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—O-1096

Notes on Snakes of the Genus Calamaria

Hymen Marx
Assistant, Division of Reptiles and Amphibians

AND

Robert F. Inger
Curator, Division of Reptiles and Amphibians

INTRODUCTION

At the generic level, the snakes of the genus Calamaria are
among the most readily recognized of the entire oriental fauna.
Yet at the species level, they present a bewildering array of super-
ficially similar forms that have been for the most part poorly defined.
These notes are offered with the hope that our analysis of the
literature and discussion of characters will define some of the species
and suggest fruitful lines of attack for a generic revision.

The species discussed below are those in the collections of Chicago
Natural History Museum (CNHM) with the addition of Bornean
forms borrowed from other institutions. The emphasis on Borneo
arises from our more general studies of the Bornean fauna. We are
indebted to the following persons for the loan of material under
their care: Mr. CM. Bogert, American Museum of Natural History
(AMNH); Mr. J. C. Battersby, British Museum (Natural History)
(BM); Mr. Arthur Loveridge, Museum of Comparative Zoology
(MCZ); Mr. M. W. F. Tweedie, Raffles Museum (RM); Miss
Margaret Storey, Natural History Museum, Stanford University
(SU); and Dr. D. M. Cochran, United States National Museum
(USNM). We are grateful to Mr. Harry G. Nelson, Roosevelt
University, Chicago, for identification of some food fragments, and to
Dr. Rainer Zangerl, Chicago Natural History Museum, for photo-
graphic assistance. The drawings are the work of Miss Phyllis
Wade, Chicago Natural History Museum.

167

168 FIELDIANA: ZOOLOGY, VOLUME 37

SPECIALIZATIONS OF THE GENUS CALAMARIA

According to published accounts (de Rooij, 1917; Taylor, 1922;
Mertens, 1930; Pope, 1935) the snakes of the genus Calamaria are
secretive, generally being found under logs and other debris on the
ground.

The following characters define the external morphological
divergence of Calamaria from the generalized, freely ranging colubrid
stock: (1) no internasals; (2) prefrontals broadly in contact with
supralabials; (3) reduction in size of nasal; (4) loreal absent (except
in C. tropica Taylor); (5) reduction or loss of oculars; (6) reduction
in size of eye; (7) reduction in number of labials; (8) parietals in
contact with supralabials; (9) reduction in overall size; (10) reduction
of tail. These characters add up to consolidation of head shields,
proportional reduction of head and tail, and reduction of overall
size; in short, the modifications commonly associated with burrowing
snakes. Many of the same characters are found in Atractaspis,
Atractus, Typhlogeophis, and Sympholis.

Different levels of specialization are shown by the various species
of Calamaria. But all forms have in common most of the characters
of the above list. Variation in the degree of reduction of the oculars
and eye, the number of labials, the length of the snout, and the
relative proportions of head shields indicate different degrees of
specialization.

Within the genus the diameter of the eye may be slightly longer
than, equal to or shorter than the distance of the eye from the
mouth. We consider that the smaller the eye the more specialized
the form. The reduction of eye size is associated with reduction
in the preoculars. We have tabulated the characters of 80 currently
recognized species (see check list) of Calamaria. Data were taken
from the literature; the original descriptions were used for species
not included in the major faunal works (de Rooij, 1917; Taylor,
1922; Pope, 1935; Smith, 1930, 1931, 1943). The relation between
the presence of preoculars and the size of the eye in all species for
which data are available is given in Table 1. It is clear from the
table that the disappearance of the preocular is associated with the
reduction in eye size.

Another character commonly mentioned in the literature is the
ratio of the width of the frontal to the width of the supraocular.
As the supraocular becomes smaller, the frontal becomes relatively
wider. This ratio is also associated with eye size. As the eye
becomes smaller the ratio increases (Table 2). The association of

MARX AND INGER: SNAKES OF GENUS CALAMARIA 169

Table 1— ASSOCIATION OF PRESENCE OF PREOCULARS WITH SIZE
OF EYE IN SPECIES OF CALAMARIA
Observed numbers of species Total Calculated numbers of species

Preocular

Preocular

Present

Absent

Present

Absent

ye*

Greater.
Equal. .
Less
Total .

.. 13

. . 23

19

.. 55

0

4

11

15

13
27
30
70

10.2
21.2
23.6
55.0

2.8

5.8

6.4

15.0

Chi square=8.62; n = 2; P=0.01

* "Greater," "equal," and "less" refer to eye diameter relative to distance
of eye from mouth.

the reduction in eye size with the reduction of the preocular and
the increase of the frontal-supraocular ratio suggests that the ocular
region represents a morphological field that is becoming smaller
as a whole.

The great majority of species of Calamaria have five or six
supralabials, with the third and fourth entering the eye. About
one-sixth of the species have four supralabials. All but one of these
have the second and third supralabials entering the eye. Since
most colubrids have more than six upper labials, those Calamaria
with four labials are considered more specialized than those with
five. The fact that the change from five to four is apparent in front
of the orbit suggests that that is the position of the reduction.
The reduction is a result either of a shortening of the snout or of a
consolidation of the shields of the snout. Each of these processes
seems to be important to burrowing snakes.

The reduction in the number of labials is independent of eye
size (Table 3). The low value of chi square in Table 3 demonstrates
that these two characters are distributed at random, relative to one
another, and that, therefore, they represent different morphological
fields with different selective relationships.

The deviation of Calamaria from the colubrid pattern of scutel-
lation is accompanied by modifications of the skull (fig. 21). Some
of these changes parallel the consolidation of the head shields. The
discussion that follows is based on one skull each of C. leucogaster,
C. gervaisi, C. leucocephala and C. septentrionalis.

In most non-specialized colubrids (as in Coluber, Natrix, Elaphe,
and Farancia) the septomaxilla has a small postero-lateral projection
that is widely separated from both prefrontal and nasal. In Cala-
maria the septomaxilla has a lateral wall, subequal in length to the

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170

MARX AND INGER: SNAKES OF GENUS CALAMARIA 171

longitudinal axis of the nasal. Posteriorly this broad wall abuts
the prefrontal and dorsally has an extensive connection with the
lateral border of the nasal. This relationship gives the septomaxilla
the position of a strut between nasal and prefrontal and, presumably,
adds rigidity to the snout. This rigidity has been noticed by G.
Haas (1930, p. 156), who comments on the reduced movability of
the transverse suture between the nasals and the frontals in Cala-
maria linnaei.

The walls of the brain case are also modified. The parietal and
supraoccipital above and the basisphenoid and basioccipital below
lack the surface sculpturing of most colubrid skulls. Instead, the
brain case has been smoothed, broadened, and flattened.

But far more radical are the changes that have taken place in
some of the bones used in engulfing food. The typical snake has a
long squamosal (about equal to the length of the skull behind the
frontal), closely applied to the dorso-lateral portion of the parietal
anteriorly but free from and superior to the occipital region pos-
teriorly. Usually (as in Coluber, Elaphe, Natrix, and Farancia) the
squamosal projects beyond the foramen magnum. The quadrate is
as long as or longer than the squamosal and, when the mouth is
closed, slopes backward and outward from its articulation with the
posterior end of the squamosal. The length, position, and freedom
of movement of these bones are necessary to the great mobility of
the mandible.

In Calamaria the squamosal is reduced to a tiny sliver no longer
than the dorsal width of the quadrate. Instead of lying on the
dorso-lateral surface of the parietal, the squamosal lies on the side
of the skull and is closely applied for its entire length to the prootic
and may even overlie part of the exoccipital. The quadrate, closely
pressed against the side of the skull and oriented vertically, is
shorter than the depth of the brain case; its dorso-ventral axis is
subequal to its dorsal width. The posterior edge of the quadrate
lies anterior to the level of the foramen magnum. The shortening
and the changed positions of the squamosal and the quadrate in
Calamaria must reduce the amplitude of jaw movement.

The pterygoid of Calamaria is also shortened, probably in asso-
ciation with the changed position of the quadrate. Whereas the
pterygoid in colubrids usually projects far beyond the level of the
foramen magnum, in Calamaria it ends at that level or anteriorly.

The shortening of these three bones may be viewed as part of
the process of consolidation. With the confinement of the ends of

Fig. 21. Skull of Calamaria leucogaster (CNHM 71598); actual length 7 mm.

172

MARX AND INGER: SNAKES OF GENUS CALAMARIA

173

the pterygoid and squamosal to the limits of the brain case and the
pulling in of the quadrate against the skull, some of the "loose
ends" of the skull have been eliminated and the three bones have
been given added protection.

Fig. 22. Maxilla of Calamaria leucogaster (CNHM 71598) ; actual length 5 mm.
First tooth missing.

One additional specialization of Calamaria is unique. The
maxillary teeth of all but two of the species examined are greatly
broadened at the base and compressed laterally (figs. 22-24). Near
the apex of the enlarged base is a small, oval, translucent area. The
pulp cavity (fig. 24) has two branches, one running out into the

Fig. 23. Maxilla of Calamaria septentrionalis (CNHM 7139); actual length
3.5 mm. Last two teeth missing.

shaft and the other curving into the posterior part of the base.
From the saddle between these two branches, fine pulp canals run
into the oval, translucent "window." The small amount of pulp in
this region is apparently responsible for its transparency.

174 FIELDIANA: ZOOLOGY, VOLUME 37

Two levels of maxillary tooth specialization are found in the
species we have examined. The less modified condition occurs in
schmidti, which has seven, widely spaced, unmodified ophidian teeth,
and in leucogaster, in which only the last two of its seven maxillary
teeth are slightly enlarged at the base (fig. 22). The more specialized
condition is characteristic of all the others. In these last, all of the
maxillary teeth are modified as illustrated. There is no space
between successive teeth, and all are enlarged at the base.

The other dentigerous bones have typical ophidian teeth (fig. 25).

Fig. 24. Maxillary tooth of Calamaria gervaisi (CNHM 15025), greatly en-
larged; cleared and stained with eosin.

The function of the specialized teeth is obscure. The diet of
Calamaria consists largely, if not exclusively, of earthworms. Four-
teen individuals, belonging to eight of the eighteen species studied,
contained earthworms. There were no marks on the food remnants
to indicate how the teeth were used. Other earthworm feeders
among oriental colubrids — Natrix nuchalis, Achalinus spinalis,
Opheodrys major, and Trirhinopholis styani (Pope, 1935) — have the
simple, conical, rather widely spaced teeth characteristic of snakes —
an indication that the unusual teeth of Calamaria may not be an
adaptation to food itself.

Instead, G. Haas' description (1930, p. 157) of the skull move-
ments of C. linnaei causes us to speculate that the modified teeth
may constitute an adaptation to facilitate food ingestion. Though
not commenting on the form of the maxillary teeth, Haas concludes
that the maxilla is capable only of very limited movement in a
longitudinal direction. Rather, during protraction of the pterygo-

MARX AND INGER: SNAKES OF GENUS CALAMARIA 175

palatal arch, the maxilla is swung laterally so that the tooth row,
which at rest is oriented obliquely inward, is rotated on its longi-
tudinal axis. Haas suspects that this swinging action of the maxilla
functions to push the prey, whose soft body tends to be squeezed

Fig. 25. Mandible of Calamaria septentrionalis (CNHM 7139) ; actual length
8.5 mm.

out laterally by the pressure of jaw and palate, in toward the center
of the mouth so that it can pass the gape.

Assuming Haas to be correct, it seems clear that the broadening of
the base of each maxillary tooth would increase the effectiveness
of the maxilla by increasing the surface it could bring to bear on
the prey. Furthermore, since Haas states that the turning of the
maxilla is stronger posteriorly, we are presented with a functional
explanation for the observation that apparently the posterior
maxillary teeth were the first to become broadened (as in leuco-
gaster).

EVALUATION OF CHARACTERS

Since 80 species of Calamaria are now recognized, part of the
difficulty of identifying these snakes lies in the number of species
to be compared. Unfortunately, some authors give the impression
of extreme haste in describing new forms; it almost seems that the
mass of forms to be compared discouraged any comparison. But
most of the trouble arises from inadequate definition of the species
in the literature. In general, the literature fails to evaluate the
characters. The significance of a range of fifty ventrals, for example,
is not examined. Indeed, many authors have neglected to sex their
snakes, thus rendering their counts even less useful.

The characters most often used to distinguish the species are:
(1) the number of supralabials; (2) the contact or separation of the
mental and first chin shields; (3) the ratio of frontal width to supra-

176 FIELDIANA: ZOOLOGY, VOLUME 37

ocular width; (4) the diameter of the eye relative to its distance from
the mouth; (5) the presence or absence of a preocular; and (6) the
number of ventrals (see Boulenger, 1894, and de Rooij, 1917).

The number of supralabials varies from four to six. The dif-
ference between five and six is merely a difference in the method of
counting and depends upon whether the small scale following the
large fifth labial is included in the count; in either case the same
labials enter the eye and the same labial (fifth) is the largest. No
significance can be attached to the difference between five and six
labials and no forms are so distinguished. The distinction between
four and five labials, however, is important. All forms with five
(or six) labials have the third and fourth scutes entering the orbit,
whereas all those with four labials have the second and third enter-
ing the orbit, with the exception of lowi in which only the third
enters it.

We have examined two or more specimens of each of fifteen
species; the two largest samples were 47 (vermiformis) and 20 (ger-
vaisi). No intra-specific variation in supralabial counts was ob-
served, nor is there any mention of such variation in the literature.
A form invariably has either four or five (or five or six) supralabials.

Almost as constant is the contact or separation of the mental
and anterior chin shields. We have seen only one instance of intra-
specific variation. Normally the first infralabials meet behind the
mental in C. leucogaster, but one of the four examined has the mental
narrowly in contact with the chin shields. Thus, this character on
the whole is reliable in distinguishing species, but caution must be
used when only a single specimen is available.

The ratio of frontal width to supraocular is subject both to in-
dividual variation and to errors in measurement. The ratio in the
large series of vermiformis varied from \Yi to 23^; gervaisi varied
from 13^ to 2; four leucocephala showed a range of 4 to 5. Clearly,
a difference of 1 in this ratio can easily be accounted for by indi-
vidual variation, but a difference of 2 or more, as in the comparison
of gervaisi and leucocephala, is a matter of species differentiation.
Keys (for example, Boulenger, 1894; de Rooij, 1917) that separate
groups of species on the basis of the frontal being twice or less than
twice as wide as the supraocular are not very helpful except in the
extremes.

Much the same can be said for eye size. There are species with
extremely small eyes and those with moderately large eyes. Some
intra-specific variation is evident but it never includes the extremes

MARX AND INGER: SNAKES OF GENUS CALAMARIA 177

of the genus. In general, only when species occupy different ends
of this scale can they be distinguished on the basis of this character.
The presence or absence of a preocular is relatively constant.
The notable exception in our experience is the series of leucocephala.
One of the four seen has no preocular on either side of the head; a
second has a preocular on one side only; the other two have a pre-
ocular on both sides. But when present the preocular is very small
and the prefrontal has a broad contact with the eye. Apparently
the preocular is in the process of disappearing in leucocephala. In
species with moderately large preoculars, the absence of this scale
is probably a rare anomaly.

The range of ventral counts reported in the literature is presented
uncritically for the most part. Even Boulenger (1894), one of the
few authors to sex his specimens, failed to analyze the counts; for
example, he gave 153-210 as the range of ventral counts for vermi-
formis but did not comment on this range or determine it for a single
series. Analysis of our sample of vermiformis (snakes from the Malay
Peninsula, Sumatra, Borneo, and the Philippine Islands) provides
an estimate of what is probably near the maximum intra-specific
variation for the genus. The total range of ventrals for our 22 males
is 53 (144-196) and for the 25 females 81 (137-217). Even if only
Bornean snakes are considered, the range for 12 females is 64
(137-200). It is only when analysis is limited to a single locality
that the variation is not unusually great; for example, our Bornean
sample includes five males and five females from Mount Kina Balu;
their ventral counts are, respectively, 154-165 and 170-180, ranges
that are very narrow in contrast to the total range for the species.
Similarly, although the four Sumatran males examined have a ventral
range of 31 (166-196), three from a single locality vary only from
191 to 196; seven Sumatran females from scattered areas have a
ventral range of 39 (179-217), whereas four from one locality have
a range of only 9 (209-217).

This pattern of wide intra-specific variation coupled with narrow
local variation is also found in other species. In gervaisi, the total
ventral range of 22 females from Luzon, Negros, and Mindanao is
39 (142-180); but nine from Montalban, Luzon, show a range of
only 9 (158-166). Five males of borneensis vary from 152 to 169,
yet four from one locality have a range of only 11 (159-169).

Given such wide intra-specific variation, ventral counts are of
little use in distinguishing forms. It is only when allopatric popu-
lations differ by more than 50 ventrals that any significance can be

178

FIELDIANA: ZOOLOGY, VOLUME 37

attached to the difference. On the other hand, in dealing with
closely related sympatric species, a difference of 20 ventrals can be
accepted as an inter-specific difference if both samples come from
the same restricted area and if the sexes are handled separately.

Although the absolute variation in subcaudal counts must be
smaller because of the numbers involved, relatively they show the
same range as the ventrals. The range of 81 ventrals in vermiformis
equals 37 per cent of the maximum count (217). The subcaudals
of our 25 female vermiformis vary from 14 to 20; this range of 7 is
equal to 35 per cent of the highest count. Yet despite this wide
intra-specific variation, there is much inter-specific differentiation
in caudal counts (see Table 4). However, because of a very sharp
sex dimorphism, subcaudal counts are worthless unless the sex is
determined.

Table 4.— POSITION OF REDUCTION TO FOUR DORSAL SCALE
ROWS IN CERTAIN SPECIES OF CALAMARIA

Number of specimens given in parentheses

Males Females

Total Total

Species Position* subcaudals Position subcaudals

leucogaster 4-8 (4) 16-18 (4)

gervaisi 6-11 (3) 16-19 (6) 2-10 (16) 11-16 (22)

griswoldi 9-12 (4) 16-17 (4) 9-10 (3) 14-16 (3)

lumbricoidea... 4-5(2) 19-22(2) 5(1) 17(1)

vermiformis. . . 2-9 (22) 19-27 (22) 0-7 (25) 14-20 (25)

grabowskyi. . . . 16-19 (2) 26-27 (2) 14-18 (2) 22-23 (2)

suluensis 9(2) 23-25(2)

pendleburyi . . . . 9(1) 18(1) 12(1) 17(1)

lautensis 8 (1) 19 (1) 6 (1) 12 (1)

leucocephala . . . 12-15(2) 34-37(2) 8-12(2) 26-29(2)

septentrionalis. 0 (1) 18 (1) 0 (2) 10-11 (2)

borneensis 6-9 (4) 22-24 (5) 4 (3) 16-19 (3)

schmidti 6-7 (2) 14 (2)

loivi 4-6 (2) 19-20 (2) 3-5 (2) 14 (2)

*Position located in terms of subcaudals, counting forward from terminal
scute.

Another character, not discussed in the literature, has been
useful in distinguishing certain species. The number of dorsal scale
rows on the tail is usually reduced to four shortly before the terminal
scute. The point at which the reduction to four scale rows occurs,
as located by the number of subcaudals counted forward from the
terminal scute, is relatively constant within species (Table 4). The
position of this reduction seems to depend more on the shape of the

MARX AND INGER: SNAKES OF GENUS CALAMARIA 179

tail than on its length, as is shown in Table 4 by a comparison of
grabowskyi with vermiformis and of lautensis with septentrionalis.
The data of Table 4 demonstrate that it would be difficult to identify
a single specimen on the basis of this character alone, but it is useful
in substantiating conclusions arrived at on the basis of other charac-
ters. We refer to the comparisons of griswoldi with vermiformis
grayi (see p. 184) and of grabowskyi with suluensis (see p. 191).

TAXONOMIC SECTION

Calamaria leucogaster Bleeker. Figures 21, 22.

Calamaria leucogaster Bleeker, 1860, Nat. Tijds. Ned. Indie, 21 : 293 — Ampat-

Lawang, Sumatra.
Calamaria brookii Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 15: 329.
Calamaria brookei de Rooij, 1917, Rept. Indo-Austr. Arch., 2: 166.

Descriptive notes. — Four females; total length 178-208 mm.; tail
0.06-0.08 of total; ventrals 146-155; subcaudals 16-18; supralabials
5, third and fourth entering eye; infralabials 5, first three pairs in
contact with anterior chin shields; oculars 1+1; position of reduction
to 4 dorsal scale rows, counting subcaudals forward from terminal
scute 4-8; maxilla with 7 conical teeth.

Discussion. — One (CNHM 67279) differs from the others in
having a narrower snout and in having the mental in contact with
the chin shields instead of being separated by the first infralabials.
The latter distinction has been thought significant in distinguishing
species of Calamaria. This is the only example of individual varia-
tion in this character that we have found. The first infralabials of
CNHM 67279 are only narrowly separated. Otherwise all four
snakes are so similar that we must place them in the same species.
Especially significant is the agreement in maxillary dentition (see
discussion, p. 174).

The differences between leucogaster and brooki as determined
from Boulenger's descriptions (1894, 1895), which were based upon
examination of the types, are: (1) frontal of leucogaster twice as
wide as supraocular, three times as wide as supraocular in brooki;
(2) snout of leucogaster pointed, no description for brooki; (3) sub-
caudals of leucogaster 14-17, of brooki 23.

As noted above, one of our snakes (CNHM 67279) has a more
pointed snout than the others. In this character, therefore, it
resembles leucogaster. But the frontal of this specimen relative to
the supraocular, said to be wider in brooki, is wider than the others.

180 FIELDIANA: ZOOLOGY, VOLUME 37

Accordingly, our material shows a recombination of these supposedly-
diagnostic characters. The differences in subcaudals can be at-
tributed to sex dimorphism; the type of brooki is a male, whereas
Boulenger (1894) gives no counts for male leucogaster.

In Boulenger's descriptions, these two forms also differ in
coloration. The difference lies in the apparent absence of longitu-
dinal dark lines in leucogaster. But de Rooij (1917), who also
examined the type of leucogaster, notes that eight dark longitudinal
lines are present. Boulenger mentions only five in brooki plus a
dark upper edge on the outer scale rows. If that dark edge be
counted as an additional line, the total in brooki is raised to seven.
The median band in our material is split into two by a light mid-
dorsal line. Otherwise the agreement between our snakes and Bou-
lenger's description of brooki is so close that we assume we are dealing
with the same form, and, therefore, that the differences in the mid-
dorsal stripe can be explained as individual variation.

Since none of the distinctions between leucogaster and brooki
stand up under analysis we must place the latter in synonymy.
This conclusion is supported by the collection of both nominate
forms from Matang, Sarawak (de Rooij, 1917, pp. 164, 166).

Localities and specimens examined. — North Borneo: Jesselton
District: Mount Kina Balu, Kiau, 3,000 feet (RM 320).

Sarawak: First Division: Santubong, Sungai Ja'ong (CNHM
71598-99). Fifth Division: Lawas (CNHM 67279).

Calamaria gervaisi DumeYil and Bibron. Figure 24.

Calamaria virgulata (non Boie) Eydoux and Gervais, 1837, in Guer. Mag.
Zool., 1837, CI. 3, pi. 16, figs. 7-10.

Calamaria gervaisi Dumeril and Bibron, 1854, Erp. Gen., 7: 76 — Java [in
error, see remarks, p. 182].

Descriptive notes. — Twenty specimens; supralabials 5, third and
fourth entering eye; infralabials 5 or 6, first pair not in contact
behind mental, first 3 pairs (18), 4 pairs (1), or % pairs (1) in contact
with anterior chin shields; oculars 1 + 1; maxilla with 8 or 9 modified
teeth; for ventral and subcaudal counts see Table 5.

Luzon: Two males; total length 190-207 mm.; tail 0.07-0.08
of total. Nine females; total length 140-269 mm.; tail 0.04-0.05 of
total. Position of reduction to 4 dorsal scale rows, counting sub-
caudals forward from terminal scute 2-7 (mean 5.1).

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182 FIELDIANA: ZOOLOGY, VOLUME 37

Negros: Two females; total length 244-296 mm.; tail 0.05-0.06
of total. Position of reduction to 4 dorsal scale rows, counting
subcaudals forward from terminal scute 5-6.

Mindanao: One male; total length 179 mm.; tail 0.09 of total.
Five females; total length 158-258 mm.; tail 0.05-0.07 of total.
One unsexed. Position of reduction to 4 dorsal scale rows, counting
subcaudals forward from terminal scute 7-11 (mean 9.1).

Discussion. — On the basis of a greater number of ventrals,
Taylor (1917) distinguished the Negros population as the subspecies
g. iridescens. Our material bears out the difference in this character.
The Mindanao population apparently differs from both the Luzon
and the Negros populations in the same character (see Table 5).
Furthermore, the Mindanao animals have a yellow collar, which is
not found in any of our Luzon or Negros snakes and is not mentioned
by either Boulenger (1894, p. 338) or Taylor (1922, p. 186). The
Mindanao series also differs from the others in the position of the
final scale row reduction (see above) .

If the Negros population is recognized as a distinct subspecies,
probably the Mindanao population should be also. But a decision
on the definition of subspecies must wait for additional material
from scattered localities.

The type locality of gervaisi is obscure. Dumeril and Bibron
(1854) state that Eydoux and Gervais collected their snake, the
type of gervaisi, in Java. Eydoux and Gervais (1837) give no
locality, but the other reptiles that they specifically state having
collected are all from Manila. The figure of Jan and Sordelli (1865)
of the type shows a Calamaria without a light collar and with a
uniform venter. This combination is characteristic of the Luzon
population, so the type probably came from Luzon. All subsequent
authors (Gunther, 1858; Boulenger, 1894; Taylor, 1922) restrict
gervaisi to the Philippines. De Haas (1950) does not include it in
the fauna of Java.

Earthworms were found in the guts of three snakes from Luzon
and two from Mindanao.

Localities and specimens examined. — Philippine Islands: Luzon:
Manila (CNHM 15025-31, 15033-35); Montalban (CNHM 15036).
Mindanao: Mount McKinley, 3,000 feet (CNHM 53374-76) ; Mount
Apo, 2,800 feet (CNHM 53377); Tagum, sea level (CNHM 53378-
80). Negros: Dumaguete (CNHM 57308); Bais, Mabaha (CNHM
61625).

MARX AND INGER: SNAKES OF GENUS CALAMARIA 183

Calamaria lumbricoidea Boie

Calamaria lumbricoidea Boie, 1827, Isis, 20: 540 — Java.

Descriptive notes. — Two males; total length 428-468 mm.; tail
0.06-0.07 of total; ventrals 186-190; subcaudals 19-22. One female;
total length 505 mm.; tail 0.05 of total; ventrals 192; subcaudals 17.

Supralabials 5, third and fourth in contact with eye; infralabials
5, first three pairs in contact with anterior chin shields, first pair
not in contact behind mental; oculars 1 + 1; position of reduction to
4 dorsal scale rows, counting subcaudals forward from terminal
scute 4-5; maxilla with 10-11 modified teeth.

In coloration these agree with Boulenger's description (1894).

Reasons for separating griswoldi from lumbricoidea are given
below.

Localities and specimens examined. — Java : Benkeng, near Serang
(AMNH 71513). No other data (MCZ 25894).

Sumatra: No other data (USNM 56439).

Calamaria griswoldi Loveridge

Calamaria lumbricoidea griswoldi Loveridge, 1938, Proc. Biol. Soc. Washington,
51: 43 — Luidan River, Bundu Tuhan, Mount Kina Balu, North Borneo.

Descriptive notes. — Four males; total length 236-425 mm.; tail
0.06 (4) of total; ventrals 166-179; subcaudals 16-17. Three females;
total length 375-467 mm.; tail 0.05-0.06 of total; ventrals 183-190;
subcaudals 14-16.

Supralabials 5, third and fourth entering eye; infralabials 5, first
pair not in contact behind mental, first three pairs in contact with
anterior chin shields; oculars 1+1; position of reduction to 4 dorsal
scale rows, counting subcaudals forward from terminal scute 9-12;
maxilla with 10 modified teeth.

Discussion. — These animals, two of which are topotypes, agree
in all significant details with Loveridge's description.

There is little difficulty in separating griswoldi from vermiformis,
lumbricoidea, or grayi. Superficially the dorsal coloration of gris-
woldi is much like that of grayi and the similarly striped "high
altitude" form of vermiformis, but in grayi and vermiformis the light
area covers only the edges of the mid-dorsal scales, forming very
narrow light lines, whereas in griswoldi the lateral fourths of each
scale are light, forming mid-dorsal stripes equal to half the width of
the dark interspaces. The immaculate yellow or white venter of
griswoldi is radically different from the boldly marked one of grayi

184 FIELDIANA: ZOOLOGY, VOLUME 37

and vermiformis. The ventral coloration of griswoldi is like that of
lumbricoidea, but the latter is uniformly dark mid-dorsally, with a
yellow stripe running the length of the outer one or two scale rows.

The eye of griswoldi is distinctly larger than the eyes of the
other forms, its diameter being at least equal to its distance from the
mouth but usually larger. The eye diameter is usually smaller than
its distance from the mouth in the other forms, but may be equal to
that distance.

The reduction to 4 dorsal scale rows occurs opposite the ninth
to twelfth subcaudal from the tip of the tail in griswoldi (mean of
7 snakes=9.7). In vermiformis the range of this point is 0-9 sub-
caudals from the tip (only one with more than 7; mean of 39=4.3),
in grayi 3-6 (mean of 8=5.0), and in lumbricoidea 4-5 (mean of
3=4.7).

Conceivably, if the total number of subcaudals were decreased,
the distance of the final reduction from the tip of the tail in terms
of subcaudals should become less and less, all other things being
equal. But the difference between griswoldi and the others in this
distance is independent of total subcaudal count as griswoldi has
fewer subcaudals.

Subcaudal counts of griswoldi are lower than those of the other
three forms. Observed ranges for males are: griswoldi 16-17 (four);
vermiformis 19-26 (seventeen), only one with less than 22; lumbri-
coidea 19-22 (two); grayi 22-27 (four). Observed ranges for females
are: griswoldi 13-16 (four, including the type); vermiformis 14-20
(twenty-one), only four with less than 17; lumbricoidea 17 (one);
grayi 15-18 (four), only one under 17.

Ten vermiformis from the type locality of griswoldi (Mount Kina
Balu) differ from the latter in ventral counts. The five males of
this series of vermiformis have ventral counts of 154-165 and the
females counts of 170-180. Two male griswoldi have 170 and 179
ventrals and five females (including the type series) a range of
183-192. This difference between the two species does not hold up
in other parts of the geographic range of vermiformis, which has a
total spread of 144-196 in the males and 137-217 in the females.

Though originally described as a subspecies of lumbricoidea,
griswoldi differs from the latter sufficiently in eye size, position of
caudal scale reduction, scale counts, and coloration to warrant
recognition as a distinct species.

The male from Lumu Lumu, Mount Kina Balu, identified by
Smith (1931) as vermiformis, is griswoldi, as is clear from its colora-
tion and counts.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 185

One snake contained an entire earthworm (78 mm. total length)
of the subfamily Megascolecinae. The gut of the earthworm was
full of wood fragments but held no mineral particles. Apparently,
the earthworm was feeding in a decaying log and was caught there
by the snake.

Localities and specimens examined. — North Borneo: Jesselton
District: Mount Kina Balu, Bundu Tuhan, 4,500 feet (USNM
130238-39); Mount Kina Balu, Lumu Lumu, 5,000 feet (BM
1929.12.22.112); Mount Kina Balu, Tenompok, 4,700 feet (RM
18949, RM no number); Mount Kina Balu (BM 95.11.7-25);
Ranau (USNM 134114).

Calamaria vermiformis Dumeril and Bibron

Discussion. — We place bruegeli in the synonymy of vermiformis
because the differences noted by Mertens are within the range of
individual variation. According to Mertens, the frontal of bruegeli
is twice as wide as the supraocular; Boulenger (1894) states that
the frontal of vermiformis is not twice as broad as the supraocular.
Our own specimens span this minor difference. The other distinction
noted by Mertens is the relatively broader rostral of bruegeli. Due
to the differences of methods of measuring and individual variation
in our specimens this distinction does not warrant recognition of a
new species. Furthermore, the counts and other characters of
bruegeli agree with the original description of vermiformis and with
our material.

The Philippine form, grayi, is obviously closely related to vermi-
formis in counts, habitus, and scale proportions. The ventral
coloration of grayi — barred with black — is just like that of Bornean
vermiformis. Most striking, however, is the agreement in juvenile
pattern. Two young grayi, measuring 149 mm. (MCZ 25781) and
216 mm. (MCZ 25778), have the yellow heads and yellow rings
typical of juvenile vermiformis, but the lined pattern of adult grayi
is clearly visible in the dark interspaces.

Minor differences do distinguish the two populations. Most
outstanding, of course, is the dorsal coloration, consisting in adult
grayi of narrow light lines on a dark background and in adult Bornean
vermiformis of a uniform, brown color. In other populations of
vermiformis, for example, from the Malay Peninsula and Sumatra,
occasional individuals have the "grayi" pattern.

The Philippine form seems to have higher ventral counts than
the Bornean and Malayan populations but agrees with Sumatran
vermiformis (see data below).

186 FIELDIANA: ZOOLOGY, VOLUME 37

Excluding the last labial, each parietal of grayi is usually bordered
postero-laterally by 4 scales; in only one of the eight seen was each
parietal bordered by 3 scales. On the other hand, only one out of
23 Bornean vermiformis and one out of 11 Sumatran vermiformis
have 4 scales in contact with each parietal; all the others have 3
scales bordering each parietal.

In view of the great similarity between grayi and vermiformis,
we must consider them conspecific. However, the differences cited
above between grayi and Bornean vermiformis justify the separation
of these as distinct subspecies.

The remaining samples of vermiformis we have seen show a
complex pattern of geographic variation. Some characters, such
as ventral counts, apparently have intense local differentiation.
The variation in ventral counts has been discussed in detail above
(p. 177). To summarize, twelve females from Borneo range from
137-200, but within a restricted area five females have a ventral
range of 11 (170-180). Similarly, five males from the same locality
have counts of 154-165, whereas twelve males from scattered parts
of Borneo have a range of 38 (144-181). Our Sumatran sample
has the same pattern of variation.

Variation in coloration is independent of that in ventral counts
and does not show the same amount of local differentiation. All
fourteen Bornean adults (over 300 mm.) are medium to light brown
above with the outer row or two of scales slightly lighter. Two out
of four snakes from the Malay Peninsula and nine out of ten from
Sumatra are dark brown or almost black above and have a sharply
defined yellow or cream stripe on the two outer scale rows. The ani-
mals of the last two regions therefore, are distinctly bicolored,
whereas those from all parts of Borneo have a uniform appearance.

The lined coloration of grayi forms the third major pattern of
vermiformis and it is characteristic of all samples from the southern
Philippines. This lined pattern also appears in the highlands of
the Malay Peninsula (Tweedie, 1950) and is the coloration of two
of the Malayan and one of the Sumatran snakes seen. We do not
have sufficient material to be sure that whole local populations of
the last two regions are characterized by this pattern.

Ventral coloration also varies geographically. The Bornean
snakes have yellow under sides crossed by black bars occupying the
complete widths of 2 to 4 ventrals; occasionally, a few of the cross-
bars are broken up into alternating black squares. The same ventral
pattern is found in two of the Malayan and three of the Sumatran

MARX AND INGER: SNAKES OF GENUS CALAMARIA 187

snakes. The two striped Malayan specimens have crossbars that
cover 4 to 6 ventrals. The other seven Sumatran animals have the
posterior four-fifths or more of the belly solid black with only a few
narrow crossbars anteriorly.

The striped snakes from the Malay Peninsula and Sumatra
have lower ventral counts (two males 154-157; one female 179)
than observed in grayi (males 165-176; females 192-207). As noted
above, grayi differs from the other striped vermiformis in the number
of scales in contact with the parietals. The similarity of grayi
to these striped individuals is probably the result of parallel mutation
in similar genetic systems.

On the basis of the data now available we recognize the following
forms of vermiformis:

Calamaria vermiformis vermiformis Dumeril and Bibron

Calamaria vermiformis Dumeril and Bibron, 1854, Erp. Gen., 7: 85 — Java.
Calamaria bruegeli Mertens, 1924, Zool. Anz., 60: 158 — central Borneo.

Descriptive notes. — Borneo: twelve males; total length 160-436
mm.; tail 0.08-0.10 of total; ventrals 144-181 (mean 161.0±3.1;
N= 12) ; subcaudals 19-26 (mean 23.2±0.5; N= 12) . Twelve females;
total length 120-642 mm.; tail 0.06-0.08 of total; ventrals 137-200
(mean 172.4±4.5; N=12); subcaudals 14-20 (mean 17.6±0.6;
N=12).

Sumatra: four males; total length 154-449 mm.; tail 0.07-0.08
of total; ventrals 166-196 (mean 186.8); subcaudals 22-23 (mean
22.8). Seven females; total length 186-503 mm.; tail 0.05-0.06 of
total; ventrals 179-217 (mean 199.1±6.1; N=7); subcaudals 16-19
(mean 17.3±0.4; N=7).

Malay Peninsula: two males; total length 168-410 mm.; tail
0.10 (two) of total; ventrals 154-157; subcaudals 22-24. Two
females; total length 253-394 mm.; tail 0.06 (two) of total; ventrals
179-183; subcaudals 17 (two).

Supralabials 5 or 6, third and fourth entering the eye; infralabials
5 or 6, first three pairs in contact with anterior chin shields, first
pair not in contact behind mental; oculars 1+1; frontal V/2 to 23^
times width of supraocular; excluding fifth labial each parietal
bordered postero-laterally by 3 (37) or 4 (two) scales; position of
reduction to 4 dorsal scale rows, counting subcaudals from terminal
scute 0-9 (mean 4.3±0.3; N=39); maxilla with 10 modified teeth.

Color (in alcohol) varying with size; small specimens (up to 250
mm.) with yellow heads and narrow yellow rings around the dark

188 FIELDIANA: ZOOLOGY, VOLUME 37

brown body; belly yellow barred with dark brown; large specimens
(over 340 mm.) various (see above, p. 186), but usually brown
above, belly with black crossbars.

Discussion.— A transition between these two color patterns is
suggested by four specimens (234-375 mm.). The smallest of these
has the yellow body rings narrowed and interrupted and the head
somewhat darkened above. The largest of the four has a lighter
head than usual for adults. Gunther (1865) suggested that the
yellow rings disappear in adults. Smith (1931) also comments on
the difference of juvenile and adult coloration.

Earthworms were found in two specimens.

Localities and specimens examined. — North Borneo: Jesselton
District: Mount Kina Balu, Bundu Tuhan, 4,500 feet (USNM
130252); Mount Kina Balu, Kiau, 3,000 feet (BM 1929.12.22.110-
111; MCZ 43564-65; RM 168-17); Mount Kina Balu, Kenokok,
3,300 feet (MCZ 43566-69). Sandakan District: Sandakan (CNHM
15003).

Sarawak: First Division: Matang (BM 1873.4.23). Second
Division: Saribas (SU 8575). Fourth Division: Baram District
(BM 97.3.4-5, 1902.11.25.16-17); Mount Batu Song, 1,000 feet
(BM 92.10.7.8). Fifth Division: Lawas (CNHM 67278); Pa
Brayong (CNHM 71646).

Borneo: Tanjong (BM 96.2.17.11-12). No additional data
(BM no reg. no.; BM 72.2.19-55; USNM 49802).

Sumatra: Kaba Wetan (USNM 70945-51); Langkat (MCZ
37769-70); Pandangsoke, Bovenlanden (MCZ 20971); unknown lo-
cality (AMNH 2879).

Malay Peninsula: Pahang: Kuala Terla, 4,500 feet (RM
4/1938); Tanah Rata, 4,600 feet (RM 1930). Selangor: Bukit
Lagong (RM 8827); Kepong Forest Reserve (RM 13880).

Calamaria vermiformis grayi Gunther

Calamaria grayi Gunther, 1858, Cat. Snakes Brit. Mus., p. 6 — Philippine
Islands.

Descriptive notes. — Four males; total length 149-358 mm.; tail
0.05-0.10 of total; ventrals 165-176 (mean 170.8); subcaudals 22-27
(mean 24.0) . Four females; total length 216-590 mm. ; tail 0.04-0.06
of total; ventrals 192-207 (mean 199.8); subcaudals 15-18 (mean
17.0).

MARX AND INGER: SNAKES OF GENUS CALAMARIA 189

Excluding fifth labial, each parietal bordered postero-laterally
by 4 scales in all but one animal; position of reduction to 4 dorsal
scale rows, counting subcaudals from terminal scute 3-6 (mean
5.0±0.4; N=8); other counts as in v. vermiformis.

Discussion. — Gunther (1858) considered grayi to be related to
lumbricoidea Boie. Reasons are given above for relating it instead
to vermiformis.

Localities and specimens examined. — Philippine Islands: Basi-
lan: near Isabela (MCZ 43649). Mindanao: Cotabato Province,
Saub (MCZ 25778-80; USNM 120811); Davao Province, Mount
Apo, 2,800 feet (CNHM 53373); Lanao Province, Lumbatan
(CNHM 15010). Leyte: Cabalian (MCZ 25781).

Calamaria grabowskyi Fischer

Calamaria grabowskyi Fischer, 1885, Arch. Naturg. Berlin, 51: 50, pi. 4, fig.
1 — Telang and Tameanglaijang, Dusson Timor District, Borneo.

Descriptive notes. — Two males; total length 320-340 mm.; tail
0.09-0.10 of total; ventrals 159-175; subcaudals 26-27. Two fe-
males; total length 278-280 mm.; tail 0.08-0.09 of total; ventrals
152-183; subcaudals 22-23.

Supralabials 5, third and fourth entering eye; infralabials 5,
first pair not in contact behind mental, first three pairs in contact
with anterior chin shields; oculars 1+1; position of reduction to 4
dorsal scale rows, counting subcaudals forward from terminal scute
14-19; maxilla with 9 modified teeth.

Color (in alcohol) above dark brown with small black spots
scattered over dorsal scales; center of scales of outer row light yellow,
lower edge of first scale row and adjacent edges of ventrals dark
brown; ventrals uniform yellowish in anterior eighth of body,
followed by ventrals with black anterior edge limited at first to a
median strip then widening to form complete dark transverse bars
across width of belly; posterior ventrals uniform yellow; under side
of tail yellow with a distinct median longitudinal dark stripe; head
above dark brown with black spots; upper lip yellowish; head below
yellowish, the anterior labials edged with black.

Discussion. — These four snakes are so similar to descriptions
(Fischer, 1885; Boulenger, 1894) of grabowskyi in coloration, in
ventral and subcaudal counts, and in the arrangement of the head
shields that we have no doubts about their identification. It should
be noted, however, that a number of named forms resemble gra-
bowskyi in these characters. Among these forms are simalurensis,

190 FIELDIANA: ZOOLOGY, VOLUME 37

elegans, prakkei, and suluensis, all of which differ from grabowskyi
in lacking dark ventral bars.

Localities and specimens examined. — Sarawak: Third Division:
Rejang River, Sarekei (CNHM 67277). Fourth Division: Bario
(RM no numbers). Fifth Division: La was District, Pa Brayong
(CNHM 71600).

Calamaria suluensis Taylor

Calamaria suluensis Taylor, 1922, Snakes Phil. Isls., p. 189 — Cagayan Sulu.

Descriptive notes. — Two females; total length 257-295 mm.; tail
0.09-0.11 of total; ventrals 148-168; subcaudals 23-25; supralabials
5, third and fourth entering eye; infralabials 5, first pair not in
contact behind mental, first three pairs in contact with anterior
chin shields; oculars 1+1; position of reduction to 4 dorsal scale
rows, counting subcaudals forward from terminal scute 9 (two);
maxilla with 7-8 modified teeth.

Discussion. — These snakes agree remarkably with the only
known specimen of suluensis, a female having 154 ventrals and 25
subcaudals. Like our specimens, suluensis is brown above with
scattered dark spots. The outer row of scales is white. A dark
brown stripe covers the lower edge of the outer scale row and the
adjacent edges of the ventrals. The head is brown with dark spots
above, the upper lip is yellow, and the lower lip and chin are yellow
with brown spots. The belly is uniformly yellow.

As indicated in the discussion of grabowskyi, suluensis is similar
to it, and to simalurensis, elegans, and prakkei. The ventral counts
of suluensis (females 148-168) are higher than those of simalurensis
(136, sex unknown), elegans (133, sex unknown), and prakkei
(126-144, sex unknown), but without additional data for sexed
animals these differences cannot be evaluated.

In addition to the striking distinction in ventral coloration,
suluensis (as we understand it) differs from grabowskyi in the fol-
lowing characters:

In grabowskyi the postocular is elongate, higher than wide, its
height equal to its distance from the mouth, and as high as the
preocular. In suluensis the postocular is squarish, its height about
one-half its distance from the mouth, and about two-thirds as high
as the preocular.

The parietals of grabowskyi are only equal in length to the
frontals, whereas in suluensis the parietals are one and one-half
times the length of the frontal.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 191

The reduction to 4 dorsal scale rows takes place in grabowskyi
opposite the fourteenth to nineteenth subcaudal and in suluensis
opposite the ninth subcaudal, in both cases counting the subcaudals
from the terminal scute.

Localities and specimens examined. — North Borneo: Sandakan
District: Sandakan (CNHM 63573). Kudat District: Bongon (BM
1906.5.23.5).

Calamaria pendleburyi Smith

Calamaria pendleburyi Smith, 1931, Bull. Raffles Mus., no. 5, p. 27 — Kiau,
Mount Kina Balu, North Borneo.

Descriptive notes. — One male; total length 241 mm.; tail 0.08 of
total; ventrals 136; subcaudals 18. Two females; total length
250-252 mm.; tail 0.07 (one) of total; ventrals 144-147; subcaudals
17 (one).

Supralabials 5, third and fourth entering eye; infralabials 5,
first pair not in contact behind mental, first three pairs in contact
with anterior chin shields; oculars 1+1; position of reduction to 4
dorsal scale rows, counting subcaudals forward from terminal scute
9-12; maxilla with 8 modified teeth.

Discussion. — Available descriptions of margaritophora, suma-
trana, and curta indicate forms similar to pendleburyi. These forms
have in common a longitudinal light stripe along the outer row of
scales, the first infralabials not in contact behind the mental, the
frontal about twice as wide as the supraocular, a moderate-sized
eye, and less than 160 ventrals.

Localities and specimens examined. — North Borneo: Jesselton
District: Mount Kina Balu, Bundu Tuhan, 4,500 feet (USNM
130237) ; Mount Kina Balu, Kiau, 3,000 feet (RM 146.17, paratype);
Mount Kina Balu, Tenompok, 4,700 feet (RM 18820).

Calamaria lautensis de Rooij

Calamaria lautensis de Rooij, 1917, Rept. Indo-Austr. Arch., 2: 163, fig. 66 —
Pulau si Laut, Cocos Island and Simalur.

Descriptive notes.— One male; total length 142 mm.; tail 0.09 of
total; ventrals 140; subcaudals 19. One female; total length 224
mm.; tail 0.06 of total; ventrals 154; subcaudals 12.

Supralabials 5, third and fourth entering eye; infralabials 5,
first three pairs in contact with anterior pair of chin shields, first
pair of infralabials in contact behind mental; oculars 1+1; position

192 FIELDIANA: ZOOLOGY, VOLUME 37

of reduction to 4 dorsal scale rows, counting subcaudals forward
from terminal scute 6-8; maxilla with 6-8 modified teeth.

Discussion.— Superficially this snake resembles many described
forms (crassa, suluensis, tropica, pendleburyi, stahlknechti, margarito-
phora, prakkei, sumatrana, elegans, simalurensis, muelleri, bicolor,
and lautensis) with low ventral and subcaudal counts, with moderate-
sized eye, and with frontal about twice the width of the supraocular.
In coloration these forms have in common an incomplete neck
band, light centers to the outer row of scales, and ventrals with
dark outer ends. Only bicolor and lautensis have the first infra-
labials in contact behind the mental. Using this character we have
tentatively limited identification to these two forms and have
eliminated bicolor on the basis of the size of the eye (smaller in
bicolor) and the relative width of the frontal (broader in bicolor) .

The gut of one specimen contained an earthworm.

Localities and specimens examined. — North Borneo: Jesselton
District: Mount Kina Balu, Bundu Tuhan, 4,500 feet (USNM
130236); Mount Kina Balu, Tenompok, 4,700 feet (RM 295).

Calamaria everetti Boulenger

Calamaria everetti Boulenger, 1893, Proc. Zool. Soc. London, 1893: 525 —
Sarawak.

Descriptive notes. — Two females; total length 168-295 mm. (latter
with incomplete tail); tail 0.11 (one) of total; ventrals 151-171;
subcaudals 20 (one); supralabials 5, third and fourth entering eye;
infralabials 5, first infralabials in contact with each other, first three
infralabials in contact with anterior chin shields; oculars 1+1;
position of reduction to 4 dorsal scale rows, counting subcaudals
forward from terminal scute 11; maxilla with 8-9 modified teeth.

These specimens agree completely with the original description.

The Bornean snake was found on the ground in a large clearing.

Localities and specimens examined. — North Borneo: Kina-
batangan District: southeast end of Dewhurst Bay (CNHM 63572).

Philippine Islands: Palawan: Brooke's Point, Kabalnecan,
1,400 feet (CNHM 53381).

Calamaria brachyura Boulenger

Calamaria brachyura Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 16: 481 —
Mount Kina Balu, North Borneo.

Descriptive notes.— One female; total length 258 mm.; tail 0.03
of total; ventrals 205; subcaudals 10; supralabials 5, third and

MARX AND INGER: SNAKES OF GENUS CALAMARIA 193

fourth entering eye; infralabials 5, first pair in contact behind mental,
first three pairs in contact with anterior chin shields; oculars 1 + 1;
position of reduction to 4 dorsal scale rows, counting subcaudals
forward from terminal scute 3; maxilla with 8 modified teeth.

In coloration agreeing with the original description except for
the absence of yellow spots on the prefrontals.

Found under a log in a cut-over forest.

Locality and specimen examined. — North Borneo: Sandakan
District: Sapagaya Forest Reserve, 50 feet (CNHM 63571).

Calamaria leucocephala DumeYil and Bibron

Calamaria leucocephala Dum6ril and Bibron, 1854, Erp. Gen., 7: 83 — type
locality unknown.

Descriptive notes. — Two males; total length 210-322 mm.; tail
0.16 (two) of total; ventrals 143-150; subcaudals 34-37. Two
females; total length 337-343 mm.; tail 0.10 (two) of total; ventrals
161-169; subcaudals 26-29.

Supralabials 5, third and fourth entering eye; infralabials 5, first
pair in contact behind mental, first three pairs in contact with
anterior chin shields; oculars 1+1 and 0+1; position of reduction
to 4 dorsal scale rows, counting subcaudals forward from tip of tail
8-15; maxilla with 10 modified teeth.

Discussion. — These specimens agree with all published descrip-
tions of this form. The preocular, when present, is very small. In
one specimen it is lacking on both sides, and in a second is present
on the left side only. Both of these snakes are females. In all
four animals the prefrontals enter the orbit.

One specimen contained the remnants of an earthworm.

Locality and specimens examined. — Locality uncertain, probably
Singapore Island (CNHM 69976-79).

Calamaria pavimentata DumeYil and Bibron

Calamaria pavimentata Dumeril and Bibron, 1854, Erp. Gen., 7: 71 — Java.

Descriptive notes. — One male; total length 244 mm.; tail 0.09 of
total; ventrals 160; subcaudals 21; supralabials 4, second and third
entering eye; infralabials 5, first pair in contact behind mental, first
three pairs in contact with anterior chin shields; oculars 1 + 1;
position of reduction to 4 dorsal scale rows, counting subcaudals
forward from terminal scute 5; maxilla with 8 modified teeth.

194 FIELDIANA: ZOOLOGY, VOLUME 37

Locality and specimen examined. — Indo-China: Annam: Bana
(CNHM 11528).

Calamaria septentrionalis Boulenger. Figures 23, 25.

Calamaria septentrionalis Boulenger, 1890, Proc. Zool. Soc. London, 1890:
34 — Kiukiang and Hong Kong.

Descriptive notes. — One male; total length 168 mm.; tail 0.07 of
total; ventrals 160; subcaudals 18. Two females; total length
277-342 mm.; tail 0.03 (two) of total; ventrals 179-180; subcaudals
10-11.

Supralabials 4, second and third entering eye; infralabials 5,
first pair in contact behind mental, first three pairs in contact with
anterior chin shields; oculars 1+1; dorsal scale rows reduce from 6
to 1 abruptly at end of tail; maxilla with 8 modified teeth.

Two specimens had eaten earthworms.

Locality and specimens examined. — China: Anhwei Province:
Ningkwo (CNHM 7139-41).

Calamaria borneensis Bleeker

Calamaria borneensis Bleeker, 1860, Nat. Tijds. Ned. Indie, 21: 296 — Sintang,
Borneo.

Descriptive notes. — Five males; total length 135-279 mm.; tail
0.09-0.11 of total; ventrals 152-169 (mean 160.8); subcaudals 22-24.
Three females; total length 184-360 mm.; tail 0.06-0.08 of total;
ventrals 173-182; subcaudals 16-19.

Supralabials 4, second and third entering eye; infralabials 4,
first pair not in contact behind mental, first two pairs in contact
with anterior chin shields; oculars 1+1; position of reduction to 4
scale rows, counting subcaudals forward from terminal scute 4-9;
maxilla with 8-9 modified teeth.

Color (in alcohol) dark grayish brown above with black crossbars
or spots mid-dorsally, lateral scale rows lighter; belly yellowish with
varying amounts of black in the form of checkers.

Discussion. — A single specimen from Kuching differs from the
above coloration in having longitudinal dark lines above and on
the lateral edges of the ventrals and in lacking the dark transverse
markings. Except for the lateral dark lines and a median longi-
tudinal dark line under the tail this snake is uniformly yellow below.

Most of this series differ from descriptions of borneensis (Bou-
lenger, 1894; de Rooij, 1917) in the presence of the dark transverse

MARX AND INGER: SNAKES OF GENUS CALAMARIA 195

markings of the back. However, the distinctive, prominent snout,
the checkered belly, and the counts support the identification. It
is clear that this form is closely related to linnaei but differs from
the latter in the shape of the snout, which in linnaei is rounded
and does not project (Boulenger, 1894) and in the subcaudal counts.
Boulenger (1894) gives the subcaudal counts of twelve female
linnaei from Java as 9-15, of which only two have more than 12.

Javan linnaei show the same variation in coloration (Boulenger,
1894) as is indicated for borneensis by a comparison of our material
with the description given by Boulenger.

Localities and specimens examined. — Sarawak: First Division:
Kuching (RM, no number). Fourth Division: Long Mujan (RM
41, 44, 48-49, 73, 75; SU 8569).

Calamaria buchi sp. nov. Figure 26.

Type. — Chicago Natural History Museum no. 71697, a female,
from Dalat, Viet Nam, Indo-China. Altitude 1,500 feet. Collected
by Father Buch in 1953.

Diagnosis. — A Calamaria with the eye about equal to its distance
from the mouth; 4 supralabials, the second and third entering the
eye; preocular present; frontal about twice as wide as supraocular;
221 ventrals.

Description. — Rostral higher than broad, visible from above;
prefrontals large, maximum length subequal to length of frontal,
posterior border angular, in contact with first and second supra-
labials and entering orbit at a point; preocular present, small,
triangular; postocular small, its height about half its distance from
mouth; supraocular broad; frontal slightly longer than broad, less
than twice width of supraocular, about two-thirds length of parietals;
4 supralabials, second and third entering eye, second larger than
first or third, fourth largest; first infralabials not in contact with
each other; mental and first three infralabials in contact with
anterior chin shields, the latter in contact for entire length; posterior
chin shields about two-thirds length of anterior ones, in contact
anteriorly only; vertebral row of nuchals smaller than adjacent
rows; maxilla with 9 modified teeth.

Scales in 13 rows; ventrals 221; anal entire; subcaudals 13; tail
ending in a blunt point; position of reduction to 4 dorsal scale rows,
counting subcaudals forward from terminal scute 3.

Color (in alcohol) purplish gray above, an obscure dark vertebral
line anteriorly; an interrupted light nuchal band; outer row of

196

FIELDIANA: ZOOLOGY, VOLUME 37

Fig. 26. Calamaria buchi sp. nov. (type, CNHM 71697); actual length of
head 7.5 mm.

scales white, edged above with black; second row of scales white in
lower half, in posterior third of body becoming dark; ventral surface
white, uniform except for dark spots on infralabials and irregular
dark speckling under tail.

Total length 389 mm.; tail length 15 mm., length of head to
end of parietals 7.5 mm.

Comparisons. — The number of labials, which we find to be a
relatively unvarying character, and the high ventral count eliminate
all but javanica lineata, gimletti, ventralis, lowi, and gracillima from
consideration. Calamaria gracillima has more than 300 ventrals, no
preocular, and a tiny supraocular; the first lower labials are in
contact. C. lowi has 198-236 ventrals but differs from buchi in the
lack of preoculars, in the smaller eye, in the relatively broader
frontal, and in coloration; the entire ventral surface of lowi is dark;
furthermore, only the third labial enters the eye in lowi. C. ventralis
has 249 ventrals, but has no preocular, a smaller eye, and dark
ventrals. C. gimletti has 200-249 ventrals, but no preocular, a

MARX AND INGER: SNAKES OF GENUS CALAMARIA 197

broader frontal, the first lower labials in contact, and the outer
rows of scales dark. C. javanica lineata has only 199 ventrals, no
preocular, a smaller eye, and light dorsal stripes.

Remarks. — The gut of the type specimen contained the remains
of an earthworm.

Calamaria schmidti sp. nov. Figure 27.

Type. — United States National Museum no. 130240. A female
from Bundu Tuhan, Mount Kina Balu, North Borneo. Collected
at 4,500 feet, on August 1, 1951, by D. H. Johnson. This species is
named in honor of Karl P. Schmidt.

Diagnosis.— A Calamaria with the eye much smaller than its
distance from the mouth; 4 supralabials, the first three subequal;
no preocular; frontal about five to six times as wide as supraocular;
maxillary teeth conical; first pair of infralabials in contact behind
mental.

Description of type. — Rostral broader than high, visible from
above; prefrontals squarish, maximum length subequal to length of
frontal, posterior border transverse, in contact with first and second
labial and eye; nasal large, slightly larger than eye; eye small, di-
ameter one-half its distance from the mouth; no preocular; one small
postocular, not as high as eye; supraocular small, about equal to
nasal; frontal about as wide as long, pentagonal, 5 to 6 times width
of supraocular, about two-thirds length of parietals; 4 supralabials,
first three subequal, fourth twice length of second, second and third
entering eye; 5 infralabials, first pair in contact behind mental,
fourth largest; two pairs of chin shields, both pairs in contact,
anterior in contact with three infralabials, anterior pair larger than
posterior pair; parietals bordered posteriorly by three nuchals;
vertebral row of nuchals distinctly smaller than para-vertebral rows.
Maxilla with 7 conical, slightly curved teeth widely spaced at the
base.

Scales in 13 rows; ventrals 144; subcaudals 14; anal entire; tail
ending in a blunt point; position of reduction to 4 dorsal scale rows,
counting subcaudals forward from terminal scute 6.

Color (in alcohol) purplish gray above, uniform; head without
markings; supralabials same color as back; anterior infralabials and
first pair of chin shields purplish gray, remainder of under side of
head yellowish; ventral surface without markings; anterior ventrals
yellowish, belly becoming increasingly more purple posteriorly, but

198

FIELDIANA: ZOOLOGY, VOLUME 37

Fig. 27. Calamaria schmidti sp. nov. (type, USNM 130240); actual length
of head 9 mm.

lighter than dorsal color; under side of tail darker than belly but
slightly lighter than dorsal surface.

Total length 253 mm., tail length 19 mm., length from tip of
snout to posterior border of parietals 9 mm.

Paratype. — Stanford University no. 8568 from Mount Kina
Balu, North Borneo. This animal is in several pieces and somewhat
dried. However, the head scutellation is distinct and unmistakably
identical to that of the type.

The following data are all that can be gleaned from this snake;
subcaudals 14; supralabials 4, second and third entering eye, first
three subequal; first pair of infralabials in contact behind mental,
first three pairs in contact with anterior chin shields; frontal 5 to 6
times width of supraocular; no preocular; postocular present; position
of reduction to 4 dorsal scale rows, counting subcaudals from
terminal scute 7; maxilla with 6, widely spaced, conical teeth.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 199

Comparisons. — Only three species of Calamaria agree with
schmidti in having four supralabials, the first infralabials in contact
behind the mental, and no preocular. The three are gimletti, gracil-
lima, and doerianense. The first two differ radically from schmidti
in ventral counts, gimletti having 202-249 and gracillima having
300-320 as compared to 144 in the type of schmidti. The frontal-
supraocular ratio of schmidti (5-6) is about twice that of gimletti
(3-3^).

Calamaria doerianense has the second and fourth supralabials
subequal in length whereas in schmidti the first three supralabials
are subequal and much smaller than the fourth. The frontal-supra-
ocular ratio of schmidti (5-6) is twice that of doerianense (2-3) .

Remarks. — The type specimen contained an earthworm.

Calamaria lowi Boulenger

Calamaria lovii Boulenger, 1887, Ann. Mag. Nat. Hist., (5), 19: 169, fig. —
Rejang River, Sarawak.

Descriptive notes. — Two males; total length 122-239 mm.; tail
0.07 (two) of total; ventrals 198-202; subcaudals 19-20. Two
females; total length 279-316 mm.; tail 0.04-0.05 of total; ventrals
223 (two); subcaudals 14 (two).

Supralabials 4, third only entering the eye; infralabials 4, first
pair not in contact behind mental, first two pairs in contact with
anterior chin shields; no preocular; postocular 1; position of reduction
to 4 dorsal scale rows, counting subcaudals forward from terminal
scute 3-6; maxilla with 8-9 modified teeth.

In all respects these snakes agree with the original description.

Locality and specimens examined. — SARAWAK: Fourth Division,
Long Mujan (RM 51, 55, no number; SU 8525).

CHECK LIST

Calamaria acutirostris Boulenger

Calamaria acutirostris Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 17: 394 —
Loka, Mount Bonthain, Celebes.

Calamaria albiventer Gray

Changulia albiventer Gray, 1834, 111. Ind. Zool., 2, pi. 86, figs. 6-9 — Penang.

Calamaria albopunctata Barbour

Calamaria albopunctata Barbour, 1908, Bull. Mus. Comp. Zool., 51: 319 —
East Indies.

200 FIELDIANA: ZOOLOGY, VOLUME 37

Calamaria alidae Boulenger

Calamaria alidae Boulenger, 1920, Ann. Mag. Nat. Hist., (9), 5: 282— Lebong
Tandai in Benkulen, Sumatra.

Calamaria apraeocularis Smith

Calamaria apraeocularis Smith, 1927, Proc. Zool. Soc. London, 1927: 224 —
Djikoro, Mount Bonthain, Celebes.

Calamaria beccari Peters

Calamaria beccarii Peters, 1872, Ann. Mus. Civ. Genova, 3: 34 — Sarawak.

Calamaria benjaminsi Edeling

Calamaria benjaminsi Edeling, 1864, Ned. Tijds. Dierk., 2: 202 — Marta-
pura, Borneo.

Calamaria bicolor Dumeril and Bibron

Calamaria bicolor Dumeril and Bibron, 1854, Erp. Gen., 7: 78 — Borneo.

Calamaria bitorques Peters

Calamaria bitorques Peters, 1872, Monatsber. Akad. Wiss. Berlin, 1872: 585
— Philippine Islands.

Calamaria borneensis Bleeker

Calamaria borneensis Bleeker, 1860, Nat. Tijds. Ned. Indie, 21: 296 — Sintang,
Borneo.

Calamaria brachyura Boulenger

Calamaria brachyura Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 16: 481 —
Mount Kina Balu, North Borneo.

Calamaria buchi sp. nov.

Dalat, Viet Nam, Indo-China.

Calamaria bungaroides Werner

Calamaria bungaroides Werner, 1901, Zool. Anz., 24: 300 — type locality
unknown, probably Java.

Calamaria ceramensis de Rooij

Calamaria ceramensis de Rooij, 1913, Bijdr. Dierk., 19: 26 — Honitetu, Ceram.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 201

Calamaria collaris Boulenger

Calamaria collaris Boulenger, 1897, Proc. Zool. Soc. London, 1897: 225, pi.
14, fig. 4 — Bone Mountains, Tomohon, Rurukan, Celebes.

Calamaria crassa Lidth de Jeude

Calamaria crassa Lidth de Jeude, 1922, Zool. Meded., 6: 248 — Gunung
Talamau, Ophir District, Padang Highlands, Sumatra.

Calamaria curta Boulenger

Calamaria curta Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 18: 62 — South
Celebes.

Calamaria doderleini Gough

Calamaria doderleini Gough, 1902, Zool. Anz., 25: 645 — Langkat, Sumatra.

Calamaria doerianense Brongersma

Calamaria doerianense Brongersma, 1928, Zool. Anz., 75: 255, 2 figs. — Pulau
Durian, Riouw Archipelago.

Calamaria egregia Barbour

Calamaria egregia Barbour, 1927, Proc. Biol. Soc. Washington, 40: 127 —
Mount Tibang, Borneo.

Calamaria electa Barbour

Calamaria electa Barbour, 1927, Proc. Biol. Soc. Washington, 40: 128 —
Pasir, Borneo.

Calamaria elegans de Rooij

Calamaria elegans de Rooij, 1917, Rept. Indo-Austr. Arch., 2: 158, fig. 64 —
Simalur Island.

Calamaria everetti Boulenger

Calamaria everetti Boulenger, 1893, Proc. Zool. Soc. London, 1893: 525 —
Sarawak.

Calamaria gervaisi Dumeril and Bibron

Calamaria gervaisi Dumeril and Bibron, 1854, Erp. Gen., 7: 76 — Java [in
error, see text p. 182].

Calamaria gimletti Boulenger

Calamaria gimletti Boulenger, 1905, Ann. Mag. Nat. Hist., (6), 15: 456 —
Kelantan, Malay Peninsula.

202 FIELDIANA: ZOOLOGY, VOLUME 37

Calamaria goringi Vogt

Calamaria goringi Vogt, 1925, Zool. Anz., 62: 64 — Java.

Calamaria grabowskyi Fischer

Calamaria grabowskyi Fischer, 1885, Arch. Naturg. Berlin, 51: 50, pi. 4, fig.
1 — Telang and Tameanglaijang, Dusson Timor District, Borneo.

Calamaria gracilis Boulenger

Calamaria gracilis Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 18: 63 —
Mount Bonthain, Celebes.

Calamaria gracillima Gunther

Calamaria gracillima Gunther, 1872, Proc. Zool. Soc. London, 1872: 594,
pi. 39, fig. A — Sarawak.

Calamaria griswoldi Loveridge

Calamaria lumbricoidea griswoldi Loveridge, 1938, Proc. Biol. Soc. Washington,
51: 43 — Luidan River, near Bundu Tuhan, Mount Kina Balu, North
Borneo.

Calamaria hollandi Taylor

Calamaria hollandi Taylor, 1923, Phil. Jour. Sci., 22: 550 — Port Holland,
Basilan, Philippine Islands.

Calamaria hosei Gunther

Calamaria hosei Gunther, 1896, Ann. Mag. Nat. Hist., (6), 17: 229 — Entoyut
River, Sarawak.

Calamaria indragirica Schenkel

Calamaria indragirica Schenkel, 1901, Verh. Ges. Basel, 13: 164, fig. 3 —
Indragiri, Sumatra.

Calamaria javanica javanica Boulenger

Calamaria javanica Boulenger, 1891, Ann. Mag. Nat. Hist., (6), 7: 279 —
Java.

Calamaria javanica lineata Brongersma

Calamaria javanica var. lineata Brongersma, 1928, Zool. Anz., 75: 256 — Long
Petah, Borneo.

Calamaria jeudei Kopstein

Calamaria jeudei Kopstein, 1926, Zool. Meded., 9: 108 — Saparua Island,
Molucca Islands.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 203

Galamaria joloensis Taylor

Calamaria joloensis Taylor, 1922, Phil. Jour. Sci., 21: 203 — Jolo Island,
Philippine Islands.

Calamaria klossi Smith

Calamaria klossi Smith, 1926, Ann. Mag. Nat. Hist., (9), 18: 79 — Siberut
Island, Mentawai Islands.

Calamaria lateralis Mocquard

Calamaria lateralis Mocquard, 1890, Nouv. Arch. Mus. Hist. Nat. Paris,
(3), 2: 136, pi. 8, fig. 4— Mount Kina Balu, Borneo.

Calamaria lautensis de Rooij

Calamaria lautensis de Rooij, 1917, Rept. Indo-Austr. Arch., 2: 163, fig. 66
— Pulau si Laut, Cocos Islands, and Simalur.

Calamaria leucocephala Dumeril and Bibron

Calamaria leucocephala Dumeril and Bibron, 1854, Erp. Gen., 7: 83 — type
locality unknown.

Calamaria leucogaster Bleeker

Calamaria leucogaster Bleeker, 1860, Nat. Tijds. Ned. Indie, 21: 293 — Ampat-
Lawang, Sumatra.

Calamaria linnaei Boie

Calamaria linnaei Boie, 1827, Isis, 20: 539 — Java.

Calamaria lowi Boulenger

Calamaria lovii Boulenger, 1887, Ann. Mag. Nat. Hist., (5), 19: 169, fig. —
Rejang River, Sarawak.

Calamaria lumbricoidea Boie

Calamaria lumbricoidea Boie, 1827, Isis, 20: 540 — Java.

Calamaria lumholtzi Andersson

Calamaria lumholtzii Andersson, 1923, Nyt. Mag. Naturk. Kristiania, 61:
123 — Tumbang Maruwei, Borneo.

Calamaria margaritophora margaritophora Bleeker

Calamaria margaritophora Bleeker, 1860, Nat. Tijds. Ned. Indie, 21: 294 —
Ampat-Lawang, Sumatra.

204 FIELDIANA: ZOOLOGY, VOLUME 37

Calamaria margaritophora gastropicta Holtzinger

Calamaria margaritophora gastropicta Holtzinger, 1920, Arch. Naturg. Berlin,
Abt. 85, p. 106 — Indonesia.

Calamaria mearnsi Stejneger

Calamaria mearnsi Stejneger, 1907, Smiths. Misc. Coll., 50: 30 — Tangob,
Mindanao, Philippine Islands.

Calamaria mecheli Schenkel

Calamaria mecheli Schenkel, 1901, Verh. Ges. Basel, 13: 165, fig. 4 — Indragiri,
Sumatra.

Calamaria melanota Jan

Calamaria linnaei var. melanota Jan, 1862, Arch. Zool. Anat. Phys., 2: 5 —
type locality unknown.

Calamaria mindorensis Boulenger

Calamaria mindorensis Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 16: 481
— Mindoro, Philippine Islands.

Calamaria mjobergi Lonnberg and Rendahl

Calamaria mjobergi Lonnberg and Rendahl, 1925, Ark. Zool. Stockholm,
17A, no. 23, p. 1 — Brastagi, Sumatra.

Calamaria muelleri Boulenger

Calamaria muelleri Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 17: 394 —
Loka, Mount Bonthain, Celebes.

Calamaria nuchalis Boulenger

Calamaria nuchalis Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 18: 62 —
South Celebes.

Calamaria occipitalis Jan

Calamaria occipitalis Jan, 1862, Arch. Zool. Anat. Phys., 2: 9 — Java.

Calamaria ornata Werner

Calamaria ornata Werner, 1909, Mitt. Naturh. Mus. Hamburg, 26: 229,
fig. 6 — Indragiri, Sumatra.

Calamaria pavimentata pavimentata DumeYil and Bibron

Calamaria pavimentata Dumeril and Bibron, 1854, Erp. Gen., 7: 71 — Java.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 205

Calamaria pavimentata formosana Maki

Calamaria pavimentata formosana Maki, 1931, Monog. Snakes Japan, p. 136,
pi. 51A, figs. 90-91 — Arisan, Formosa.

Calamaria pendleburyi Smith

Calamaria pendleburyi Smith, 1931, Bull. Raffles Mus., no. 5, p. 27 — Kiau,
Mount Kina Balu, North Borneo.

Calamaria pfefferi Stejneger

Calamaria pfefferi Stejneger, 1901, Proc. Biol. Soc. Washington, 14: 191 —
Miyako Shima, Yayeyama group, Riu Kiu Archipelago.

Calamaria polillensis Taylor

Calamaria polillensis Taylor, 1923, Phil. Jour. Sci., 22: 549 — Polillo, Polillo
Island, Philippine Islands.

Calamaria prakkei Lidth de Jeude

Calamaria prakkei Lidth de Jeude, 1893, Notes Leyden Mus., 15: 252 —
Sandakan Bay, North Borneo.

Calamaria quinquetaeniata Despax

Calamaria quinquetaeniata Despax, 1912, Bull. Mus. Paris, 18: 203 — East
Indies.

Calamaria raveni Cochran

Calamaria raveni Cochran, 1923, Proc. Biol. Soc. Washington, 36: 92 —
Sungai Merah, Borneo.

Calamaria rebentischi Bleeker

Calamaria rebentischi Bleeker, 1860, Nat. Tijds. Ned. Indie, 21: 293 — Singa-
wang, Borneo.

Calamaria schlegeli Dumeril and Bibron

Calamaria schlegeli Dumeril and Bibron, 1854, Erp. Gen., 7: 81 — Borneo.

Calamaria schmidti sp. nov.

Bundu Tuhan, Mount Kina Balu, North Borneo.

Calamaria semiannulata Boettger

Calamaria semiannulata Boettger, 1898, Kat. Rept. Senck. Mus., 2: 84 —
Borneo.

206 FIELDIANA: ZOOLOGY, VOLUME 37

Calamaria septentrionalis Boulenger

Calamaria septentrionalis Boulenger, 1890, Proc. Zool. Soc. London, 1890:
34 — Kiu Kiang and Hong Kong.

Calamaria simalurensis de Rooij

Calamaria simalurensis de Rooij, 1917, Rept. Indo-Austr. Arch., 2: 159,
fig. 65 — Simalur Island.

Calamaria smithi Dunn

Calamaria smithi Dunn, 1923, Jour. Malayan Asiat. Soc, 1 : 3 — Long Mujan,
Sarawak.

Calamaria sondaica Barbour

Calamaria sondaica Barbour, 1908, Bull. Mus. Comp. Zool., 51: 319 — Buiten-
zorg, Java.

Calamaria stahlknechti Stoliczka

Calamaria stahlknechti Stoliczka, 1873, Jour. Asiat. Soc. Bengal, 42: 119,
pi. 11, fig. 2— Dilli, Sumatra.

Calamaria suluensis Taylor

Calamaria suluensis Taylor, 1922, Snakes Phil. Isls., p. 189 — Cagayan Sulu,
Philippine Islands.

Calamaria sumatrana Edeling

Calamaria sumatrana Edeling, 1870, Nat. Tijds. Ned. Indie, 31: 379 —
Redjang, Sumatra.

Calamaria tropica Taylor

Calamaria tropica Taylor, 1922, Snakes Phil. Isls., p. 194 — near Naujan,
Mindoro, Philippine Islands.

Calamaria ulmeri Sackett

Calamaria ulmeri Sackett, 1940, Not. Nat., no. 41, p. 2 — northwest of Blang-
kedjeren, Atjeh Province, Sumatra.

Calamaria uniformis Smith

Calamaria pavimentata var. uniformis Smith, 1921, Proc. Zool. Soc. London,
1921 : 426 — Langbian peaks, Annam, Indo-China.

Calamaria ventralis Cochran

Calamaria ventralis Cochran, 1923, Proc. Biol. Soc. Washington, 36: 91 —
Sungai Merah, Borneo.

MARX AND INGER: SNAKES OF GENUS CALAMARIA 207

Calamaria vermiformis vermiformis Dumeril and Bibron

Calamaria vermiformis Dumeril and Bibron, 1854, Erp. Gen., 7: 85 — Java.

Calamaria vermiformis grayi Gunther

Calamaria grayi Gunther, 1858, Cat. Snakes Brit. Mus., p. 6 — Philippine
Islands.

Calamaria virgulata Boie

Calamaria virgulata Boie, 1827, Isis, 20: 540 — Java.

Calamaria zamboangensis Leviton

Calamaria zamboangensis Leviton, 1952, Jour. Washington Acad. Sci., 42:
239, fig. 1 — Zamboanga, Mindanao, Philippine Islands.

REFERENCES
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1894. Catalogue of the snakes in the British Museum (Natural History).
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Dumeril, A. M. C. and Bibron, G.

1854. Erpetologie general ou histoire naturelle des reptiles. 7, pt. 1, xvi+780
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Eydoux, Fortune and Gervais, Paul

1837. Voyage de la Favorite. Reptiles. Guerin's Mag. Zool., 1837, CI. 3,
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Fischer, J. G.

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Gunther, Albert

1858. Catalogue of colubrine snakes in the collection of the British Museum.

xvi+281 pp. London, Taylor & Francis.
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Museum. Ann. Mag. Nat. Hist., (3), 15: 89-98, pis. 2, 3.

Haas, C. P. J. de

1950. Checklist of the snakes of the Indo-Australian Archipelago. Treubia,
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Haas, Georg

1930. tJber die Kaumuskulatur und die Schadelmechanik einiger Wiihl-
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Jan, Georgio and Sordelli, Fernando

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Mertens, Robert

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Pope, C. H.

1935. Reptiles of China. Nat. Hist. Central Asia, 10: Hi +604, 27 pis., 78
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Roou, Nelly de

1917. The reptiles of the Indo-Australian Archipelago. II: Ophidia. xiv+334
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Smith, M. A.

1930. The Reptilia and Amphibia of the Malay Peninsula. Bull. Raffles
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208

MARX AND INGER: SNAKES OF GENUS CALAMARIA 209

1931. The herpetology of Mt. Kinabalu, North Borneo, 13,455 ft. Bull.

Raffles Mus., no. 5, pp. 1-32, pis. 1, 2.
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3, xii+583 pp., 166 figs.

Taylor, E. H.

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and new subspecies. Phil. Jour. Sci., 12: 353-379, pis. 1, 2, text figs. 1, 2.
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Tweedie, M. W. F.

1950. Notes on Malayan reptiles, no. 2. Bull. Raffles Mus., no. 23, pp. 191-
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