The Classification of Snakes

Author(s): E. D. Cope
Source: The American Naturalist, Vol. 28, No. 334 (Oct., 1894), pp. 831-844
Published by: The University of Chicago Press for The American Society of Naturalists
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 THE

AMERICAN NATURALIST
VOL. XXVIII. October, 1894. 334

THE CLASSIFICATION OF SNAKES.

By E. D. COPE.

Owing to the absence of limbs and other points in which

diversity is usually apparent, the classification of the snakes
has always presented difficulties to the zoologist. An order
which dates from Cretaceous time and has spread over the en-
tire world, must have differentiated in structure, if its history
has been like that of other orders of Vertebrata. Yet the re-
searches of anatomists have only resulted in finding characters
which define five suborders, and about a dozen families. Of
the natural groups thus defined, one family, the Colubridoe,
embraces three-fourths of the species, and is of cosmopolitan
distribution. So long as this was the principal result attained,
it remained clear that the stronghold of the order had not yet
been taken.
The primary divisions above referred to, are defined by
peculiarities of the skeleton, and these were mostly originally
described by Johannes Mffller. In the preparation of their
Herpetologie Gennrale, Dumeril and Bibron made a full study
of the dentition. The results they obtained were important,
but they were very far from expressing an exact and clear cut
classification. The greatest defect of their definitions based on
the teeth is that they too often fail to define. One type passes
by easy gradations into another, so that in many cases it is im-
55

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832 The American Naturalist. [October,
possible to determine what type a given dentition represents.
In most cases it is clear that, among Colubrid snakes at least,
no higher groups than genera can be predicated on dentition,
and frequently not even these. Under such circumstances
further structural characters had to be sought for if we are to
have any clear idea of the affinities and phylogeny of this
curious branch of the Reptilia. In any case no systematic
arrangement can be regarded as final until the entire anatomy
is known.
In 18641 I pointed out that certain snakes, notably the water
snakes, have the vertebral hypapophyses continued to the tail,
as in the truly venomous forms. Boulenger has since found
this character in a good many forms which I had not examined,
and which have no affinity to the water snakes. This char-
acter, while important, presents the same evanescent stages in
certain types that the dental characters before noticed exhibit.
It had long appeared to me that the only prehensile organs pos-
sessed by serpents, the hemipenes, might probably present
structural variations expressive of affinity or diversity. In
18932 I examined these structures in many of the leading
types, and was gratified by the discovery of a great many
structural characters. In fact these organs exhibit a variety
of ornamentation and armature beyond any part of the
anatomy in the Ophidia, and I am satisfied that they furnish
more important indication of near affinity than any other part
of these reptiles yet examined. No one hereafter can be sure
of the place of a serpent in the system until the hemipenis has
been examined.
Still another part of the structure remained to be studied.
The assymmetry of the lungs of snakes had often been noted
by anatomists, but very little was known as to the range of
variation. Accordingly the present year,3 I undertook a study
of the pulmonary organs. I was able to confirm observations
previously made by Schlegel and Stannius, and to correct some
others, and to add a great number of facts as to species not
I Proceedings Academy of Natural Sciences, Philada.
2 American Naturlist, 1893, p. 477.

33Proceeds. Amer. Philos. Soc., 1894, p. 217.

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-13o.4 The Classification of Snakes. 833
previously examined. I cannot give here all the details ob-
served, for which I refer to the papers quoted, but I give a
general view of the results. One of these is that I am able to
confirm the conclusion of Boulenger; i. e, that the Colubriform
venomous snakes, the Proteroglypha, (cobras, Elapes, etc.), do
not differ in any fundamental respect from the non-venomous
Colubridae, and that they can not be characterized as a sub-
order. The suborders then are:
Catodonta (Type Glauconia).
Epanodonta (Type Typhlops).
Tortricina (Ilysiidae and Rhinophidse).
Colubroidea (Peropoda, Asinea, and Proterogylpha).
Solenoglypha (Typical venomous forms).
The hemipenis is a projectile organ in the form of a hollow
tube whose base is on one side of the middle line, and which
opens into the anus. When retracted it lies beneath the tail,
extending for a greater or less distance, and terminating in a
cylindrical muscle. This has considerable length, and is
finally inserted on a caudal vertebra. When the organ is pro-
jected this muscle is drawn forwards, so as to evaginate the
tubular organ. Thus the inside of the tube becomes the out-
side, and the entire organ projects freely from its base ante-
riorly. It finds its way into the corresponding oviduct of the
female (Plate XXVIII, v), and when once in place it cannot be
tracted in most species, without invagination. This is per-
formed by the contraction of the now internal retractor muscle.
This is inserted on the internal face of the apex, and draws it
inwards, so that it soon assumes the original ensheathed posi-
tion beneath the tail. It cannot be withdrawn from the
oviduct without invagination, because it is generally set
with strong bony spines which diverge backwards. They
have a perfect grip on the walls of the oviduct, and would
in some instances lacerate that organ if the two bodies
should be forcibly drawn apart. In other cases the hemi-
penis would be torn off at the base. Snakes sometimes partially
project this organ, apparently in some instances for defence,
as the spines are very pungent, and are sometimes curved
like cats claws. Such at least would seem to have been the

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834 The America'e Natiuralist. [October,
case with two Heterodonptatyrhivus, (spotted adders), which were
brought to me with the organs projected so as to present the
spines. They were caught by a cat, and were represented to
me as fighting their captor in this and other ways. Snakes
are, however, very careful not to present these organs fully
evaginated so-as to expose the delicate structures near the apex.
I have never seen this to be the case in an alcoholic specimen,
(with one possible exception), and I should judge that this was
the general experience, from the figures given by authors. It
is said that male snakes may be compelled to project the hemi-
penes by holding them before a fire, but I have not seen this.
The hemipenis of the Ophidia is traversed by a groove which
divides the superficial investment to the internal integument
(or external integument when the organ is retracted), which
commences at the base internally, and soon turns to the exter-
nal side of the organ and continues to its extremity. This is
the sulcus spermaticus (s s in Plate xxvii). This sulcus is
always bifurcated in venomous snakes, and I find it to be
equally bifurcated in many harmless snakes (Figs. 2, 3, 7).
The investing tissues may or may not correspond with this
bifurcation. Thus the hemipenis may be more or less bifur-
cate (Figs. 1, 2, 7, 9, 10, 11). Schlegel states that it is bifurcate
ih venomous snakes, but it is not so in the sea-snake Hydro-
p his hardwickii, nor in Bmmngarus setifasciatus, Hoplocephalus
coronatts, ete., while it isbifurcate in many non-venomous forms.
Next to the bifurcation of the sulcus in importance, is the
nature of the surface of the external investment (internal when
retracted). In the most perfect types both venomous and non-
venomous, this surface is reticulate like tripe, the enclosed
areas forming calyces, which may have a suctorial function
(Figs. 6, 9, 10, 11). Their borders are often papillose, and are
sometimes so deeply divided into papilla as to lose their
original character. These papille may be the seat of osseous
deposit, becoming bristles or spines, (sp), which become larger
toward the middle of the length, and lose their mutual mem-
branous connections. These isolated spines may extend to the
apex, but they rarely extend to the base. The surface may,
however, be laminate and not reticulate, and the laminme may

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1894.] The Classification of Snakes. 835
be longitudinal (Figs. 4, 7) or transverse (Figs. 1, 2, 3, 5). In
either of these cases they may not be spiniferous. The apex
or apices of the organ may be furnished with a rigid papilla
(Fig. 5) or awn.
In the Tortricina and Peropoda (the constrictors), the hemi-
penis is not spinous, and the sulcus is bifurcate (Figs. 1, 2, 3),
and in the Boidae the hemipenis is bifurcate also, although in
some genera (Xiphosoma, Ungualia), the branches are very
short. The external integument is never reticulate, but is
always laminate with elongate papillae at the extremities, in
Epicrates (Fig. 2), Xiphosoma, and Ungualia. The laminate are
pinnate from the sulcus as an axis, in Morelia, Enygrus, Lich-
anura and Eryx, and are transverse (flounced), in Charina
(Fig. 3). In Ilysia they are pinnate (Fig. 1), with a few Ion-
gitudinal plicw below.
Similar gradations in the characters of the hemipenis are to
be seen in the types of venomous snakes. Thus in the Pro-
teroglypha this organ is spinous to the tip, on a calyculate
basis, in Hydrophis, Elaps, (surinamensis); Dendraspis. It is
reticulate at the extremities and spinous below, in Callophis
(bivirgatus); Naja (Fig. 9); Acanthophis; Bungarus and Sepe-
don; the apex smooth in the two genera last named. In Elaps
rkigrocitnctus the organ is smooth below, with spines at the
apex.
In Solenoglypha the genus Atractaspis is spinous to the
apex, apparently on a longitudinally laminate basis. In the
Viperide and Crotalide the spines are on a flounced basis.
The apices are calyculate in Bitis, Clotho (Fig. 10), and Vip-
era, and spinous in Cerastes. They are calyculate in (Crota-
lidoe in Bothrops, Ancistrodon, Crotalophorus, Crotalus and
Uropsophus (Fig. 11). In Crotalus (dwrissus of the Neotropical
fauna), the papillae are not ossified; in all the other genera
they are spinous.
The condition of knowledge as to the lungs of snakes was
stated by Stannius, in 1856, as follows: " The detailed ac-
counts as to the single or double character of the lungs.
leaves much to be desired. Among Ophidia Angiostomata
there possess a single sack, Rhinophis and all Typhlopidae

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836 The American Naturalist. [October,
which have been. examined; as to the Tortricidue [Ilysiidae],
there are apparently species with two lungs (T. xenopeltis)
[a Xenopeltis unicolor], and others with a single lung (T.
scytale) [=-llysia scytale]. Among Eurystomata, all the Per-
opoda (Boa, Python, Eryx) possess. apparently two lungs.
The Calamarina that have been investigated have one lung.
Among Colubrina and Glyphodonta, there are great varia-
tions. All the Coronelle of Schlegel possess, according to
Schlegel, a single lung. I find the lung single in Rhachiodon
scaber [Dasypeltis]. Tropidonotus natrix [Natrix vulgaris] has a
very small rudiment of a second lung. Coluber [Spilotes] var-
abilis possesses, according to Schlegel, the rudiment of a second
lung. According to the statement of Meckel, this rudiment is
common in Coluber. The Xenodons have, according to
Schlegel, a single lung (X. severus and X. rhabdocephalus). In
Heterodon I find a rudimental second lung. The Lycodons,
according to Schlegel, possess a single lung; as also do Psam-
mophis and Homalopsis. In Dendrophis colubrina Schlegel
found the rudiment of the second lung. In Dipsas, according
te Schlegel, there are variations; but he states that D. multi-
maculata, D. 1xvis and D. annulata [Sibon annulatum], have but
one lung. The Achrochordina have but one lung. Among
Hydrophide I found in three species of Hydrophis the lung-
sack simple. Meckel states that Platurus has a very small
rudiment of a second lung. Among the remaining poisonous
snakes there is an insignificant rudiment of the second lung
in the Elapina and Crotalina; while the Viperina possess an
entirely simple lung."
An examination of about one hundred and fifty species of
nearly all types yielded the following results.
The snakes, with rudimental posterior limbs (Peropoda),
show in the character of their lungs, what they show in the
rudimental limbs themselves, and in the hemipenis, the near-
est relationships to the Lacertilia. They possess, with an
exception to be noted later, two well-developed lungs, one of
which is larger than the other. The smaller lung lies to the
right side and ventrally, while the larger one lies to the left
side and dorsally. In some species the dorsal and ventral

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1594.j The Classification. of Snakes. 837
relation is more pronounced than in other. In the Colubroidea
the right or ventral lung is generally present, but of very
much reduced proportions, the usual size being from two to
five millimeters in length (Plate XXVIII RL). It is connected
with the other lung by a foramen which perforates the tracheal
cartilage at a point a little beyond the apex of the heart, and
opposite to the proximal part of the dorsal lung. It is some-
times connected to the dorsal lung by a short tube, in which
cartilaginous half rings are seen in but two of the genera ex-
amined, viz., Heterodon and Conophis. The lumen of the
rudimental lung may be lined by the same reticulate structure
as is seen in the dorsal lung, or its walls may be smooth. In
some Colubroidea the rudimental lung is absent, but such
species are relatively few.
The dorsal lung may present proximally alongside of the
trachea an auricle or pocket, and this is so developed in the
genus Heterodon (Plate XXVIII), as to reach to the head, with-
out communication with the trachea, other than that furnished
by the normal portion of the lung. In the Solenoglypha,
without exception, this extension of the dorsal lung is present,
and extends to the head, and its lumen is continuous with the
trachea throughout its length. The same structure exists in
the genera Hydrus and Hydrophis; and also in the West
Indian peropodous genus Ungualia, which differs besides from
other Peropoda in having but one posttracheal lung. Finally
the tracheal lung, as I shall call it, is distinct from the true
lung in the water snakes Platurus and in Chersydrus. In the
former of these genera the trachea is not separate from the
lumen, while in Chersydrus it is distinct. It, however, com-
municates with the cells of which the lung consists in this
genus by a series of regularly placed foramina on each side.
There is no lumen in the tracheal lung of Chersydrus. In the
blind burrowing Typhlops we have a still further modifica-
tion of the tracheal lung. It is without lumen, and is com-
posed of coarse cells of different sizes. These have no com-
munication with the trachea or lung that I can discover. It
has occurred to me that this structure, which extends from the
heart to the throat, may not be a pulmonary organ.

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838 The American Naturalist. [October,
I have referred to the dorsal and ventral positions of the two
lungs. The rudimental lung is to the right of the dorsal lung
in the Colubroidea, but in the Ilysiidae it is to the left. It is quite
questionable which lung this rudiment in this family really
represents. In the Typhlopidae, the single lung is on the right
side and extends from the heart to the liver. It has the posi-
tion of the rudiment lung of the Colubroidea, and may repre-
sent it. I cannot decide this question without further material.
In Glauconia there is but one true lung, and this is ventral in
position, and originates to the right of the heart, so that in
this genus also it may represent the rudimental lung of the
Colubroidea. There is here no tracheal lung or organ.
I have no doubt of the propriety of the separation of the
Ungualiide from the other Peropoda, on. account of its pul-
monary characters. Nor is there anly doubt in my mind of
the necessity of the separation of the Leptognathinae from the
Xenodontinoe, on account of its large tracheal lung. The genus
Heterodon differs very much from other Xenodontine, in the
possession of an enormous diverticulum of the lung, but as it is
not present in the allied genus Lystrophis Cope, its wider dis-
tinction may be a questionable proceeding. The very marked
characters of the genus Chersydrus characterize the family, as
well as the osteological characters. It remains to be seen
whether the family I termed the Nothopide, but which Bou-
lenger unites with the Chersydridae, agrees with it in pulmo-
nary characters. The remarkable tracheal lung or gland dis-
tinguishes the Epanodonta from the Catodonta, emphasiz-
ing the differences observed in the osteology of the skull.
The value of the rudimental right lung as a character of the
Colubroidea is increased by my investigations. In only two
genera have I found it present or absent, viz., Halsophis and
Pityophis. I am not sure but that I may yet find it in the
P. melanoleucus, where I have failed hitherto, but I am sure
that it is present in some species of Halsophis and wanting in
others. A natural group of American Colubrinoe, appears to
be characterized by its absence, viz., bRhinochilus, Cemophora
and Ophibolus; all genera with an entire anal shield. The
development of cartilages in the bronchial foramen or tube of

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1394.] The Classification. of Snakes. 839
the rudimental lung is not a constant character. I found it
in one Heterodon platyrhinus and not in another; it is present
in Ctonophis pulcher, but absent in 0. sumichrastii.
The rudimental lung is often concealed from view and diffi-
cult to discover. The best test of its presence is the foramen
which connects it with the trachea, which will generally be
found piercing the cartilage of the latter near the apex of the
heart. The rudimental organ may then be found by inserting
a bristle, and observing its destination through the more or
less transparent tissues. In but one instance have I found a
rudimental lung without a connecting foramen, viz., in the
Mexican Ficimia olivacea. On the other hand, the foramen
may terminate in a small blind sac.
The pulmonary characters may be determined without
much dissection. The position of the heart must be first as-
certained, and a longitudinal median incision made in' the
abdominal wall. In all forms except the Epanodonta and
Catodonta, the trachea will be found passing to the left side of
the heart, and entering the lung near its apex. By splitting
the trachea, not too near its abdominal border, on turning the
free margin upwards as the snake lies on its back, the foramen
bronchiale will be seen and its lumen can be explored. The
trachea is concealed by the Esophagus, which must be drawn
to the left side of the body in order to make the examination.
The examination of the tracheal lung requires the division
of the abdominal wall farther towards the head.
The tracheal lung greatly extends the surface available for
blood aeration. This is necessary to snakes for the reason that
the huge masses of food which they ingest, so compress the
true lung that another organ is necessary. Most snakes
whether they have a tracheal lung or not, have the pulmonary
organ greatly elongated, so that while one portion is compressed
by the contents of the alimentary canal, another part is free to
function. The tracheal lung enables the snake to inflate the
anterior part of the body. This is conspicuous in the true
venomous species (Solenoglypha). In the same way Heterodon
inflates its huge diverticulum. In the marine water snakes
Cherwydrus and the Hydrophidae, these organs serve as floats.

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840 The American Naturalist. [Octobers,
In the fresh-water snakes (Natricinae) there is no tracheal
lung. The hemipenis of this group is very characteristic,
(Plate XXXVII fig. 8).
As an illustration of the modifications in classification
necessary in view of the characters which I have observed, I
give an analysis of the genera of the group which I have
called the Xenodontinae.4 These genera belong mainly to the
southern Hemisphere, and chiefly to the Neotropical Realm, a
few genera occcuring in Africa and North America. The
characters of the division are as follows.
Hemipenis with bifurcate sulcus spermaticus, and armed
with well developed spines, which are developed from the
marginal papillae of calyculi, when the latter are present.
Hypapophyses of the vertebra generally present only anter-
iorly.5
A. Lung without large proximal diverticulum...
I. Apex of hemipenis without calyces or spines but with a.
membranous disc. (Disciferi Fig. 7),
(. Rostral plate not recurved.
Hemipenis undivided, no scale-pits; Aporophis Cope..
Hemipenis divided; no scale-pits; Opheomorphus Cope.
Hemipenis divided; one scale-pit; Xenodon 6Boie.
vty. Rostral plate recurved.
Heinipenis divided; one scale-pit; Lystrophis Cope..
II. Hemipenis transversely plicated (divided); (Flabellati).
Plicae not pappillose; diacranterian; Helicops Wagl...
Plicse not pappillose; isodont; Pseudoeryx7 Fitz.
Plicxe pappillose; isodont; Rhabdosoma8 D. & B.
III. Calyculate, and not capitate (Calyculati).
5s. Hemipenis undivided.
Fusiform; isodont; Carphophiops Gerv.
Colubriform; isodont; two nasals; Diadophis B. & G.
Colubriform; diacranterian; one nasal; Amastridium Cope.
4American Naturalist, 1893, p. 481.
5 In Helicops they are continued to the tail.
.6 Including Liophis Wagl.
7 Dimades Gray.

8 Catostoma and Adelphicus are closely allied.

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1894.1 The Glasaifeation of Snakes. 841
Colubriformr; diacranterian; two nasals; Hypsirhynchus Gthr.
. lHemipenis double.
Fusiform; isodont; Parancia Gray.
Colubriform; diacranterian; no scale pits; Dromicus Bibr.
Colubriform; diacranterian; one scale pit; Mlonobothris9 Cope.
Colubriform; diacranterian; two scale pits; Halsophis Cope.
IV.. Capitate (or pocketed) (Capitati).
. Hemipenis undivided.
Scale pits single; scales smooth; Pliocercus Cope.
No scale pits; scales smooth; Rhadinmea Cope.
Scales keeled; prenasals in contact; Tretanorhinus D. & B.
of. Hemipenis divided.
Rostral normal; isodont; Minia B. & G.
V. Pappillose at apex.. (African) (Papillati).
Hemnipenis single; Grayia Gthr.
Hemipenis bifurcate; Tieleusl0 Cope.
VI. Calyculate with spinous bands to apex. (Calycispinosi).
Subisodont; attenuate; Uromacer D. & B.
VII. Exclusively spinous to apex; (diacranterian). (Spinosi).
Anterior teeth wanting; Enulius Cope.
Anterior teeth present; anal divided; no scale pits;
Echinantherall Cope..
Anterior teeth present; anal entire; one scale pit;
Acanthophallus Cope.
A A Left lung with a proximal diverticulum, extending to
the throat.
VIII. Calyculate and capitate.
Rostral recurved; hemipenis divided; diacranterian;
Heterodon Beauv.
Any one familiar with these genera will perceive that they
are not represented in a linear series in the table. I-e will
also observe that genera of probably iiot very close affinities

9 Gen. nov. Type, Drormictas chaamissonis Auct.

10Amer. Naturalist, 1893, p. 482.
11 Gen. nov.; type Aporophis cyanoplernts Cope. This species is thought by
Boulenger to be Natrix melanostigrma Wagl; bat that. species is represented
as unicolor above. The present species has three loingittudiiial bands, one median
and one on each side.

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842 The American Naturalitd. [October,.
are placed close together, as for instance Tretanorhinus and
Helicops12 and their associates. This is, however, a necessity
of an artificial key and is not new in zoology. The charac-
ters presented by the hemipenis are more readily determinable,
and are more constant that those to be found in any other
part of the structure.
InA further illustration of -the same subject I present a
synopsis of another tropical group, this time entirely Amer-
ican, which only differs from the Xenodontinac in the grooving
of the posterior maxillary tootb, i. e., the Scytalinme.
I. Apex without calyces or spines, but with a membranous
disc. (Disciferi).
Hemipenis divided; Erythrolamprus Boie.
II. Hemipenis transversely or obliquely plicate; (divided).
(Flabellati).
No calyces; rostral plate normal; Jaltris Cope.
Calyces at apex; rostral plate produced; Conophis Peters.
III. Calyculate and not capitate. (Calyculati).
v. Hemipenis divided.
Rostral recurved; Rhinostoma Wagl.
Rostral normal; pupil erect; Oxyrrhopus Wagl.
Rostral normal; pupil round; Philodryas Wagl.
99. Hemipenis undivided.
Rostral normal; 'Thamnodynastes Wagl.
IV. Capitate (also calyculate). (Capitati).
Hemipenis undivided; colubriform; Coniophanest" Hallow.
Hemipenis undivided; fusiform; Hydrocalamus Cope.
V. Spinous to apex; (divided). (Spinosi).
Two nasal plates; Tachymenis Wiegm.
One nasal plate; Tomodon D. & B.
AVI. Apex smooth, or with one row of spines; (divided).
(Levi).
Urosteges one rowed; a band of minute calyces; Scytale Wagl.
Urosteges two rowed; no calyces; Lyqophis14 Tsch.
12 Helicops is certainly to be placed in this family and has no relationship to the
Natricine with which it has been hitherto associated.
13 The penial characters of Coniophanes distinguish it from Erythrolamprus, with
which I have proposed to unite it.
14 Type Lygophis elegans Tsch. =Dryophyla. poecilostornus Cope.

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 PLATE XXVII.

I,~~~~~~~~~~~~~~~~~~~~~~~~~~~,

I ci~~~~~~~~~~~~~~~~~~~~~~~~~~~~~RIA

4 ~~~~w~

4IIA-~~~~~~~~~~~I

Ilemipenes of Ophicdia.

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 PLATE XXVIII.

OcPO

Heteroclon platyrhinus Latr.

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1894.] The Classtifation of Sitkes. 843
Comparison of this table with that of the genera of Xeno-
dontinme, shows that both present identical modifications
of structure in the case of five of the subdivisions. Only
two types, (V and VI), of the Xenodontinae have not been
found in the Scytalinle; and one, (no. VI), of the latter group
has not been found in the Xenodontinae."5

EXPLANATION OF PLATES.
PLATE XXVII.
(From an unpublished Bulletin of the U. S. National
Museum). Hemipenes of distinct types of Ophidia. The
organ is split and the entire surface exposed. The student
must remember that the lateral borders are artificial, and are
continuous on the middle line behind the center of the figure
in the projected organ. When the organ is bifurcate, but one
branch is split; (figs. 1- 2-7-9-10-11).
Fig. 1. ilysia scytale L. Brazil.
Fig. 2. Epicrates arngutifer D. & B. Cuba.
Fig. 3. Charina bottle Blv. Oregon.
Fig. 4. Holarchus ancorus Gird. Philippine Ids.
Fig. 5. Oligodon sub quadratus D. & B. Java.
Fig. 6. Bascanurn constrictor L. N. America.
Fig. 7. Opheomorphus alticohts Cope. Peru.
Fig. 8. Natrix fasciata sipedon L. N. America.
Fig. 9. Naja haje L. melanoleuca Hallow. W. Africa.
Fig. 10. Clotho arietans L. S. Africa.
Fig. 11. Uropsopus confluentus Say. Texas.

LETTERING.

ss. Sulcus spermaticus; f, flounces; p, papillke; cl, calyces

or calyculi (ruches); 1, laminwa; sp, spines; spi, spindles.

PLATE XXVIII.
(From the Proceedings of the American Philosophical
Society, 1894). Viscera of Heterodon platyrhinus Beauv. The
15 Reflection has caused me to drop the major division Xenodontidse, and to refer
its two subfamilies to the Colubridae.

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844 The American Naturaliat. [Octber,
heart is turned partly over, and the esophagus is separated by
being drawn to the left of the other viscera. One oviduct is
split at the base so as to disclose the vaginal portion. In con-
sequence the rectum is displaced to the right. The lettering
is as follows.
Tr, trachea; Car, Carotid artery; Hy sheath containing
hyoid cornua; Oe, oesphagus; Vr, vertebral artery; A. P,
arteria pulmonalis; L. L, left lung; R L, right, rudimentall)
lung; H, heart; A R, left aorta root; V 0, vena cava ascend-
*ens; L, liver; St, stomach; G B, gall bladder; Sp, spleen;
F, fontanelle of oviduct; I, intestine; Ov, ovary; C A, corpus
adiposum; K, kidney; Od, oviduct; R, rectum; U, ureter;
V, vaginal portion of oviduct; Cl, cloaca.

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