univ. Cal. : 4 , $, Berkeley, Cal. Coll. Stanford Univ., collected by
C. H. Gilbert:' it3 S J. 43 $ $, Napa Creek, Calistoga, Napa Co., Cal..
May 31, 1897. Coll. Cal. Acad. Sd. : 2 $ ?, i 4 , Haywards, Alameda
Co., Cal.; I $, Alameda Co., Cal.; 2 4 S , 2 $ $. Santa Rosa Isl., Cal.;
4 $ $, Sierra el Taste, Baja Cal.; i ?, St. Helena, Napa Co., Cal.; 1 ?
Kings River, Fresno Co., Cal.; i S , Lime Point, Marin Co., CaL; i S,
San Miguel de Comondu, Baja Cal.; i $,5 i i , locality not marked, but
probably fi-om California ; i $ , Batde Mountain, Nevada.
A careful examination of this large number of specimens
shows that the Chordodes gordloides Montg. of my preced-
ing paper was based upon immature specimens of C. occi-
dentalis Montg., so that gordioides is not a tenable species.
tmmmmmm
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• vmijfffi-x.r, ' raawwwWi
ZooL.— Vol. i.]
MONTGOMER Y—GORD/ACEA.
337
^m^
Owing to the great amount of individual variation afforded
by these specimens from California (123 ^ j>55 9 ^ ,) ^
description of them is necessary, and this description may
be compared with that of occidentalis and ^* gordioides*^
given in my preceding contribution.
Form.—Th& form of the male has already been described and figured, so
that it is only necessary to note that the anterior portion of the body is the
most slender, the head (fig. 10) pointed, and the body either cylindrical or
flattened (depending upon the state of development of the genital products).
The females resemble the males in general form, and in them also the ante-
rior portion (fig. 6) is the most slender, they are stouter than the males ; the
posterior end (figs. 9, 13) is enlarged and more or less knob-shaped as in all
females of this genus ; seen from the ventral surface (fig. 23) this posterior
end appears disc-shaped, with the cloacal aperture in its centre.
C»/i<r/^.— Thirty-two specimens were examined, cleared in cedar oil, and
transverse sections were made of twenty-four of them. These preparations,
with those which formed the basis of my previous descriptions, show how
variable the configuration of the cuticle is in this form, and present all inter-
mediate phases between the figs. 115-117, PI. XV, of C. occidentalis of my
previous description, and figs. jo6, 107, PI. XIII, 1. c, of C. '■^ gordioidei"
and leave no doubt that the latter is only the young of the preceding.
On surface view the areoles may be irregularly polygonal and well sepa-
rated from one another, or they may be elongate and confluent so as to pro-
duce transverse rows (figs. 7, 13-30). The areoles of the same individual
may be all of the same color, which appears to be usually the case, or groups
(of a few each) of larger areoles may be darker than the others (fig. 18). The
cuticle of one specimen had the appearance of that of Gordius pUskei Camer. ,
owing to the incrustation of minute dirt particles between the areoles (fig. 13).
Small, refractive, circular or oval pits occur in varying number on the cuticle ;
they lie for the main part between areoles, sometimes on their surfaces ; their
most fi^uent mode of occurrence is between two areoles which together
present a dumb-bell shape ; their number varies greatly, and they are found
either singly, in pairs, or in threes (figs. 7, 15-20).
On transverse section (figs. 8, 11, 31, 33) the areoles are usually of the
same height but varying diameter in the same individual, but are never as
high as long, though in the males they are usually higher than in the females ;
they are either close together or well separated. In a few specimens, as in
the type first figured, tower areoles with irregularly serrated summits lie
between the higher ones, but such a relation was found in only three or four
cases. Between the areoles, occurring only at wide distances, are delicate
finger-shaped or clut>-shaped hyaline processes, of greater height than the
areoles. In a few specimens were noticed hyaline processes of a stcdt coni-
cal form, sunk into the summits of certain of the areoles (fig. 23). In some
males a cupn^haped depression occurs on the summits of a few of the areoles.
The circular or oval pits seen on sur&ce views are found on section (fig. 8)
to represent structures, the like of which has not yet been described for any
Gordiacean ; they were overlooked in my description of the type spedmen,
i«^^ ' :j'-,yi^y!i^^i'i-^"' ' " •' "•"'•"* 'i--^^ssiur
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338
CALIFORNIA ACADEMY OF SCIENCES. [Proc. 30 Ssm.
owing to their paucity in that individual, but I have found them on re-exam-
ination. They were largest and most abundant in a large female, and appeared
to be absent in only three specimens. Each pit has the form of a sack of
spherical or ovoid shape (some are flattened parallel to the surfece of the cuti-
cle), with an exterior opening on the surface of the cuticle ; the length of
these sacks varies from about one-fourth to two-fifths the diameter of the
cuticle. On section it would appear that almost all of them lie between
arcoles, sunk into the fibrous cuticle, though on sur&ce view they frequently
seem to lie on the summits of areoles. The outline of each pit is rounded
and sharply demarcated, though no special limiting membrane appears to be
present. In each lie two (rarely three) refractive spines, which are usually
sitiuted vertical to the siuiace of the cuticle, but in some cases parallel to it.
Each spine is somewhat rod-shaped, thickest basally, the base obtusely
rounded and usually in close contact with the wall of the pit. At its distal
end each spine has a short, pointed process which projects out of the narrow
opening of the pit ; the longitudinal axis of this pointed portion does not
always coincide with that of the basal portion. Thus these spines are some-
what similar in form to the central stilets (with their bases) of the Metane-
mertean proboscis.
Color. — The males vary from a light yellowish, through rufous or chocolate
or olive-brown, to a nearly pitch black color, smaller individuals being usually
but not always lighter than larger ones. In most of them the anterior portion
of the body is lighter than the posterior ; the head is lightest in color, very
frequendy yellowish white, while the rest of the body is chocolate-brown.
Just behind the light, head portion is usually a more or less pronounced dark
neck ring, which is seldom wholly absent. The females averse much
lighter in color than the males, and are usually a yellowish or yellowish
brown color ; a few are a dark olive-brown, but none are black. In them,
too, there is usually a darker neck ring, and often a brownish area around
the doacal aperture. In both sexes the mouth aperture is usually marked
by a small black spot, easily seen with a hand lens : thus of the 154 individ-
uals collected on the same day at the same locality (coU. Stanford University),
96 males showed this spot, 15 males were without it, and in a males die
whole head was black ; and all the 43 females showed the spot. The black
mouth spot would thus appear to be pretty constant (though it was absent
in the type specimens of this species).
DinuHSums.—\MC%<eiA male, 460 mm.; largest female, 420 mm.
Comparison. — C. gordioides Montg. becomes a synonym
of this species. This species differs from C. bedriaga and
C. fardalis Camer. , to which the configuration of its cuti-
cle appears to be otherwise quite similar, in the presence
of the remarkable cuticular pits, the structure of which,
however, can only be determined by sections. Such pits
have never heretofore been described : but clear, rounded
cavities situated just below tubercles or aeroles of the cuticle
'Sa r**
■-^ »-
iMHMiMM
ZooL.— Vol. 1.]
MONTGOMER Y— GORDIA CEA.
339
' S* !**
would seem to be frequent in Chordodes, at least I find
them in my species of C. morgani, C. puerilis, and C. albi-
barbatus, though not in C.furnessi.
General diagnostic characters. — The union of the follow-
ing characters serves to distinguish this from any other
North American form: the low aeroles with the remark-
able interareolar pits, the attenuation of the anterior portion
of the body, and the black mouth spot. The females of
this species may easily be mistaken for females of G. den-
sareolatus Montg., since in some specimens of the former
the interareolar pits may be very few in number (or entirely
wanting ?). But the form of the posterior end is a good
distinguishing character : in the female of C. occidentalis it
is enlarged — somewhat knob-shaped, while in the female
of G. densareolatus it is not of greater size than the preced-
ing portion of the body ; there is at least no difficulty in
distinguishing them when one has a series of specimens of
both before him.
■^ ►•
Key to the Species of Gordiacea of the North
American Continent, North of Mexico.
I. Posterior end trilobed Paragordius varius (Lbidy), $ .
II. Posterior end bilobed, spirally inrolled..(/)iraj'<wdrfW and Gordius, i 6),
A. A sharp, V-shaped cuticular ridge behind the cloacal aperture.
a. The cuticle dotted with viWX&si^Xs.. Gordius aguaticus Linn., i .
b. The cuticle not dotted with white spots.
1. A parabolic line of hairs on the tail lobes.
G. a. difficilis Montg., i .
2. No line of hairs on the tail lobes ..G, a. robustus (Lbidy), i .
B. No sharp, V-shaped cuticular ridge behind the cloacal aperture.
a, A iongitudhial line of long hairs on each side of the cloacal
aperture ..G. lineatus Leidy, i .
b. No line of long hairs on each side of the cloacal aperture.
1. Head end obliquely truncated.
Paragordius varius (Lbidy), i .
2. Head end not obliquely truncated.
a. Conical spicules behind the cloacal aperture.
I. Tail lobes short, nearly conical.
Gordius densareolatus Montg., i .
3. Tail lobes neariy cylindrical.
G. longareolatustllovriG., i .
CALIFORNIA ACADEMY OF SCIENCES. [P»oc. 3D Sbr.
*. No conical spicules behind the cloacal aperture.
G. pla/ycepAalus MouTG., S.
Posterior end not lobed, spirally inroUed, with a depression or
groove on its ventral surface, narrower than preceding portion
of body (Chordodes, S i).
A. Cuticular areoles longer than high, on and between them small
circular pits Chordodes occidentalis Montg., <J .
B. Cuticular areoles higher than long.
a. Spines on the summits of the highest papillae.
C. pueritis Montg., .5 .
b. No spines on the summits of the highest papilke.
C. morgani Montg., i .
. Posterior end not lobed, not spirally inroUed.
A. Posterior end not noticeably enlarged (Gordius, ? ? ).
a. With elevated cuticular areoles on the whole sur&ce of the
body.
I. With paired dark stripes in the median lines.
Gordius leidyi Montg., ? .
a. No dark stripes in the median lines.
Areoles elongated in the long axis of the body, well
separated G. longareolatus Montg., ? .
*. Areoles not elongated in the long axis of the body.
1. Areoles closely apposed, tending to produce
longitudinal ridges G. lineatus Lbidv. ? .
a. Areoles more or less confluent, tending to pro-
duce transverse rows, head usuaJly cylin-
drical G. densareolaitts Montg., ? .
3. Areoles usually separated, interareolar groups
of fine hairs, head usually flattened.
G. platycephtUus Montg., $.
*. Without elevated cuticular areoles on the whole surface of the
body.
I. No cuticular areoles.
a. Cuticle dotted with white spots. .C aquaticus Linn., ?.
*. Cuticle without white spots. ..G. a. robuslus (Lbidy), ? .
a. Cuticular areoles at the ends of the body only.
G. a. difficilis Montg., $ .
Posterior end swollen, somewhat knob-shaped, slightly con-
stricted off.. (Chordodes, ? ?).
Cuticular areoles longer than high, on and between them
small circular pits Chordodes occidentalis, Montg., $ .
Cuticular areoles higher than long.
I. Spines on the summits of the highest areoles.
C. puerilis Montg., $ .
a. No spines on the summits of the highest areoles.
C. morgani Montg., 9 .
Wistar Institute of Anatomy,
Philadblphia, Pbnnsvlvania.,
June ao, 1898.
W
•'*■»-
liiiiiii
342 CALIFORNIA ACADEMY OF SCIENCES. [P«oc. 30 8b».
l:i
EXPLANATION OF PLATE XIX.
(All the outlines were drawn with the aid of the camera ludda and a Zete
microscope and lenses. The cuticle is represented as seen in Canada bal-
sam, after clearance in cedar oil.)
Gordius lot^areolatus, sp. nov., type. Head end (otq. A, oc. a).
Idem. Ventral view ofposterior end (idem).
Idem. Lateral view of the posterior end, the ventral side to the
left (idem). ^ . ^u v t
Idem. Surface view of the cuUde, the arrow denoting the hne ot
the longitudinal axis of the body (obj . C, oc. 4)-
Idem. Transverse section of the cuticle (homog. immers. i/ia,
OC 2)
Chordodes occidentalis Monto., dorsal view of the head of a female
(obj. A, oc. 2).
Idem. Surface view of the cuticle (obj. C, oc. 4).
Idem. Transverse section of tiie cuticle (homog. immers. 1/12, oc.
4, tube lengtii j8o mm.). . .j . »u
Idem. Lateral view of the posterior end, the ventral side to the
right (obj. A, oc. a). , ^. x. a
Chordodes occidentalis Montg., male. Lateral view of the head
end (obj. A, oc. 2). .
Idem. Transverse section of tiie cuticle ; in tins figure, as in the
following, dirt granules are imbedded between the areoles
(homog. immers. 1/12, oc. 2).
Idem. Surface view of the cuticle (otg. C, oc. 2).
Gordius aquaticus Linn., male from Berkeley, California. Sur-
face view of the cuticle ; the intersecting clear lines transverse
the white spots (obg. C, oc. 2). ^
Idem. Surface view of tiie body, as seen witii low na^ification
(obj. A, oc. 2), to show tiie arrangement of the white spots.
The continuous lines represent the outiines of the body, the
dotted lines the area to which the drawing was limited.
Fig.
Fig.
Fig.
I.
2.
3-
Fig.
4-
Fig.
5-
Fig.
6.
Fig.
Fig.
8.
Fig.
9-
Fig.
10.
Fig.
II.
Fig.
Fig.
12.
13-
Fig. 14
-.H
•?"?^
PRnr„nALAcAiJ,-5cT,3°5EP.Znai,VDi,l
fM'i:!T:''3^'H':] PIjATE XIX.
1.
7.
9.
\^
10.
MowrcauEiv.ila..
S.
v_/
4.
5.
6.
11.
■ •■.'Htlf.'
m
13.
^^^.
^..A V-- — »•■■ ■■■>>...-■■-'
t:---'
12.
..0..-
14.
LlTFU-wmnK 5 tthV, a.i
344
CALIFORNIA ACADEMY OF SCIENCES. [Piioc. 3D Skr.
EXPLANATION OF PLATE XX.
All figures refer to Califomian specimens of Chordodes occidentalis Montg.
Figs. 15-30. Surface views of the cuticle of different individuals, fig. 20 from
a female, the others from males (obj. C, oc. 4).
Figs. 21, 33. Transverse sections of the cuticle of a male and a female
(homog. immers. i/i3, oc. 3).
Fig. 33. Female, ventral view of the posterior end (obj. A, oc. a).
r
0ljK)!Miniiai«<»A^^M'<nk' T
M
^> ^
;a::.Sci.3°5i;r.Zi].i;.V::,I,
21.
Mam-aaMEiTf. Hel.
23.
[MoNTQDMEm'! P:
22.
ins aiUTTDN « HEY. S.F.
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