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FIELDIANA: ZOOLOGY
A Continuation of the ZOOLOGICAL SERIES of
FIELD MUSEUM OF NATURAL HISTORY
VOLUME 49
NATURAL HISTORY ae
FOU Sy M ees ¢\ ee ARSHALL
CHICAGO NATURAL HISTORY MUSEUM CHICAGO, U.S.A. 1965
THE SYSTEMATICS AND EVOLUTION OF THE ORIENTAL COLUBRID SNAKES OF THE GENUS
CALAMARIA
ROBERT F. INGER
Curator AND HYMEN MARX Associate Curator
Division of Amphibians and Reptiles
FIELDIANA: ZOOLOGY VOLUME 49
Published by CHICAGO NATURAL HISTORY MUSEUM
NOVEMBER 29, 1965
Edited by EDWARD G. NASH
Library of Congress Catalog Card Number: 65-2467 1
PRINTED IN THE UNITED STATES OF AMERICA BY CHICAGO NATURAL HISTORY MUSEUM PRESS
CONTENTS
PAGE INTRODUCTION « «.% 2 % 0 ¢-s 4 3 4 jee & oH eG ee Hee eS 5
GENERIC DESCRIPTION. . . . . . 2. 1 1 ee ee TAXONOMIC PROCEDURE ... 2... 60 8 ee ee ee ee ee ee ew) 8 RECOGNITION OF SPECIES .......... 2... 2. 2... 2.2... 8 VARIATION OF CHARACTERS ............... 2... ... 14 SPECIES:ACCOUNTS «4: 4 @ 4.6 4 6 6 WE 2B bw eR ae RRR Ee ~ OO EVOLUTION AND SPECIATION. ........... 2... 2... 2... 241 Phylogenetic relations of the genus Calamaria ............ 241 Evolutionary trends within the genus Calamaria ........ . . 252 Patterns of evolution. ......... uo Oe ee A ee me DOYSIDUTION. «. con « 2 oe Gd ee ee Ee Ree wee ee oe BE Centers of evolution and dispersal. .............. . . 275 Time of dispersal and speciation. ............... . . 279 DUNMAR Ys 4s ae va Bo wea ee eee ee ee ee ole wee See APPENDIX A—Formation of Dendrogram ............... . 285 APPENDIX B—Faunal List ............... 2... 2... . 298 REPWRENCES 5 4 « 4&6 24 % © & KR w Re & Sd og ewe eS a & » ZOD TINDIES ¢ ie. boos we Oe me Be & Bw ee eo ee ea a we 1. we Be: OE
INTRODUCTION
The genus Calamaria, distributed from southern China and the Riu Kiu Islands southward through the East Indies to the Philip- pines and Celebes, is one of the two or three most successful genera of colubrids in the eastern Oriental tropics. These highly specialized, burrowing, forest-dwelling snakes are easily recognized as members of a single genus. Very few species of colubrid snakes have as few scale rows (13) as Calamarza; none have more extensive fusion of head scales. In this highly restricted morphological framework, no amount of speciation can disguise the generic identity of a form. One result of this homogeneity is that only four generic names have been pro- posed for the 148 forms named since 1826.
Definition of species limits, however, has been a difficult problem in the past because of the homogeneity of the group and because so few specimens have been available to most workers. Since the last revision (de Rooij, 1917) of a large section of the genus, two enormous collections of a few species have been made at several localities in Java (Kopstein, 1941; de Haas, 1941). These collections housed in the Rijksmuseum van Natuurlijke Historie, Leiden, provide a critical means of evaluating variation.
Without an opportunity to examine these Javanese snakes in our own laboratories, our study would have been seriously hampered. We are, therefore, especially grateful to Drs. L. G. Brongersma and M. Boeseman of the Rijksmuseum for having loaned us these very large collections. The curators of other museums have also been generous and in many cases took the most liberal view of adminis- trative rules in lending us specimens. As a result we were able to see more than 2600 specimens.
We wish to express our thanks to the following colleagues and institutions for permission to study specimens in their collections: Gordon Conway and Chin Phui Kong, Agriculture Department, Tuaran, Sabah; Charles M. Bogert, American Museum of Natural History (AMNH); James E. Bohlke, Academy of Natural Sciences of Philadelphia (ANSP); J. C. Battersby (retired) and Alice G. C. Grandison, British Museum (Natural History) (BMNH); Alan E.
5
6 FIELDIANA: ZOOLOGY, VOLUME 49
Leviton, California Academy of Sciences (CAS); Neil D. Richmond, Carnegie Museum (CM); Chicago Natural History Museum (CNHM); Edward H. Taylor (EHT-HMS); Bengt Hubendick, Naturhistoriska Museet Goteborg (GM); A. Capart, Institut Royal des Sciences Naturelles de Belgique (IRSNB); Enrico Tortonese and Lilia Capocaccia, Museo Civico Genova (MCG); Ernest E. Williams, Museum of Comparative Zoology (MCZ); Villy Aellen, Museum d’Histoire Naturelle, Genéve (MHNG); Jean Guibé, Museum Na- tional d’Histoire Naturelle, Paris (MHNP); Robert C. Stebbins, Museum of Vertebrate Zoology (MVZ); F. Gouin, Musée Zoologique de l’Université, Strasbourg (MZUS); Josef Eiselt, Naturhistorisches Museum Wien (NHMW); Lothar Forcart, Naturhistorisches Mu- seum Basel (NMB); Eric R. Alfred, National Museum of Singapore (NMS); Greta Casteegley, Naturhistoriska Rijksmuseum Stockholm (NRS); L. D. Brongersma and Mart Boeseman, Rijksmuseum van Natuurlijke Historie (RMNH); Tom Harrisson, Sarawak Museum (SM); A. Kleinschmidt, Staatliches Museum fiir Naturkunde in Stuttgart (SMNS); Robert Mertens and Konrad Klemmer, Sencken- berg Museum (SNG); George S. Myers and Walter C. Brown, Nat- ural History Museum of Stanford University (SU); Norman Hartweg and Charles F. Walker, University of Michigan Museum of Zoology (UMMZ); Doris M. Cochran, United States National Museum (USNM); Leif R. Natvig, Universitetets Zoologiske Museum, Oslo (UZMO); Dirk Hillenius, Zoologisch Museum Amsterdam (ZMA); Heinz Wermuth (formerly) and Giinther Peters, Zoologisches Museum Berlin (ZMB); Werner Ladiges, Zoologisches Museum Hamburg (ZMH); Walter Hellmich, Zoologische Sammlung des Bayerischen Staates, Munchen (ZSBS); M. L. Roonwal, Zoological Survey of India (ZSI).
We are also grateful to Dr. G. B. Rabb, and Dr. T. M. Uzzell for critical comments on various parts of the manuscript. The drawings, charts, and maps were prepared by Miss Marion Pahl, Miss Phyllis Wade, and Miss Janet Wright. The photographs were taken by Mr. H. V. Holdren. The several versions of the manu- script were typed by Mrs. Marion Anderson and Miss Janet Wright.
The research was partially supported by National Science Foun- dation Grant G-6257.
CALAMARIA H. Boie
Calamaria H. Boie in F. Boie, 1827, Isis, 20, col. 519 and 539—type species Calamaria lumbricoidea Boie by subsequent designation of Brongersma, Inger, and Marx, ms.
Changulia Gray, 1834, Ill. Ind. Zool., 2, pl. 86, figs. 6-9—type species Chan- gulia albiventer Gray by original designation.
Typhlocalamus Gunther, 1872, Proc. Zool. Soc. London, 1872, p. 589—type species T'yphlocalamus gracillimus Gunther by original designation.
Keiometopon Taylor, 1962, Univ. Kansas Sci. Bull., 43, p. 257—type species Keimetopon booliatt Taylor by original designation.
Application for conservation of the generic name has been made to the International Commission on Zoological Nomenclature (Bron- gersma, Inger, and Marx, MS).
Habitus vermiform; head not distinct from neck; tail short.
Eye distinct, diameter greater than, equal to, or smaller than eye- mouth distance; rostral rounded; paired prefrontals fused with inter- nasals and in contact with supralabials; frontal 114 to 8 times width of supraocular; parietal meeting last supralabial; an enlarged para- parietal (Fig. 1); 4 or 5 supralabials, second and third, third, or third and fourth touching eye; nasal single and surrounding nostril; no loreal; a single preocular present or absent; one postocular; no an- terior temporal; mental touching or separated from anterior chin shields; 5 infralabials, first two or three touching anterior chin shield; two pairs of chin shields, posterior ones meeting or separated in mid- line; 3 or 4 gular scales in midline between chin shields and first ventral.
Dorsal scales in 13 longitudinal rows throughout body length; scales smooth; no apical pits; anal single; subcaudals paired.
Posterior vertebrae without hypapophyses; 6 to 12 maxillary teeth, modified (Fig. 3) in most species; palatal and mandibular teeth conical.
This genus can be distinguished from all other snakes on the basis of the combination of the following characters: dorsal scales in 18 rows throughout body; internasals and prefrontals fused; parietal broadly in contact with supralabials.
7
8 FIELDIANA: ZOOLOGY, VOLUME 49
The genus Keiometopon Taylor was based on a single specimen collected in Malaya. Through the courtesy of Dr. E. H. Taylor we have been able to examine the type specimen. It is clearly an anom- alous specimen of Calamaria schlegeli. The frontal shield is fused with the prefrontals and a median suture runs the length of the dorsal surface of the head. The scales on the side and ventral surface of the head are normal. Several fusions of the longitudinal scale rows occur opposite the first fifteen ventrals. Between the twentieth and the twenty-sixth ventrals the second scale row on each side splits returning the dorsal scale count to thirteen, which number continues to the vent. The maxillary teeth have the characteristic modifica- tion of Calamaria.
TAXONOMIC PROCEDURE In establishing taxa and their interrelations, we have adopted the following procedure: 1) examine all characters practically accessible; 2) analyze variation of characters within sympatric samples; 3) correlate all characters within individuals within samples; 4) establish sympatric taxa on basis of (2) and (8); 5) extend analysis of variation to allopatric samples; 6) on basis of (4) and (5) establish limits of species;
7) compare Calamaria with other genera of Colubridae to deter- mine distribution of certain characters in the family;
8) on basis of (7) decide which characters of Calamaria are primitive (within the genus) and which derived (or ad- vanced) ;
9) on basis of (5) and (6) establish relations between species (or species groups) ;
10) on basis of (8), (9), and zoogeographic possibilities, work out
phylogeny within the genus.
The validity of the procedure can be tested through examination of the logic and with fresh material. Perhaps the latter is the best test, for in effect by our taxonomy we are making predictions con- cerning associations of characters in snakes not yet examined.
RECOGNITION OF SPECIES
Starting with large sympatric samples, such as one from Tjikad- jang, Java (Table 1), we have examined as many characters as pos-
INGER AND MARX: THE SNAKE GENUS CALAMARIA 9
sible and then analyzed the distribution of the total sample with respect to these characters. As shown in Table 1, the sample from Tjikdajang may be treated as two isolated populations (linnaez and lumbricoidea) that differ in ventral counts (both sexes), subcaudal counts (females), tail ratio (males), supralabials, three aspects of coloration, and ornamentation of hemipenes.
Treated as one population the Tjikadjang sample consists of two clusters of individuals in a multidimensional space generated by the 14 characters. Fragmentation of a species into two such disparate groups at one locality can occur only under very limited circum- stances. The separation may be based on age differences. In this case juveniles (less than 200 mm.) and adults (females with large ova) occur within each group, thus eliminating this explanation. The separation could reflect sexual dimorphism, but the table shows both sexes are well represented in each group. Seasonal changes affect some animals profoundly and produce cyclomorphosis, as in Daphnia (Brooks, 1946). Seasonal variation does not account for the division in this sample as half of the ‘“lumbricoidea”’ group and all of the ‘“‘lin- naev’ group were collected in months January—May; the range of variation (Table 1) within the two groups is too narrow for one-half of the ‘“‘lwmbricoidea’’ group to cause the wide gap between the groups.
The only possible explanation remaining for the split in the sam- ple under the single-species hypothesis is polymorphism. Because of the characters used in the analysis (see below), such polymorphism would involve large segments of the genotypes, which in turn in- creases the possibility of recombinations, crossovers, and hence inter- mediate phenotypes. As no intermediate phenotypes appear if we consider coloration, ornamentation of hemipenis, number of supra- labials, and number of ventrals (which almost certainly have a multi- ple factor mode of inheritance), the polymorphism explanation can be rejected.
Thus the split single-species hypothesis can be rejected for lack of a sound biological explanation. This hypothesis also runs con- trary to all experience with species of snakes. Clearly the only rational interpretation of the correlated distribution of characters in the Tjikadjang sample is that two reproductively isolated popu- lations are involved.
Another large sample, this one from Wonosobo, Java, was ana- lyzed in the same manner (Table 1). In this case three clusters of individuals develop in the multidimensional space. Again the most
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12 FIELDIANA: ZOOLOGY, VOLUME 49
reasonable biological interpretation is that we have isolated popula- tions. Similarly the sample from Tjibodas, Java, consists of three separate populations.
If we combine all three of these local samples, we arrive at four populations, two of which occur at all three localities. These pop- ulations differ from one another in more than a half dozen characters. We have considered sympatric populations that differ to this extent to be distinct species.
A similar analysis of the sample from Mount Bonthain, Celebes, yields three populations that differ as much as the Javan populations though not in exactly the same characters. Not mentioned in the table is the peculiar, pointed snout of acutirostris. The rostral shield of acutirostris does not curve over the dorsal surface of the snout as it does in curta and muellert (compare Figs. 19 and 26). The pre- frontal meets the first three supralabials of acutirostris but only the first two of curta and muellerz.
The sample from Mount Kina Balu, North Borneo (Table 1) is more complex than the preceding ones. Our analysis shows it to consist of nine species, any two differing in a number of characters. The difference between such pairs does not always involve the same characters. For example, swlwensis and schlegeli males do not differ in ventral counts but do in the relation of mental and chin shields; schlegeli and virgulata males differ in ventral counts but not in the relation of mental and chin shields. The table does not do justice to some of the differences. Three species are listed as having a striped dorsal pattern; lateralis has one broad white stripe, vzrgulata several narrow dark stripes, and griswoldi five narrow light stripes on each side.
In analyzing these sympatric groups, we have been striving for a measure of the genotypic divergence associated with genetically de- termined reproductive isolation. We have assumed that reproductive isolation depends on extensive differentiation of genotypic patterns. It is impossible in our present state of knowledge to speak directly of genes and chromosomes of Calamaria. Instead we are obliged to use the phenotype and to assume that it bears approximately the same correspondence to the genotype of Calamaria as the phenotype of Drosophila bears to its genotype.
Some of the characters used in the preceding analysis of sympatric species involve metamerism (counts of ventrals and subcaudals), some conformation (tail reduction, shape of teeth, frontal ratio, or- namentation of hemipenis), and some chemistry (coloration). Some
INGER AND MARX: THE SNAKE GENUS CALAMARIA 13
involve skeleton (teeth), some skin (color and scale proportions), one an accessory reproductive organ, and others (the metameric char- acters) skeleton, muscles, blood vessels, and nerves. It is unlikely that any simple change in the genetic background of developmental patterns will affect all of these types of characters and tissues in pre- cisely the same way.
The characters mentioned in this section do not exhaust the list of those used in the total work. Nonetheless, they represent a good sampling of the phenotype. We therefore assume that we have also sampled the genotype.
By virtue of their independent existence together, which demon- strates their reproductive isolation, these groups of sympatric species provide a rough standard of the kinds and extent of differences that distinguish species of Calamaria. ‘This standard cannot be applied too rigidly, however, for several factors may alter its applicability. For example, differences between sympatric forms may be exag- gerated, compared to those distinguishing allopatric forms, by such phenomena as character displacement (Brown and Wilson, 1956). In the matter of degree of difference, we are far from certain that genotypic differences are strictly proportional to phenotypic differ- ences. We cannot say that in the imaginary case of species A, B, and C having ventral counts, respectively of 140-150, 160-170, and 180- 190, that the genotypic differences between A and C are twice those between A and B or between Band C. Yet there are probably posi- tive correlations between phenotypic and genotypic differences and between the extent of genotypic difference and reproductive isolation, which is our basie criterion of specific differentiation.
In general we have applied the rough standard derived from the comparison of sympatric species to comparisons of allopatric popu- lations. Despite the limitations of that standard, we can be reason- ably sure that allopatric populations differing morphologically to the same extent as sympatric species are at least as different genotypi- cally as are sympatric species.
Serious difficulties arise when allopatric samples are not identical morphologically, yet do not differ as much, or in as many characters, as do the sympatric species listed in Table 1. Every sample of snakes is part of a species. In a revision one must allocate each to a species or run the risk of advancing a classification that does not classify many of the available samples. If one treats all characters as taxo- nomically equivalent, as advocated by Sneath and Sokal (1962), a decision in many of these cases is impossible without arbitrary action.
14 FIELDIANA: ZOOLOGY, VOLUME 49
We have felt obliged to judge the differences, in such difficult cases, In terms of the characters involved. That is, we have not considered all characters equally significant or equally helpful in as- sessing potential reproductive isolation, but have instead attributed varying taxonomic significance to each character. We have not graded taxonomic significance in a rigorous fashion such that char- acter a has a value of 1, character 6 a value of 2, and so on. Instead, we have adopted a crude ordinal system that in effect states that a difference between samples in character a is more likely to reflect genotypic difference of a fundamental sort (i.e., one associated with reproductive isolation) than is a difference in character 6. This type of weighting is based on a generic-wide analysis of the extent and nature of variation in each character. Such analysis of characters is the subject of the next section.
VARIATION OF CHARACTERS
Relative width of frontal.—The width of the frontal as a multiple of the width of the supraocular has been used in diagnosing species of Calamaria for at least seventy years (ef. Boulenger, 1894). Within local samples the ratio of frontal to supraocular width varies, the magnitude of the range of variation usually being about one-half multiple of supraocular width (e.g., 14% to 2, see Table 1).
Intraspecific variation itself varies with sample size. As the num- ber of specimens increases, the range of variation in the frontal- supraocular ratio does likewise (Table 2). Similarly, as the maximum value of the ratio within a species increases, the range of variation also increases (Table 3). The last result is not surprising since the physically possible magnitude of variation in a wide frontal is greater than that in a narrow one.
TABLE 2.—Frequency Distribution of Species of Calamaria with Respect to Sample Size and Range of Variation in Frontal : Supraocular Ratio.
Sample Range of variation Total size 1 or more! species 2-5 4 13 6-15 4 13
16-40 3 Uh
41-637 9 ila
! Multiple of supraocular width. A range of 1 may be 114-24, 2144-34, ete.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 15
TABLE 3.—Frequency Distribution of Species! of Calamaria with Respect to Range of Variation in Frontal : Supraocular Ratio and Intraspecific Maxima of the Ratio.
Intraspecific maximum of Range of variation Total ratio 1 or more? species under 2 0 2 2-214 0 14 216-2! /s 4 9 3-3 4 10 13 4 or more 6 6
1 Species with sample size of 1 are omitted. 2 Multiple of supraocular width. A range of 1 may be 114-214, 214-3, ete.
/
The wide range of intraspecific variation characteristic of the frontal ratio makes interpretation of observed differences between samples difficult. The safest criterion for interspecific variation is absence of overlap in the observed ranges of the species being com- pared. Some consideration should also be given to sample size. The difference between schlegeli and linnaei at Wonosobo (Table 1) is an example of reliable interspecific variation; the ranges do not overlap and the samples are large. The difference between acutirostris and curta at Mount Bonthain (Table 1) probably represents interspecific variation despite the small number of curta involved; the gap be- tween the two ranges gives confidence in the decision. We doubt that the same interpretation can be placed on the difference between the Kina Balu samples of griswoldi and suluensis; the ranges almost overlap and relatively few specimens were examined.
Oculars.—Species of Calamaria have one or no preocular and one postocular. Intraspecific variation in ocular count was found in only two species. In one, schlegelt, a preocular is almost always present in the populations from Malaya and Singapore, almost always absent in the population from Java, and present or absent in snakes from Borneo and Sumatra. When a preocular is present in a snake from a population characterized by the absence of preoculars, the scale is small.
We examined only three specimens of the second variable species, lumholtzi. Two of the three lacked preoculars but the third had a small one.
Labials.—Species of Calamaria have either four or five supra- labials. The third and fourth labials border the eye in those species having five supralabials, whereas the second and third do when four
16 FIELDIANA: ZOOLOGY, VOLUME 49
supralabials are present. The only exception to this rule is low7 lowz, which has four supralabials and only the third one bordering the eye in some populations.
Intraspecific variation in supralabial counts is negligible. Of 276 specimens of lwmbricoidea counted, 270 had the supralabial formula (2-3)
5(8-4), three had the formula 5(2-3), two had the formula 5 (3-4)’
—3) 5(3-4)" Of 2538 gervaist, 251 had the formula 5(8-4). Of 223 schlegeli, 220 had the formula 5(8-4). Of 199 modesta, 198 had 5(8-4).
4 (2-3) (2-3) Of 178 linnaet, 171 had 4(2-8), two had —~>5-,., three had 5-5—.
5(3-4)’ (8-4)’ and two had 5(8-4).
and one
Supralabials, because of the limited intraspecific variation, are useful in distinguishing species.
Infralabials show less interspecific variation than supralabials. Five is the usual number, the first three touching the anterior chin shield. Calamaria borneensis invariably has four infralabials of which only the first two touch the anterior chin shield. In C. lowz the infra- labial formulas are 4(2) or 5(8).
Paraparietals—All species of Calamaria have an enlarged scale or shield along the posterolateral margin of the parietal and behind and slightly above the level of the last supralabial (Fig. 1). This shield, which we call the paraparietal, is usually surrounded by five or six scales counting the parietal and supralabial.
The functional significance of the difference between these counts is obscure, if indeed there is any. Certain specimens of lumbricoidea have only four scales bordering the paraparietal; in these specimens the paraparietal is short, squarish, and does not reach the end of the parietal. In other lumbricoidea, the paraparietal is oblong as in Fig. 1 and reaches the end of the parietal or slightly beyond. Although the paraparietals of the two snakes shown in Fig. 1 differ in length, spe- cies having five scales bordering the paraparietal do not have con- sistently longer or shorter paraparietals than those with six.
The difference in counts reflects the number of scale rows meet- ing the paraparietal. The fourth, fifth, and sixth rows always touch the paraparietal. Species in which the third row also meets the para- parietal have six scales bordering that shield.
INGER AND MARX: THE SNAKE GENUS CALAMARIA Li
Fic. 1. Position of paraparietal (Ppa) and number of scales surrounding it in Calamaria buchi (top) and C. schmidti (bottom). Pa=parietal. SL=last supra- labial.
Unfortunately, we discovered this variation in scutellation after we had returned part of the material loaned to us. As a result, we have almost no information on this character in some species (e.g., pavimentata, septentrionalis) otherwise examined in large numbers. Despite this handicap, the data in Table 4 which lists the larger samples indicate the patterns of variation in this character.
Individual variation within a local population is relatively insig- nificant. For example, all of 25 linnaei from Bogor have counts of
18 FIELDIANA: ZOOLOGY, VOLUME 49
TABLE 4.—Frequency Distribution with Respect to Number of Scales Bordering the Paraparietal in Large Samples of Calamaria.
Species Seales bordering paraparietal 4/4 4/s 5 /s 5/s 6/s 6/4 7 /, COUNT OSI TSes Aa odode GS woriad 1 3 24 1 VCO 2.0 neta pee ie See aes 1 9 DHOTOUES 66.6652 % 0nd bees es iw! it DOTNCCNSIS: ca sc ce SL Se as 14 gervaist (Manila)........... 1 94 5 gervaist (remainder)........ 2 155 12 3 GPA0OWS CDE ons % 6 x Das sesteaccac 1 1 18 2 OQUTSWOWD 25:8 ose es He Dae a plat leucogaster................ 15 1 1 TOC es aan as a ae A at 2 61 i WOW oso ex oon Sho ee ae 34 lumbricoidea (Philippines)... 9 4 5 lumbricoidea (remainder)... . 2 151 9 Z modesta (Borneo).......... 4 modesta (remainder)........ 3 151 i MUCHET. vcs Shee ba when ss 1 62 1 MUCHOS <6 onc Hie eee Y wee 1 7 schlegeli (Sumatra)......... 1 6 schlegeli (remainder)...... ; 12 1 SIUNCISTS cg once ck weal | 13 1 i! sumatrana............005. 17 virgulata (Celebes)......... 14 virgulata (Java)............ a virgulata (Sumatra)........ 1 i virgulata (Borneo).......... 4 virgulaia (Sulu). .... 6.0.0.5 5 virgulata (Mindanao)....... 4 nt virgulata (Palawan)........ 3
6/6, all 11 griswoldi from Mount Kina Balu have 5/5, and all but 6 per cent of gervaist from Manila have 5/5.
The restriction of local variation gives reliability to the geographic variation observed (Table 4). The extent and sharpness of geo- graphic variation is not the same in all species. Calamaria gervaisi, which ranges throughout the Philippine Islands except for Palawan, shows no geographic variation. Although lumbricozdea from the Phil- ippine Islands differ in this character from the rest of the species’ sample, the frequency distributions overlap considerably. Roughly the same amount of variation occurs in vrgulata, but differences between populations are sharper.
Our observations indicate that, despite some intraspecific varia- tion and despite limited interspecific variation, this character must be used in any taxonomic work on the genus Calamaria.
Relation of mental to anterior chin shields.—The mental shield may touch the anterior chin shields or be separated from them by the first infralabials. We found no intraspecific variation in 31 species; 13
INGER AND MARX: THE SNAKE GENUS CALAMARIA 19
species varied to some extent; nothing can be said about the six spe- cies of which we saw only one specimen each. Intraspecific variation (Table 5) may take the form of a relatively rare anomaly as in lin- naer or schlegelt. In pavimentata, lowi, and virgulata the variation is distinctly geographic. In alidae and modesta variation, though more frequent than in some species, is not geographic.
TABLE 5.—Frequency Distribution with Respect to the Relation Between the Mental and Anterior Chin Shields in Variable Species of Calamaria.
Asym- Species Touching Separated metrical!
oO ~ isr) i) ~~ j~) ~ —_ iw)
CUCLON oakea Syne bum oe He sd Be Re 0 GTQDOWSIYE on cc ceed dea ew ba bes 21 leucogaster é UOC eS os estas ws x vi xcs bahen ade We eS Dae lowt (Borneo)................0.. 57 low. (Malaya). ..004¢56040 e004 ones WOCHENL «eee eb ered = we oe wes we MOdESIA... 2. ee ee eee ees palavanensis......... 00.0 eee pavimentata (Laos)............... pavimentata (remainder).......... SCWICGOU < okk-da seeks Bae ee hes ak virgulata (S. Celebes)............. virgulata (N. Celebes)............ virgulata (remainder).............
oe) —_ Co e we) SOO HALPODwWeOAOCcrh
bo oD AD
— SCoOonrnronNnrKo RPOONNORNOCCOrFOrFH Oo
Qe ono
1 One of first infralabials reaching mid-line and thus separating mental from chin shield of that side.
The snakes listed as asymmetrical in Table 5 have been instruc- tive. In these specimens the first infralabial of one side reaches the midline and thereby separates the mental from the chin shield of that side. This variant thus forms an intermediate between two extreme types. Nominate species have been described that differ from others only in the relation of the mental to the chin shields.
Calamaria simalurensis de Rooij, for example, differs from mo- desta only in having the mental touching the anterior chin shields. Yet in the publication (de Rooij, 1917) describing simalurensis, the illustration of the holotype of another new species, elegans (also from Simalur Island), clearly shows the asymmetrical arrangement. Three other specimens from Simalur we have examined have the mental separated from the chin shields as in Javan modesta and agree with modesta in all other characters. The large Javan sample (167 speci- mens) includes six snakes that have the mental touching the chin
20 FIELDIANA: ZOOLOGY, VOLUME 49
shields and six that have the asymmetrical pattern. This intermedi- ate arrangement helped us to interpret the variation in this character.
The holotype of Calamaria schlegeli differs from the types of leuco- cephala and agamensis only in this character. This difference was apparently the reason why Boulenger (1894) separated the last two from schlegelt. The holotype is the only specimen of the more than 200 schlegeli we have seen having the mental touching the chin shield. The discovery of two snakes having the asymmetrical arrangements supports our decision that this sole difference between schlegeli and the two other nominate forms reflects individual rather than inter- specific variation.
Table 5 shows that species do differ in this character. The lack of intraspecific variation in 81 species and the low frequency of vari- ants in additional species enable one to use the relation of mental and chin shields as a diagnostic or key character. However, when two samples differ only in this character, probably intraspecific rather than interspecific variation is involved.
Chin shields and gulars.—Every species of Calamaria has two well-developed pairs of chin shields. The anterior chin shields are usually larger than the posterior ones and always reach the mid- ventral line. The posterior chin shields also meet in the mid-ventral line in most species, but are separated by a small gular scale in some.
Three or four gulars precede the first ventral in the mid-ventral line. If the posterior chin shields do not meet in the mid-line, invari- ably there are four gulars in the mid-line. Only one species (apraeoc- ularis) having the posterior chin shields meeting in the mid-line has four gulars in the median row.
The relationship of the posterior chin shields to each other and, consequently, the number of median gulars vary geographically in lumbricoidea (‘Table 21). They also vary in schlegeli, but the varia- tion is not geographic.
Mazxillary teeth.—In our earlier paper on Calamaria, we described the distinctive, basally expanded maxillary teeth of many species of Calamaria. We have extended our survey to every species in the genus. Although we examined enough specimens of most species to determine the scope of major intraspecific variation, we did not measure enough teeth to permit statistically valid analysis of minor variations in size and shape. This survey was adequate, however, to confirm our earlier finding that there are two sharply different types of maxillary teeth.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 21
Fic. 2. Maxilla and maxillary teeth of Calamaria leucogaster (top) and C. sep- tentrionalis (bottom). Length of maxilla in leuwcogaster 5 mm., in septentrionalis 3.5 mm.
The peculiarly modified teeth shown in the lower halves of Fig- ures 2 and 8 have bilobed pulp cavities. The pulp cavity in the un- modified, conical type of tooth (upper halves of Figs. 2 and 8) is conical and slightly curved, paralleling the shape of the tooth.
Serial variation is limited to minor changes in size and, in the unmodified dentition, to slight basal thickening of the most poste- rior teeth. This thickening does not result in a tooth resembling the modified ones. Thus, a snake has but one type of maxillary tooth (Figs. 2 and 8).
Apparently local, individual, and ontogenetic variation do not affect the form of the maxillary teeth. We examined nine luwmbri- coidea from Tjikadjang, Java, and although two were shorter than 200 mm. and two longer than 475 mm., tooth shape in this series was essentially uniform.
The maxillary teeth of lumbricoidea from five land masses, of leu- cogaster from three land masses, of gervaisi from three, of virgulata from four, and of modesta from four were examined. We found no intraspecific variation in tooth shape.
Fic. 3. Unmodified maxillary teeth of Calamaria acutirostris (top) and modi- fied ones of C. linnaei (bottom).
22
INGER AND MARX: THE SNAKE GENUS CALAMARIA 23
The number of teeth in a maxilla varies from 6 to 12. In most species the observed range of variation is 2 or 3 (Table 6). Local individual variation in tooth number in certain species may be as great as that over the entire geographic range of most species. For example, acutirostris from southern Celebes has 10 to 12 maxillary teeth; lowz lowt from western Borneo has 8 to 10. Other species, such as lumbricoidea, show almost no local variation. Geographic variation appears in lowi and virgulata. This form of intraspecific variation may be more common than our limited data indicate.
Sympatric species differ in tooth counts in varying degrees. Bor- nean specimens of modesta have lower counts (6-8) than do Bornean lumbricoidea (10). The counts of Bornean sulwensis and grabowskyt overlap but do not coincide. Of the species from southern Celebes, curta has distinct counts whereas those of the other three species overlap (Table 6).
The tooth counts are not related to the form of the teeth. The species having unmodified maxillary dentition have 6 to 12 teeth, which is almost identical to but slightly exceeds the range (6 to 11) in species having modified teeth.
All but six species of Calamaria have modified maxillary teeth. Because only two types of maxillary teeth occur in this genus, the
TABLE 6.—Variation in Maxillary Dental Counts in Species of Calamaria.
No. of teeth 7 8 9 10 11 12
Species Region No. of specimens curta southern Celebes 3 acutirostris southern Celebes 3 Z 3 muelleri southern Celebes 2 4 2 nuchalis southern Celebes i! 2 lumbricoidea Philippine Islands 3 lumbricoidea Borneo 6 lumbricoidea Sumatra 1 7 lumbricoidea Java Z 2 grabowskyi Borneo Z Z 1 suluensis Borneo 2 2 virgulata Borneo 2 virgulata Java 1 virgulata Philippine Islands a 1 virgulata Celebes Z 3 schlegeli Borneo 4 schlegeli Sumatra y| schlegeli Malaya 3 schlegeli Bangka 1d. schlegeli Java 9 4 lowi lowt Borneo 1 2 4 lowi gimletti Malaya 2 3 prakkei Borneo 5 1 eiselti Sumatra 3 crassa Sumatra (i
24 FIELDIANA: ZOOLOGY, VOLUME 49
usefulness of tooth shape in distinguishing between species is limited despite the absence of intraspecific variation. The number of maxil- lary teeth is potentially of greater value in the definition of species because of the wider intrageneric variation.
The maxillae of species having modified teeth are deeper relative both to the teeth and to the size of the snake. The snakes shown in Figure 2 have teeth of the same length relative to total length. Yet the maximum depth of the maxilla (again relative to total length) of the snake having modified teeth is 1.7 times that of the one having unmodified teeth. The unmodified teeth illustrated are clearly longer relative to the maxilla than are the modified ones.
Similar differences are shown in other specimens. In one of acutz- rostris (447 mm.), which has unmodified teeth, the maximum depth of the maxilla was 42 micrometer units, the length of the longest tooth 45 units. In a linnaet (292 mm.), which has modified teeth, the maxilla was 45 units and the tooth 22. In one lumbricoidea (393 mm.), which also has modified teeth, the maxilla was 52 units and the tooth 32. The depth of the maxilla relative to total length of these three specimens stand in the relationship of 1.0 to 1.6 to 1.4.
Ventrals—The generic range in ventral counts is 119 (joloen- sis, &') to 804 (gracillima, 2). The maximum counts of most spe- cies fall between 146 and 185. The first ventral is preceded by three longitudinal rows of small gular scales and is about as wide as these three rows combined.
Sexual dimorphism in ventral counts is characteristic of Cala- maria. Females of most species have higher counts than males. Unless intraspecific comparison of the sexes is restricted to speci- mens collected at one locality, geographic variation complicates the picture. For example, the counts of males of luwmbricoidea from Tjikadjang, Java completely span the counts of females from Mount Kina Balu although sexual dimorphism is sharp at each locality (Table 7).
Some of the complexities of variation in ventral counts are shown in Table 7 which lists those species represented by at least 17 specimens.
In general, ranges of variation within local populations are rela- tively narrow, all but three of the 30 examples in Table 7 being less than 16. Neither sample size nor absolute number of ventrals exerts much influence on local ranges of variation (Figs. 4 and 5).
Intraspecific ranges, considering entire species samples, are highly variable. Part of the differences between species in the ranges of
range of variation in ventrals
140 150 160 170 180 190 200 210 220 230
maximum number of ventrals within local samples
Fic. 4. Relation between maximum number of ventrals and range of variation in ventral counts within local samples of Calamaria.
26 FIELDIANA: ZOOLOGY, VOLUME 49
local sample size
4 8 10 12 4 16 18 20 “22 24 (26° 28° -30° 432 34
local variation in ventrals
Fic. 5. Relation between local intraspecific variation in ventral counts and local sample size in species of Calamaria.
ventral counts can be explained by the maximum individual ventral counts (Fig. 6). It is self-evident that a species having a maximum count of 230 could have, for example, a range of almost 100 ventrals (as in lumbricoidea), whereas one having a maximum count of 150 could not.
Sample size has little effect on total intraspecific variation (Fig. 7). The ranges of linnaez are about half as extensive as those of grabow- skyt which is represented by less than a fifth as many specimens (Table 7).
Species of terrestrial snakes distributed in several islands might be expected to show greater ranges of variation than those occurring in only one island because of the geographic isolation of the popula- tions of the first group. However, the ranges of variation in species such as grabowskyi and borneensis, which are known only from Bor- neo, may be as great as the ranges of species such as leucogaster and schlegeli, which occur on three or four land masses including Borneo.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 27
yh (eo)
NO je)
oO
range of variation 1n ventrals Ww (o)
140 150 160 170 180 190 200 210 220 230 240 250
maximum number of ventrals within’ species
Fic. 6. Relations between maximum number of ventrals and intraspecific ranges of variation in ventral counts in Calamaria.
The range of ventral counts in gervaisz, which occurs on nine of the Philippine Islands, is no greater than that of schlegeli (known from four land masses) and is less than those of low? and modesta (each known from three land masses). The range of ventral counts in females of lumbricoidea from Borneo is 64, or greater than the range of variation in all other species except pavimentata. Thus the fragmen- tation of the geographic distribution into separate land masses does not affect the range of ventral counts significantly.
Narrow local variation makes it possible to rely on ventral counts for interspecific comparisons, provided the samples being compared are from the same locality (first four lines of Table 7). Interspecific comparisons of ventral counts based on allopatric samples are not reliable because of the wide intraspecific ranges of variation in some species; such interspecific comparisons must be bolstered by evidence from other characters.
Subcaudals.—The observed generic range in these counts is six (septentrionalis, 2) to 44 (schlegelt, ~), the maximum within spe-
28
FIELDIANA: ZOOLOGY, VOLUME 49
TABLE 7.— Intraspecific Ranges of Variation in Ventral Counts of Calamaria in Local and Species-wide Samples.
Species
lumbricoidea grabowskyi
suluensis griswoldi
lumbricoidea
linnaer
linnaei schlegeli
schlegeli
pavimentata
borneensis
lowr
gervaist
modesta
modesta
acutirostris
mueller bicolor
sumatrana
melanota
leucogaster septentrionalis
cies falling between 15 and 30 in 89 species. variation are variable (Table 8).
Single locality Males Females
, TS
No. Range No. Range
[Mount Kina Balu, Borneo]
7 154-165 7 170-180 6 176-181 5 180-190 2 131-136 3 144-147 4 166-179 5 183-192 [Tjikadjang, Java]
26 179-187 30 197-209 25 182-142 25 149-159 [Wonosobo, Java]
12 184-146 19 150-164 44 145-155 54 162-173 [Penang Island]
4 144-151 4 160-167 [Suisharyo, Formosa]
4 148-155 10 166-176 [Long Mujan, Borneo]
9 155-169 5 170-182 3 198-202 5 221-226
[Manila, Luzon| 25 147-158 26 157-190
[Bandung, Java]
28 165-179 19 191-202 [Tjibodas, Java]
18 155-166 25 165-194
ilar to those of ventral counts. Sexual dimorphism in subcaudal counts (see Table 8) follows the
pattern common in the Colubridae.
Entire geographic range
Males
Range
144-196 150-186 129-138 155-179
144-196 130-149
130-149 129-161
129-161
125-168
126-169 163-213
132-164
131-179
131-179
148-161 129-155 139-169 126-157 121-142 126-146 148-166
Females io
No.
101
Range
137-229 164-190 142-168 183-192
34-229 148-166
148-166 136-180
136-180
137-206
159-192 215-256
142-190
158-202
158-202
163-174 155-178 151-160 164-175 131-154 129 157 168-188
Intraspecific ranges of The patterns of variation are sim-
Males have more subcaudals
INGER AND MARX: THE SNAKE GENUS CALAMARIA 29
130]
120
110
100
90
80
70
60
sample _ size
50
40
30
species - wide
20
10
10 20 30 40 50 60 70 80 90
species- wide variation 1n_ ventrals
Fic. 7. Relation between intraspecific variation in ventral counts and species- wide sample size in Calamaria.
than females in all species of Calamaria except possibly suluensis (Table 36, p. 128) and everetti (p. 183). As in the case of ventral counts comparison of the sexes should be based on specimens from a single locality because of geographic variation. Females of lwmbri-
30 FIELDIANA: ZOOLOGY, VOLUME 49
TABLE 8.—Intraspecific Ranges of Variation in Subcaudal Counts of Calamaria in Local and Species Wide Samples.
Single locality Entire geographic range Males Females Males Females Go, eS SE See Species No. Range No. Range No. Range No. Range [Mount Kina Balu, Borneo] lumbricoidea 7 238-24 7 17-19 184 17-27 118 = 13-21 grabowskyt 6 26-29 5 21-24 15 23-29 18 20-28 suluensis Z 18-20 2 17-20 6 18-20 13 14-26 griswoldi 4 16-17 4 138-16 6 16-18 4 18-16 [Tjikadjang, Java] lumbricoidea 26 17-21 30 =. 18-16 184 17-27 118 13-21 linnaer 25 15-19 yas 8-12 74 15-22 95 7-13 |Wonosobo, Java] linnaet 11 17-22 19 9-13 74 15-22 95 7-13 schlegeli 42 28-338 52 23-28 99 25-44 113 19-87 [Penang Island] schlegeli 4 29-32 4 24-25 99 25-44 113 19-87 [Suisharyo, Formosa] pavimentata 4 20-23 10 +=16-18 38 18-33 50 8-20 [Long Mujan, Borneo] borneensis 8 20-22 5 13-19 24 20-26 28 13-21 low2 3 19-22 5 12-14 26 14-26 37 ~=610-18 [Manila, Luzon] gervaist 41 15-19 34 10-14 110 15-21 129 #£10-18 [Bandung, Java] modesta 31 25-29 19 15-19 100 += 19-81 19 W222i [Tjibodas, Java] modesta 14 24-28 18 13-18 100 +=19-81 19) W220 acutirostris 18 20-24 16 13-17 bicolor 12 21-28 7 Le=Zil leucogaster 14 17-26 22. 7 W219 melanota 9 238-26 9 16-20 muelleri 29 16-21 38 9-15 septentrionalis 28 15-19 30 6-11 sumatrana 6 14-20 tal 10-14
coidea from Mount Kina Balu have almost as many subcaudals as males of lwmbricoidea from Tjikadjang, Java (Table 8), though sexual dimorphism is clear at each locality.
The evidence for lack of dimorphism in sulwensis and everettz is equivocal. At Mount Kina Balu the few males and females seen of
INGER AND MARX: THE SNAKE GENUS CALAMARIA 31
suluensis had almost identical subecaudal counts, whereas in south- eastern Borneo males seem to have slightly higher counts (Table 8). In the case of everettt, none of the males came from the localities of the females.
Within local populations the intraspecific range of variation is usually less than six subcaudals. Neither sample size nor maximum number of subcaudals affects the local range of variation (Figs. 8 and 9).
When entire species samples are analyzed, intraspecific ranges in- crease primarily because of geographic variation, which is relatively common (see Tables 26, 30, 45). Intraspecific ranges at this sam- pling level vary according to the maximum individual counts within species (Fig. 8); a species such as schlegeli (maximum subcaudal count 44) can have a total range of variation of 20 subcaudals whereas a species such as septentrionalis (maximum subcaudals 19) cannot. The size of the total species sample had little effect on range of vari- ation (Fig. 9).
Because of narrow local variation, subcaudal counts are helpful in differentiating between sympatric species. Their usefulness in in- terspecific comparisons when allopatric samples are involved is not as great, though some species are decidedly short-tailed and others long-tailed.
Reduction of the number of dorsal scale rows on the tail.—The num- ber of dorsal scale rows on the tail is reduced to four in most species of Calamaria. As in our previous study, we have located the posi- tion of the reduction to four rows by means of the subcaudal oppo- site which it occurs. In Table 9 we have listed the position of the reduction in terms of subcaudals between the level of the reduction and the vent and between the reduction and the terminal scute.
Individual variation is extensive in most species and in both sexes, yet not so extensive as to obscure sex dimorphism and interspecific variation (Table 9).
The position of the reduction to four rows is affected by at least two factors. Whether measured from the vent or from the terminal scute, the position of the reduction is correlated with the total num- ber of subcaudals. It is self-evident that, as the total subcaudal count decreases, the maximum number of subcaudals separating the level of the reduction from either end of the tail must decrease. Yet the total number of subcaudals accounts for only slightly more than half of the variation in position of the reduction (or between 0.39 and 0.82; see Table 10).
32 FIELDIANA: ZOOLOGY, VOLUME 49
The other factor influencing the position of the reduction is tail shape. This influence is reflected in sex dimorphism and in inter- specific variation. Males of Calamaria are like those of most other snakes in having thicker tails basally than females. This difference is evident in the consistently greater mean distance of the level of the reduction from the vent in males (Table 9). Only four of the 21 means of males are less than 11 subcaudals from the vent and hence in the section of the tail sheathing the hemipenes. By contrast 13 of 21 means of females lie within 11 subcaudals from the vent. Sim-
variation
range of
14. 16 18 20 22 24 26 28 30 32
maximum number of subcaudals within local samples
x 20 x O 18}— 16 14 oO xO 612 Xx os xX Lavi “S211 oO X = 6%) co 8 re) x x ey fe) (@) Oo x X » rr i eRe re nk eo 6 9 x o = “0 0. +O KC xO X x= o 4 fe) fe) x :0=Q
I2514 16" VBS 20" 92204245 826728) 305482 34 SG) S8ae40l e425 eas
maximum number of subcaudals within species
Fic. 8. Relation between intraspecific variation in subcaudal counts and maxi-
mum number of subcaudals in local (top) and species-wide samples (bottom) of Calamaria.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 33
130 120 re) — 110 x = 100 x x
a a
70
sample size oO ro) i
20 |= Q cone
30 x“ ® ° pees
Species - wide my ro) © { | local sample size
10
4 6 8 10 12 14 16 18 20
species-wide variation in subcaudals local variation in subcaudals
Fic. 9. Relation between intraspecific variation in subcaudal counts and size of species-wide (left) and local (right) samples of Calamaria.
ilarly the minimum distance from the vent in the 21 species listed in Table 9 is less in females than in males; the minimum distance in females of all 21 species falls within 10 subeaudals (11 of them within 5 subcaudals) whereas only 14 of the minima of males lie in that range (only three within five subcaudals).
Sex dimorphism in the number of subcaudals between the level of the reduction and the terminal scute is, with few exceptions, not significant statistically or otherwise (Table 9).
The importance of shape in determining the position of the re- duction is also shown by interspecific differences. In species having slender tails the reduction takes place much farther from the terminal scute than in thick-tailed species. Males of septentrionalis, linnaer, and griswoldi have approximately the same number of subcaudals
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34
INGER AND MARX: THE SNAKE GENUS CALAMARIA 35
(means 16.5-17.5). In septentrionalis (Fig. 10A) the tail does not taper and six scale rows reach the terminal scute. The tail of lin- naet is slightly tapered and the mean number of subcaudals between the reduction rows and the terminal scute is 5.7 (83 per cent of the
Fic. 10. Ventral views of tails in males of Calamaria septentrionalis (A), C. griswoldi (B), C. modesta (C), and C. grabowskyi (D).
36 FIELDIANA: ZOOLOGY, VOLUME 49
TABLE 10.—Correlation Between Total Number of Subcaudals and Position of Reduction to Four Dorsal Scale Rows in Species of Calamaria.
Reduction Reduction
from vent from terminal scute
r Pp r P Males: ¢ 2s «xx despa eas +0.49 0.02 +0.56 <0.01 Females... .4c26e te, 54 +0 .39 >0.10 +0.82 <0.001
total). The tail of griswoldi (Fig. 10B) is one of the most slender in the genus and the reduction is 11.2 subcaudals (67 per cent) from the terminal scute. Males of modesta and grabowskyi have equally long tails (means of subcaudals 26.3 in both), but that of grabowskyi (Fig. 10D) is tapered near the base and that of modesta (Fig. 10C) only near the tip. The mean position of the reduction in grabowskyz males is 18.5 subcaudals (70 per cent of the total) from the terminal scute and 3.7 (14 per cent of the total) in males of modesta.
Thus despite individual variation and sexual dimorphism, the position of the reduction to four rows can be used as a valid taxo- nomic character and in some cases helps in distinguishing between species. Because sexual dimorphism in the number of subcaudals between the reduction and the terminal scute is less pronounced than in the number between the vent and reduction, we have given the position of the reduction relative to the end of the tail in the descrip- tions of species.
Hemipenis and cloaca.—The retracted hemipenes of slightiy more than half of the species were examined (Table 11). The material available was not sufficient for adequate sampling of each species. We dissected the hemipenis of a single male in nine species, of two males in 11 species, of three males in four species, of four males in two species, of five males in one, of 20 males in one, and of 46 males in one. Our analysis of variation is, therefore, of limited value.
TABLE 11.—Frequency Distribution of Species of Calamaria with Respect to Form of Hemipenis. Body of hemipenis
Forked or Forked Non-forked non-forked Unknown
Calyces SMO OU vc hee 17 ik Papillate:::. 0. ere eee. 6 1 Smooth or papillate...... i 1
Witknowine- eee ree: 2 21
INGER AND MARX: THE SNAKE GENUS CALAMARIA 37
The most common forms of individual variation involve the posi- tion of the fork and the length of the hemipenis. The fork of the retracted hemipenis usually lies opposite the fourth to sixth sub- caudal. In 24 males of lwmbricoidea, the fork was opposite the second subcaudal in two specimens, opposite the third in three, the fourth in 11, the fifth in seven, and the sixth in one. In 12 males of schlegela the fork lay opposite the fourth subcaudal in six, the fifth in five, and the sixth in one. Similar variation occurs in 14 of the 16 other species which have forked hemipenes and of which we examined two or more males.
TABLE 12.—Relations of Length of Hemipenis to Total Number of Subcaudals in Two Species of Calamaria.
C. lumbricoidea Total number of subcaudals
End of hemipenis! 19 20 21 22 28 24 25 26 27 Number of specimens 6 1 1 7 1 8 1 3 1 1 9 3 2 2 2 10 1 3 1 i 1 li 1 C. schlegela Total number of subcaudals 27 28 29 30 31 34 39 40 42 Number of specimens 7 1 8 1 1 9 2° J 1 iL 10 1 1 1 11 1
1 Determined by subeaudal opposite which retractor muscle inserts on hemi- penis.
We have measured the length of the hemipenis in terms of sub- caudals, letting the insertion of the retractor muscle mark the end of the hemipenis. The retractor may insert as close to the vent as the sixth subeaudal and as far as the fourteenth; in most males ex- amined the insertion is opposite the eighth to tenth.
In the two large series examined, lwmbricoidea and schlegeli, the length of the hemipenis varies. But that variation bears little rela- tion to tail length as measured in terms of subcaudals (Table 12). Similar minor variation occurs in other species.
38 FIELDIANA: ZOOLOGY, VOLUME 49
What may be called major, in contrast to the preceding minor, variation appears in sulwensis and lumbricoidea. The calyces of one suluensis male are smooth-edged, those in a second papillate. The hemipenes of lwmbricoidea are simple (non-forked) with smooth-edged calyces, forked with smooth calyces, or forked with papillate calyces. These variants are geographically localized (Inger and Marx, 1962; see also p. 85 of this paper).
As we examined at most two males in 20 of the 29 species dis- sected, the true extent of major intraspecific variation in hemipenes cannot be evaluated from our data. Intraspecific variation as strik- ing as that of lwmbricoidea should not be expected in every species, for only minor variation was found in 20 males of schlegeli from all parts of that species’ wide range.
Most species of Calamaria have deeply forked hemipenes having smooth-edged calyces (Table 11). Although the characteristics of many of the species in the table are based on the dissection of one or two males, the previous generalization seems valid. Even if major intraspecific variation is much more common than our data indicate, it is unlikely that we would have found such a high proportion of snakes falling in one cell of Table 11—provided that our selection of specimens for dissection was random with respect to hemipenial form. As we found no ontogenetic variation in hemipenes and as we could not know the form of the hemipenis prior to dissection, randomness in our selection was assured.
We dissected two males of each of the two species characterized by non-forked hemipenes and two or three males of four species char- acterized by papillate hemipenes. Despite the small samples in- volved, we believe that much of the interspecific variation indicated in Table 11 will be substantiated by future work.
The cloacas of females have three general forms (Inger and Marx, 1962). Cloacas called bulbous have oviducts opening into the cham- ber posterior to the anterior wall of the cloaca. If the anterior end of the cloaca has an indentation and the oviducts open posteriorly, the cloaca is called cardioid. A bilobed cloaca has the oviducts open- ing into the anterior ends of short cloacal lobes.
Females of 20 species were dissected. We examined one female of each of seven species, two of each of 10 species, three of one, four of one, and 27 of one (lwmbricoidea). Only four of the two-specimen samples show no variation. The cloacas of the other multi-speci- men samples fall in at least two categories.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 39
Variation in cloacal form is apparently not related to stage of the reproductive cycle (Inger and Marx, 1962). Possibly the shape of the cloaca is transitory and modified by differential contraction of muscles.
In view of the wide intraspecific variation, we have not utilized cloacal form in our taxonomic decisions.
Thickness.—Species of Calamaria clearly differ in body thickness. Great variation in quality of preservation made it impractical to measure the thickness of the body directly. The only satisfactory method of determining bulk was by water displacement which gave us the volume of each animal. Obviously shrunken and broken ani- mals were not used. Unfortunately these measurements were not made until relatively late in our study after a good part of the bor- rowed material had been returned.
A graduated cylinder was partly filled so that the meniscus was at a given line. A snake was submerged and the increase in volume (to the nearest 0.5 ml.) gave us the volume of the snake. The volume was divided by the total length of the individual giving the volume per millimeter of length. This measure is an index of thickness.
Despite the limitations of small samples, these measurements give clear indication of ontogenetic changes in body thickness, weaker in- dications of sexual dimorphism, and good indications of interspecific variation (Fig. 11).
We have made volume measurements for 43 species (Table 13); 21 species are represented by enough measurements to indicate at least the pattern of individual variation. Eighteen of the 21 show ontogenetic variation clearly. Each of these eighteen species shows increase in the volume per millimeter of total length as total length increases. The three species that do not show clear ontogenetic change in this character are lewcogaster, lowi, and sumatrana; the last two species are slender and the first is moderate in thickness.
In lumbricoidea, gervaist, schlegelt, and virgulata the males seem to be somewhat bulkier than females of corresponding lengths. Even in these species, however, the data are too few to substantiate this relationship. Differences between species in thickness show up with even quick visual examination. For example, evselti is several times the diameter of lowz; these two species represent extremes of thick- ness in this genus (Fig. 11). The results of visual inspection are con- firmed by our measurements of volume (Table 13). Even relatively subtle differences between species (e.g., evselta vs. crassa or lumbri- coidea) can be detected by these measurements.
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INGER AND MARX: THE SNAKE GENUS CALAMARIA
4]
TABLE 13.—Body Volume and Ratio of Volume to Total Length and Numbers of Ventrals and Subcaudals in Calamaria.
Museum number
lowt
SM ‘14’ CNHM 109974 CNHM 72373 CNHM 129001 CNHM 109975 CNHM 129002 CNHM 109976 CNHM 129003
alidae NHMW 16996
apraeocularis
MCZ 25265 MCZ 25300
forcarti
N MB 8958 ZMA 10073 NHMW 16710
buchi CNHM 71697
pavimentata
CNHM 100869 CNHM 11528
virgulata
ZMA 10240 RMNH — RMNH -— SU 22405 RMNH —— RMNH —— RMNH -— NHMW 16695 NMS 4
NMS 4 CNHM 63571
NHMW 16716:2
RMNH —— RMNH 39
NHMW 16716:1
MHNP 5780 SNG 19441
joloensis CAS 60901
boesemani GM 3230
Locality
Borneo oF
Sumatra
Celebes
Nias 9:
Sumatra
Indo-China
Tonkin Annam
Sumatra Celebes
Mindanao
Celebes
+9
Java Celebes
Borneo Celebes
Java Celebes
no data Celebes
Jolo
Celebes
Sex
40 40 10 QV 40 Q, 40 40
40 Qy
Qy 40 A
Qy 40
40 40 Qy Ay 40 40 10 10 10 Qt OQ A110,
Qy
Total Length Vol. mm.
238 238 239 268 276 289 291 304
254
148 237
216 235 239
389
268 244
100 150 173 193 195 219 225 237 254 257 258 271 273 283 286 307 336
144
116
ml.
DO bo bo
ovc
WPF EN WHh DD
BDEUAnNIEP EE PWWWNNR ©
OVdtAN On
OUOr
Vol./TL ml./mm.
.008 .008 .018 .009 .014 .014 .014 .010
cocococococo°o
0.012
0.007 0.013
.009 .009 .008
ooo
0.021
.O11 .012
oo
.008 .010 .014 .010 .015 .014 .018 O17 .016 .016 .016 .018 .023 .021 .019 .015 .018
cooocoooooooooocochwc
0.010
0.009
Ven- trals
226 225 202 238 196 246 233 228
203
178 220
176 200 177
221
186 160
195 180 182 227 160 182 192 197 208 208 205 216 194 189 192 236 209
119
170
Sub- caudals
TABLE 13 (cont.)
Museum
number Locality
brongersmar GM 3229
everettz
RMNH -—
CNHM 63572 BM 1902 a CNHM 109971 os
javanica CNHM 109791 _ Billiton
Celebes
Borneo be ]
lateralis
MCZ 43582 Borneo
sumatrana
MCG 30382 Sumatra ZMA 10238 ie ZMH 3996 : AMNH 2873 ” ZMH 3996 uP RMNH 4860 fs ZMH 2466 m2 ZMA 10237 oe ZMH 2307 22
abstrusa
ZMB 5986 ZMB 5986 BM 64.4.7.11
no data 99
Sumatra?
borneensis
CNHM 109973 Borneo SMNS 4584 u CNHM 76295 a SMNS 4584 = CNHM 72372 ” CNHM 109972 sd
suluensis
SMNS 34.98 RMNH — RMNH — ‘ MHNP 57-812 2. NHMW 16708:2 ZMH 4212 is NMS 146.17 oe NHMW 16708:1 NHMW 16708:3 Ss CNHM 76294 ae
Borneo ’)9
palavanensis
BM 94.6.30.51 CAS 62151 BM 94.6.30.50 +4
Palawan 9)
gervaist
SU 17931 Negros SU 18229 Cebu
10 70 QA
+O
+O
40 40 Q)40 1040 Q AQ,
40 Ay A
40 40 40 40 QQ Q, Q, 40 40 40 40 QV 40 10 410A,
40 QV 40
Qy 40
Total
Length, Vol. Tol:
mm.
225
144 168 187 208
169
190
168 169 174 184 200 208 222 227 234
162 162 209
209 210 228 244 275 353
109 165 190 203 221 227 252 259 264 287
134 280 323
ee 120
42
COD re
St
Loe et SS)
WPWNNMRND W
Dee
OOnOnOFPEPWWHNHE
CIN OVE
Pah) .75
.20
.75
Vol./TL ml./mm.
0.018
o°ococo
cococooooococc
ooo
ooococococococoe ocooocooo
ooo
.014 .009 .O11 .010
.012
.O11
.018 .015 .014 .010 .013 .010 .014 .018 .015
O11 Od .012
.014 .014 .013 .016 .022 .031
.009 .012 .016 .015 .019 .020 .020 .026 .023 .028
.007 .021 .025
.009 .008
Ven- trals
155
157 151 152 153
168
150
126 130 149 165 167 173 157 164 174
130 129 152
138 175 171 176 159 177
147 142 158 129 138 133 147 154 161 164
178 174 180
170 149
Sub- caudals
18
TABLE 13 (cont.)
Museum number
CM 2530
SU 19874 CAS 60471 CNHM 53377 CM 2470
CM 2540 MCZ 37699 MCZ 25775 CM 2485
SU 17929 CNHM 96620 SU 18910
CM 2537 CNHM 533880 MCZ 25752 CM 2538
SU 15965
SU 15963
SU 15947
nuchalis USNM 61201 RMNH -— RMNH 83882 MCZ 45486 NHMW 16715 RMNH -— USNM 61193
leucogaster RMNH -— RMNH -— RMNH -— CNHM 71598 RMNH -— SMNS 4585 RMNH 3994
curta MCZ 25302 MCZ 25301
septentrionalis CNHM 7141 ZSBS 169/47 CNHM 7140 CNHM 7139
déderleini MZUS ——
schlegeli ZSBS 1933/0
NHMW 16712:6
CNHM 121033 SMNS 4586
NHMW 16712:3 NHMW 16696:1 NHMW 16712:2
CNHM 121039 MHNP 4417
Locality
Luzon Negros Basilan
Mindanao
Luzon %99
Mindoro Polillo Luzon Negros
Mindanao
Negros Luzon
Mindanao
Tablas Luzon Panay
Negros
Celebes
Sumatra Borneo Sumatra Borneo Sumatra Borneo Sumatra
Celebes
China 99 >?
99
Sumatra
Borneo Sumatra Java Borneo Sumatra Java Sumatra Java Billiton
TM i) ~
40 40 40 40 40 40 10 QV QV QQ 4H WO QAAAAQAQ+O
AAW AAQ\MAQQA,
40 40 Qh 40 40 40 40
40 40 409, 40 40
%
Ay 40 QV A 40 AA 40 40
Total Length, mm.
135 149 170 179 186 186 190 191 196 199 204 210 245 258 262 263 277 281 321
123 184 195 200 249 280 294
151 162 177 195 198 205 223
195 280
168 263 277 342
288
163 189 198 255 275 283 287 317 326
43
Vol.
SOy,e fF owH
_ BoB Roo Do
o one)
CONBCNNMDWWP
DHARAAHAIWEHPWNNMNNMNMNNH 5 ° . —
Vol./TL ml./mm.
007 .013 12 O11 O11 .013 O11 .010 .015 .020 .020 .014 .020 .023 .023 .023 .022 .030 .025
coooooooococoooqoooqocoocno
oooococoo
oo oooocooo
oooco
So
oooooooooa
.008 .016 .026 .020 .016 .029 .034
.015 .006 .028 .026 .020 .020 .018
.015 .029
.021 .024 .029 .047
.038
.012 .016 .018 .024 .025 .027 .028 022 .026
Ven- trals
162 159 133 145 153 146 172 148 155 154 147 148 160 150 163 169 168 177 177
146 142 133 145 156 143 145
126 147 127 148 146 151 143
170 156
160 178 180 172
163
148 152 146 129 152 159 138 169 135
Sub- caudals
TABLE 13 (cont.)
Total Museum Length, Vol. Vol./TL Ven- Sub- number Locality Sex mm. ml. ml./mm. trals caudals griswoldi NMS -— Borneo ot 236 4 0.017 166 16 CNHM 72434 be of 334 i Bs 0.039 170 17 USNM 134114 és Q 376 9 0.024 190 14 NMS 18949 - ot 379 14 0.037 val 16 MCZ 438580 ” Q 488 18 0.037 186 13 grabowskyr CNHM 76293 — Borneo o 182 2 0.011 186 28 NHMW 16998 % oh 209 6 0.023 150 23 RMNH -— ” of 287 8 0.028 160 25 RMNH -— e ‘eh 307 i) 0.029 162 24 MCZ 43573 e ot 316 8 0.025 176 2 ZMA 10230 ‘ Q 340 10 0.029 180 Oe, NHMW 16693 ‘is Q S00 14 0.039 168 21 MHNP 89-192 ” of 362 11 0.030 181 28 MHNP 89-191 i Q 388 10 0.026 189 VAIL RMNH 568 i Q 451 28 0.062 190 20 ceramensis CNHM 83464 Ceram 2 266 ij 0.026 154 19 ZSBS 947/20 - Q 301 11 0.037 165 20 lautensis RMNH 4716B _ Kokos Ids. ee 215 6 0.028 123 24 modesta MHNP 3299 Java Q io 2 0.013 192 18 Z5BS 1935/0 Pe Q 201 3 0.015 174 21 ZSBS 2635/0 fe Q 224 33) 0.016 199 19 ZSBS 2646/0 is of 269 8 0.030 154 31 ZSBS 2646/0 « Q 269 7 0.026 166 il ZSBS 1934/0 Z ot 286 rh 0.024 iL 26 ZSBS 2646/0 a of 304 bt 0.036 154 29 ZSBS 2675/0 ‘ Q 319 9 0.028 173 18 CNHM 83164 i o 343 12 0.035 162 25 ZSBS 2675/0 is fot 352 12 0.034 171 7A bicolor SM “164” Borneo oy 164 2 0.012 167 23 NHMW 16997 :2 i of 178 3 0.017 139 Oe, MCG 748.36 4 Q 181 4 0.022 159 PAu RMNH 68 ‘ en 281 i 0.025 152, 24 RMNH 1678 a Gh 284 9 0.032 145 21 RMNH -— iJ oh Slat 10 0.032 142 26 RMNH 10542 is 2 3515) 14 0.042 156 20 SM Cd.5.26.7a ‘a Q 391 21 0.054 159 20 muelleri MCZ 25332 Celebes Q 146 il 0.007 fall 13 NHMW 16692:7 té Q 208 3 0.014 165 13 USNM 1208138 a ot 214 A.) 408021 152 18 ZMB 156385 a Q 218 4 0.018 170 18} MCZ 25322 x of 235 6.5 0.028 148 18 SNG 19443 ‘s Q 242 8 0.033 158 13 SNG 19442 fo Ct 254 8 0.031 141 23 MCZ 25317 i Q 298 8.5 02029 169 14 MCZ 25315 te 2 320 9 0.028 2 14 ZMB 13946 ce Q 355 15 0.042 167 2
TABLE 13 (cont.)
Museum number
hilleniusi ZMA 10078 SM 66 1 99
acutirostris
NHMW 187138:11 NHMW 1387138:10 NHMW 1387138:7 NHMW 1387138:9 NHMW 138718:5 NHMW 138713:3 NHMW 1387138:6 NHMW 187138:1 NHMW 1387138:4 NHMW 187138:2
albiventer CAS 14940
crassa NHMW 16994 RMNH —— RMNH 34 RMNH 34 RMNH 34 ZMB 5229 RMNH 34 RMNH 34
melanota
ZSBS 2039/0 ZSBS 2039/0 SMNS 4529 RMNH 1152 CNHM 109964
lumbricoidea NHMW 16992 :2-3 NHMW 16992 :2-3 NMS —— RMNH 932 SMNS 3497 RMNH — RMNH 4865 NMS -—— RMNH 1264 CAS 15289 ZMA 10234 RMNH 5806 NMS 1932 MCZ 20971 RMNH 5806 MCZ 43568 ZMH 2466 RMNH 5806 NMS —— NHMW 16700:1 ZMB 13110
Locality
Borneo ’)
Celebes
Singapore
Sumatra 99
Borneo bE
Malaya Java Borneo Sumatra Nias Malaya Java
Mindanao
Nias Sumatra Malaya Sumatra
Borneo Sumatra de)
Malaya Sumatra Borneo
Sex
40 40 FQ, 40 AQ, 40 QVQ,40Q, one e
+O
40 Q9 40 40 10 40 QQ,
Ay +0 40 QA,
WI AAAAAWOAAYO OW AAAAAIO A
Total Length, Vol. mm. ml. SLT 13 368 19 138 2 150 2, 173 5 193 4 291 14 296 14 330 14 341 24 370 20 447 18 200 6 131 2 168 vA 227 5 330 13 300 14 Bo4 1s B42 16 ota 19 182 4, 189 4 202 3 218 6 230 6 170 2 176 3 177 3 186 3 190 5 193 o 246 6 249 4 254 7 254 5 259 6 277 fa 294 ‘- 301 11 306 14 347 15 351 9 353 1} 367 14 379 bh 382 17
45
Vol./TL ml./mm.
oooocococoocoo oo
So
ooocococooco
ooococoe
ooococoooococooooococnhwmc So
041 .052
.013 .018 .029 021 .048 .047 .042 .070 054 .040
.026
.015 s08Z
.045
Ven- trals
Sub- caudals
18 16
TABLE 13 (cont.)
Museum number
RMNH 618 NHMW 16701:2
RMNH 588 MHNP 89-193 NHMW 16702
margaritophora ZSI 149386 MHNP 39186 NHMW 16707 RMNH 4689 RMNH — ZMB 8462 USNM 70952
bitorques
MCZ 25776 ZMB 7444 ZMB 3778 MHNP 00-364 CAS 15295 SNG 19394 MHNP 00-3865
linnaet
RMNH -—— RMNH — RMNH — RMNE =— RMNH — RMNH —— RMNH — RMNH —— RMNH 52
RMNH ——
prakkei
ZSI 13306 RMNH 4360
erseltz
NHMW 16701:1 NHMW 16703:4 NHMW 16703:1 NHMW 16703:3 CNHM 134723
NHMW 16703:2
Locality
Java Sumatra Malaya Java Malaya Java Borneo Sumatra
Java +2 4 9:3 Sumatra 99
+9
99
Phil. Ids.
Luzon b He
no data Java
Singapore
Borneo
Sumatra ”
NM f¢°) ~
40 40 40 40 40 40 40 40 Q 40 4 AAA Ay 40 Ay AQ QY A QA
40 Ay Q, 40 10 10 10 AQ +0
Qy %
40 40 40 QV Qa 40
Total Length, Vol. mm. ml. 409 21 410 10 413 16 451 18 454 22, 460 34 472 ZZ 490 23 200 4 234 8 255 6. 256 10 259 6 261 9 358 16 250 4 329 10 329 16 328 12 398 Zi. 412 26 420 24 122 Bt 171 2 Zoo 6 242 5 245 6 281 9 292 12 307 20. 336 20 396 38 204 4 243 6. 244 q 333 22 343 2A 365 24 381 28 424 42
Vol./TL ml./mm.
.051 .024 .039 .040 .048 .074 .047 .047
oocooocoooc]oa
.020 .034 .025 .039 .023 .034 .045
ooococeoo
.016 .030 .049 .037 .053 .063 .057
cococococeoe]o
.008 O17 .026 .021 .024 .032 .041 067 .060 . 100
ocooocococoooo
.020 .027
oo
.029 .066 .061 .066 .073 .099
oo°ococoo
Ven- trals
182 191 162 187 160 181 179 169
152 150 157 152 160 159 161
172 175 157 170 174 167 168
159 140 143 157 154 154 159 137 150 154
126 130
153 137 137 151 151 151
Sub- caudals
As with many other characters, the many known species of Cala- maria can be arranged in order of increasing thickness and form a continuous gradient from one extreme to the other.
Coloration.—Taken as a whole, the genus Calamaria is highly vari-
able.
46
The dorsal pattern may consist of broad or narrow stripes,
INGER AND MARX: THE SNAKE GENUS CALAMARIA 47
narrow or broad transverse bands, and spots of varying sizes. Or the dorsum may be unmarked or divided into dark upper and light lower portions. The individual scales, where they are not involved in the spots, stripes, and bands of the pattern, may be a solid color (e.g., Fig. 40), or have a fine network or speckling on a contrasting background (e.g., Fig. 33). In many species the scales of the first row have light centers and dark edges and form a continuous light stripe or a row of light dots. Dorsally the head usually has the ground color of the body, often with obscure, irregular spots. The upper lip and the underside of the head are usually light and often have dark spots on scale sutures. In a few species the head and trunk colors contrast sharply.
The ventrals of most species are yellow or whitish. The common- est pattern consists of a dark spot at the lateral corners of each ven- tral. In some species (e.g., ezseltz) the anterior ventrals are yellow whereas those farther back have a black area that becomes progres- sively larger on successive plates until it covers the entire width of the belly. In some species each ventral has a dark anterior band and a light posterior one (Fig. 56). A pattern consisting of wider dark bands on a light background occurs in two species. A checkered pattern (Fig. 13) occurs in several species.
The ventral surface of the tail usually has a continuation of the belly coloration, often with the addition of a dark mid-ventral stripe.
Some variation occurs in almost all species. But the type and extent of intraspecific variation differ from species to species. In some species, for example linnaez, both dorsal and ventral colorations vary widely (Figs. 12 and 13). Others, e.g., griswoldi (Fig. 23) do not vary at all. Although we have seen many more linnaez (659) than griswoldi (12), the differences between these two species in ex- tent of variation is not attributable to sample size because rarely do any two specimens of linnaez resemble one another as much as do all 12 of the griswoldz.
The patterns of the different parts of the body may vary in differ- ent ways. Generally, the coloration of the head varies less than the trunk. In alidae (p. 236), for example, the trunk may be striped, spotted, or banded, whereas the head varies only in the number of dark spots. By contrast, the trunk of schlegeli (835 examined) is always dark brown or black above the unspotted, yellow first two scale rows, whereas the head may be completely yellow or completely dark brown or any intermediate bicolored condition.
7 from
linnae
dorsal pattern of Calamaria
1n
.
lation
il var Specimen at bottom of middle row from Bandung, Java.
12. Individue ang, Java.
‘ Ie
Fi
Tjikad)
48
Fic. 18. Individual variation in ventral pattern of Calamaria linnaei from Tjikadjang, Java.
49
50 FIELDIANA: ZOOLOGY, VOLUME 49
Ontogenetic variation characterizes lumbricoidea (p. 82) and prob- ably bicolor (p. 153). In luwmbricoidea both head and trunk are in- volved but though darkening of the color is the age trend, the change is not exactly parallel in the two areas (Figs. 21 and 22). In bzcolor the entire dorsal surface, except for the tail, becomes darker with age.
Intraspecific variation may be common in a local population. All of the snakes (linnaet) in Figs. 12 and 13 except one are from Tji- kadjang, Java. At Bukit Tinggi, Sumatra, seven schlegeli have three of the types of head coloration listed in Table 44. Other species may show little individual or local but conspicuous geographic variation. All specimens of lumbricoidea from Borneo are uniformly dark brown above the first two scale rows; those from the Philippine Islands are striped; those from eastern Sumatra are non-striped and those from western Sumatra striped (Table 21).
Despite local and geographic intraspecific variation, coloration frequently is one of the characters differentiating similar species. Striped griswoldi, for example, is immediately distinguishable from the sympatric, non-striped population of lwmbricoidea. Even striped lumbricoidea differ from griswoldi as the light lines of the former are confined to the very edges of the mid-dorsal scales whereas in gris- woldi they cover at least the lateral fourths of each scale (Fig. 28). Furthermore, the belly of griswoldi is immaculate yellow and that of lumbricoidea is almost always banded. Another example of interspe- cific variation involves gervaist, which is non-banded, and bitorques which is. Calamaria crassa and eiseltt form another closely related pair differing in coloration (cf. Figs. 46 and 47).
Interspecific differences are not universal, however, and some spe- cies (e.g., grabowskyi, suluensis, and palavanensis) cannot be distin- guished on the basis of coloration. Because of such interspecific similarity and because of wide intraspecific variation in other cases, color differences have to be carefully analyzed in conjunction with other characters before interpretations are made. But this caveat applies to most of the characters used in this study.
Correlation of head characters.—Although a certain amount of in- dividual variation occurs in most characters, we consider a species to be variable (e.g., preocular present or absent; mental touching or not touching chin shields) for the purposes of this section only if the variation is geographic or if, as in the case of lumholtzi, the two to four specimens examined were variable.
Association of pairs of cephalic characters was tested by means of contingency tables. Chi-square values reached significant levels
INGER AND MARX: THE SNAKE GENUS CALAMARIA 51
TABLE 14.—Association of Presence of Preocular with Size of Eye in Species of Calamaria (Numbers in body of table are numbers of species).
Preocular Observed Calculated Present Present or or Eye size! Present absent Absent Total Present absent Absent Greater 16 0 0 16 12.5 0.6 2.8 Equal 19 0 3 22 17.2 0.9 4.0 Less 4 ve 6 12 9.4 0.5 Zee chi-square=20.1; n=4; P <0.001 1 “Greater,” ‘“‘equal,’’ and “‘less’’ refer to eye diameter relative to distance of
eye from mouth. Species varying from “less” to “equal’’ or from ‘‘equal’’ to “sreater’’ are listed as “equal.”
(P< 0.05) in only five tests: preocular and eye size (Table 14) ; frontal ratio and eye size (Table 15); preocular and frontal ratio (Table 16) ; preocular and relation of mental to chin shields (Table 17); number of scales surrounding paraparietal and number of supralabials (Ta- ble 18). Results of tests between all pairs of the cephalic characters are summarized in Table 19.
The association of frontal ratio, eye size, and preocular based earlier on data taken from literature (Marx and Inger, 1955), is con- firmed by the material we have since examined (Tables 14-16). These three topographically related characters tend to change as a unit and are what Cain and Harrison (1960) call ‘‘necessary corre-
TABLE 15.—Association of Frontal Ratio with Size of Eye in Species of Calamaria (Numbers in body of table are numbers of species).
Eye size Observed Calculated
—_
os =
Frontal — ratio! Greater Equal Less Total Greater Equal Less
—2 3 0 0 3 1.0 1.3 0.7 2-214 10 5 i) 16 5.1 7.0 3.8 216-21/s 3 5 3 Ly. 3.5 4.8 2.6 3-34 0 10 t 14 4.5 6.2 3.4 4— 0 2 4 6 1.9 2.6 1.4
Chi-square=27.2; n=8; P <0.001
1 Ratio of frontal width to supraocular width. Species categorized by intra- specific maxima.
52 FIELDIANA: ZOOLOGY, VOLUME 49
TABLE 16.—Association of Presence of Preocular with Frontal Ratio in Species of Calamaria (Numbers in body of table are numbers of species).
Preocular Observed Calculated Present Present Frontal or or ratio Present absent Absent Total Present absent Absent —2 3 0 0 3 Ape: 0.1 0.5 2-214 16 0 0 16 12.5 0.6 2.9 214-21/s 9 1 i) i 8.6 0.4 21.0 3-3 14 0 D 14 10.9 0.6 2.5 4— 2 1 3 6 Anat 0.2 at Total 39 2 9 39.1 ies) 9.0
Chi-square= 18.2; n=8; P=0.05
lates.” Their association prohibits the use of them as three inde- pendent estimates of similarity or divergence. Yet it would be an oversimplification to say that we are dealing with a single character, the ocular region, in taxonomic evaluation of a comparison. For as the association is not perfect, the parts of this unit have changed at different rates. A small ocular region may be one in which the pre- ocular has disappeared while the eye has remained moderate in size (e.g., javanica), one in which the eye has become small while the pre- ocular has not disappeared (e.g., helleniusz), or one in which both eye and preocular have become reduced (e.g., schlegeli cuviert). Thus despite their association, each of these characters has to be consid- ered in evaluating a comparison of two samples.
TABLE 17.—Association of Presence of Preocular with Relation of Mental to Chin Shields in Species of Calamaria
(Numbers in body of table are numbers of species).
Preocular Observed Calculated Present Present
Mental and or or chin shield Present absent Absent ‘Total Present absent Absent Separated 12 2 5 19 14.8 0.8 3.4 Separated
ortouching 2 0 2 4 Ghai Ot 0.7 Touching 25 0 2 PAE Dileewll sal 4.9
Chi-square=9.5; n=4; P=0.05
INGER AND MARX: THE SNAKE GENUS CALAMARIA 53
TABLE 18.—Association of Number of Scales Surrounding Paraparietal with Number of Supralabials in Species of Calamaria.
(Numbers in body of table are numbers of species)
Seales surrounding paraparietal
Observed Calculated Supra- és se labials 5 S5or6e 6 Total 5 5 or 6 6 4 1 0 10 Ld a 0.4 G0 5 22 2 15 39 17.9 1.6 19.5
Chi-square=9.4; n=2; P=0.01
The association of number of scales surrounding the paraparietal with number of supralabials does not have an obvious explanation. The reduction from five to four supralabials occurs in the preorbital region (Marx and Inger, 1955, p. 169) and has no apparent functional relation to an increase in number of scales at the rear of the head. Correlations of this sort, which does not involve necessary correlates, are valuable in attempts to evaluate the relations of populations as they provide additional evidence of common genotypes.
Association of the preocular character with the relation between mental and chin shields has no obvious explanation. Separation of mental and chin shields occurs proportionately oftener with the absence of a preocular than one would expect on the basis of chance alone. Contact between mental and chin shields is probably the primitive condition in this genus (p. 254) and absence of preocular advanced (p. 254). Thus the association shown in Table 17 does not result from progressive modification of both characters. Loss of the
TABLE 19.—Summary of Chi-Square Tests of Association Between Pairs of Cephalic Characters in Species of Calamaria. (Value of chi-square in upper right half of table, with degrees of freedom as superscripts. Corresponding values of P in lower left hand. Character states as in Tables 14-18)
Supra- Para-
labials Preocular Frontal Eyesize Mental parietal Supralabials 2, o.0- 14 3.92 2.1? 9.42 Preoculars S04 ; 18 .28 20.1! oe 5.64 Frontal 1022 0.05 ae 27 ..2° 13.08 13.98 Eye size >0.1 <0.001 <0.001 ai 5.94 2.64 Mental > 0.3 0.05 > 0.1 0.2 i 6.64 Paraparietal 0.01 >0.2 0.09 >0.5 a O
54 FIELDIANA: ZOOLOGY, VOLUME 49
preocular is correlated with general reduction in the ocular region. The other characters associated with the preocular reduction of the ocular region, namely eye size and frontal ratio, are not associated with the chin character (Table 19). Consequently the association of preocular and mental appears not to have a functional basis. As these characters are not necessary correlates, both may be used as independent estimates of similarity or divergence of populations.
KEY TO FORMS OF CALAMARIA
1. One sipralabial enter? Or0b ts on «ac 2s Oead sdah ers wees lowi lowi (p. 222) Two supralabials entering orbit.....0.0. «620964 s 64o08es44.ndae nine aes 2 2. Second and: third supralabials entering orbit) 3.252444. eas Se eee 3 Third and fourth supralabials entering orbit........ 20.4.6. .5-s0.@9s400- ee 16 3: dP reocular absent...«cctie. DURE By lees ek aes 1 es a eee 4 Preocular PVESENS 6s: 65.40 dices desu Woe ih iste wd oe ase ee a oe ae 9 4. Mental not touching anterior chin shields... ... .¢..24.. 0.0. sheuuinls oe oe 5 Mental touching anterior chin shields... 23.0 .22.50. 3. 42) nae oe eee 7 5. Supraocular and postocular fused into a single shield...... gracillima (p. 229) Supraocular distinct from postocular .. 4.....<4.0hed«..4 06008 2s ee eee 6
6. Frontal 5 to 6 times width of supraocular; nasal larger than postocular; maxil- lary teeth unmodified (fig. 2 top)..................0.0.. schmidti (p. 74)
Frontal 214 to 4 times width of supraocular; nasal equal to, or smaller than postocular; maxillary teeth modified (fig. 2 bottom). . .lowi gimletti (p. 226)
7. Eye much smaller than eye-mouth distance; posterior chin shields meeting in
MIGHNE Sc 6 ic ied pow dere oa ae iS RSG OE ES BE ORE OO Eee 8
Eye equal to eye-mouth distance; posterior chin shields not meeting in mid-
DDN Oh eae gentss dato. See acd a: ani o Oe Sep imes: eas date tea javanica (p. 209)
S.. Posterior ventrals dark browite...% <an543. eee eee lowi lowi (p. 222) Posterior ventrals cream: COlOTeEd | .j5.a4 u00). os me aieen oe lowt wermuthi (p. 225)
9. Two infralabials touching anterior chin shields........... borneensis (p. 194) Three infralabials touching anterior chin shields... ......2...4. 49335 10
10. Mental touching anterior chin shields... . «...042.-t54an0.000.0 eee 11 Mental not touching anterior chin shields... 0.425. .5.:05 1.) pee 15
11. Each ventral with a dark anterior half and a light posterior half......... 12 Ventrals never transversely divided into light and dark halves........... 13
12. Narrow longitudinal stripes mid-dorsally; tail tapering abruptly at end. battersbyz (p. 208) No longitudinal stripes mid-dorsally; tail tapering in posterior half. melanota (p. 198) 13. Length of parietal greater than distance from rostral to posterior tip of frontal. buchi (p. 211) Length of parietal less than distance from rostral to posterior tip of frontal. .14 14. Ventrals with black rectangular markings; or if not, lateral corners of succes- sive ventrals not identical in color (fig. 18)............. linnaei (p. 200) Lateral corners of every ventral identical, dark; dark mid-ventral pigment, if present, confined to a very few dots irregularly scattered. pavimentata (p. 212)
15.
16,
i
18.
19.
20.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 55
Tail as thick as body, not tapering, blunt (fig. 10A).. .septentrionalis (p. 218)
Tail not as thick as body, tapered, pointed............ pavimentata (p. 212) PreOCUl ar BOSC ae Shred ee he on ata eae tems Poe eee Eames 17 Preotular present. oi nx xad 6 655-64. RGAE aH EdRS RSS HA HONE AAR Ow 23
A continuous dark mid-dorsal stripe occupying scales in vertebral row. mecheli (p. 233)
Without continuous dark vertebral stripe......................0...0.. 18 Each ventral with a dark anterior half and a light posterior half or completely
CATS ice bode Ab haek we OSES i ae oe cen OSes Oe apraeocularis (p. 287) Ventrals cream-colored centrally or with a few irregularly scattered dark
SOG ss hah 6 cd we EA A ee WA Ge eee be ee a ea ee eee 19 A light longitudinal stripe on second and third scale rows, bordered below by
a.dark stripe On first TOW «co. 5 e046 sesh bbs ad SSPE SHOES lumholtzt (p. 148) Stripes, if present, Not as ADOVE. «6 ios ce cee vewavevs neues tuavesca cues 20 Tail thick, not tapered, end broadly rounded (fig. 14A)....... alidae (p. 235) Tail tapered at least in posterior fourth (fig. 14B).............00..0.0... va |
B
Fic. 14 Ventral views of tails in males of Calamaria alidae (A) and C. schle-
geli (B).
ZA.
22.
23.
24.
Most of dorsal scales dark brown with a light network, or yellowish with a
dark network S41 f1G733) oc a0 ierda dod nn cad OBE eae a eww hone aie ee bars 22 Each dorsal scale above the second row dark brown without a light network
ATO ich eat tac, gms SRN ac Pe ha Sade a nse 8 me schlegeli (p. 154) A narrow, light. nuchal collar.....:.......22.2h.e0e..: ceramensis (p. 232) Nomuchal*collairenn nes oe, wate ae eee rebentischi (p. 230) Mental not touching anterior chin shields........................00040. 24 Mental touching anterior chin shields.................. 00.0000 cee eau 42 Maxillary teeth unmodified: (fig. 2 top). ...¢..0..0000002.086 sees ceded. 29
Maxillary teeth modified (fig: 2 bottom) ........... 6.0.00 000ae ce eeeees Zi
30.
31.
FIELDIANA: ZOOLOGY, VOLUME 49
Six shields and scales surrounding paraparietal (as in fig. 1 bottom). lautensis (p. 70)
Five shields and scales surrounding paraparietal (as in fig. 1 top)........26 No dark pigment On venivals: <:.24.22244.2 ieee aoe leucogaster (p. 68) Dark pigment on outer edges of ventrals.................4.. ulmert (p. 68)
Dorsum with wide, dark, transverse crossbands four or more dorsal scales wide. bicolor (p. 149)
Dorsum lacking crossbands, or if present, one or two scale rows wide...... 28
Ventral coloration as dark as dorsal, the two areas separated by a white stripe. lateralis (p. 146) Coloration of dorsum different from that of ventrum................... 29
A single, light continuous stripe on second and third scale rows. lumholtzi (p. 148)
If stripe is present, not occupying above position on lateral scale rows... .30 Five scales and shields surrounding paraparietal (as in fig. 1 top)........ ol Six scales and shields surrounding paraparietal (as in fig. 1 bottom)...... 37 All ventrals behind first fourth of body with dark pigmentation at least on
their lateral tis .s s,s. 20% ase en we oes Se Re eo eee 33 Almost all ventrals without pigmentation. 6 iad waco aca nee eee 32 Prefrontal not touching third supralabial................. schlegeli (p. 154) Prefrontal touching third supralabial...................... bicolor (p. 149)
Reduction to four dorsal scale rows on tail more than 13 subcaudals from tip. everetti (p. 183)
Reduction to four dorsal scale rows on tail 18 or fewer subcaudals from tip. .34 Oblique black stripe running from eye to corner of mouth .brongersmaz (p. 118)
No oblique black stripe running back of éye... 2...) ae © ee 35 Diameter of eye longer than ventral border of prefrontal .palavanensis (p. 134) Diameter of eye not longer than ventral border of prefrontal. ........... 36 art DIONE (hes LOAN .2 4 acon. oe ni ae els = HES ee virgulata (p. 186) Mail Taperine& to: a. pomt (ie. 1B) nese ee taeend aa, ee modesta (p. 138) Dorsum with narrow dark crossbands.. ..a.. ci... .<s. sec. ee. eee 38 Dorsum without narrow dark crossbands;’. ......<.«.. so. oh .a eee eee 39 Hye less than: eye-mouthidistance. .. so. tcc. av aicanace: déderleini (p. 177) Eye 1% times eye-mouth distance.....................-. boesemani (p. 171) Eye 11% times eye-mouth distance..................... boesemani (p. 171) Eye, at its largest, slightly greater than eye-mouth distance............. 40
Fic. 15. Ends of tails of Calamaria virgulata (A) and C. modesta (B).
49,
50.
D1.
58.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 57
Tail ending bluntly ie. LOA) wis 6-cs Gener cee oe daa ceeds virgulata (p. 186) Tail tapering to a point (figs. 14B, 15B) .¢o<.iecéssaeeee beens ae bene Bes 41
. Ventrals with dark pigment at least at lateral tips.......... modesta (p. 138) Ventrals without dark pigment.......................... schlegeli (p. 154) Maxillary teeth unmodified (fig. 2 top). ....... 000000000 0.00000...00.. 43 Maxillary teeth modified (fig. 2 bottom)..........0.000...0..0....00.. 44
. Posterior chin shields meeting in midline..................... curta (p. 71) Posterior chin shields not meeting in midline............. acutirostris (p. 73)
. Paraparietal surrounded by 6 shields and scales (as in fig. 1 bottom)...... 45 Paraparietal surrounded by 5 shields and scales (as in fig. 1 top)......... 53
& Ventrals: 176-00 MOPe..<3sa<.4e8 Shao dp ehore seed need ok Oho sea 4bot Son 46 Ventrals 175: OF 168s ¢ pea wha awe heed GER eek OS SPE ES oe BAe bs 47
. No dark transverse bands behind head.................. virgulata (p. 186) At least 2 transverse bands behind head................... forcarti (p. 184)
. Dark pigment of ventrals restricted to lateral corners................... 48 Ventrals in posterior half of body with dark pigment across entire width... .51
. A continuous dark vertebral stripe formed by spots on each vertebral scale Nie. Olson G4 heed a 2 ed dade etme Aes thane eee es sumatrana (p. 181)
If vertebral spots present, not forming continuous stripe for entire length of DOG Vcc or buen ah be oe eee ey eee oo hd hee he gees at oak Be Gee 49 Subcaudals of males 24 or more, of females 14............. abstrusa (p. 179) Subcaudals of males 20 or less, of females 11 or less.................... 50 Dark spots on dorsal scales forming short lines 2 to 5 scales long (fig. 44); frontal shorter than parietal................00... margaritophora (p. 166) Dark spots on dorsal scales not forming lines; frontal equal to or longer than WaRICT Als a\s.04 a8 ae xi deo 48 a eA Oe Srh oh oh py cae Ae» a nuchalis (p. 170)
Underside of tail with dark crossbands more than one subcaudal wide. eiselti (p. 175) Underside of tail without crossbands, or if present, less than one subcaudal
WUC a cs 3 ace. 2 Bee Sue lave se te Rte O49 FAG aed edt ape ne & Coa ndee cok a eas eae x 52 Sides of neck with triangular extensions of light ventral coloration; diameter of eye greater than eye-mouth distance.................. crassa (p. 173) Sides of neck without triangular extensions of ventral coloration; diameter of eye equal to or smaller than eye-mouth distance...... virgulata (p. 186)
. Two light stripes on each side of body (fig. 24)............. albiventer (p. 94) Not colored as above... 0... cece nee ence nner eeeeees 54 Four or five light stripes running length of body on each side (fig. 23)... .. 55 No light stripes above first scale row... 0.0... ee eee 56
. Belly immaculate yellow. .......... 0.2.00 0 ete eee aes griswoldi (p. 92) Belly with dark crossbands or almost completely black. .lumbricoidea (p. 75) Dorsal scales above third row solid dark color without network (fig. 25)... .57 Most of dorsal scales above third row dark with light network or light with dark Heb WwOrk (HG io Nan ttis. aea ssc sche k ode e area areas ee eee wee ae a 58
. Eye small, its diameter about two-thirds eye-mouth distance (fig. 25).
hilleniusi (p. 96) Eye larger, at its smallest slightly less than eye-mouth distance. lumbricoidea (p. 75) Two to six dark, black-edged, wide crossbands behind head (fig. 27). bitorques (p. 104) No stich crossbandsy (t4.c0c foe ee Se ae es bs 59
58
64.
Or
~
10.
fa
FIELDIANA: ZOOLOGY, VOLUME 49
59. Scales of first row with yellow or white centers forming a stripe or row of light
dots running length of body... .O24. dseG os 4 ee eee 60 No row of light dots or light stripe on first scale row................... 64 A continuous or narrowly interrupted light nuchal collar................ 61 No light nuchal Collar. 326. <iv. S22 Be sees widest noc oe eee 62 Subcaudals in males more than 30, in females more than 20. -prakket (p. 120) Subcaudals in males less than 25, in females less than 20..... gervaist (p. 106) Light centers of scales in first row forming row of light dots. . .gervaisi (p. 106) Light centers of scales in first row forming uninterrupted stripe.......... 63
Ventrals in males 150 or more, in females usually more than 164. grabowskyi (p. 00)
Ventrals in males less than 140, in females usually less than 164.
suluensis (p. 123) Eye small, its diameter about two-thirds eye-mouth distance.
joloensis (p. 102) Eye larger, at its smallest slightly less than eye-mouth distance..........65 Head pointed, distinctly tapered in front of eye (fig. 26)..... muelleri (p. 99)
Head rounded, not conspicuously tapered in front of eyes (fig. 28). gervaisi (p. 106)
KEY TO SPECIES OF CALAMARIA FROM BORNEO
Third or second and third supralabials entering orbit................... 2 Third and fourth supralabials entering orbit.4. e.2 24.0 a see f( Preocular ADSEN Gas. c+ wie ores oes ov oe an Oe eS Re ee eee 3 Preociwlar presents ly 8 cg nD Bee Mi Oe &,. eee ee 5 Supraocular and postocular fused into a single shield.... . gracillima (p. 229) Supraocular distinct from postoculary. 6). 8 a ee ae eee 4 Mental touching anterior chin shields....................... lowi (p. 221) Mental not touching anterior chin shields................. schmidti (p. 74) . Two infralabials touching anterior chin shield............ borneensis (p. 194) Three infralabials touching anterior chin shield. ......................-. 6
Narrow longitudinal stripes mid-dorsally; tail short, tapering abruptly at end. battersbyz (p. 208)
No longitudinal stripes mid-dorsally; tail longer, tapering in posterior half. melanota (p. 198)
A light longitudinal stripe on second and third scale rows bordered below by a
darks Stripe-On first row: 22.25. cece ns ee ee lumholtzi (p. 148) Stripes, if present, not as above: .....2...5.550 081.6008 eee 8 Preocular- absent’ ca). aie noice bdo ails eta: ee ee 9 Preocular, presemtt:.672 23 cst ooi fens oe Go aecs oe eee 10 Most of dorsal scales dark brown with a light network or yellowish with a
ON (0) en ae en ee 28, ken, rebentischi (p. 230)
Each dorsal scale above the second row dark brown without a light network.
schlegeli (p. 154) Mental not touching the anterior chin shields Mental touching anterior chin shields.........................-.-2. 18
Dorsum with wide dark transverse crossbands four or more scale rows wide. bicolor (p. 149) Dotsum lacking crossbands), 208.0... 005 8). Se eee 12
i2.
13.
14,
15.
16.
17.
18,
19),
20.
21.
22.
10.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 59
Ventral and dorsal coloration dark brown with a bold continuous lateral white BOC t os abe ocr heen 4 aa oe a eure) BES lateralis (p. 146) Ventral coloration different from dorsal coloration..................... 13 Almost all ventrals without dark pigmentation........................ 14 All ventrals behind first fourth of body with dark pigmentation at least on GHECI TACeCKAl G06. « i dhe a-a-e any Sack Kw DMA sO nw ee bh eda aekas whee Ban 16 Eye larger than eye-mouth distance..................... leucogaster (p. 63) Eye smaller than eye-mouth distance..............000 0000.0... 00000.4. 15 Prefrontal not touching third supralabial................. schlegeli (p. 154) Prefrontal usually touching third supralabial............... bicolor (p. 149) Reduction to four dorsal scale rows on tail 14 or more subcaudals from tip. everetti (p. 133) Reduction to four dorsal scale rows on tail 13 or fewer subcaudals from tip. .17 Tail ending in a blunt tip (fig. 15A)..............000000.. virgulata (p. 186) Tail tapering 10 2 POMt.< \ 6646404 c ce mneee Seeevenss beatae modesta (p. 138) Belly with dark crossbands, each more than one ventral wide. lumbricoidea (p. 75) Belly not cross-banded or with bands narrower than one ventral......... 19 Eye two-thirds eye-mouth distance...................... hilleniusi (p. 96) Eye equal to or greater than eye-mouth distance....................... 20 Dorsum with longitudinal dark and light stripes........... griswoldi (p. 92) Mid-dorsal region not striped............. 0.000000 ccc eee ee, 21 Tail thick, tapering abruptly near end toablunttip......... prakkei (p. 120) Tailstapering cradually . cc as eaacade eka deacdvaddeak Pe eho aah be DEO ea RS: 22 Ventrals of males under 145, of females usually less than 164. suluensis (p. 123) Ventrals of males 145 or more, of females usually more than 164. grabowskyi (p. 129) KEY TO SPECIES OF CALAMARIA FROM SUMATRA Peeocular absent: 20cs 65.4445 6%8 5004 op ehas Sd bein Os 66d sede PaDE AL ae 2 Preocular present. << 6.454 ose anv eb eR ae bY eROD EEDA Mew EEK ORE ORES DEEES:A 4 Dark stripe on vertebralrow..............0... 00.00.0000. mecheli (p. 233) Without dark stripe on vertebral row...........00.00 0000000000000 000. 3 Wentrals, 199 OF MOle 0's 3 nade ooh Sot h dee bowen Pars ow eRe ak alidae (p. 235) Ventrals, 150 OF Les 8 ice acd-snn alle Re ati Ree PR reeds lo ecw none schlegeli (p. 154) Mental not touching anterior chin shields............................. 5 Mental touching anterior chin shields...............0...00 00.00.0000. 10 Dorsum with narrow dark crossbands................... déderleini (p. 177) Dorsum without erossbands.. ... ¢. e644 <c.44 3858s. ER Sea boli 144 6 Ventrals without dark pigment on lateral edges........................ 7 Ventrals with dark pigment at least on lateral edges.................... 8 . Eye smaller than eye-mouth distance..................... schlegeli (p. 154) Eye greater than eye-mouth distance.................... leucogaster (p. 63) . Unmodified maxillary teeth (fig.2 top).................00... ulmeri (p. 68) Modified maxillary teeth (fig. 2 bottom)..............0.... 0.000000 ee 9 s Tail thick and blunt (ig. 15A) o37 00 Men Se ste eee virgulata (p. 186) ail ¢apering toa, pomidiles 5B ind eno 2 PO So modesta (p. 138) Paraparietal surrounded by 6 scales and shields (as in fig. 1 bottom)....... 11
Paraparietal surrounded by 5 scales and shields (as in fig. 1 top)...........16
bo
FIELDIANA: ZOOLOGY, VOLUME 49
Ventrale 1 7G-Or nore. PI eee eee forcarti (p. 184) Ventals 175 or less... ..esieds. ce be ded 14 bh ods oR ea os ee ee 12 Ventrals in posterior half of body with dark pigment across entire width... .18 Ventrals at most with dark pigment restricted to lateral corners........... 14 Waderside or tail cross-panded cae ac ee ee eee sea eiselti (p. 175) Underside of tail not-erdss-banded ss o:5.o:s aihurdiaw o-4 Saws are Lee crassa (p. 173) A continuous dark vertebral stripe formed by spots on each vertebral scale.
sumatrana (p. 181)
If ‘vertebral stripe present, interrupted: 0. ).-:4 25.0024. 06 i eee oe 15 . Subcaudals of males 24 or more, of females 14............. abstrusa (p. 179) Subcaudals of males 17 or fewer, of females 11 or fewer. margaritophora (p. 166) . “BE Wo pare OF HONG S06 ¢ o.c5, ns 2A 10+ oe Odo whe cS albiventer (p. 94) No stripes, or more than two pairs of light stripes........ lumbricoidea (p. 75)
KEY TO SPECIES OF CALAMARIA FROM ASIATIC MAINLAND
Third and fourth supralabials enter: orbit. .7o.0::.24u se ee ee eee 2 Second.and third supralabials enter orbit... o.seesecda aol vee 5 Mental not touching anterior chin shields................ schlegeli (p. 154) Mental. touching anterior.chin. sulelds ic: quake nctlocetiweid sedi os aeons 3 Bellywith dark:icrossbands:~ .. dvctiueul, a. cea 28 Se lumbricoidea (p. 75) Belly yathout crosshands. vidoes Hawa. aun. erect oe 4 Light dorsolateral stripe present (fig. 24)................. albiventer (p. 94) No light dorsolateralistrines t23e¢ cee ae oe ean ae prakkei (p. 120) Preocular absent. 42) s3 Aes we ee eee lowi (p. 221) PECOCUIAT PRCSOO Gs 4.5544 kas 6 OARS aRaa KE OD ROMO ae le eee 6 Ventrals vellow, immaculate. 4 i-0...¢5 2466. Pea Ps hd he buchi (p. 211) Ventrals yellow; with dark lateral tips... 205.0 acoe) 4ea5 34 +o eee eee if . Tail as thick as body, not tapering, end broadly rounded (fig. 10A).
septentrionalis (p. 218) Tail not thick as body, tapered, pointed............... pavimentata (p. 212)
KEY TO SPECIES OF CALAMARIA FROM JAVA
; Second and third supralabials entering orbit... .....0 20.64 oa. eee 2 Third and fourth supralabials entering orbit..; a; <..i4.5< cs. a0 tae eee 4 Preocularpresent:viosn... sh awars nen acento cae See linnaei (p. 200) Preoclarawsent i. 6-142 Sua das oS ee ees ie ens eee 3 Posterior chin shields meet in midline............... lowt wermuthi (p. 225) Posterior chin shields do not meet in midline............. javanica (p. 209) Mental not touching.anterior chin shields, +. .a2a).2 2.415 6205 eee 5 Mental touching anterior chin shields.................. lumbricoidea (p. 75) Freocular absent; a2 «wc vaste. ous ole nde ee schlegeli (p. 154) Preocuilar Presents softeg: icsch cco eet ests, Ripe okie cee 6 Five scales and shields surrounding paraparietal (as in fig. 1 top)........ 7 Six scales and shields surrounding paraparietal (as in fig. 1 bottom)...... 9 Ventral coloration as dark as dorsal, the two areas separated by a white stripe.
lateralis (p. 146) Ventral coloration different from dorsal coloration
CoO
INGER AND MARX: THE SNAKE GENUS CALAMARIA 61
Eye greater than eye-mouth distance................... leucogaster (p. 63) Eye less than eye-mouth distance......................... bicolor (p. 149) Tail tek and: blunt (ie. TOA) | i 20: 60s aad 20480% Ne baaws virgulata (p. 186) Tail tapering to a point (fig: 15B) . 2... 06 cca ee denne eden modesta (p. 138)
KEY TO SPECIES OF CALAMARIA FROM CELEBES AND
MOLUCCAS Preocular Absent, poate te ose dew eka se dikes hd OS ee kona eee Ree Adhd 9 Preocular DECsOUl.. 4.4 a4 4-45 osad Bak che bab Receacaa ig eee oo PG be alee ck es Z Mental not touching anterior chin shields.........................00.. 3 Mental touching anterior chin shields.................... 00.0.0... 000. 5) Paraparietal surrounded by six scales and shields (as in fig. 1 bottom)...... 4
Paraparietal surrounded by five scales and shields (as in fig. 1 top). brongersmai (p. 118)
. Eye almost 11% times eye-mouth distance............... boesemani (p. 171)
Hye, at its largest, slightly greater than eye-mouth distance .virgulata (p. 186)
First gular touching anterior chin shields................ acutirostris (p. 73) First gular not touching anterior chin shields.......................... 6
. Paraparietal surrounded by six scales and shields (as in fig. 1 bottom).... 7
Paraparietal surrounded by five scales and shields (as in fig. 1 top). muelleri (p. 99)
Tail thick, tapering abruptly at end (fig. 15A)............. virgulata (p. 186) Tail tapering gradually from base............000 00000000... eee eee 8 . Ventrals yellow, immaculate except for dark lateral tips... .. nuchalis (p. 170) Ventrals dark, yellowish on posterior edges only............... curta (p. 71) Mental touching anterior chin shields................ apraeocularis (p. 237) Mental not touching anterior chin shields............... ceramensis (p. 282)
KEY TO SPECIES OF CALAMARIA FROM PHILIPPINE ISLANDS
1.
Mental not touching anterior chin shields..........................00. Z Mental touching anterior chin shields.............................4.. 3 Subcaudals of males 21 or less, of females 14 or less........ virgulata (p. 186) Subcaudals of males 23 or more, of females 16 or more. . .palavanensis (p. 134) Belly with dark crossbands......... 00.6 s0440ue00ca0s lumbricoidea (p. 75) Belly awithouterossbands vctsousc eke dare nee Pewee de eee Redwa deen eee eed 4
Two to six dark, black-edged crossbands behind head (fig. 27). bitorques (p. 104)
No black-edged, dark crossbands behind head......................... 5
Eye smaller than eye-mouth distance.................... joloensis (p. 102) Eye equal to or greater than eye-mouth distance........... gervaisi (p. 106)
SPECIES ACCOUNTS
Calamaria leucogaster Bleeker. Figure 16.
Calamaria leucogaster Bleeker, 1860, Nat. Tijds. Ned. Indie., 21, p. 293—-Ampat Lawang, Sumatra; Gtinther, 1865, Ann. Mag. Nat. Hist., (3), 15, p. 89; Edeling, 1870, Nat. Tijds. Ned. Indie, 31, p. 381; Boulenger, 1894, Cat. Snakes Brit. Mus., 2, p. 341; Bartlett, 1895, Sarawak Note Book, no. 1, p. 84; Baumann, 19138, Zool. Jahrb., (Syst.), 34, p. 271; Mertens, 1924, Zool. Anz., 60, p. 159; Jong, 1928, Treubia, 10, p. 147; Werner, 1929, Zool. Jahrb., (Syst), 57, p. 172; Angel, 1941, Bull. Mus. Nat. Hist. Nat., (2), 13, p. 412; de Haas, 1950, Treubia, 20, p. 569; Marx and Inger, 1955, Fieldi- ana, Zool., 37, p. 179.
Calamaria arcticeps Giinther, 1866, Ann. Mag. Nat. Hist., (3), 18, p. 25, pl. 4, fig. C—Borneo; Mocquard, 1890, Nouv. Arch. Mus. Paris, (3), 2, p. 120.
Calamaria beccari Peters, 1872, Ann. Mus. Genova, 3, p. 34—Sarawak; Moc- quard, 1890, loc. cit., p. 120; Boulenger, 1894, op. cit., p. 343; Werner, 1929, loc. cit., p. 172; de Haas, 1950, loc. cit., p. 564; Marx and Inger, 1955, loc. cit., p. 200.
Calamaria brookit Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 15, p. 329— Matang, Sarawak; Boulenger, 1894, op. cit., 3, p. 647; Bartlett, 1895, op. cit., p. 85; 1896, ibid., no. 2, p. 116; Werner, 1929, loc. cit., p. 172; de Haas, 1950, loc. cit., p. 565.
Calamaria smithi Dunn, 1923, Jour. Malayan Asiat. Soc., 1, p. 3—Long Mu- jan, Sarawak; Werner, 1929, loc. cit., p. 172; de Haas, 1950, loc. cit., p. 574; Marx and Inger, 1955, loc. cit., p. 206.
Calamaria leucocephala (non Duméril and Bibron), Boettger, 1898, Kat. Rept. Samm. Senck. Naturf. Ges., p. 84.
Calamaria bicolor (non Duméril and Bibron), Vogt, 1925, Zool. Anz., 62, p. 64.
Syntypes.—British Museum (Natural History) 63.12.11.141; Rijksmuseum van Natuurlijke Historie 3994.
Taxonomic notes.—The holotypes of arcticeps, beccari, brooki, and smithi are identical to the syntypes of lewcogaster except that the last are unstriped.
Giinther’s description (1866) did not mention the oculars of art?- ceps. However, Mr. J. C. Battersby has kindly examined the holo- type at our request and informs us that a preocular is present.
63
64 FIELDIANA: ZOOLOGY, VOLUME 49
Peters (1872) was in error in reporting that the holotype of beccarz had 2 postoculars. It is a typical Bornean lewcogaster with a single postocular.
Diagnosis.—Maxillary teeth unmodified (Fig. 2, top); third and fourth supralabials enter orbit; mental not touching anterior chin shields; paraparietal surrounded by 5 shields and scales; no dark pigment on ventrals.
Description.—Rostral usually higher than wide, portion visible from above less than 1% length of prefrontal suture; prefrontal sub- equal to frontal, touching first 2 supralabials; frontal hexagonal, 134 to 41% times width of supraocular, about 34 length of parietal; parietal 114 times length of prefrontal; paraparietal usually (19 of 23) surrounded by 5 shields and scales; nasal smaller than postocular; preocular present; neither ocular as high as eye; eye greater than eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, second next largest, other 3 subequal; mental tri- angular, separated from anterior chin shields in all but 2 specimens; 5 infralabials, first 3 touching anterior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.006-—0.028 (7 specimens); tail tapering near tip to a sharp point; dorsal scales reduce to 4 rows on tail oppo- site third to sixteenth subcaudal anterior to terminal scute.
Hemipenis forked opposite fourth subeaudal, retractor muscle beginning opposite tenth subcaudal, sulcus bifurcate, calyces smooth (2 specimens). Cloaca of female bilobed (2 specimens).
Seven to 8 unmodified maxillary teeth (8 specimens).
Ventrals: males, 126-146 (mean 125.4; N=12); females, 129-157 (mean 145.6; N=21). Subcaudals: males, 17-26 (mean 21.2; N=14); females, 12-19 (mean 15.4; N=22).
Total length: males, 75-219 mm.; females, 82-223 mm. Ratio of tail to total length: males, 0.085—-0.135 (mean 0.106; N=12); females, 0.056-0.092 (mean 0.071; N=18).
Ground color dark brown, dorsal scales usually with a light or dark network; body with or without dark longitudinal stripes; the striped pattern consisting of a narrow dark stripe along the edges of the vertebral and paravertebral scale rows, a wider stripe on adja- cent halves of the fourth and fifth scale rows, a stripe along adjacent halves of the second and third scale rows, and a thin dark line along the upper edge of the first scale row; lower two-thirds of the first scale
INGER AND MARX: THE SNAKE GENUS CALAMARIA 65
Fic. 16. Calamaria leucogaster (CNHM 67279).
row cream-colored; a black collar 2 to 5 scales wide, separated from parietals by 1 to 3 scales and sometimes bordered posteriorly by a narrow yellow ring; a dark ring around body at root of tail, usually a second black ring near tip; head dark brown middorsally with a light streak running through the center of the prefrontal, the lateral edge of the supraocular, and the lateral portion of the parietal; a dark longitudinal stripe along adjacent portions of prefrontals and supralabials, continuing behind eye along adjacent edges of parietal and supralabial; remainder of upper lip and underside of head cream- colored; ventrals and subcaudals cream-colored; tail with a mid- ventral dark stripe in some individuals.
Indwidual variation.—Only one specimen (SNG 19410) has a dark ventral surface. One (RMNH unnumbered, from Upper Mahakam) has the lateral edges of the ventrals dark.
Two specimens (CNHM 67279, MCZ 43576) have the mental meeting the anterior chin shields at a point. The snake from Upper Mahakam has the mental in broad contact with the anterior chin shields.
One specimen (MHNP 389-188) has the preocular and prefrontal fused on the left side.
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67
68 FIELDIANA: ZOOLOGY, VOLUME 49
Geographic variation.—The striped pattern (Fig. 16) is character- istic of most of the specimens from Borneo, although in 8 the stripes are faint and in 4 they are absent. All Sumatran specimens are un- striped. The 2 presumed to be from Java (see below) have faint stripes. No other geographic variation (Table 20) was found.
Distribution.—Sumatra and Borneo (Fig. 17).
The only specimens reported from Java (MHNP 39-188 and 39- 189) were obtained by Boudart. Other specimens Boudart reportedly obtained at Batavia (see p. 170) probably were not collected there. This, in turn, casts doubt on the authenticity of the Javanese local- ity for leuwcogaster.
SUMATRA (RMNH 8 unnumbered; ZMB 6298, 31408, 33610). Sumatera Utara: Deli (SNG 19410; ZMA 10077). Sumatera Sela- tan: Ampat Lawang (BM 63.12.11.141-syntype; RMNH 3994-syn- type).
SARAWAK (BM 1908.5.28.69; MCG 30456-holotype of beccarz). Kindi District (BM 1911.1.80.25). First Division: Matang (BM 95.2.28.30-holotype of brookiz; SM unnumbered), Kuching (SM un- numbered), Santubong (CNHM 71598-99), Bidi, near Bau (SM unnumbered). Second Division: Lubok Antu (SM unnumbered). Third Division: Long Mujan (BM 1922.11.24.5-holotype of smith). Fourth Division: Niah (CNHM 129004, 131601-04). Fifth Divi- sion: Lawas (CNHM 67279).
NORTH BORNEO. Kota Belud District: Kiau, 915 meters (NMS 320), Mount Kina Balu (BM 94.6.30.58; MCZ 43575-76). Labuan District: Labuan (BM 94.6.30.52).
INDONESIAN BORNEO: Long Petah (ZMA 10110); Upper Maha- kam (RMNH unnumbered).
BORNEO (BM 68.1.27.22; SM 3 unnumbered; ZMB 8012); central Borneo (SMNS 4585).
? Java. Jakarta (MNHP 39-188, 39-189).
Specimens examined.—A42.
Calamaria ulmeri Sackett. Figure 18.
Calamaria ulmeri Sackett, 1940, Not. Nat., no. 41, p. 2—northwest of Blang- kedjeren, Atjeh Province, Sumatra; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 206.
Holotype.—Academy of Natural Sciences of Philadelphia 21535.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 69
Diagnosis.—Maxillary teeth unmodified; third and fourth supra- labials entering orbit; mental not touching anterior chin shields; paraparietal surrounded by 5 shields and scales; lateral corners of ventrals with dark pigment.
Fic. 18. Holotype of Calamaria ulmert.
Description.—Rostral wider than high, portion visible from above 28 length of prefrontal suture; prefrontal shorter than frontal, touch- ing first 2 supralabials; frontal hexagonal, 214 times width of supra- ocular, about 34 length of parietal; parietal 114 times length of prefrontal; paraparietal surrounded by 5 shields and scales; nasal smaller than postocular; preocular present; neither ocular as high as eye; eye twice the eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 3 subequal and slightly larger than fourth; mental triangular, not touching anterior chin shields; 5 infralabials, first 8 touching anterior chin shields; both
70 FIELDIANA: ZOOLOGY, VOLUME 49
pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Tail incomplete but tapering gradually from base; reduction to 4 dorsal scale rows, if it occurs at all, must take place near the end of the tail. Six scale rows are present opposite the twenty-third sub- caudal of this incomplete tail.
Ten unmodified maxillary teeth (1 specimen).
Ventrals 186 in female; subcaudals 23+ in female. Snout-vent length, 284 mm.; incomplete tail, 28 mm.
Body and tail brownish above, each scale with a dark network and a dark central spot; these spots unite to form dark stripes run- ning the length of body and tail; head brown above with small dark spots; upper halves of supralabials dark, lower halves yellowish with dark sutures; underside of head yellowish with small dark spots; lat- eral corners of ventrals and subcaudals dark brown, otherwise entire ventral surface immaculate yellow.
Distribution.—Northern Sumatra (Fig. 17).
SUMATRA. Sumatera Utara: Atjeh Province, about 40 km. north- west of Blangkedjeren, 2080 meters altitude (ANSP 21535-holotype).
Calamaria lautensis de Rooij
Calamaria lautensis (part) de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 1638, fig. 66—Pulau Si Laut, Cocos Islands; Werner, 1929, Zool. Jahrb., (Syst)., 57, p. 172; de Haas, 1950, Treubia, 20, p. 569; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 203. Lectotype.-—Rijksmuseum van Natuurlijke Historie 4716A, here designated.
Taxonomic notes.—As the Amsterdam Museum syntype (ZMA 10111) from Cocos Islands is damaged, we are selecting one of the Rijksmuseum syntypes as the lectotype. See page 139.
Diagnosis.—Maxillary teeth unmodified; third and fourth supra- labials entering orbit; preocular present; mental not touching ante- rior chin shields; paraparietal surrounded by 6 shields and scales.
Description.—Rostral slightly wider than high, portion visible from above ¥% length of prefrontal suture; prefrontal slightly shorter than frontal, touching first 2 supralabials; frontal hexagonal, 214 to 21% times width of supraocular, about 3/; length of parietal; parietal 1% to 1% times length of prefrontal; paraparietal surrounded by 6 shields and scales; nasal smaller than postocular; preocular present; neither ocular as high as eye; eye less than eye-mouth distance;
INGER AND MARX: THE SNAKE GENUS CALAMARIA 71
5 supralabials, third and fourth entering orbit, fifth the largest, first, third, and fourth subequal, second slightly larger; mental triangular, not touching anterior chin shields; 5 infralabials, first 3 touching an- terior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.028 (1 specimen); tail thick, tapering near end to a sharp point; dorsal scales reduce to 4 rows on tail opposite third to fifth subcaudal anterior to terminal scute.
Hemipenis forked opposite fifth subcaudal, calyces smooth (1 specimen).
Eight to 9 unmodified maxillary teeth (8 specimens).
Ventrals: males, 123-1380 (N=2); female, 146. Subcaudals: males, 29-32 (N=2); female, 14+ (incomplete).
Total length: males, 215-244 mm.; female, incomplete. Ratio of tail to total length, 0.181-0.184 (N=2).
Body and tail dark brown above, each scale with a light network; most scales with a dark central spot; spots frequently unite to form short longitudinal stripes; anteriorly 2 or 3 light spots or vertical bars on side of body; scales of first row whitish, the light area becom- ing progressively reduced toward rear of body; head dark brown above with obscure lighter spots; dark pigment reaching lip on at least the first 2 supralabials; posterior supralabials with varying amounts of dark pigment; underside of head yellowish with dark brown spots; ventrals and subcaudals with dark lateral margins; tail with a dark mid-ventral streak; otherwise ventral surface immacu- late yellow.
Distribution.—Cocos Islands (Fig. 17).
PULO SI LAUT (RMNH 4716A-lectotype, 4716B-syntype; ZMA 10111-syntype).
Calamaria curta Boulenger
Calamaria curta Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 18, p. 62—South Celebes; 1897, Proc. Zool. Soc. London, 1897, p. 224, pl. 14, fig. 2; de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 160; Smith, 1927, Proc. Zool. Soc. London, 1927, p. 224; Werner, 1929, Zool. Jahrb., (Syst)., 57, p. 171; de Haas, 1950, Treubia, 20, p. 566; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 201.
Holotype.—British Museum (Natural History) 96.4.29.35.
Diagnosis.—Maxillary teeth unmodified; third and fourth supra- labials entering orbit; mental touching anterior chin shields; second pair of chin shields meeting in midline.
FIELDIANA: ZOOLOGY, VOLUME 49
—~l bo
Description.—Rostral higher than wide, portion visible from above 14 to 24 length of prefrontal suture; prefrontal shorter than frontal, touching first 2 supralabials; frontal hexagonal, 1°/s to 21% times width of supraocular, about ‘/; length of parietal; parietal 114 to 124 times length of prefrontal; paraparietal surrounded by 6 shields and scales; nasal smaller than postocular; preocular present; neither ocular as high as eye; eye equal to or greater than eye-mouth dis- tance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal or second slightly larger than others; mental triangu- lar, touching anterior chin shields; 5 infralabials, first 3 touching an- terior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.015-0.029 (2 specimens); tail tapering gradually from base to an extremely long terminal scute; dorsal scales reduce to 4 rows on tail opposite third to sixth subcaudal an- terior to terminal scute.
Cloaca of female cardioid (1 specimen). Seven unmodified maxillary teeth (8 specimens).
Ventrals: male, 156; females, 153-170 (mean 162; N=4). Sub- caudals: male, 28; females, 14-17 (mean 15.3; N=4).
Total length: male, 1538 mm.; females, 195-807 mm. Ratio of tail to total length: male, 0.105; females, 0.067—-0.072 (mean 0.073; N=4).
Color dark brown above, most of dorsal scales with a fine dark net- work; dorsal scales darker in anterior corners; head dark brown above and on sides with obscure darker spots; supralabials with dark sutures and yellowish ventral borders; head yellow below with dark brown spots; ventrals dark, usually brown in anterior 24 and yellowish on posterior edges; small yellowish spots usually present in dark portion of each ventral; subcaudals yellow and brown, usually yellow con- fined to posterior halves.
Ecological notes.—Smith (1927) found four specimens in and under rotting logs. He stated that their food consisted of earthworms.
Distribution.—Southern Celebes (Fig. 17). CELEBES: peak of Mount Bonthain, 1500-1800 meters (BM 1926.
8.20.167-168; MCZ 25301-02) ; south Celebes, 610 meters (BM 96.4. 29.35—holotype).
Specimens examined—d.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 73
Calamaria acutirostris Boulenger. Figure 19.
Calamaria acutirostris Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 17, p. 394— Loka, Mount Bonthain, Celebes; 1897, Proc. Zool. Soc. London, 1897, p. 223, pl. 13, fig. 2; Boettger, 1898, Kat. Rept. Samm. Senck. Naturf. Ges., pt. 2, p. 86; de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 161; Smith, 1927, Proc. Zool. Soc. London, 1927, p. 224; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 171; de Haas, 1950, Treubia, 20, p. 563; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 199.
Lectotype.—Naturhistorisches Museum Basel 1686, here desig- nated.
Fic. 19. Paratype of Calamaria acutirostris (NMB 1688).
Diagnosis.—Maxillary teeth unmodified; third and fourth supra- labials entering orbit; mental touching anterior chin shields; second pair of chin shields not meeting in midline.
Description.—Rostral higher than wide or as wide as high, scarcely visible from above; prefrontal subequal to frontal, touching first 3 supralabials; frontal hexagonal, 224 to 31% times width of supra- ocular, about 24 length of parietal; parietal 11% times length of pre- frontal; paraparietal surrounded by 6 shields and scales; nasal slightly smaller than postocular and larger than preocular; preocular present, neither ocular as high as eye; eye less than or equal to eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, first smallest, next 3 subequal; mental triangular, touching anterior chin shields; 5 infralabials, first 3 touching anterior chin shields; posterior pair of chin shields not meeting in midline; 4 gulars in midline between anterior chin shields and first ventral.
Body thickness index 0.013-0.070 (10 specimens) ; tail thick, either tapering gradually or abruptly near end to a point; dorsal scales re-
74 FIELDIANA: ZOOLOGY, VOLUME 49
duce to 4 rows on tail opposite first to sixth subcaudal anterior to terminal scute.
Hemipenis forked opposite fifth to seventh subcaudal, retractor muscle beginning opposite ninth to twelfth subcaudal, sulcus bifur- cate, calyces smooth (5 specimens). Cloaca of female bilobed (2) or bulbous (1).
Ten to 12 unmodified maxillary teeth (9 specimens).
Ventrals: males, 148-162 (mean 154.6; N=18); females, 163-174 (mean 167.6; N=16). Subcaudals: males, 20-24 (mean 21.5; N=18); females, 18-17 (mean 15.2; N=16).
Total length: males, 138-415 mm.; females, 150-447 mm. Ratio of tail to total length: males, 0.086-0.111 (mean 0.094; N=17); females, 0.057-0.069 (mean 0.068; N=16).
Color reddish brown above without markings; dorsal scales with- out network; head dark brown above; dark pigment ending on a more or less oblique line on supralabials, usually reaching lower edge of an- terior supralabials; underside of head yellow with dark spots on ante- rior infralabials and chin shields; ventrals yellow, usually with a brownish speckling across their anterior edges; subcaudals yellow, edged with brown.
Ecological notes—Smith (1927) found ten specimens under stones on Mount Bonthain.
Distribution.—Southwestern Celebes (Fig. 17).
CELEBES (NMS 8 [2]; ZMB 15636): Mount Bonthain, Djikoro (CNHM 88161; MCZ 25292-96, 25298; USNM 120810), Loka (BM 96.12.9.61-63—paratypes; NHMW 16714; NMB 1681—lectotype, 1684, 1688-89, 5228-29—all paratypes); Bua Praeng (NHMW 16713: 1-11; SNG 19440).
? Java (ZMB 138947). This specimen was collected by the same collector (Fruhstorfer) and on the same date as SNG 19440. The snake probably came from Celebes.
Specimens examined.—s4.
Calamaria schmidti Marx and Inger. Figure 1.
Calamaria schmidti Marx and Inger, 1955, Fieldiana, Zool., 37, p. 197, fig. 27 — Bundu Tuhan, Mount Kina Balu, North Borneo.
Holotype.—United States National Museum 130240.
Diagnosis.—Maxillary teeth unmodified; four supralabials, sec- ond and third supralabials entering orbit.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 75
Description.—Rostral broader than high, portion visible from above about 14 length of prefrontal suture; prefrontal squarish, sub- equal to length of frontal, touching first 2 supralabials; frontal pentag- onal, 5 to 6 times width of supraocular, about 24 length of parietal; parietal 124 times length of prefrontal; paraparietal surrounded by 6 shields and scales; nasal larger than eye or postocular; no pre- ocular; postocular not as deep as eye; eye small, diameter 14 eye- mouth distance; 4 supralabials, second and third entering eye, fourth longest, third 34 or more length of second, first and third equal; mental triangular, not touching anterior chin shields; 5 infralabials, first 83 touching anterior chin shields; both pairs of chin shields meet- ing in midline; 3 gulars on midline between second pair of chin shields and first ventral.
Tail ending in a blunt point; dorsal scales reduce to 4 rows on tail opposite the seventh to eighth subcaudal anterior to terminal scute.
Six or seven unmodified maxillary teeth (2 specimens).
Ventrals: males unknown; female, 144 (1 specimen). Subcaudals: females, 14 (2 specimens). Total length of complete female 253 mm.; ratio of tail length to total length: female, 0.075 (1 specimen). The paratype is badly damaged and cannot be measured.
Color purplish gray above, uniform, scales without network; head without markings; supralabials same color as back; anterior infra- labials 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 posteri- orly, but lighter than dorsal color; under side of tail darker than belly but slightly lighter than dorsal surface.
Food.—The holotype contained an earthworm. Distribution.—Mount Kina Balu, North Borneo.
NorTH BoRNEO: Mount Kina Balu, Bundu Tuhan, 1370 meters (USNM 130240—holotype), Mount Kina Balu (SU 8568—paratype)
Calamaria lumbricoidea H. Boie. Figure 20.
Calamaria lumbricoidea H. Boie in F. Boie, 1827, Isis, 20, p. 540—Java; Schlegel, 1837, Phys. Serp., pt. 2, p. 27, pl. 1, figs. 14-16; Duméril and Bibron, 1854, Erp. Gén., 7, p. 89; Giinther, 1858, Cat. Colubrine Snakes Brit. Mus., p. 5; Jan, 1862, Arch. Zool. Anat. Phys., 2, p. 8; 1865, Icon. Ophid., 10, pl. 2, fig. 2; Lidth de Jeude, 1890, Notes Leyden Mus., 12, p. 254; Boulenger, 1894, Cat. Snakes Brit. Mus., 2, p. 333; Boettger, 1886, Ber. Senck. Naturf. Ges., 2, p. 105; 1894, 72 Semon, Zool. Forsch. Austr., p. 125; 1898, Kat. Rept. Samm. Senck. Naturf. Ges., pt. 2, p. 83; de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 153; Werner, 1929, Zool. Jahrb.,
FIELDIANA: ZOOLOGY, VOLUME 49
(Syst.), 57, p. 170; Kopstein, 1930, Treubia, 12, p. 274; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 183; Inger and Marx, 1962, Syst. Zool., 11, Di oc.
Calamaria lumbricoidea lumbricoidea, de Haas, 1950, Treubia, 20, p. 570.
Calamaria vermiformis Duméril and Bibron, 1854, Erp. Gén., 7, p. 85—“‘Java”’ [in error]; Jan, 1862, loc. cit., p. 8; 1865, loc cit., pl. 2, fig. 3; Lidth de Jeude, 1890, in Weber, Zool. Ergebn., p. 182; 1922, Zool. Meded., 6, p. 247; Boulenger, 1894, op. cit., p. 333; 1896, Cat. Snakes Brit. Mus., 3, p. 646; 1912, Fauna Malay Penin., p. 155; Bartlett, 1895, Sarawak Note Book, no. 1, p. 83; Werner, 1896, Verh. Zool. Bot. Ges. Wien, 46, p. 17; 1929, loc. cit., p. 170; Boettger, 1898, loc. cit., p. 838; Brown, 1902, Proc. Acad. Nat. Sci. Philadelphia, 1902, p. 180; Cohn, 1905, Zool. Anz., 29, p. 545; Baumann, 1913, Zool. Jahrb., (Syst.), 34, p. 270; Smith, 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 162; 1925, Sarawak Mus. Jour., 3, p. 4; 1930, Bull. Raffles Mus., no. 3, p. 58; 1931, zbid., no. 5, p. 27; de Rooij, 1917, op. cit., p. 153; Holtzinger-Tenever, 1920, Arch. Naturg., 85, p. 86; Mertens, 1924, Zool. Anz., 60, p. 158; Lonnberg and Rendahl, 1925, Ark. Zool., 17A, p. 2; de Haas, 1950, loc. cit., p. 575; Tweedie, 1953, Snakes Malaya, p. 49; Marx and Inger, loc. cit., p. 185; Taylor and Elbel, 1958, Univ. Kansas Sci. Bull., 38, p. 1044.
Calamaria vermiformis vermiformis, Marx and Inger, 1955, loc. cit., p. 187.
Calamaria temmincki Duméril and Bibron, 1854, Erp. Gén., 7, p. 87—‘‘Su- matra or Borneo’; Giinther, 1858, loc. cit., p. 5.
Calamaria. grayi Giinther, 1858, Cat. Colubrine Snakes Brit. Mus., p. 5— Philippine Islands; Boettger, 1886, loc. cit., p. 105; Boulenger, 1894, loc. cit., p. 08; Taylor, 1922, Sndkes' Philip. Ids:, p. 184% 1922," Philipwviour: Scl.;,21,/p; 204; Werner, 1929, loc.cctjap 170.
Calamaria dimidiata Bleeker, 1860, Nat. Tijds. Ned. Indie, 21, p. 295—Java.
Calamaria melanorhynchus Bleeker, 1bid.—Ampat Lawang, Sumatra.
Calamaria alkeni Bleeker, 7zbid.—Ampat Lawang, Sumtra.
Calamaria flaviceps Giinther, 1865, Ann. Mag. Nat. Hist., (8), 15, p. 90— Sarawak and Borneo.
Calamaria philippinica Steindachner, 1867, Verh. Zool. Bot. Ges. Wien, 17, p. 514, pl. 8, figs. 4-6—Philippine Islands; Boettger, 1886, loc. cit., p. 105.
Calamaria stahlknechtii Stoliczka, 1873, Jour. As. Soc. Bengal, 42, p. 119, pl. 11, fig. 2—near Deli, Sumatra; Boulenger, 1885, Ann. Mag. Nat. Hist., (5), 16, p. 388; 1894, loc. cit., p. 885; Modigliani, 1889, Ann. Mus. Stor. Nat. Genova, (2), 7, p. 119; de Rooij, 1917, loc. cit.,.p. 154; Werner, 1929; loc. cit., p. 170; de Haas, 1950, loc. cit., p. 574; Marx and Inger, 1955, loc. cit., p. 206.
Calamaria variabilis Lidth de Jeude, 1890, in Weber, Zool. Ergebn., p. 1838, pl. 16, fig. 8—Buitenzorg, Java.
Calamaria vermiformis var. sumatranus Lidth de Jeude, 1890, Notes Leyden Mus., 12, p. 18—Deli, Sumatra.
Calamaria bungaroides Werner, 1901, Zool. Anz., 24, p. 300—type locality unknown; 1929, loc. cit., p. 178; de Rooij, 1917, loc. cit., p. 173; de Haas, 1950, loc. cit., p. 565; Marx and Inger, 1955, loc. cit., p. 200.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 77
Calamaria bruegeli Mertens, 1924, Zool. Anz., 60, p. 158—central Borneo; de Haas, 1950, loc. cit., p. 565.
Calamaria géringt Vogt, 1925, Zool. Anz., 62, p. 64—Java; de Haas, 1950, loc. cit., p. 567; Marx and Inger, 1955, loc. cit., p. 202.
Calamaria goeringi, Werner, 1929, loc. cit., p. 171; Brongersma, 1930, Treubia, 12, p. 302.
Changulia lumbricoidea, de Haas, 1941, Treubia, 18, p. 369.
Calamaria gracilis (non Boulenger), Angel, 1941, Bull. Mus. Nat. Hist. Nat., (2), 13,, p. 411.
Calamaria vermiformis grayi, Marx and Inger, 1955, loc. cit., p. 188; Leviton, 1968, Proc. Calif. Acad. Sci., 31, pp. 379, 389, 393, 404.
Lectotype.-—Rijksmuseum van Natuurlijke Historie 105438 (ex 42).
Taxonomic notes.—Dumeéril and Bibron did not compare verm- formis with the very similar species lwmbricoidea. Boulenger (1894) differentiated these two forms solely on the basis of eye size, which he said was smaller than the eye-mouth distance in lwmbricoidea and equal to that distance in vermiformis. De Rooij (1917) merely copied Boulenger’s descriptions. In his descriptions Boulenger stated that the ventral coloration of lumbricoidea was whitish whereas that of vermiformis was highly variable, ranging from uniform yellow through increasing amount of black transverse bands to uniform black.
Our examination of 375 specimens from all parts of the combined ranges of lumbricoidea and vermiformis convinces us of their conspeci- ficity. We confirm Boulenger’s observation of the range of ventral coloration. We cannot, however, confirm Boulenger’s observations on eye size; Javanese snakes agreeing in all details with the type series of lumbricoidea have eye diameters ranging from slightly smaller to larger than the eye-mouth distance. Snakes from Borneo, indistin- guishable from the type series of vermiformis, show the same varia- tion in eye size. Both sets of-specimens have the same distinctive round snout and a relatively thick body. Especially significant is the fact that only these two nominal forms out of the entire genus show ontogenetic variation in coloration of the head (see below).
The type locality of both vermiformis and lumbricoidea is Java. All of Boie’s material was from Java and subsequent collecting has turned up all of his species of Calamaria on Java. Consequently the type locality of the older name, lumbricoidea, is correctly given. Numerous specimens agreeing with the types of lwmbricoidea in those few characters by which these two type series differ (see Geographic variation) have been collected over a long period of time on Java by various collectors.
78 FIELDIANA: ZOOLOGY, VOLUME 49
Seven specimens with the characteristics of vermiformis (1.e., banded ventrally) have been reported from Java. Of these only one (MHNP 89-187) is reported from a specific locality, Batavia (=Ja- karta). As none of the recent large Javanese collections of Calamaria include vermiformis, which is one of the common forms in other parts of the generic range, we doubt that this color form occurs on Java.
The detailed original description and figures of stahlknechti agree in counts and coloration with typical Sumatran specimens of luwm- bricoidea. Snakes from Nias identified by Boulenger (1894) and de Rooij (1917) as stahlknechtt have been examined and are also lumbricoidea.
The holotype of géringi does not differ from Javanese lumbricoidea.
Our examination of the holotype of bruegelz confirms our previous opinion (Marx and Inger, 1955) that it is synonymous with vermi- formis and therefore with lumbricoidea.
The original description of Calamaria bungaroides fits juvenile Bornean and Sumatran lumbricoidea exactly.
We have examined the types of temmincki, dimidiata, melano- rhynchus, alkent, flaviceps, vermiformis var. sumatranus, and variabilis and agree with Boulenger (1894) that they are synonymous with lumbricoidea and vermiformis of Boulenger.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; paraparietal surrounded by 4 or 5 scales and shields; belly yellow with black crossbars wider than width of one ventral.
Populations of lwmbricoidea from Java and parts of Sumatra have immaculate light bellies. Javanese lumbricoidea are distinguished from all other species known from that island by the other characters mentioned in the diagnosis. Those characters also distinguish the light-bellied Sumatran lumbricoidea from all other Sumatran Cala- maria except albiventer. The last has two lateral light stripes which differentiate it from lumbricordea.
Description.—Rostral wider than high, or higher than wide, por- tion visible from above about half length of prefrontal suture; pre- frontal equal to or slightly shorter than frontal, touching first 2 supralabials; frontal hexagonal, 14% to 214% times width of supra- ocular, about 34 to °/s length of parietal; parietal 114 times length of prefrontal; paraparietal surrounded by 4 or 5 shields and scales; nasal smaller than postocular; preocular present, higher than post-
INGER AND MARX: THE SNAKE GENUS CALAMARIA 79
ocular; postocular usually not as high as eye; eye smaller than, equal to, or greater than eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal or third the smallest; mental triangular, touching anterior chin shields; 5 infra- labials, first 8 touching anterior chin shields; both pairs of chin shields usually meeting in midline; usually 3 gulars in midline be- tween posterior chin shields and first ventral (see Geographic varia- tion below).
Fic. 20. Calamaria lumbricoidea (NHMW 16698:3).
Body thickness index 0.012-0.074; tail thick, tapering abruptly near tip to sharp point; dorsal scales reduce to 4 rows on tail opposite first to eleventh subcaudal anterior to terminal scute.
Hemipenis variable (see Geographic variation). Cloaca of female bilobed, bulbous, or cardioid, not varying geographically (Inger and Marx, 1962).
Nine (1 specimen), 10 (19) or 11 (2) modified maxillary teeth.
Ventrals: males, 144-196 (mean 172.0; N=182); females, 137-229 (mean 190.8; N=116). Subcaudals: males, 17-27 (mean 21.5; N= 134); females, 18-21 (mean 16.8; N=118).
Total length: males, 149-498 mm.; females, 120-642 mm. Ratio of tail to total length: males, 0.068-0.114 (mean 0.080; N=126); females, 0.089-0.0838 (mean 0.056; N=1138).
Color dark brown or black above, with or without narrow, light stripes; scales without network; first one or two scale rows yellow;
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82 FIELDIANA: ZOOLOGY, VOLUME 49
ventral surface yellow, with or without black crossbands (see Onto- genetic variation and Geographic variation).
Ontogenetic variation.—The heads of juveniles (up to about 200 mm., total length) are yellow in preservative whereas those of adults are dark brown or black. Darkening of the head begins on the snout and gradually spreads to cover the entire head (Fig. 21).
Juveniles of most populations have yellow bands across the dor- sum. These bands are one-half to two scales wide (Fig. 22).
Geographic variation.—Figure 21 suggests that Bornean juveniles tend to have lighter heads than do those from Java. Adults from Java often retain a narrow light band or pair of light spots on the rear of the parietals. Such spots or bands rarely occur in adults in other parts of the range (Fig. 21).
The yellow dorsal crossbands are usually less than one scale wide and usually interrupted in juveniles from western Sumatra and west- ern Malaya. In juveniles from Borneo and the Philippine Islands these bands are usually continuous and one to two scales wide. Yel- low crossbands are absent in juveniles from Java, eastern Sumatra, and central Malaya (Fig. 22).
Tweedie (1953) states that longitudinally striped lwmbricoidea are confined to high altitudes and uniform brown snakes to low altitudes. The striped Malayan snakes examined by us come from 1,800 to 1,400 meters above sea level and non-striped ones from 500 to 1,350 meters. Limiting observations to Malaya might lead to the conclu- sion that the variation is altitudinal. However, all lumbricozdea from Java (350 to 1,400 meters) and Borneo (50 to 1,480 meters) are non- striped though they span the altitudinal range of the Malayan speci- mens. All lumbricoidea from the Philippine Islands (sea level to 850 meters) are longitudinally striped.
Clearly this aspect of dorsal pattern varies geographically rather than altitudinally (Table 21). The non-striped Malayan snakes occur along the western edge of the peninsula and the striped ones farther inland. In Sumatra non-striped snakes occur along the east- ern coast and striped ones on the west.
The ventrum in lwmbricoidea varies from uniform yellow to almost solid black. Black on the belly usually takes the form of black cross- bands that occupy 2 to 4 adjacent ventrals separated by varying numbers of yellow scales. In three specimens the belly is entirely black. Some specimens without black crossbands have black speck- ling that varies from a few tiny spots on scattered ventrals to a more or less continuous dark stripe midventrally. Geographic variation
INGER AND MARX: THE SNAKE GENUS CALAMARIA 83
TABLE 21.—Frequency Distribution of Dorsal Patterns and Number of Gulars in Samples of Calamaria lumbricoidea.
Dorsal pattern No. of gulars Localities Striped Uniform 3 4 TRAUONC < «, 2:446-428 da-wee ewe ore we 1 I Malaya: Panbane< sed seen ds8 aes 8 7 1 Pele, ced OY VR Ss 4 4 SClANGOr ecu cease sewn 4 3 JONOLEs «5.5 bok Same owecks 2 2 Sumatra: Padang area........... 20 3S 25 WG es: Oe dono ee ose de 8 1 cs Palembang..... nes 1 1 S! Rambé.uscsaccenadus ii 1 Bindjay <4 62cc ccd deces 2 2 LONGED Gs nao as ea eew eed 2 2 Ophir District.......... 1 j 1 ii INAS doe Se ha ee ih pe eRe ee 6 1 5 Borneo: Kina Balt. wi: sis eedees Le 13 LaWas. .dvecedeiatonens 1 I Pa Brayou es ic.0< 64 awregrs 1 1 Baram District......... 4 2 2 TN rs od, ants cca haceanee A 2 1 1 M AVON os «vd aeensane ds 1 1 Batu Song............. 1 1 Tandon? ic ccddssnencex Z 2 Mahakam............. 2 Z Sandakan..... cea s 1 a PO ee olny odd sow Seta Ged Gow, eee 125 125 NSUUNG.4 92 Sedo awed iene thes poet 1 1 Wid aNAGso &hxedeinee ons oe 8a nes 14 13 1 TOOL 4 & &-esnennint & a ine ae eee oa 1 1 BAST ov x. sites Se Si dn ae dae 1 1 LF: eee a or 1 1
of ventral coloration is shown in Table 22. The belly pattern shows not only inter-island variation, but also variation within Sumatra as a comparison of the Deli and Padang specimens indicates.
Populations from the Philippine Islands, Borneo, Java, western Sumatra, and central Malaya have 3 mid-ventral gulars whereas 4 is the common number elsewhere (Table 21).
Within islands the number of ventrals varies to a remarkable ex- tent (Tables 28 and 24). Yet at any given locality, lumbricoidea follows the common pattern of this genus by having relatively little individual variation (see p. 24). For example, in the Kina Balu sam- ple the range of ventrals is 170-180 in seven females and 154-165 in seven males or only a fraction of the range for the entire Bornean sample. Despite much overlap, the populations from Java and the Philippine Islands have higher ventral counts than do those from other areas.
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INGER AND MARX: THE SNAKE GENUS CALAMARIA 85
Similar geographic variation—slight differences between areas with extensive overlap—is shown by subcaudal counts, the position of the reduction to four dorsal scale rows, and the ratio of tail length to total length (Tables 25 and 26).
The hemipenis of lumbricoidea shows variation not seen in other species of Calamaria (Inger and Marx, 1962). Males from the main- land, Sumatra, Nias, and Java are alike in having the hemipenis forked beginning between the second and sixth subcaudals (as is characteristic of the genus) and in having both rami covered with papillate calyces; the sulcus forks at the base and a branch runs to the apex of each fork (as in Fig. 4F of Dowling and Savage, 1960).
The same type of hemipenis was found in males from North Borneo and Sarawak. However, males from widely separated local- ities in Borneo have undivided hemipenes that have smooth calyces; the sulcus is bifurcate in these simple hemipenes. Aside from the range of variation among these Bornean specimens, the striking aspect is the lack of variation within a local population, such as that of Mount Kina Balu.
Characters that vary geographically in lumbricoidea are not asso- ciated in any fixed way with the result that geographic variation in one character does not coincide with that of another (Table 27). If subspecies were named they would have to be based on a single character and the choice of that character would be arbitrary.
Distribution.—From southern Thailand to Java, Borneo, and Leyte (Fig. 72).
THAILAND: Bangkok (UMMZ 65331).
MALAYA: Perak: Gunong Kledong (NMS unnumbered), Larut Hills (BM 1900.6.14.22), 1870 meters (NMS unnumbered), Taiping (ZMB 26457). Pahang: Fraser’s Hill (NMS 30229, 4 unnumbered), Cameron Highlands, Kuala Terla (CNHM 723874), Cameron High- lands, Tanah Rata, 1400 meters (NMS 2 unnumbered). Selangor: Kepong Forest Reserve (NMS 18880, 30462), 640 meters (NMS 9474), Kepong, Bukit Lagong (NMS 8827). Johore: Gunong Pulai (NMS 2 unnumbered).
Nias (BM 84.12.81.7, 84.12.31.8/2]; RMNH 4865). Gunong Sitoli (MCG 30881); west Nias (ZMA 10284).
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88 FIELDIANA: ZOOLOGY, VOLUME 49
Si Rambe (MCG 30376). Sumatera Utara: Bindjai (ZMH 2466/2]), Deli (BM 89.12.16.108; NHMW 16817; RMNH 1228[2]—syntypes of vermiformis var. sumatrana, 6636, 1 unnumbered; SNG 193890, 19392; ZMA 10227, 10229, 10235), Langkat (MCZ 37769-70). Su- matera Tengah: Bukit Tingi (BM 1928.2.18.238-380; RMNH 5806[2]; ZMA 10228[3]), Padang (BM 88.5.24.4-6; MCZ 20971; NHMW 16700[2], 16701-02; RMNH 511, 535), Batu Sangkar (RMNH 4861), Solok (NMB 1708), Ophir district, Sukamananti, 200 meters (RMNH 5806), Ophir district, Tanano-Talu, 1000 meters (RMNH 5806), Kaba Wetan (USNM 70945-51). Sumatera Selatan: Palembang (ZMA 10232), Ampat Lawang (BM 68.12.11.184; RMNH 3976— syntype of melanorhynchus). ?Sumatra (RMNH 41).
GREAT NATUNA (BM 95.5.1.76).
TABLE 25.—Frequency Distribution with Respect to Certain Characters in Males of Calamaria lumbricoidea.
Subcaudals
iy iS 19 20 21 22 23 2% 25 26 27 Mainland....... 1 1 3 si 1 1 1 NIAS:. 66 ek wank oe i Ze 2 Sumatra 1 7 8 14 6 1 Borneo......... 2 AD, 7 D 1 Z Philippine Islands....... 2 i 2, 1 1 She: ne re | 1 14. 20: % 1
Position of reduction to four dorsal seale rows!
1O di 12 i8 I) 15 .-16 17. 18 19 20 2F “2225
Mainland....... Z 1 3 3 |! Ee a aer nO ) 1 3 iL Sumatra 1 2. 3 3 8 7 4 3 2 3 Borneo: 624.4244 I 1 4 2 6 3 3 al if 1 Philippine Islands....... il I 8 74 WAV Aas 6 hub hie a 2 6 i eomeen 65) 9 2 Tail ratio? Class centers 54 63 68 73 <78 83-88 93 98105. 108s Mainland....... 1 3 1 1 2 las. . il 74 2 SUMAatra).. eee 74 7 {Cle} 7 7 Borneo. 4... 5 1 4 4 74 a 2 1 Philippine Islands. ...... 3 2 lt "lc hig: ieee Ocean Ai Ore 4 Z
' Position given in terms of subecaudals counted from vent. * Ratio in thousandths of total length.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 89
TABLE 26.—Frequency Distribution with Respect to Certain Characters in Females of Calamaria lumbricoidea.
Subcaudals
13 14 15 16 17 18 19 20 21 Mainland........... 1 2 3 yA 2 INAS? 25-4444 once 1 UMA Sa: « bak Sees c ss 2 2 1 10 9 4 ] BOImnGOs 24/08 4-40 ede a 2 3 T 1 2 Al Philippine Islands 2 3 -2 3. a 1 SAVE fog oxigen se wk ee 2 ll 16-18 1
Position of reduction to four dorsal scale rows!
6 % 8 9 10 di 12 i3 if 15 16 17 18
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Tail ratio?
Class centers
39-40 43 48 53 58 63 68 73 78 83
Mainland........... 4 2 Z Z 1 1 Nias. . 1
SUNITA A sco scared de it 9 10 7
Borneo ces Se fos il 1 3 4 4 2 3 Philippine Islands.... 1 2 % 4
DONG ove ahaee cakes 2 15 18 38
1 Position given in terms of subcaudals counted from vent. > Ratio in thousandths of total length.
SARAWAK (BM 45.10.2.45—syntype of flaviceps): First Division: Bidi (SM Cd.5.26.1¢; Cd.5.26.15b), Kuching (SM Cd.5.26), Matang (BM 73.3.4.23). Second Division: Saribas (SU 8575). Fourth Divi- sion: Niah, Rumah Sigi (CNHM 129005), Niah, Kuala Sakoloh (CNHM 129006), Niah (CNHM 181620-27), Mount Batu Song, 300 meters (BM 92.10.7.8), Baram district (BM 97.3.4.5, 1902.2.11. 25.16-17; NHMW 16992:1). Fifth Division: Lawas (CNHM 67278), Lawas, Sungei Usop (SM Cd.5.26.1d), Pa Brayong (CNHM 71646).
NORTH BORNEO: Ranau District: Mount Kina Balu (MHNP 89- 193), Bundu Tuhan, 1370 meters (USNM 180252). Kota Belud District: Mount Kina Balu, Tenompok, 1480 meters (MCZ 43568), Kiau, 915 meters (BM 1929.12.22.110-111; MCZ 48564-65; NMS 16817), Kenokok River, 1000 meters (MCZ 43566-69; UMMZ 82829[2]). Sandakan District: Sandakan (CNHM 15008).
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INGER AND MARX: THE SNAKE GENUS CALAMARIA 91
INDONESIAN BORNEO: Boven Mahakam (RMNH 2 unnumbered), Tandjong (BM 96.2.17.11-12), southeast Borneo (SMNS 3498, 3498a, 3497; ZMH 2421).
BORNEO (BM 56.9.27.9—syntype of flaviceps, 72.2.19.55, 1 un- numbered; USNM 49802); central Borneo (SNG 19889—paratype of bruegeli); north Borneo (NHMW 16992:2-3; ZMB 18110).
BASILAN: near Isabela (MCZ 43469).
MINDANAO: Zamboanga Province: Zamboanga City (CAS 15291), Zamboanga (City?) (CAS 620380). Cotabato Province: Mount Malin- dang, Gumay (CNHM 96529), Cotabato (NMB 8518), Saub (MCZ 25778-80; USNM 120811), Cotabato coast (ZMA 10236). Misamis Province: Lake Lanao (CAS 15286-89), Lanao, Lumbatan (CNHM 15010). Davao Province: Mount Apo, Todaya, 855 meters (CNHM 53013).
NEGROS: Negros Oriental: Dumaguete (SU 15952). BOHOL (NHMW 16815).
LEYTE: Cabalian (MCZ 25781).
SAMAR: Osmera (USNM 122215).
PHILIPPINE ISLANDS (BM 1946.1.2.17, 1946.1.2.21—-syntypes of grayi; ZMB 5446).
JAVA (BM 44.2.32.65, 66.8.14.344, 70.6.7.46, 73.5.8.26; CNHM 83163; MHNP 165; NHMW 16698(6], 16816, 2 unnumbered; NMS 258; RMNH 10419[4], 10548—lectotype of lumbricoidea, 42[2]—para- types of lwmbricoidea; SNG 193888, 19393; ZMA 10233[2]; ZMB 1552, 4069, 4117, 6936, 8545—holotype of géringi, 8758, 10261; ZMH 4651- 4652, 5880). M. Pangherango (MCG 30457); ?Tjikorai (NMS 5). Djawa Barat: Benkang near Serang (AMNH 71513), Djakarta (NMS 544), Bogor (RMNH 4827—holotype of varzabilis; SNG 193886, Med- amendoeng Peak, between Bogor and Sindanglaja (MHNP 95-52[2}), Sindanglaja (MCZ 7523), Bandung, 800 meters (RMNH 7463/3], 1 unnumbered), Mount Gede, Tjibodas, 1450 meters (RMNH 8604), Rarahan, near Tjibodas (MCG 6780, 30878), Mount Garut (NHMW 16697:1), Mount Garut, plantation Dajeuhmanggoeng (RMNH un- numbered), Tjikadjang, 900 meters (RMNH 6833/4], 126 unnum- bered), Tjusurupan (SNG 19887, 19391), Tasikmalaja, 350 meters (NHMW 16697:2). Djawa Tengah: Wonosobo, Kepil, 550 meters (RMNH 8555), Wonosobo, Selomojo, 600 meters (RMNH unnum- bered), Wonosobo, Tlogodjati, 1000 meters (RMNH unnumbered).
92 FIELDIANA: ZOOLOGY, VOLUME 49
Questionable localities: Java (BM 61.8.12.44, 63.12.11.171—holo- type of dimidiata; MHNP 174 and 3300[2]—syntypes of vermiformis; ZMB 4118—syntype of vermiformis). Java: Batavia (MHNP 39- 187; RMNH 80—holotype of temmincki). Java: Bandung (RMNH co.118). Dutch Indies (MNHP 12-51, 12-52, 12-54; RMNH 3975). Malayan Archipelago (NHMW 16694). Ternate (BM 78.1.381.8-9). No data (RMNH unnumbered).
Specimens examined.—s886
Calamaria griswoldi Loveridge. Figure 23.
Calamaria lumbricoidea griswoldi Loveridge, 1938, Proc. Biol. Soc. Washing- ton, 51, p. 43—Luidan River, Bundu Tuhan, Mount Kina Balu, North Borneo; de Haas, 1950, Treubia, 20, p. 570.
Calamaria griswoldi, Marx and Inger, 1955, Fieldiana, Zool., 37, p. 1838. Holotype.-—Museum of Comparative Zoology 48580.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; color blackish brown above with a narrow white line between successive scale rows; immaculate yellowish white below.
Description.—Rostral wider than high, portion visible from above 24 length of prefrontal suture; prefrontal */¢ length of frontal, touch- ing first 2 supralabials; frontal hexagonal, 114 to 2 times width of supraocular, about 24 to 34 length of parietal; parietal 124 to 2 times length of prefrontal; paraparietal surrounded by 5 shields and scales; nasal smaller than postocular; preocular present; neither ocu- lar as high as eye; eye equal to or slightly greater than eye-mouth dis- tance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal; mental triangular, touching anterior chin shields; 5 infralabials, first 8 touching anterior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.017-0.089 (5 specimens); tail thick, tap- ering from base to a sharp point (Fig. 10B); dorsal scales reduce to four rows on tail opposite ninth to thirteenth subcaudal anterior to terminal scute.
Hemipenis forked opposite fourth subcaudal; suleus spermaticus forked; calyces papillate (2 specimens).
Nine to 10 modified maxillary teeth (4 specimens).
Ventrals: males, 155-179 (mean 168.5; N=6); females, 188-192 (mean 187.2; N=5). Subcaudals: males, 16-18 (mean 16.7; N=6); females, 18-16 (mean 14.5; N=4).
INGER AND MARX: THE SNAKE GENUS CALAMARIA 93
Fic. 28. Calamaria griswoldi (NMS R18949).
Total length: males, 192-425 mm.; females, 875-488 mm. Ratio of tail to total length: males, 0.059-0.068; (mean 0.063; N=6); fe- males, 0.048-0.056 (mean 0.052; N=4).
Color dark brown above; dark portion of scales without network; blackish brown stripes occupying central 24 of each scale row above the first, yellowish stripes on edges on adjacent scale rows; scales of first row yellow, immaculate in anterior part of body, usually each scale with a small dark spot in posterior half of body; head dark brown above; supralabials yellow in lower 24; head below immaculate yellow; an oblique light bar running forward from gular region onto rear of parietals; ventrals immaculate yellow; subcaudals yellow, usually a faint zig-zag dark line mid-ventrally.
Distribution.—Highlands of northwestern Borneo (Fig. 72).
NORTH BORNEO: Ranau District: Ranau (USNM 134114), Mount Kina Balu (BM 5.11.7-25), Lumu Lumu, 1525 meters (BM 1929.12.22.112), Bundu Tuhan, 1370 meters (CNHM 72434; USNM 180288), Luidan River, near Bundu Tuhan (MCZ 48580—holotype,
94 FIELDIANA: ZOOLOGY, VOLUME 49
43581—-paratype). Kota Belud District: Tenompok, 1430 meters (NMS 18949, 1 unnumbered). Western North Borneo (SM Cd. 5.26.c-d).
SARAWAK. Fourth Division: Kelabit Plateau (SM unnumbered). Specimens examined.—12.
Calamaria albiventer Gray. Figure 24.
Changulia albiventer Gray, 1834, Ill. Ind. Zool., 2, pl. 86, figs. 6-9—Penang Island.
Calamaria albiventer, Giinther, 1858, Cat. Colubrine Snakes, p. 4; 1864, Rept. Brit. Ind., p. 197; Boulenger, 1894, Cat. Snakes Brit. Mus., 2, p. 336; 1912, Fauna Malay Penin., Rept. and Batr., p. 156; Flower, 1896, Proc. Zool. Soc. London, 1896, p. 886; 1899, zbid., p. 674; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 170; Smith, 1930, Bull. Raffles Mus., no. 3, p. 58; Tweedie, 1953, Snakes Malaya, p. 50; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 199; Batchelor, 1958, Malaya Nat. Jour., 12, p. 107.
Calamaria indragirica Schenkel, 1901, Verh. Ges. Basel, 13, p. 164, fig. 3— Indragiri, Sumatra; de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 154; Werner, loc. cit., p. 170; de Haas, 1950, Treubia, 20, p. 568; Marx and Inger, loc. cit. p. 202.
Calamaria ornata Werner, 1909, Mitt. Naturh. Mus. Hamburg, 26, p. 229, fig. 6—Sungei Lalak, Indragiri, Sumatra; de Rooij, loc. cit., p. 178; de Haas, loc. cit., p. 572; Marx and Inger, loc. cit., p. 204. Syntypes.— British Museum (Natural History) 1946.1.2.10, 1946. eZee.
Taxonomic notes—The striking color patterns of indragirica and ornata are identical to that of albiventer. As the counts and scale proportions also agree, we are putting the two Sumatran names in the synonymy of albiventer.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; body with 4 narrow light stripes.
Description.—Rostral broader than high, portion visible from above about 24 length of prefrontal suture; prefrontal 34 length of frontal, touching first 2 supralabials; frontal hexagonal, 124 to 2 times width of supraocular, about ’/s length of parietal; parietal 114 times length of prefrontal; paraparietal usually surrounded by 5 shields or scales; nasal smaller than or equal to postocular; preocular present, neither ocular as high as eye; greater than eye-mouth dis- tance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal; mental triangular, touching anterior chin shields;
5 infralabials, first 3 touching anterior chin shields; both pairs of
INGER AND MARX: THE SNAKE GENUS CALAMARIA 95
chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.026 (1 specimen); tail tapering gradually from base, abruptly tapering at tip to a sharp point; dorsal scales reduce to 4 rows on tail opposite fifth to eighth subcaudal anterior to terminal scute.
Cloaca of female bilobed (1 specimen). Nine modified maxillary teeth (8 specimens).
Ventrals: males, 1483-144 (N=2); females, 147-162 (mean 156.4; N=10). Subcaudals: males, 21-22 (N=2); females, 15-19 (mean 17.0; N=10).
Fic. 24. Calamaria albiventer (CAS 14940).
Total length: males, 205 mm. (N=2); females, 170-361 mm. Ratio of tail to total length: males, 0.088-0.098 (N=2); females, 0.047-0.088 (mean 0.066; N=9).
Color of body brown above; two pairs of light longitudinal stripes; lower half of first scale row and adjacent edges of ventrals brown; adjacent halves of first and second scale rows yellowish; a broad dark stripe on upper half of second scale row, all of third and fourth
96 FIELDIANA: ZOOLOGY, VOLUME 49
rows, and lower half of fifth row; this dark band sometimes split by a lighter brown stripe formed by light speckling on adjacent halves of third and fourth scale rows; yellowish stripe on adja- cent halves of fifth and sixth scale rows; a dark stripe on vertebral scale row and adjacent halves of sixth rows; top of head light brown with a dark network or dark spots; upper third or half of supralabials brown; remainder of supraliabals and underside of head yellowish, usually with dark spots on infralabial sutures; ventrals yellow except for dark lateral edges; underside of tail yellowish with a dark median stripe.
In life reddish brown above; upper light stripe red, lower light stripe bluish white; both light stripes edged with black; underside of head lemon yellow shading into red on neck; underside of body and tail bright red (Flower, 1899, p. 674).
Geographic variation.—Two Sumatran specimens (syntypes of indragirica) have 147 and 148 ventrals and 19 subeaudals. One of these snakes is a female; the second was too desiccated to determine the sex. The third Sumatran animal (holotype of ornata) has 148 ventrals and 17 subcaudals (Werner, 1909b, p. 229); sex was not given. If these 3 snakes are females, Sumatran females have lower ventral and higher subcaudal counts than females from Malaya (156-160; 15-17).
Distribution.— Western Malaya and east-central Sumatra (Fig. fO2\
MALAYA: Penang Island (BM 1946.1.2.10 and 1946.1.2.18—syn- types, 60.8.19.1269[2], 98.9.22.39). Perak: Larut Hills, 1870 meters (NMS unnumbered). Province Wellesley (BM 96.6.25.28).
SINGAPORE (CAS 14940).
SUMATRA: Sumatera Tengah: Indragiri (NMB 1697-98—syn- types of zndragirica), Indragiri, Sungei Lalak (holotype of ornata; not examined).
?India (BM 2 unnumbered).
Specumens examined.—12.
Calamaria hilleniusi new species. Figure 25. Holotype.—Zoologisch Museum Amsterdam 10078, a male from Samarinda, Indonesian Borneo.
Paratypes.—Sarawak Museum unnumbered, a female from north- western Borneo; Agriculture Department, Sabah, No. 72, (male) and,
INGER AND MARX: THE SNAKE GENUS CALAMARIA of
Chicago Natural History Museum 142088, (female) both from Tua- ran, Tuaran District, Sabah (=North Borneo).
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; paraparietal surrounded by 5 shields and scales; eye about 24 eye-mouth distance; dorsal scales without network.
Description.—Rostral wider than high, portion visible from above equals half length of prefrontal suture; prefrontal equals length of frontal, touching first 2 supralabials; frontal hexagonal, 21% to 3 times width of supraocular, about 34 length of parietal; parietal 114 to 1% times length of prefrontal; paraparietal sur- rounded by 5 scales and shields; nasal oriented laterally, equal to postocular; preocular present; both oculars as high as eye; eye about 24 eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, fourth almost as long as fifth along ventral border, dorsal margin of fourth labial half of its ventral margin; first 4 supralabials subequal along labial border; mental triangular, touching anterior chin shields; 5 infralabials, first 8 touching anterior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.041-0.052 (2 specimens); tail tapering gradually from base to a point; dorsal scales reduced to 4 rows on tail opposite third to ninth subcaudal anterior to terminal scute.
Hemipenis in holotype too small for accurate description, sex determined by examination of gonads.
Ten modified maxillary teeth (8 specimens).
Ventrals 147-155 in males (holotype 155), 154-161 in females; subcaudals 18-21 in males (holotype 18), 14-16 in females; total length males 317-830 mm. (holotype 317), females 368-370 mm.; ratio of tail length to total length 0.072-0.106 in males (holotype 0.072), 0.078-0.076 in females.
Color of holotype brown above, yellow below, the dark pigment ending abruptly on center of third scale row; dorsal scales without network; head brown above, the dark pigment ending in an oblique line from upper edge of second supralabial to lower third of fifth; supralabials yellow, the first dark brown dorsally and anteriorly; a small yellow spot near lateral margin of parietal; head immaculate yellow below; ventrals immaculate yellow; dorsal brown color ex- tending ventrally on tail to cover most of subcaudals, leaving a short yellow stripe along central part of each row of subcaudals.
98 FIELDIANA: ZOOLOGY, VOLUME 49
Fic. 25. Holotype of Calamaria hilleniusi.
Color of paratypes dark brown above, one with 21 narrow (2-8 scales wide) crossbands of lighter brown, towards rear of body cross- bands become lighter; other paratypes uniformly dark brown above; dark pigment ending abruptly along second scale row; in anterior half of body first two scale rows immaculate yellow; in posterior half scales of first two rows with dark anterior edges; head dark brown above; supralabials immaculate yellow except for their dorsal edges; head immaculate yellow below; ventrals yellow, first 10 to 25 immaculate; other ventrals with dark spots at antero-lateral cor- ners, the dark area becoming wider in posterior part of body and occupying entire width of ventrals just before vent; subcaudals yellow, edged with dark brown.
Comparisons.—Calamaria hilleniusi is similar to lumbricoidea in the relation of mental to chin shields, paraparietal and supralabial counts, and the absence of a network in the dorsal seales. It differs from lwmbricoidea in the presence of light crossbands in some adults
INGER AND MARX: THE SNAKE GENUS CALAMARIA 99
and the reduction of the ocular region, which is reflected in a smaller eye and wider frontal. The reduction to four dorsal scale rows takes place closer to the tip of the tail and the number of subcaudals is lower in male hilleniusi than in males of Borneo lumbricoidea (com- pare above description with Table 27).
Calamaria hilleniusi differs from other species related to lum- bricoidea, e. g., griswoldi and albiventer, in coloration and in the size of the ocular region.
The patterns of hilleniusi are duplicated in Borneo in C. bicolor. Calamaria hilleniusit also resembles this species in its small eye and relatively wide frontal, but differs in having a laterally oriented nasal, the prefrontal separated from the third supralabial (compare Figs. 25 and 40), the mental touching the anterior chin shields, lower subeaudal counts (18-21 versus 21-28 in males; 14-16 versus 18-21 in females), and shorter tail ratios (0.072-0.106 versus 0.106-148 in males; 0.073-0.076 versus 0.079—-0.093 in females).
Calamaria hilleniusi resembles schlegeli closely in body form, wide frontal, and small eye. The pattern of the holotype of hilleniusi is identical to that of schlegelt. Calamaria hillencusi differs from schleg- eli in having the mental touching the anterior chin shields, lower subcaudal counts (18-21 versus 27-42 in males; 14-16 versus 28 in females), smaller tail ratios (0.072-0.106 versus 0.120-0.2138 in males, 0.073-0.076 versus 0.099 in females), and thicker body diameter. The holotype of hilleniusi is known to be sympatric with schlegeli at Samarinda. The specimen of schlegeli from Samarinda ZMA 10079) is a male with the following characteristics: 186 ventrals, 34 subeaudals, and tail ratio 0.156. The difference in ventral counts between this specimen of schlegeli and the holotype of hilleniusi suggests that the latter probably has higher ventral counts.
Distribution.— Borneo (Fig. 72).
Calamaria muelleri Boulenger. Figure 26.
Calamaria muelleri Boulenger, 1896, Ann. Mag. Nat. Hist., (6), 17, p. 894— Loka, Mount Bonthain, Celebes; 1897, Proc. Zool. Soc. London, 1897, p. 223, pl. 14, fig. 1; Boettger, 1898, Kat. Rept. Samm. Senck. Naturf. Ges., pt. 2, p. 86; de Rooij, 1917, Rept. Indo-Austr. Arch., 2, p. 161; Smith, 1927, Proc. Zool. Soe. London, 1927, p. 224; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 171; de Haas, 1950, Treubia, 20, p. 272; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 204.
Lectotype.—Naturhistorisches Museum Basel 1690, here desig- nated.
100 FIELDIANA: ZOOLOGY, VOLUME 49
Taxonomic notes.—Species of Calamaria on Celebes fall into three morphological groups, one of which is characterized by modified maxillary teeth and relatively low ventral counts. This group of four species shows no particular specialization though two of the species, nuchalis and muellert (Fig. 26), have narrowed snouts.
Calamaria muellert and C. nuchalis are sympatric in southwestern Celebes in the vicinity of Mount Bonthain (Fig. 29). In that region the two differ in ventral coloration (immaculate in nuchalis, speckled in muellerz,) in lateral coloration (white stripe on first scale row in nuchalis, absent in muellerz), and in the number of scales and shields surrounding the paraparietal (6 in nuchalis, 5 in mueller). These differences also distinguish these species in central Celebes where they occur though they have not been collected at the same localities.
Calamaria muelleri is apparently sympatric with brongersmaz, the third Celebesian species having low ventral counts, in the Lake Posso area. At that place the two differ in the relative lengths of frontal and parietal shields (frontal shorter than parietal in bron- gersmar, usually longer in muellerz), relation of the mental and an- terior chin shields (separated in brongersmaz, touching in muelleri,) shape of snout (broadly rounded in brongersmai, narrow and with an enlarged rostral in muellerz), and ventral coloration (bold black squares in brongersmaz, speckled in muellerz).
The fourth of these species, boesemant, is not sympatric with muellert. The latter differs from boesemani in having the mental touching the chin shields and the frontal longer than the parietal and in lacking a light stripe on the first scale row.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; 5 shields and scales surrounding paraparietal; dorsal scales with a light or dark network; subcaudals 21 or less in males and 15 or less in females; portion of rostral visible from above at least equal to length of prefrontal suture; head distinctly tapered in front of eyes.
Description.—Head distinetly narrowed anteriorly; rostral as high as wide, portion visible from above equal to or longer than prefrontal suture; prefrontal 24 length of frontal, touching first 2 sup- ralabials; frontal hexagonal, 134 to 3 times width of supraocular, equal to or longer than length of parietal; parietal 11% times length of prefrontal; paraparietal surrounded by 5 shields and scales; nasal smaller than postocular; preocular present; neither ocular as high as eye; eye equal to or slightly greater than eye-mouth distance; 5
INGER AND MARX: THE SNAKE GENUS CALAMARIA 101
supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal or second slightly larger; mental triangular, touching anterior chin shields; 5 infralabials, first 3 touching anterior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.007-0.042 (10 specimens); tail gradually tapering to a moderate point; dorsal scales reduce to 4 rows on tail opposite second to eighth subcaudal anterior to terminal scute.
Hemipenis forked opposite fifth or sixth subcaudal; retractor beginning opposite seventh to ninth subcaudal, calyces smooth (4 specimens). Cloaca cardioid (1) or bulbous (1).
Nine to 11 modified maxillary teeth (8 specimens).
Ventrals: males, 129-155 (mean 146.7; N=80); females, 155-178 (mean 167.7; N=388). Subcaudals: males, 16-21 (mean 18.5; N=29); females, 9-15 (mean 12.8; N=88).
Total length: males, 116-262 mm.; females, 186-855 mm. Ratio of tail to total length: males, 0.072-0.101 (mean 0.089; N=29); females, 0.041-0.062 (mean 0.050; N=87).
Color brown above, each scale with a fine dark network; scattered scales with dark central spot; spots not forming lines; first scale row usually lighter than those above but without light longitudinal stripe; head brown, densely spotted with blackish brown; a dark stripe usually running from nasal back through eye along lower por- tion of prefrontal and upper margins of supralabials, descending slightly to include upper half or two-thirds of last supralabial and lower edge of parietal; remainder of supralabials yellow with a dark stripe along lower edge or with dark sutures; underside of head yellow with dark spots; ventrals highly variable but always with dark brown pigment, in the form of small spots in longitudinal rows, in the form of speckling, or as transverse bands along anterior half of each ven- tral; underside of tail with a broad median brown stripe and a nar- rower brown stripe along outer edges of subcaudals, the two stripes separated by a yellow stripe; in some specimens underside of tail dark brown with a few isolated spots.
Distribution.—Central and southwestern Celebes (Fig. 29).
CELEBES (NMS 2 unnumbered; ZMB 15688), south Celebes (BM 96.4.29.82-33): Mount Bonthain (NHMW 16691; ZSBS 545/20), Djikoro (AMNH 68381-82; BM 1926.8.20.159-166; CNHM 83162; MCZ 25815-24, 25326-33; USNM 120818), Loka (CNHM 104619; NMB 1690—lectotype, 1691-93—-paratypes, 5227), Idrulaman(BM
102 FIELDIANA: ZOOLOGY, VOLUME 49
96.4.29.27[2]), Bua Praeng (NHMW 16692: 1-10; SNG 19942-43) ; Luhu (BM 96.12.9.67), between Lake Posso and Gulf of Tomini (NMB 1694).
Fic. 26. Paratype of Calamaria muelleri (NMB 1692).
No data (BM 1946.1.1.82-84; MZUS 2 unnumbered).
?Java (ZMB 13946, 13948/3]). As these snakes were collected by the same person (Fruhstorfer) and on the same date as SNG 19942-438, they certainly must have come from Celebes.
Specimens examined.—b68
Calamaria joloensis Taylor
Calamaria joloensis Taylor, 1922, Phil. Jour. Sci., 21, p. 203—central Jolo Island, Philippine Islands; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 170; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 203; Leviton, 19638, Proc. Calif..Acad. Sci., 31, p. 387.
Holotype.—California Academy of Sciences 60901.
Taxonomic notes.—The holotype has the lowest ventral count we have observed in the genus. The number of supralabials, eye size, position of nasal, and relation of mental and chin shields recall hilleniusi of Borneo. But joloensis differs from that species in having relatively longer parietals (equal to half of the head length), a blunter tail, fewer ventrals, and light network on each dorsal scale. Except for the much smaller eye and lower ventral count, 7oloenszs is similar to gervaist of the Philippine Islands.
As the unique specimen of joloensis is not sympatric with either hilleniust or gervaisi, a certain decision as to the specific differentia- tion of joloensis is not possible now. However, because we have not
INGER AND MARX: THE SNAKE GENUS CALAMARIA 103
observed intraspecific variation in eye size that could account for the difference between gervaisi and joloensis, we recognize the latter as a full species. For a similar reason, the shape of the head suggests that joloensis is specifically distinct from halleniusze.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labial entering orbit; preocular present; mental touching anterior chin shields; eye about two-thirds eye-mouth distance; dorsal scales with a light network.
Description.—Rostral broader than high, portion visible from above about half length of prefrontal suture; prefrontal shorter than frontal, touching first 2 supralabials; frontal hexagonal, 3 times width of supraocular, about 24 length of parietal; parietal 114 times length of prefrontal; paraparietal surrounded by 5 shields and scales; nasal larger than postocular; preocular present; neither ocular as high as eye; eye ’/1) of eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, first, third, and fourth sub- equal and smaller than second; mental triangular, touching anterior chin shields; 5 infralabials, first 3 touching anterior chin shields; both pairs of chin shields meeting in midline; 8 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.010; tail short, thick, tapering near end to a blunt tip; dorsal scales reduce to 4 rows on tail opposite sixth subcaudal anterior to terminal scute.
Hemipenis bifurcate. Eight modified maxillary teeth.
Ventrals 119 in male; subcaudals 138; total length 144 mm.; ratio of tail length to total length 0.069.
Color faded in preservative but dark brown above, each scale with a light network; head dark brown above and on sides; head below evidently with brown spots on labials; ventrals yellowish, those in anterior part of body with dark pigment in antero-lateral corners, increasing amounts of dark pigment on ventrals farther back on body; in posterior two-thirds of body most ventrals with a dark band across anterior halves; underside of tail with dark pigment but pattern too faded for accurate observation.
Distribution.—Jolo Island, Philippine Islands (Fig. 68). CENTRAL JOLO ISLAND (SU 60901—holotype).
104 FIELDIANA: ZOOLOGY, VOLUME 49
Calamaria bitorques Peters. Figure 27.
Calamaria bitorques Peters, 1872, Monatsber. Akad. Wiss. Berlin, 1872, p. 585 —Philippine Islands; Boulenger, 1894, Cat. Snakes Brit. Mus., 2, p. 338; 1896, Cat. Snakes Brit. Mus., 3, p. 646; Boettger, 1886, Ber. Senck. Naturf. Ges., 1886, p. 105; Taylor, 1922, Snakes Phil. Ids., p. 185; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 170; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 200; Leviton, 1968, Proc. Calif. Acad. Sci., 31, p. 390.
Calamaria gervaisi (non Duméril and Bibron), Boettger, 1898, Kat. Rept. Samm. Senck. Naturf. Ges., pt. 2, p. 83 (part). Holotype.—Zoologisches Museum Berlin 7444.
Taxonomic notes.—C. bitorques is very similar to C. gervaisz in counts and shape of head scales. The two differ in coloration and size. The former has black edged crossbands on the anterior part of the body (Fig. 27). C. gervaist may have narrow yellow rings behind the head forming a wide cross band, but the band thus formed does not have dark edges. The difference in color pattern is not an effect of ontogenetic change for the characteristic pattern of betorques is present in specimens in the size range 150-260 mm., which is well within the size range of gervazst.
The largest gervaist we have seen measured 821 mm. None of the more than 80 gervaist from Luzon exceeded 290 mm. By con- trast 11 of 20 b¢torques (known from Luzon only) were longer than 300 mm., with a maximum of 420 mm.
We have examined specimens of both species from Manila, Mount Isarog (Camarines Sur), and Bulusan (Sorsogon). The size and coloration differences just noted distinguish these two species at each of these localities.
These species also differ in average number of subcaudals. Fe- male gervaisi from Luzon have 10-15 subeaudals (mean 12.67, mode 13, N=78); female bitorques have 12-17 (mean 15.3, mode 15, N=14). Male gervaisi from Luzon have 15-20 subeaudals (mean 16.68, mode 16, N=65); male brtorques have 17-20 (mean 18.83, N=6). Female gervaist from Manila have 10-14 subcaudals (80 specimens) and fe- male bitorques 15-17 (2). Female gervaisi from Mount Isarog have 11-18 subcaudals (4 specimens) and female bitorques 14-15 (2). A single female of gervaisi from Bulusan has 16 subcaudals and a single female of betorques 14. Males of both species were not available from these localities.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; paraparietal surrounded by 5 shields and scales; 2 to 6 dark, black-edged, crossbands behind the head.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 105
Description.—Rostral as wide as high, portion visible from above about 24 length of prefrontal suture; prefrontal shorter than frontal, touching first 2 supralabials; frontal hexagonal, 114 to 2 times width of supraocular, about °/; length of parietal; parietal 11% times length of prefrontal; paraparietal surrounded by 5 scales and shields; nasal much smaller than postocular; preocular present; postocular not as high as eye; eye equal to or slightly greater than eye-mouth distance;
Fic. 27. Holotype of Calamaria bitorques.
5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal; mental triangular, touching anterior chin shields; 5 infralabials, first 3 touching anterior chin shields; both pairs of chin shields meeting in midline; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.016—-0.063 (7 specimens); tail thick, ta- pered near end to blunt point; dorsal scales reduce to 4 rows on tail opposite sixth to eleventh subcaudal anterior to terminal scute.
Cloaca of females bilobed (1) or cardioid (1). Nine modified maxillary teeth (2 specimens).
106 FIELDIANA: ZOOLOGY, VOLUME 49
Ventrals: males, 150-157 (mean 154.0; N=5); females, 157-197 (mean 173.4; N=14). Subcaudals: males, 17-20 (mean 18.8; N=6); females, 12-17 (mean 15.3; N=14).
Total length: males, 150-833 mm.; females, 222-420 mm. Ratio of tail to total length: males, 0.066-0.087 (mean 0.076; N=5); fe- males, 0.039-0.086 (mean 0.057; N=14).
Color light to dark brown above; dorsal scales with fine dark network; 2 to 6 dark-edged saddles on anterior part of body reaching the second or third scale row; behind area of saddles dorsum usually with scattered small, dark spots; scales of first row usually immac- ulate yellow at least in anterior half of body; head brown above with obscure dark markings; supralabials usually cream-colored without dark spots; a dark streak extending from nasal across lower edges of prefrontal and parietal and upper edge of last labial; underside of head yellowish, usually immaculate; ventrals immaculate yellow or with dark lateral edges; underside of tail immaculate yellow or with narrow dark mid-ventral streak.
Distribution.—Luzon (Fig. 68).
LUZON (BM 72.8.20.49; ZMB 3778, 7444-holotype, listed in Ber- lin Museum catalogue as from “‘Luzon’’), central Luzon (SNG 19394). Cagayan Province: Cape Engano (BM 95.9.21.4). Isabela Province (CAS 15295). Rizal Province: Manila (MHNP 00-864- 65), area between Subic Bay south to Clark Air Force Base (MVZ 73672). Camarines Sur Province: Mt. Isarog, Curry, Pili (CNHM 142515-16). Sorsogon Province: Bulusan (CNHM 142510-14).
PHILIPPINE ISLANDS (BM 72.10.11.16, 72.10.11.20, 1 unnum- bered; MCZ 25776).
Specimens examined.—20.
Calamaria gervaisi Duméril and Bibron. Figure 28.
Calamaria gervaisii Duméril and Bibron, 1854, Erp. Gén., 7, p. 76—‘‘Java’”’ {in error]; Gtinther, 1858, Cat. Colubrine Snakes Brit. Mus., p. 4; Jan, 1862, Arch. Zool. Anat. Phys., 2, p. 8; 1865, Icon. Ophid., 10, pl. 2, fig. 1; Fischer, 1885, Jahrb. Wiss. Anst. Hamburg, 2, p. 80; Boettger, 1886, Ber. Senck. Naturf. Ges., 2, p. 105; 1898, Kat. Rept. Samm. Senck. Naturf. Ges., pt. 2, p. 83; Boulenger, 1894, Cat. Snakes Brit. Mus., 2, p. 338; Taylor, 1922, Snakes Philippine Ids., p. 186; Werner, 1929, Zool. Jahrb., (Syst.), 57, p. 171; Marx and Inger, 1955, Fieldiana, Zool., 37, p. 180.
Calamaria gervaisii gervaisti, Taylor, op, cit., p. 187; Leviton, 1963, Proc. Calif. Acad. Sci., 31, pp. 390, 394, 407.
INGER AND MARX: THE SNAKE GENUS CALAMARIA 107
Calamaria mindorensis Boulenger, 1895, Ann. Mag. Nat. Hist., (6), 16, p. 481 Mindoro; 1896, Cat. Snakes Brit. Mus., 3, p. 646; Taylor, op. cit., p. 190; Werner, loc. cit., p. 171; Marx and Inger, loc. cit., p. 207.
Calamaria gervaisii iridescens Taylor, 1917, Phil. Jour. Sci., 12, p. 360— Can- laon Volcano, Negros; 1922, op. cit., p. 188; Brown and Alcala, 1961, Ecology, 42, p. 632; Leviton, loc. cit., pp. 384, 398, 402.
Calamaria tropica Taylor, 1922, Snakes Philippines Ids., p. 194—-near Naujan,
Mindoro; Marx and Inger, loc. cit., p. 206.
Calamaria polillensis Taylor, 1923, Phil. Jour. Sci., 22, p. 549—Polillo, Polillo
Island; Werner, loc. cit., p. 171; Marx and Inger, loc. cit., p. 205.
Calamaria hollandi Taylor, 1923, Phil. Jour. Sci., 22, p. 550—Port Holland, Basilan; Werner, loc. cit., p. 171; Marx and Inger, loc. cit., p. 202.
Calamaria gervaisi hollandi, Leviton, loc. cit., pp. 379, 393.
Calamaria gervaisit polillensis, Leviton, loc. cit., p. 403.
Syntypes.—Museum National d’Histoire Naturelle ¢2314~-7202, two specimens labeled “‘Java’’. No author subsequent to Duméril and Bibron (1854) has reported this species from outside the Philip- pine Islands. We now believe, contrary to our previous opinion (Marx and Inger, 1955), that the type locality cannot be restricted to a particular island within the Philippines except arbitrarily.
Taxonomic notes.—We have examined the holotype of mindorensis Boulenger and cannot distinguish it from gervazst. Boulenger (1895) did not compare mindorensis with any other Calamaria. Although our counts (ventrals 177, subcaudals 17) on the holotype differ from Boulenger’s (ventrals 193, subcaudals 15) we are certain that the specimen examined is the holotype of mindorensis because of the agreement in coloration, size (total length 232 mm., tail length 13 mm., according to our measurements; 240 and 13 mm., according to Boulenger), and shield arrangement. Furthermore it is the only Cala- maria from Mindoro in the collection of the British Museum.
Calamaria tropica Taylor was distinguished from all other species on the basis of the presence of loreals. These “‘loreals’ are tiny, triangular scales much smaller than even the nasal and are wedged between prefrontal, preocular, and second supralabial. Though one of these scales appears on each side, they differ in size and we can only conclude that they represent fragments of the prefrontals. An additional specimen of gervaist (SU 21804) has a similar fragment on one side of the head. The holotype of tropica is certainly conspecific with the other Mindoro specimens of gervaist we have examined.
Calamaria polillensis was distinguished by Taylor (19238) from gervaist on the basis of its lower ventral and subcaudal counts and the presence of a neck ring. As may be seen in Tables 28-31, the
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110 FIELDIANA: ZOOLOGY, VOLUME 49
counts of the holotype and paratype we have seen fall within the ranges of gervaisi from other islands; the two paratypes not examined lie within the same range of variation.
Taylor (ibid.) compared C. hollandi with polillensis, which we have just shown to be a synonym of gervazsz, but not to gervaisi. The character—extent of the rostral visible from above—by which Taylor distingiushed holland: from polillensis shows considerable variation in gervais?. The direct comparison of the holotype of hollandi to various populations of gervaisz does not reveal any differences of diagnostic value.
Taylor (1917) distinguished his Negros form, C. gervaisz iridescens, from gervaist from other islands in the Philippines by ventral and subeaudal counts and by size. Marx and Inger (1955) substantiated the count differences on the basis of 17 specimens from Luzon and Mindanao. However, with larger series available these differences do not appear to be taxonomically significant (Tables 28-31).
Only the holotype of zridescens is unusually long (306 mm.). The largest of the 5 paratypes measured 240 mm. Of the gervaisi we measured, 18 of 97 from Luzon, 1 of 2 from Tablas, 4 of 8 from Panay, 1 of 9 from Mindanao, and 10 of 49 from Negros exceeded 250 mm.
Fic. 28. Calamaria gervaisi (CNHM 150381).
INGER AND MARX: THE SNAKE GENUS CALAMARIA 111
As approximately the same proportion of the Luzon and Negros samples fall in this size range, the supposed difference between 77r7- descens and other gervaisz has no taxonomie significance.
Diagnosis.—Maxillary teeth modified; third and fourth supra- labials entering orbit; preocular present; mental touching anterior chin shields; paraparietal surrounded by 5 shields and scales; sub- caudals of males under 28, of females under 20; no light stripes above second scale row; a dark-edged, interrupted, light stripe on first scale row usually present; no dark-edged dorsal saddles; head not conspicuously tapered before eyes.
Description.—Rostral broader than high or higher than broad, portion visible from above 14 to equal to prefrontal suture; pre- frontal about */; length of frontal, touching first 2 or 3 supralabials: frontal hexagonal, 114 to 21% times width of supraocular, about 34 to */; length of parietal; parietal 114 to 124 times length of prefrontal; paraparietal surrounded by 5 shields and scales (rarely 3, 4, or 6); nasal smaller than postocular; preocular present; neither ocular as high as eye; eye slightly less than or slightly greater than eye-mouth distance; 5 supralabials, third and fourth entering orbit, fifth the largest, first 4 subequal, or fourth narrower than others; mental triangular, touching anterior chin shields; 5 infralabials, first 3 touch- ing anterior chin shields; both pairs of chin shields meeting in mid- line; 3 gulars in midline between posterior chin shields and first ventral.
Body thickness index 0.007-0.030 (21 specimens); tail thick, tapering gradually to a point; dorsal scales reduce to 4 rows on tail opposite third to twelfth subcaudal anterior to terminal scute.
Hemipenis forked opposite fourth to sixth subcaudal; retractor muscle beginning opposite ninth or tenth subcaudal; sulcus bifurcate; calyces smooth (2 specimens). Cloaca of female shallowly bilobed (3) or cardioid (1).
Eight or 9 modified maxillary teeth (9 specimens).
Ventrals: males, 182-164 (mean 151.2; N=81); females, 142-190 (mean 166.1; N=101) (Tables 28 and 29). Subcaudals: males, 15- 21 (mean 17.2; N=110); females, 10-18 (mean 13.2; N=129) (Tables 30 and 81).
Total length: males, 79-251 mm.; females, 104-321 mm. Ratio of tail to total length: males, 0.063-0.096 (mean 0.075; N=81); females, 0.041-0.070 (mean 0.051; N=98).
Color light or dark brown above; each scale above first row yellowish with a dark network; some scales with a dark spot forming
112 FIELDIANA: ZOOLOGY, VOLUME 49
longer or shorter dark stripes; scales of first row dark or with clear yellow centers forming a row of yellow dots; head above same color as body with irregular dark spots; lower half of supralabials yellowish, occasionally sutures dark brown; 1 or 2 complete or incomplete yellow rings on head and neck, present or absent (Table 32); under- side of head yellowish, usually with small dark spots; ventrals