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An inconspicuous, conspicuous new species of Asian pipesnake, genus Cylindrophis (Reptilia: Squamata: Cylindrophiidae), from the south coast of Jawa Tengah, Java, Indonesia, and an overview of the tangled taxonomic history of C. ruffus (Laurenti, 1768)

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Abstract and Figures

We describe a new species of Cylindrophis currently known only from Grabag, Purworejo Regency, Jawa Tengah Province (Central Java), Java, Indonesia. Cylindrophis subocularis sp. nov. can be distinguished from all congeners by the presence of a single, eponymous subocular scale between the 3rd and 4th or 4th and 5th supralabial, preventing contact between the 4th or 5th supralabial and the orbit, and by having the prefrontal in narrow contact with or separated from the orbit. We preface our description with a detailed account of the tangled taxonomic history of the similar and putatively wide-ranging species C. ruffus, which leads us to (1) remove the name Scytale scheuchzeri from the synonymy of C. ruffus, (2) list the taxon C. rufa var. javanica as species inquirenda, and (3) synonymize C. mirzae with C. ruffus. We provide additional evidence to confirm that the type locality of C. ruffus is Java. Cylindrophis subocularis sp. nov. is the second species of Asian pipesnake from Java.
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Accepted by Z. Nagy: 12 Jan. 2016; published: 21 Mar. 2016
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4093 (1): 001
An inconspicuous, conspicuous new species of Asian pipesnake,
genus Cylindrophis (Reptilia: Squamata: Cylindrophiidae),
from the south coast of Jawa Tengah, Java, Indonesia, and
an overview of the tangled taxonomic history of C. ruffus (Laurenti, 1768)
Department of Animal Evolution and Systematics and Zoological Collection Marburg, Faculty of Biology, Philipps-Universität Mar-
burg, Karl-von-Frisch-Straße 8, 35032 Marburg, Germany
Faculty of Sciences and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton, WV1 1LY, United Kingdom;
and West Midland Safari Park, Bewdley, Worcestershire DY12 1LF, United Kingdom
Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and Department of Verte-
brate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA
Corresponding author. E-mail:
Co-first authors, listed in alphabetical order
We describe a new species of Cylindrophis currently known only from Grabag, Purworejo Regency, Jawa Tengah Pro-
vince (Central Java), Java, Indonesia. Cylindrophis subocularis sp. nov. can be distinguished from all congeners by the
presence of a single, eponymous subocular scale between the 3
and 4
or 4
and 5
supralabial, preventing contact be-
tween the 4
or 5
supralabial and the orbit, and by having the prefrontal in narrow contact with or separated from the
orbit. We preface our description with a detailed account of the tangled taxonomic history of the similar and putatively
wide-ranging species C. ruffus, which leads us to (1) remove the name Scytale scheuchzeri from the synonymy of C. ruf-
fus, (2) list the taxon C. rufa var. javanica as species inquirenda, and (3) synonymize C. mirzae with C. ruffus. We provide
additional evidence to confirm that the type locality of C. ruffus is Java. Cylindrophis subocularis sp. nov. is the second
species of Asian pipesnake from Java.
Key words: Cylindrophis subocularis sp. nov., C. ruffus, Serpentes, Cylindrophiidae, Asian pipesnakes, species complex,
morphology, Central Java, Indonesia, Greater Sunda Islands
Wir beschreiben eine neue Art der Gattung Cylindrophis, die gegenwärtig nur aus Grabag, Purworejo, Jawa Tengah
(Zentral-Java), Java, Indonesien, bekannt ist. Cylindrophis subocularis sp. nov. unterscheidet sich von allen anderen Arten
dieser Gattung durch das Vorhandensein einer einzelnen, namensgebenden Subokular-Schuppe, die sich zwischen das
dritte und vierte oder das vierte und fünfte Supralabial-Schild schiebt, und den Kontakt zwischen dem vierten oder fünften
Supralabiale und dem Auge verhindert. Zudem steht das Präfrontale in minimalem Kontakt mit dem Auge oder ist von
diesem separiert. Wir stellen unserer Beschreibung einen detaillierten Überblick über die verworrene Taxonomie-Ge-
schichte der ähnlichen und scheinbar weit verbreiteten Art C. ruffus voran, was uns dazu veranlasst (1) den Namen Scytale
scheuchzeri aus der Synonymie von C. ruffus herauszunehmen, (2) C. rufa var. javanica als species inquirenda zu
betrachten, und (3) C. mirzae mit C. ruffus zu synonymisieren. Wir liefern weitere Hinweise für die Berichtigung der
Typuslokalität von C. ruffus auf Java. Bei Cylindrophis subocularis sp. nov. handelt es sich um die zweite auf Java
vorkommende Asiatische Walzenschlange.
Schlüsselwörter: Cylindrophis subocularis sp. nov., C. ruffus, Serpentes, Cylindrophiidae, Asiatische Walzenschlangen,
Art-Komplex, Morphologie, Zentral-Java, Indonesien, Große Sundainseln
Zootaxa 4093 (1) © 2016 Magnolia Press
The genus Cylindrophis. The henophidian snake genus Cylindrophis Wagler, 1828 currently comprises 13
secretive, semifossorial species, including C. aruensis Boulenger, 1920; C. boulengeri Roux, 1911; C. burmanus
Smith, 1943; C. engkariensis Stuebing, 1994; C. isolepis Boulenger, 1896; C. jodiae Amarasinghe et al., 2015; C.
lineatus Blanford, 1881; C. maculatus (Linnæus, 1758); C. melanotus Wagler, 1828; C. mirzae Amarasinghe et al.,
2015; C. opisthorhodus Boulenger, 1897; C. ruffus (Laurenti, 1768); and C. yamdena Smith & Sidik, 1998 (see
Wallach et al. 2014; Amarasinghe et al. 2015). These snakes are collectively referred to as Asian pipesnakes due to
their cylindrical appearance, with a body of near-uniform diameter. Members of the genus are small- to medium-
sized (total length 125–857 mm), rather stout-bodied snakes that may be defined on the basis of the following
eidonomic characters: (1) a relatively blunt head with minute eyes, head not distinct from neck, bearing a mental
groove; (2) absence of true gastrosteges, with ventral scales only slightly larger than or equal in size to dorsal
scales; (3) presence of a pair of pelvic spurs (= cloacal spurs) in both sexes; (4) a very short tail, often with
conspicuous ventral coloration; and (5) contrasting light and dark ventral blotching (e.g., de Rooij 1917; Smith
1943; Taylor 1965; Greene 1973; pers. obs.). The conspicuous ventral color pattern plays a vital role in the
defensive behavior of Cylindrophis species. When threatened, pipesnakes will flatten the posterior portion of their
body and arch it above the ground to display their ventral pattern, while the head remains concealed among the
body coils (e.g., Flower 1899; Barbour 1912; Smith 1927, 1943; Campden-Main 1970; Deuve 1970; Greene 1973).
Distribution. Cylindrophis is a widely distributed genus (Flower 1899; de Rooij 1917; Smith 1943; Lal Hora
& Jayaram 1949; Taylor 1965; Campden-Main 1970; Deuve 1970; McDowell 1975; in den Bosch 1985; Stuebing
1991; Adler et al. 1992; Iskandar 1998; Zug et al. 1998; McDiarmid et al. 1999; Orlov et al. 2000; de Lang 2011)
with species occurring from Sri Lanka (one species) throughout the continental and insular parts of Southeast Asia
(12 species currently recognized). In Southeast Asia the genus is distributed from southern China and Hong Kong
through Vietnam, Laos, Cambodia, Thailand, Myanmar, Peninsular Malaysia, and Singapore including Singapore,
south to the Greater Sunda Islands (Borneo, Sumatra, Java, as well as some of their offshore islands), Sulawesi, the
Lesser Sunda Islands (Lombok, Komodo, Flores, Sumbawa, Timor), and east to the Maluku Islands (Halmahera,
Wetar, Damar, Babar, and into the Tanimbar Archipelago); the eastern distributional limit, the Aru Islands, was
considered questionable by Iskandar (1998). However, within this vast range, smaller-scale zoogeographic
patterns, phylogenetic relationships, and even the true species richness of the genus remain poorly known.
Many species of Cylindrophis, especially those from the eastern end of the distribution (e.g., C. aruensis, C.
boulengeri, C. isolepis, C. yamdena), are known from very few specimens (McDowell 1975; Iskandar 1998; Smith
& Sidik 1998). This is likely due to both the remoteness of the eastern Indonesian islands and the secretive lifestyle
of these snakes, and Cylindrophis diversity in this region may still be underestimated. Even on Borneo, an island
with a relatively well-studied herpetofauna (Das 2004), Stuebing (1994) discovered C. engkariensis, a species with
a potentially very restricted range. More recently, Amarasinghe et al. (2015) described two new species (one from
Singapore and one from Vietnam) that had been masquerading under the name C. ruffus. However, the descriptions
and redescriptions (including of C. ruffus) presented by these authors contain some inaccuracies, including
descriptive errors, which unfortunately increase the complexity of an already intricate taxonomic situation.
The problematic nature of Cylindrophis ruffus. Compared with other members of the genus, the species
Cylindrophis ruffus sensu historico (e.g., Schlegel 1837b, 1837−1844; de Rooij 1917; Smith 1943; for a definition
of the term sensu historico see below) exhibits an extraordinarily wide distribution, extending from mainland
Southeast Asia across most parts of the Greater Sunda Islands into eastern Java (de Rooij 1917; Smith 1943; Taylor
1965; McDiarmid et al. 1999; Wallach et al. 2014). It was already identified as a species complex (Smith & Sidik
1998) and it appears to include several undescribed taxa (Amarasinghe et al. 2015; Mecke et al., in prep.). Despite
its redescription by Amarasinghe et al. (2015), both the morphological definition and the geographic range limits
of C. ruffus sensu stricto remain unsettled. Cylindrophis ruffus sensu historico appears to be common, frequently
encountered (Smith 1943; Taylor 1965; Campden-Main 1970; Kupfer et al. 2003), and well represented in museum
collections, but a comprehensive taxonomic revision of this group has never been conducted. While it is evident
that the taxonomy of C. ruffus is flawed, its complex taxonomic history, the absence of a type specimen, and an
incorrect type locality (“Surinami”) have stood in the way of developing a stable taxonomic hypothesis (Boie 1827;
Schlegel 1837a, b; McDiarmid et al. 1999; Wallach et al. 2014). Furthermore, due to the age of available museum
specimens in general, and of type material in particular, it is only through a thorough morphological study
encompassing the entire range and variation of C. ruffus that its taxonomy can be resolved.
Zootaxa 4093 (1) © 2016 Magnolia Press
History necessitates three working definitions of Cylindrophis ruffus. As part of our comprehensive review
of the genus Cylindrophis, we examined several hundred museum specimens listed by the available collection data
as C. ruffus. We noted that, given the long history of C. ruffus in the literature and the morphological diversity of
examined specimens, three definitions of C. ruffus as a taxonomic unit became necessary to permit a complete
understanding of how different authors through time dealt with the taxon. Our most inclusive definition for the
taxon is ‘C. ruffus sensu historico
,’ which includes all forms historically considered to be part of C. ruffus at one
time or another, but before the revision of Amarasinghe et al. (2015). This definition includes C. burmanus as well
as the forms that were recently described as C. jodiae and C. mirzae by Amarasinghe et al. (2015); it essentially
covers forms from all over Southeast Asia and into the Indonesian archipelago. The second, more specific
definition is ‘C. ruffus sensu lato,’ which excludes C. burmanus and C. jodiae, but still includes the weakly defined
C. mirzae as well as populations from Borneo, Java, Sumatra, and Peninsular Malaysia. Specimens north of
Peninsular Malaysia belong either to C. burmanus or C. jodiae (pers. obs.). Our third definition is ‘C. ruffus sensu
stricto,’ by which we refer to the true species C. ruffus.
An unusual population from Java. As we progressed with our study, we noticed that a particular specimen
series was sufficiently different from C. ruffus sensu historico to warrant recognition as a distinct species, even
while our review of C. ruffus was still in progress. Specifically, our work in the collections at the Naturalis
Biodiversity Center in Leiden, the Netherlands (formerly the Rijksmuseum van Natuurlijke Historie; RMNH), the
Natural History Museum in Vienna, Austria (NMW), and the Museum für Naturkunde Berlin, Germany (ZMB),
revealed several specimens labeled as C. ruffus that had apparently been collected at a single, isolated locality on
the Indonesian island of Java, and which allowed easy differentiation from all other forms of Cylindrophis by a
unique character: the presence of a subocular scale. We here describe this species, which is currently only known
from Grabag, Purworejo Regency, Jawa Tengah Province (Central Java), Indonesia, and provide an historical
overview of C. ruffus taxonomy.
Material and methods
Morphological characters. For each specimen of the new species (n = 8) and all specimens used for comparison
(n = 451), we recorded data for 52 morphological characters. Of these, 37 were metric, eight meristic, and seven
qualitative. In the list below, character names are provided in bold, followed by their definitions.
The following metric characters were obtained (characters used for the calculation of ratios are abbreviated for
convenience): snout-vent length (SVL), measured from tip of snout to cloaca; tail length (TL), measured from
cloaca to tip of tail; body diameter (BD), calculated as the mean of body height and body width at midbody; head
length (HL), measured from tip of snout to articulation of quadrate bone; head width (HW), measured at level of
anterior margin of parietals; snout length (SL), measured from tip of rostral to anterior margin of orbit; snout
width (SW), measured at level of nares; eye diameter (ED), measured as length of orbit; interorbital distance
(IOD), measured as shortest distance between orbits across head; naso-orbital distance (NOD), measured from
posterior margin of naris to anterior margin of orbit; internarial distance, measured between interior margins of
nares; length of prefrontal-eye contact (PrefO), measured at prefrontal margin bordering orbit. We also measured
the following head scale characters (dimensions of these scale characters are expressed as the maximal length,
height, or width): rostral height and width; nasal length and height; prefrontal length and width; frontal length and
width; parietal length and width; supraocular length and width; postocular length and height; anterior temporal
length and height; upper posterior temporal length and height; mental height and width; anterior chin shield length
and width; posterior chin shield length and width; and mental groove length. SVL and TL were measured to the
nearest 1 mm by gently straightening the respective specimen along a metric ruler. All other metric characters were
measured to the nearest 0.1 mm under a stereomicroscope using digital calipers and a measuring magnifier. We
also calculated the following ratios: TL/SVL, BD/SVL, HL/SVL, HW/HL, SL/HL, SW/SL, ED/HL, IOD/HL,
NOD/HL, and PrefO/ED.
1. The term sensu historico has been used by scholars in the classical sciences (specifically of the languages of Ancient Greece and Ancient Rome)
to indicate that a term is used within an historical context, as opposed to a direct translation. We borrow this term to distinguish between a taxon
as historicall y defined and one based on the most current taxonomy.
Zootaxa 4093 (1) © 2016 Magnolia Press
The following meristic characters were counted: number of dorsal scale rows, counted in an inverse ‘V’ shape
(to include all dorsal scales developmentally associated with a single pair of ribs) at (A) one head length behind
head, (B) at midbody, and (C) one head length before cloaca (displayed in a formula as A/B/C); ventrals,
beginning with the gular scale bordered by posterior chin shields; subcaudals, counted from cloaca to end of tail,
excluding terminal spine (this count included, if present, a single row of multiple small scales bordering cloaca,
counted as one subcaudal); postoculars; temporals, including (a) number of anterior temporals and (b) number of
posterior temporals, expressed in a formula as a + b; number of supralabials; number of infralabials; and number
of light transverse ventral blotches present along body, beginning with first blotch behind head to last blotch
anterior to cloaca. Head scales occurring bilaterally were counted on (a) the right and (b) the left side of the body.
We use the formula a|b when counts are different on either side of the body; a single value for a bilaterally
occurring head scale character indicates that counts on both sides of the body resulted in an identical value.
The system of counting ventral scales described by Dowling (1951) is not applicable to anilioid snakes
(Aniliidae, Anomochilidae, Cylindrophiidae, Uropeltidae) because these, unlike more advanced snakes, have no
true gastrosteges and no preventral scales. Gower & Ablett (2006) therefore proposed a ventral-counting system for
these snakes that includes every scale between the mental and cloacal scute. We did not apply their system, because
all members of the genus Cylindrophis possess a mental groove formed by the first pair of infralabials and two
pairs of enlarged chin shields, with the latter morphologically distinct from the smaller scales bordering them
posteriorly. Consequently, ventral scales were counted from the first unpaired scale positioned medially behind the
mental groove to the, often slightly enlarged, scale anterior to the divided cloacal scute.
In terms of qualitative characteristics, we recorded the specific supralabials contacting the orbit; the specific
infralabials contacting the chin shields; the condition of the cloacal scute (divided or entire); and pattern and
coloration of head, dorsum, venter, and tail. For descriptions of pattern and coloration we applied the terminology
of Köhler (2012). Numbers in parentheses behind the respective capitalized color name refer to the coding therein.
Sex was determined by the presence of testes or ovaries and oviducts and only if ventral incisions into the body
cavity already existed.
Comparative material. Comparative morphological data were obtained primarily from museum specimens
examined by the authors. Only for comparisons with Cylindrophis aruensis and C. yamdena did we use data from
the original species descriptions or other relevant literature.
We compared the new species to 451 specimens from across the range of Cylindrophis, housed in the following
collections (abbreviations follow Sabaj Pérez [2014]): AMNH, MHNG, MTD (= MTKD), NMB, NMBE, NMW,
RMNH, SMF, ZMA (now in Naturalis, Leiden; RMNH), ZMB, ZMH, and ZRC. Since the examined material used
for species delineation included (1) very distinct species not easily confused with the new species, and (2) 231
specimens of C. ruffus sensu lato, our Appendix includes only a relevant subsample of museum specimens used for
direct comparisons herein, most notably specimens of C. ruffus sensu lato from Java, including 53 specimens with
precise localities (e.g., towns, regencies) and 60 lacking exact locality data (specimens labelled only as collected on
‘Java’). Although C. mirzae might ultimately be considered a valid species, we herein refrain from differentiating
between C. mirzae and C. ruffus for reasons outlined in the taxonomic history section.
Statistical analyses. Since our new species is sufficiently distinct from congeneric taxa by a multitude of
characters (see Results: Comparisons), and with a revision of C. ruffus in progress, our statistical analyses for this
study focused exclusively on revealing characters to distinguish between the new species and C. ruffus from Java
(the type locality of C. ruffus; see Results: History leads to the type locality of Cylindrophis ruffus). Meristic
characters that were constant between the groups or exhibited two expressions only were excluded from all
statistical analyses.
For statistical tests, the data analysis software R (R-Core Team, version 3.1.3) was used. The normality
assumption for individual variables (i.e., of the metric and meristic characters, and ratios defined above) was tested
with a Shapiro-Wilk statistic. Prior to variance analyses (see below), tested metric variables were adjusted to the
mean SVL across all groups, in order to minimize variance due to possible ontogenetic variation between different
populations (e.g., Thorpe 1975, 1983; Turan 1999; Vogel et al. 2007; van Rooijen & Vogel 2008, 2010, Mecke et
al. 2013). The equation for the adjustment of data follows Vogel et al. (2007), van Rooijen & Vogel (2008, 2010),
and Mecke et al. (2013):
= Y
– ß * (SVL
Zootaxa 4093 (1) © 2016 Magnolia Press
In this formula, Y
is the value of the respective, allometrically adjusted variable of the i
specimen, Y
is the
original value of this variable of the i
specimen, β is the pooled regression coefficient of Y against SVL, SVL
the SVL of the i
specimen, and SVL
is the overall mean SVL of all specimens.
Subsequently, adjusted metric characters, meristic characters, and ratios were tested for statistically significant
differences between the two Cylindrophis forms occurring on Java (our new species and C. ruffus sensu lato). We
used one-way ANOVA (analysis of variance) if a variable fulfilled normal distribution, and a Mann-Whitney U-test
if a variable was not normally distributed. When the respective statistical test yielded significant outputs (i.e.,
statistically confirming differences between the two compared forms), these are shown in the Results section with
superscripted asterisks indicating probability levels as follows: * < 0.05; ** < 0.01; *** < 0.001.
Comments on the taxonomic history of Cylindrophis ruffus (Laurenti, 1768)
Early beginnings: Johann Jakob Scheuchzer’s (1672–1733) Physica Sacra Illustrata. Scheuchzer (1735) was
probably the first author who, in his pre-Linnæan treatise entitled Physica Sacra Illustrata, depicted snake
specimens referable to Cylindrophis ruffus sensu historico, presenting three different illustrations (Tabulae
DCXXIX-F, DCLX-3, DCCXLVIII-6; illustrated in Fig. 1A–C herein) of specimens from the Linck collection
(Merrem 1820; Boie 1827; Wagler 1828–1833; see also Bauer & Wahlgren [2013] for an overview of the Linck
collection). A precise identification of the specimens depicted, including their allocation to C. ruffus, C. burmanus,
or C. jodiae, however, is difficult.
Albertus Seba (1665–1736) and his Cabinet of Natural Curiosities. In the second volume of his Thesaurus,
Seba (1735: Tabulae VII-3, XXV-1; illustrated in Fig. 1D–E herein), described and figured two snakes based on
specimens housed in his cabinet of natural curiosities. These were identified as the taxon Cylindrophis ruffus by
subsequent authors (e.g., Merrem 1820). Seba’s short diagnosis indicates that both snakes originated on Ambon, an
island in the Moluccas. However, in the main description (following the diagnosis) and referring to Tabula XXV-1
(illustrated in Fig. 1E herein), Seba (1735: 26) assigned a larger area of distribution to the respective specimen,
namely “Les Grandes & […] les Petites Indes” [i.e., Asia and the American Continents]. Since the figures in Seba
leave little doubt as to the identity of the specimens (C. ruffus sensu historico), it is evident that they must have
originated in Asia. The taxon, however, does not appear to occur on Ambon (de Lang 2013), an island with a five-
centuries-long history of commercial and strategic importance for Europe, with specimens both collected or merely
shipped from there (e.g., Weijola & Sweet 2015).
Laurens Theodorus Gronovius (1730–1777) and the first detailed account of Cylindrophis ruffus. In his
Musei Ichthylogici, a detailed, descriptive catalogue of fish, amphibian, and reptile specimens housed in his Leiden
cabinet of curiosities, Gronovius (1756) introduced under the heading “6. ANGUIS squamis abdominalibus
CLXXIX, & squamis caudalibus VII” [6. SNAKE with 179 ventral scales and seven subcaudal scales] a taxon that
Merrem (1820) listed as Tortrix rufa (= Cylindrophis ruffus). Gronovius’s fairly detailed description of his species
“6. ANGUIS” (Gronovius 1756: 54; see also Adler et al. 1992) matches C. ruffus sensu historico, based on the
following morphological characters: 179 ventrals; seven subcaudals; small eyes; ventrals slightly enlarged,
hexagonal; stout, short, conical tail; reddish coloration with white transverse ventral bands. Although Gronovius
stated that his specimen originated in “Surinamam” [sic] [= Suriname], a thorough literature survey revealed that
there is no snake taxon known from Suriname (nor a species from outside Asia) that would match his description.
The only Asian species matching the listed characteristics are C. ruffus sensu lato and C. jodiae, and we therefore
conclude that Gronovius’s specimen must have been collected in Asia.
Josephus Nicolaus Laurenti (1735–1805) and the species description of Cylindrophis ruffus. The valid
species name ruffa was coined by post-Linnæan author Laurenti in 1768, who placed this taxon from a location he
listed as “Surinami” (Laurenti 1768: 71) into the genus Anguis Linnæus, 1758. As was common practice during
Zootaxa 4093 (1) © 2016 Magnolia Press
FIGURE 1. Historical drawings of Cylindrophis ruffus sensu historico. Illustrations from: (A–C) Scheuchzer (1735); and (D–
E) Seba (1735). Illustrations are not to scale. Plate prepared by Hinrich Kaiser and Mark O’Shea.
that time, Laurenti only provided exceedingly short descriptions of the known amphibian and reptile species that,
taken on their own, would hardly permit a proper diagnosis of specific taxa. However, in the case of his taxon
Anguis ruffa, Laurenti (1768: 139) stated “hospitatur in Museo Gronoviano” [housed in the collection of
Gronovius], thereby apparently referring to Gronovius’s 1756 catalogue (and hence to Anguis species number 6). A
Zootaxa 4093 (1) © 2016 Magnolia Press
comparison of Gronovius’s and Laurenti’s texts shows that Laurenti’s description is, by virtue of its wording, a
shortened version of that provided by Gronovius, with both authors providing the same erroneous information
regarding the specimen’s provenance. This leaves little doubt that the species identity of Laurenti’s A. ruffa is the
same as Anguis species number 6 of Gronovius (1756). It is unfortunate that the type specimen of A. ruffa appears
to be lost (Iskandar & Colijn 2002). Gronovius’s private collection was partly incorporated into the collection at the
BMNH (e.g., dried fish-skins; Gray 1854), but the rest of his collection probably became dispersed. His
herpetological collection cannot be traced to any larger museum collection extant today (Aaron M. Bauer, in litt.).
Johann Friedrich Gmelin (1748–1804) and his new species name. In his enhanced edition of the Linnæan
Systema Naturae, Gmelin (1789) listed Anguis rufus (nomen emendatum) and attributed this taxon to Laurenti
(1768). Gmelin (1789) also coined a new species name, A. striatus, and attributed this species to the pre-Linnæan
Gronovius by referencing the publication of the latter, directly referring to Anguis species number 6 (“A. Gron.
mus. 2. p. 53. n. 6.”). Since Laurenti (1768) clearly refers to Gronovius (1756) in his description of A. ruffa as well,
the connection between the descriptions published by Gmelin (1789), Laurenti (1768), and Gronovius (1756)
leaves little doubt, that A. striatus can be regarded as an objective junior synonym of Cylindrophis ruffus. Daudin
(1803) also listed Gmelin’s accounts of A. striatus and A. rufus, and Gronovius’s description of Anguis species
number 6 in his references for his description of Eryx rufus (comb. nov. for Anguis ruffa Laurenti, 1768).
Contributions by Patrick Russell (1726–1805). Russell (1801) used the preoccupied name Anguis scytale
Linnæus, 1758 (current name Anilius scytale) to refer to a Cylindrophis ruffus specimen he received from Java
(Russell 1801: Plate XXVII; illustrated in Fig. 2A herein). Hence, Anguis scytale Russell, 1801 is a junior
homonym of Anilius scytale (Linnæus, 1758) and a subjective junior synonym of C. ruffus (Laurenti, 1768).
George Shaw (1751–1813) and the confusion over Anguis scytale. Shaw (1802) depicted a Cylindrophis
ruffus specimen as part of his description of Anguis Corallina, using a figure (Shaw 1802: Fig. 131; illustrated in
Fig. 2B herein) undoubtedly based on Seba (1735: Tabula XXV-1; see Fig. 1E herein). In his references prefacing
the description of A. Corallina, Shaw listed Gmelin (1789), although in his own account of A. corallinus (nomen
emendatum) Gmelin referenced Laurenti (1768) as his source for that name. Laurenti (1768), Gmelin (1789), and
Shaw (1802) list the same plate in Seba (1735: Tabula LXXIII-2) as a reference. Alas, the specimen in this Tabula
is not a Cylindrophis at all, but an individual of Anilius scytale (a South American species), and hence, Laurenti’s
Anguis corallina and Gmelin’s A. corallinus have been regarded as synonyms of Anilius scytale (e.g., Wallach et
al. 2014). We agree and therefore do not follow Boulenger (1893) in regarding Shaw’s Anguis Corallina as
synonymous with C. rufus (nomen emendatum). We believe that the C. ruffus figure in Shaw (1802), the sole
indication supporting synonymy of C. ruffus with A. Corallina, was used by mistake; it does not correspond to
Seba’s Tabula LXXIII-2.
Blasius Merrem (1761–1824) and the problem with Scytale scheuchzeri. In his Versuch eines Systems der
Amphibien, Merrem (1820) listed Tortrix rufa (nomen emendatum) and described a new species, Scytale
scheuchzeri. As part of this description, Merrem referred to an illustration in Scheuchzer (1735: Tabula 647-1;
illustrated in Fig. 2C herein). The name S. scheuchzeri was considered synonymous with Cylindrophis ruffus by
subsequent authors (e.g., Boie 1827; Schlegel 1837b; Duméril & Bibron 1844; Gray 1849; McDiarmid et al.
1999; Bauer & Wahlgren 2013; Wallach et al. 2014). However, it is evident from both Scheuchzer’s illustration
and Merrem’s description of his genus Scytale (non Scytale Latreille in Sonnini and Latreille, 1802) that S.
scheuchzeri is not conspecific with C. ruffus. Despite similarities in coloration, the specimen depicted by
Scheuchzer has enlarged gastrosteges and a tapering tail. Merrem (1820) also listed enlarged gastrosteges in his
generic description of Scytale. Hence, the name S. scheuchzeri does not refer to an anilioid snake but most likely
to a colubroid snake, and we therefore remove this name from the synonymy of C. ruffus.
Contributions by Friedrich Boie (1789–1870) and Hermann Schlegel (1804–1884). Boie (1827) was the
first author to correct the distribution of Cylindrophis ruffus (under the name Tortrix rufa) to Java (not Schlegel
1837a, b, as commonly believed
; see e.g., Wallach et al. 2014). Schlegel (1837a: 128) then revised the
distribution of C. ruffus (as T. rufa) to “Java et de Célèbes” [Java and Sulawesi], but already indicated that the
Sulawesi form was distinct, later (1837b: 11) referring to it as Tortrix melanota (= C. melanotus; see also
Wal lac h et al. 2014). Schlegel (1837b) provided distribution records for the genus Cylindrophis (as Tortrix) from
2. Both Amarasinghe et al. (2015) and Uetz & Hošek (2015) list Schlegel (1844) as the reference for the type locality correction for C. ruffus to
Java. However, Schlegel (correctly cited as 1837−1844), in the explanatory text supplementing the plates in his Abbildungen Neuer oder
Unvollständig Bekannter Amphibien, does not provide such a correction (but see Schlegel 1837a, b).
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FIGURE 2. Historical drawings of Cylindrophis ruffus sensu historico (A, B & D−G) and Scytale scheuchzeri (C). Illustrations
from: (A) Russell (1801); (B) Shaw (1802); (C) Scheuchzer (1735); (D) Wagler (1828–1833); and (E–G) Schlegel (1837–
1844). Illustrations are not to scale. Plate prepared by Hinrich Kaiser and Mark O’Shea.
India: (1) “Tranquebar” (Tharangambadi, State of Tamil Nadu, SE India; see Russell 1801: 33), which was an
important seaport during Russell’s time; and (2) “Bengale” (NE India and Bangladesh). However, Smith (1943)
indicated that the genus Cylindrophis did not occur on the Indian subcontinent, and hence the distributional
records listed above appear to be in error and a reflection of maritime trade routes as opposed to natural
Johann Georg Wagler (1800–1832) and Cylindrophis resplendens. A new species from Java was
described and figured by Wagler (1828–1833: Tabula V-1; illustrated in Fig. 2D herein) under the name
Cylindrophis resplendens Wagler, 1828. Although Wagler (1828–1833) provided a figure of C. resplendens in
life (see Fig. 2D herein), capably illustrated by Kaspar Georg Karl Reinwardt (1733–1854) (see also Schlegel
1837b), in the Observationes following the species description, he explicitly referenced Russell (1801) for
additional illustrations of that taxon.
Cylindrophis resplendens, the type species of the genus Cylindrophis (Wallach et al. 2014), has since been
synonymized with C. ruffus (e.g., Schlegel 1837b; Duméril & Bibron 1844; Gray 1849; Boulenger 1893; Smith
1943; McDowell 1975; McDiarmid et al. 1999; Wallach et al. 2014; Amarasinghe et al. 2015). Wagler’s
description of C. resplendens was based on specimens housed in the “Museo Parisiensi” [now MNHN],
Lugdunensi Bat.” [now RMNH], and “in collectione mea” [in my collection; probably referring to the ZSM
collection]. One or more type specimens may still exist in the collection of the MNHN, but we failed to locate
specimens from the time of the original description matching Wagler’s Tabula V-1 in the collections of either
John Edward Gray (1800–1875) and Cylindrophis rufa var. javanica, the name of a taxon from Borneo.
Gray (1849: 112) described Cylindrophis rufa var. javanica in a simple two-line listing for a single specimen from
Borneo (not from Java, as stated by Amarasinghe et al. 2015), donated by Sir James Brooke (1803–1868), the first
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White Rajah of Sarawak. This specimen is still extant in the collection of the BMNH
. Gray (1849) referred to
figures in Schlegel (1837–1844: Plate 33, Figs 5–10; illustrated in Fig. 2E–G herein), which according to
Schlegel’s own statement were drawn from a single Javanese specimen. However, Schlegel (1837–1844) also
mentioned similarities between the Java “race” (Schlegel’s term) and a specimen the RMNH received from
Borneo. This may have led Gray, who was clearly familiar with Schlegel’s works, to apply the geographically
incongruous name javanica (referring to the island of Java) to a specimen from a locality on Borneo. Gray’s taxon
was synonymized with C. ruffus by Amarasinghe et al. (2015).
Malcom Arthur Smith (1875–1958) and a valid species from Myanmar. Smith (1943) described a
subspecies of Cylindrophis ruffus from “Tenasserim and Burma as far North as Myitkyina” (today’s Myanmar) as
C. rufus burmanus. This taxon was accepted as a subspecies with the spelling C. r. burmanicus (nomen
emendatum) by Lal Hora & Jaya Ram (1949), and in its original form by Taylor (1965). McDiarmid et al. (1999)
and Wallach et al. (2014) included subspecies in their synonymy lists of species, but these lists allow no conclusion
regarding the validity of the listed subspecies. Recently, Amarasinghe et al. (2015: 41) raised C. r. burmanus to
species level (see also Iskandar & Colijn 2002) and provided a redescription of that species based on “the presumed
type series.” However, among the six paralectotypes designated by Amarasinghe et al. (2015) is one specimen
(cited as ZMB 3094) that these authors considered to “probably” be a paralectotype, based on Iskandar & Colijn
(2002). The ZMB accession number of this specimen actually identifies a neotropical frog (Frank Tillack, in litt.)
and hence cannot possess “the same characters as the lectotype” (Amarasinghe et al. 2015: 41). Iskandar & Colijn
(2002) stated that ZMB 3094 originated at “Bhamo,” Myanmar. The only Cylindrophis specimen from Bhamo
housed in the ZMB collection has the accession number ZMB 11619, and it was collected by Leonardo Fea (1852–
1903) in the late 1880s. We doubt that this specimen could have belonged to the original type series used by Smith
(1943) to define C. r. burmanus. We consider the designation of ZMB 3094 as a paralectotype of C. burmanus to be
Amarasinghe et al. (2015) also presented conflicting data on the shape of the collar of Cylindrophis burmanus.
In their Table 2 (see also their Figs. 2 & 3), the band around the neck was listed as “dorsally interrupted” in that
species, yet it was described as complete when referring to C. burmanus in their diagnoses of both C. ruffus (“a
complete and narrow ring encircling the nape in C. burmanus,” p. 38) and C. burmanus (“a complete and narrow
ring encircling the nape,” p. 41). As seen in the illustration of the C. burmanus lectotype (Amarasinghe et al. 2015:
Fig. 3A), the band is actually separated by a single, dark brown vertebral scale. Our unpublished data show that this
character is quite variable in both C. burmanus and Javanese C. ruffus and not useful to diagnose either taxon.
Likewise, there is incongruity in the description of the pattern of dorsal blotches in C. burmanus. Whereas in their
Table 2 Amarasinghe et al. (2015) indicated that C. burmanus had alternating dorsal blotches, they also stated that
the species had paired (or “constant”; their term, p. 41) dorsal blotches. In a group of snakes where the true level of
intra- and interspecific morphological variability has not been fully explored, such contradictions may lead to a
similar level of instability as has resulted from the original descriptions (Laurenti 1768; Smith 1943).
History leads to the type locality of Cylindrophis ruffus. As a consequence of our careful review of the
historical literature, we agree with Amarasinghe et al. (2015) that the type locality of Cylindrophis ruffus sensu
stricto should be restricted to Java. The taxonomic history of the species shows that specimens in historical times
were most often collected on Java (e.g., Russell 1801; Boie 1827; Wagler 1828–1833; Schlegel 1837–1844), which
was an important trading hub for the Dutch Empire. With the establishment of the Dutch East India Company (in
Dutch: Vereenigde Oostindische Compagnie, VOC) in Batavia (now Jakarta) in 1611, trade to Europe from
Southeast Asia became heavily influenced by shipping conducted on behalf of the VOC (Boxer 1965). After the
disbanding of the VOC in 1799, the various administrations of the Netherlands continued trading with their
Southeast Asian colonies during the Napoleonic upheaval, although contacts with these colonies were often
3. In his published snake catalogue, Gray (1849) listed six specimens of C. rufa, three (ac) from Penang (presented by General Hardwick e), one (d)
from Borneo listed as “Var. 1. Javanica” (presented by Sir James Brooke), and two additional ones (ef) listed as “Var. 2.” without providing a
Latin name. However, in the extant handwritten catalogue at the BMNH, the entry for the particular specimen from Borneo presented by Sir
James Brooke, is found under the number IV.23.2.a, which is also how it is listed in the collection’s online database. We have ascertained that the
specimen identified in the collection by a jar label as IV.23.2.a (“Penang. Gen. Hardw icke”) is unquestionably co nspecific with C. jodiae and
therefore cannot have originated on Borneo. Furthermore, the specimen in the jar labelled “IV.23.2.d. Borneo. Sir J. Brooke” possesses l arge
blotches on the prefrontals, as mentioned in Gray ’s description. The error is therefore not in Gray’s published snake catalogue, but appears to be
an error that might have happened when the entries in Gray’s catalogue were transferred to the extant BMNH catalogue. Thus, the holotype of C.
rufa var. javanica really does have the number IV.23.2.d. It is not currently indicated as a ty pe specimen in the BMNH collection.
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blockaded by the British. Shortly after The Netherlands were annexed by France in 1810, the last Dutch colony in
Southeast Asia, Java, fell to Britain in 1811. However, the Netherlands regained independence and became a
kingdom in 1813, restoring their authority over the islands of Southeast Asia in 1816. The Dutch presence lasted
until a protracted dispute with Indonesia in the 1960s
, and trade continued throughout this time (e.g., Motadel
As highlighted above, the pipesnake specimen on which Laurenti (1768) based his description was housed in
Gronovius’s extensive natural history collection located in Leiden. Laurens Theodorus Gronovius and his father,
Jan Frederik Gronovius (1686–1762), were both renowned naturalists who were tied into early global trade, and
both would have received specimens from America and Asia via their trade connections (e.g., Margócsy 2014).
Based on the historic and economic circumstances that place Java as the nexus of Dutch trade with Southeast Asia,
along with the fact that Javanese Cylindrophis are the form most reliably described and illustrated in historical
accounts, we regard the type locality restriction Java as conforming with Recommendation 76A.1.4 of the
International Code on Zoological Nomenclature (ICZN 1999). For a neotype designation (Mecke et al., in prep.),
we believe that the type locality should be further restricted to northwestern Java, where the main trade port was
located at the time the original type specimen would have been collected (before 1756); most other parts of Java
remained undeveloped during that time as indicated by historic maps (e.g., “Nouvelle Carte de l’Isle de Java” by
Baussard 1756).
Amarasinghe et al. (2015) offered another hypothesis to demonstrate that the original type specimen originated
in Java: the possible confusion between the town of Batavia, Saramacca District, Suriname, and Batavia (Jakarta),
Java Island, Indonesia. While this is an interesting hypothesis, historical evidence appears to contradict this line of
reasoning. Firstly, shipments of specimens to private collectors in Leiden from mid-18
century Suriname would
have included only the name of the colony (i.e., Suriname) and possibly the main port (Paramaribo), but not the
name of a strategically irrelevant, small settlement (Marinus Hoogmoed, in litt.). Secondly, the settlement in
present-day Suriname near the confluence of the Coppename and Saramacca Rivers called Batavia was founded
only in 1790 (Anonymous 2015), several decades after the specimens Laurenti described would have had to have
reached Leiden in order to become integrated into Gronovius’s collection. Thus, it appears that the problem with
the type locality of C. ruffus sensu stricto really is a documentation error and not due to confusion with the
geographic identity of a place.
Synonyms. Based on the careful survey of early literature accounts and descriptions, we have determined that
the following names are synonyms of Cylindrophis ruffus (with type locality in Java): (1) Anguis striatus Gmelin,
1789 and, until evidence to the contrary becomes available, (2) A. scytale Russell, 1801, and (3) C. resplendens
Wagler, 1828. Gray’s (1849) C. rufa var. javanica should be regarded as species inquirenda until a formal revision
of C. ruffus is conducted. Gray’s name javanica would be available for the purposes of nomenclature for a
Cylindrophis species from Borneo, and if combined with the masculine generic name would need to be emended to
javanicus. Even though C. engkariensis and C. lineatus are Bornean taxa, they are clearly distinct from C. ruffus
and from the javanica type specimen held at the BMNH (BMNH IV.23.2.d.) and therefore not impacted by the
availability of the name javanica.
Comments on Amarasinghe et al. (2015). In their recent publication, Amarasinghe et al. (2015) redescribed
Cylindrophis ruffus based on 14 specimens from Java. However, the characters used in their diagnosis do not allow
either unequivocal species identification, nor are they suitable to establish stable species boundaries. Our
unpublished data from 113 Javanese specimens indicate that C. ruffus sensu lato includes sympatric forms with
specimens that (1) possess 19 or 21 dorsal scale rows at midbody, (2) show great variability in the number of
ventrals (179−225), (3) have either a complete or interrupted collar, and (4) may or may not possess dorsal blotches
that are, if present, either paired or alternating, and either complete or interrupted. We are currently in the process
of determining the taxonomic status of Javanese C. ruffus populations (Mecke et al., in prep.) and to resolve which
of these forms are conspecific with the specimen described by Gronovius (1756).
Amarasinghe et al. (2015) also described two new species of Cylindrophis, C. jodiae and C. mirzae. This
publication exists in two versions, an earlier one, in which Fig. 8 lists the names of the new species as C. jodii and
C. mirzai, and a revised version in which these errors have been corrected. These versions are otherwise
4. Indonesia gained independence in 1949 after a period of Japanese occupation during World War II (1942–45), but Dutch New Guinea did not
become part of I ndonesia until international pressure and Indo nesian military infiltration forced the Netherlands to relinquish control in 1962
(Gruss 2005).
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indistinguishable, and it appears that the revised version was simply exchanged on the journal’s website for the one
with the errors. This is evident from the URL
used to download the revised file. However, having been validly
published in the first version of the paper, the names C. jodii and C. mirzai must be considered objective junior
synonyms of C. jodiae and C. mirzae, respectively.
While the pholidotic characters of Cylindrophis jodiae, a species widely distributed on mainland Southeast
Asia (pers. obs.), conform to our unpublished data, qualitative color characters vary both intraspecifically and
ontogenetically (Kieckbusch et al., unpublished data). The definition of C. mirzae, on the other hand, appears to be
problematic. One of the key characteristics listed by Amarasinghe et al. (2015: Table 3) to differentiate C. mirzae
from C. ruffus was an invariable dorsal scale row count of 21 at midbody in C. mirzae. However, some specimens
we have examined from Singapore (the type locality of C. mirzae) have 19 dorsal scale rows, and the ratio of
Singaporean specimens with 21 vs. 19 scale rows in our data set is 8:8, with both forms possessing a similar range
of ventrals. In their Table 3, Amarasinghe et al. (2015) also list color pattern characteristics to distinguish C. mirzae
from C. ruffus. A complete narrow nape band and complete narrow dorsal crossbands, however, can occur in
specimens from Singapore with either 19 or 21 dorsal scale rows. These bands may also be interrupted in either 19-
or 21-row specimens, and are hence not useful to distinguish among species. Furthermore, C. ruffus with collection
localities on Java (the type locality of that species) may have 19 or 21 dorsal scale rows at midbody, and these
forms are equally variable in dorsal color pattern as specimens from Singapore. While we agree with Amarasinghe
et al. (2015) that C. ruffus sensu stricto is a taxon with an invariable number of middorsal scale rows, and that
forms with 19 dorsal scale rows should be distinct at species level from those with 21 rows (this difference being
the main character these authors used to differentiate C. mirzae from C. ruffus), the lack of a type specimen for C.
ruffus makes it at this point uncertain whether the 19-row or the 21-row morphotype represents C. ruffus sensu
stricto, and this hinders a diagnosis and renders their definitions of both C. mirzae and C. ruffus unsuccessful.
Lastly, Amarasinghe et al. (2015: 38) stated that “C. ruffus could extend beyond Java, e.g., Borneo and Peninsular
Malaysia,” which would include Singapore and overlap with the distribution of C. mirzae, but they failed to
demonstrate this zoogeographical scenario using voucher specimens. Given the problems outlined above, we see
no alternative than to place C. mirzae in the synonymy of C. ruffus until it can be unequivocally defined and
differentiated from that species.
Species description
Having ascertained the history of Cylindrophis ruffus sensu historico in general, and the history and morphology of
C. ruffus sensu lato in particular, we are confident when we propose that a population from south-central Java with
morphological features that allow unequivocal identification should be recognized taxonomically. We formally
describe this species below.
Cylindrophis subocularis sp. nov.
(Figs. 3−5; Table 1)
Holotype. RMNH.RENA 8785 (Figs. 3−4; Table 1), an adult female, collected in Grabag, Purworejo Regency
(formerly Koetoardjo), Central Java Province (Jawa Tengah), Java, Indonesia, by Felix Kopstein in February 1937.
The original label for this specimen states “Grabag, Koetoardjo, Midden Java. +10 m.”
Paratypes. All RMNH.RENA specimens were collected by Kopstein at the type locality. RMNH.RENA 8958
(Fig. 5A), a gravid female, was collected in October 1937; RMNH.RENA 8959 (Fig. 5B), an adult female, was
collected in November 1937; RMNH.RENA 11257 (Fig. 5C), an adult male, was collected in August 1937;
RMNH.RENA 11263 (Fig. 5D), an adult male, was collected in August 1937; RMNH.RENA 47929 (Fig. 5E), an
adult male, was collected in November 1937. NMW 21559.1 (Fig. 5F), an unsexed adult specimen from Java (no
precise locality provided), was also collected by Kopstein, presumably during 1937, but the date is unknown.
Referred specimen. ZMB 53459, an unsexed adult with no further collection data.
Definition. A species of the genus Cylindrophis that can be readily distinguished from all congeners by the
following combination of characters: (1) presence of a single subocular scale, positioned between 3
and 4
or 4
5. A Googl e search for the paper by Amarasinghe et al. (2015) by title leads to a downloadable pdf at the URL
51488619/nsd_014410685_corrected.pdf. This URL features the term “corrected,” implying that an uncorrected version existed for download at
least temporarily.
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and 5
supralabial, contacting postocular and separating 4
or 5
supralabial from orbit (Fig. 4B); (2) prefrontal in
very narrow contact with or separated from orbit; (3) 19 smooth dorsal scale rows at midbody; (4) 6−7 supralabials;
(5) 6−7 infralabials; (6) 190−196 ventrals; (7) 6−7 subcaudals; (8) 40−48 transverse light ventral blotches, and (9)
light blotches on lateral surfaces of prefrontals (Fig. 3A, 4A & B).
FIGURE 3. Holotype of Cylindrophis subocularis sp. nov. (RMNH.RENA 8785) in (A) dorsal and (B) ventral view.
Numbered units on ruler are in centimeters. Photos by Sven Mecke.
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FIGURE 4. Holotype of Cylindrophis subocularis sp. nov. (RMNH.RENA 8785). (A) Dorsal, (B) lateral, and (C) ventral view
of the head. (D) Lateral view of a midbody section (left side). Scale bar = 2.0 mm. Drawings by Felix Mader based on
photographs by Sven Mecke.
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FIGURE 5. Paratypes of Cylindrophis subocularis sp. nov. in dorsal view. (A) RMNH.RENA 8958; (B) RMNH.RENA 8959;
(C) RMNH.RENA 11257; (D) RMNH.RENA 11263; (E) RMNH.RENA 47929; (F) NMW 21559.1. All RMNH specimens
were collected at the type locality, Grabag, Purworejo (formerly Koetoardjo) Regency, Central Java Province (Jawa Tengah),
Java, Indonesia. NMW 21559.1 is from Java, Indonesia, without detailed locality data. Numbered units on ruler are in
centimeters. Photos by Sven Mecke.
Comparisons. Cylindrophis subocularis sp. nov. can be easily distinguished from all congeners by the
presence of a single subocular, positioned between the 3
and 4
(rarely between the 4
and 5
contacting the postocular and separating the 4
(or 5
) supralabial from the orbit (e.g., Fig. 4B). In the following
comparisons, ranges are followed by mean ± standard deviation and sample size (n), with the measures and counts
for C. subocularis provided in parentheses. Whenever range and mean ± standard deviation are not provided, the
respective character was invariable within a species.
Cylindrophis aruensis possesses 23 (19, n = 8) dorsal scale rows at midbody and 173–182 (190–196, 193.7 ±
2.0, n = 8) ventrals (Boulenger 1920; McDowell 1975; Amarasinghe et al. 2015). Cylindrophis boulengeri
6. While the general, relative position of the subocular is fixed, it may be bordered by the 4
and 5
supralabial, resulting from a vertical division of
the 3
upper labial.
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possesses 197–204, 200.3 ± 3.5, n = 3 (190–196, 193.7 ± 2.0, n = 8) ventrals; and wavelike markings on
supralabials, which may run onto prefrontals (uniformly dark supralabials and light blotches on prefrontals).
Cylindrophis burmanus possesses 201−210, 208.3 ± 7.7, n = 6 (190–196, 193.7 ± 2.0, n = 8) ventrals. Cylindrophis
engkariensis possesses 17, n = 1 (19, n = 8) dorsal scale rows at midbody; 230
, n = 1 (190–196, 193.7 ± 2.0, n = 8)
ventrals; rugose (smooth) dorsals on tail; a dorsal pattern of two paravertebral rows of spots (dorsal pattern of
transverse, light, dorsolateral blotches); and uniformly colored prefrontals (light blotches on prefrontals).
Cylindrophis isolepis possesses 21, n = 2 (19, n = 8) dorsal scale rows at midbody; and nasals separated by rostral
(nasals in contact). Cylindrophis jodiae possesses 21, n = 77 (19, n = 8) dorsal scale rows at midbody; and wavelike
markings on supralabials (uniformly dark supralabials). Cylindrophis lineatus possesses 21, n = 1 (19, n = 8) dorsal
scale rows at midbody; 210
, n = 1 (190–196, 193.7 ± 2.0, n = 8) ventrals; 9, n = 1 (6–7, 6.6 ± 0.5, n = 8)
subcaudals; and a dorsal pattern of stripes (dorsal pattern of transverse, light, dorsolateral blotches). Cylindrophis
maculatus does not possess light blotches on prefrontals (present); has a relatively longer snout, with SL/IOD =
1.03–1.25, 1.13 ± 0.06, n = 34 (0.94–1.03, 1.00 ± 0.03, n = 7); and a dorsal pattern of reddish-brown, large and
round blotches (dorsal pattern of transverse
, light, dorsolateral blotches). Cylindrophis melanotus (including its
synonyms Tortrix rufa var. celebica Schlegel, 1844, T. rufa var. celebensis Gray, 1849
, C. celebensis Smith, 1927,
and C. heinrichi Ahl, 1933) possesses 230–268, 245.3 ± 10.5, n = 35 (190–196, 193.7 ± 2.0, n = 8) ventrals; and
predominantly light-colored supralabials, including a characteristic dark bar running down the supralabials below
eye (completely dark supralabials and light blotches on prefrontals). Cylindrophis opisthorhodus possesses 23, n =
6 (19, n = 8) dorsal scale rows at midbody; and has a light dorsum with dark speckles forming two paravertebral
rows and occasionally a discontinuous vertebral line (dorsal pattern of transverse, light, dorsolateral blotches).
Cylindrophis ruffus sensu lato (including its synonyms Anguis striatus Gmelin, 1789, A. scytale Russell, 1801, C.
resplendens Wagler, 1828, and C. mirzae), and C. rufa var. javanica Gray, 1849 (inferred from the relevant
descriptions, drawings, figures, or examination of type material) do not have a subocular scale (present). Javanese
C. ruffus sensu lato have the prefrontal usually in broad contact with the orbit (Fig. 6; Table 1), with PrefO/ED =
0.28–0.60, 0.38 ± 0.08, n = 51 (prefrontal in narrow contact with or separated from the orbit [Fig. 4B]; with PrefO/
ED = 0.0–0.27, 0.11 ± 0.11, n = 8); results of Mann-Whitney U-test: Z = 0.29, p < 0.001
. Cylindrophis yamdena
possesses 21 (19, n = 8) dorsal scale rows at midbody, and a pale light dorsum without any pattern (Smith & Sidik
1998) (dorsal pattern of transverse, light, dorsolateral blotches).
Description of the holotype: metrics (in mm) and pholidosis. An adult female; SVL 385; tail very short, TL
10 (2.6 % of SVL); head not distinct from body; body cylindrical, body diameter 12.0 (3.1 % of SVL); head
rounded in dorsal view; HL 11.9 (3.1 % of SVL); HW 8.7 (73.1 % of HL); snout rounded in dorsal and lateral
view; SL 5.1 (42.8 % of HL); SW 3.4 (66.7 % of SL); ED 1.3 (10.9 % of HL); pupil round; IOD 5.0 (42.0 % of
HL); NOD 3.7 (31.1 % of HL); PrefO/ED 0.04; internarial distance 2.5; pelvic spurs not visible externally but
hidden in pouches situated laterally of cloacal plate, covered by scales; 21/19/17 dorsal scale rows, scales smooth,
apical pits absent; 196 ventrals; six subcaudals + one terminal spine; cloacal plate divided; rostral clearly visible
from above, triangular, wider than high (rostral height 2.0, rostral width 2.2); two pentangular nasals, height 1.9,
length 2.6; nasal suture sinistral in respect to prefrontal suture; naris positioned close to the suture of nasal with
first supralabial; postnasal absent; loreal absent; prefrontal in contact with 2
and 3
supralabial; preocular absent;
rectangular subocular scale present, length 1.0, height 0.9; one pentangular postocular (length 1.1, height 1.4);
temporal formula 1 + 2, anterior temporal larger than each posterior temporal (anterior temporal length 2.5, height
2.6; upper posterior temporal length 2.6, height 2.1); 6|7 supralabials: on right side of head: 1
smallest, 3
, 4
, 5
, and 6
equal in size, 2
and 3
in contact with prefrontal, 3
in contact with orbit; on the left side: 1
smallest, 3
largest, 4
, 5
, and 6
equal in size, 2
, 3
and 7
equal in size, 2
, 3
, and 4
in contact with
prefrontal, 4
in contact with orbit; six infralabials, 3
in contact with first pair of chin shields; first pair of
infralabials in contact, preventing contact of mental with first pair of chin shields; mental triangular, wider than
high, width 2.2, height 1.5; two pairs of chin shields, anterior chin shield length 2.1, width 2.0, posterior chin shield
length 2.6, width 1.3; mental groove present, length 3.5; one hexagonal prefrontal, length 2.9, width 3.2; one
pentangular supraocular, length 2.7, width 2.6; frontal rectangular, length 3.2, width 3.8; one pentagonal parietal,
length 2.9, width 2.7.
7. Stuebing (1994) reported 234 ventrals for the holotype of C. engkariensis. A re-examination of the specimen by one of us (HK) showed that there
are only 230 ventral s present.
8. Blanford (1881) reported 215 ventrals for C. lineatus and Smith & Sidik (1998) pro vided a ventral range of 210−215.
9. Tortrix rufa var. celebensis Gray, 1849 is a nomen emendatum for T. rufa var. celebica Schlegel, 1844 and should currently be regarded a
junior synonym of Cylindrophis melanotus Wagler, 1828. It is also a junior secondary homony m of C. celebensis Smith, 1927.
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TABLE 1. Data for the individual type specimens of Cylindrophis subocularis sp. nov., and a comparison of this species
with C. ruffus sensu lato from Java (data of specimens with precise collection locality shown only). Metric characters are
given in mm. Ranges are followed by mean ± standard deviation (indicated in parentheses). An ‘X’ indicates a fusion
between the subocular and the postocular.
TABLE 1. (continued).
Description of the holotype: coloration and pattern in preservative (after 78 years in ethanol). Dorsal
surface of head Sepia (279) with a Pale Buff (1) blotch on each prefrontal, extending from center of scale at about
half scale’s width to lateral edge of scale; most upper head scales with lighter edges; supralabials Sepia (279);
ventral surface of head Sepia (279) with lighter edges of scales and a Pale Buff (1) ‘X’-shaped marking beginning
at level of lower edges of 3
infralabial, extending to throat (Fig. 4C); neck with a two scale broad Pale Buff (1)
collar, interrupted medially in vertebral region, located one dorsal scale behind parietals; dorsal surfaces of trunk
and tail Burnt Umber (48); dorsal surface of trunk with paired, occasionally slightly alternating, transversely
arranged Pale Buff (1) blotches, approximately one scale broad, well-developed anteriorly and posteriorly, very
faint or absent at central part of trunk; dorsal surface of tail with a Pale Buff (1) band that continues to the ventral
surface, demarcating a Raw Umber (48) tail tip; ventral surface of trunk Raw Umber (280), with 43 transverse,
alternating ventrolateral Pale Buff (1) blotches (two ventral scales broad at midbody); cloacal region and ventral
surface of tail Pale Buff (1), with a Raw Umber (280) tail tip (from 4
subcaudal to terminal caudal spine), and
Raw Umber (280) blotches on scales covering the cloacal spurs.
Status Holotype Paratype Paratype Paratype
Sex F F F M
SVL 385 394 326 451
TL 10 9 10 11
Dorsals 21/19/17 21/19/18 20/19/18 21/19/17
Ventrals 196 194 192 195
Subcaudals 6 7 7 7
Supralabials 6|76 6 6
Infralabials 6 6 7 6
Ventral bands light 43 40 48 43
Ventral bands dark 43 40 48 43
Subocular scale length 1.0|0.8 0.8|1.0 0.6|0.9 1.6|1.8
Subocular scale height 0.9|0.6 0.6|1.1 0.6|0.9 1.7|1.9
PrefO/ED 0.04 0 0.02 0.27
C. ruffus sensu lato
(n = 53)
Status Paratype Paratype Paratype
Sex M M unsexed -
SVL 331 353 288 148–737 (356.1±143.8)
TL 7 10 10 4–19 (9±3.3)
Dorsals 21/19/17 20/19/17 21/19/17 19–23/19−21/15–19
Ventrals 196 194 190 179–225 (194.5±8.9)
Subcaudals 7 7 6 5–7 (5.9±0.7)
Supralabials 6 7 6 6
Infralabials 6 6|76 6
Ventral bands light 40 43 45 33–59 (45.9±6.0)
Ventral bands dark 40 43 44 32–59 (45.2±5.8)
Subocular scale length X|1.3 1.0|1.1 1.1|1.1 -
Subocular scale height X|1.0 1.0|0.9 0.9|1.0 -
PrefO/ED 0.25 0.21 0 0.28–0.6 (0.38±0.08)
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FIGURE 6. Head of a Cylindrophis ruffus sensu lato specimen from Bogor, Java (SMF 16980), in lateral view. Note the broad
contact of the prefrontal with the orbit. Scale bar 2.0 mm. Drawing by Felix Mader based on a photograph by Gunther Köhler.
Intraspecific variation. Our assessment of the variation is based on the holotype and six paratypes (three
males, three females, one unsexed specimen; Figs. 3 & 5; Table 1), with measurements provided in mm and listed
including range and mean ± standard deviation and specimen numbers (n) in parentheses: SVL 288–451 (361.1 ±
53.7, n = 7); TL 7–11 (9.6 ± 1.3, n = 7); 21/19/17 (n = 5), 20/19/18 (n = 1), and 20/19/17 (n = 1) dorsal scale rows;
190−196 (193.8 ± 2.2, n = 7) ventrals; 6–7 (6.7 ± 0.5, n = 7) subcaudals; six (n = 5), seven (n = 1) or 6|7 (n = 1)
supralabials; six (n = 5), seven (n = 1), or 6|7 (n = 1) infralabials; 4
supralabial in contact with orbit in specimens
with seven supralabials (n = 2); subocular present on both sides of head in all specimens (n = 7); subocular may be
fused with postocular (n = 1); subocular in contact with postocular, orbit and 3
and 4
supralabial (in the case of
the presence of six supralabials) or 4
and 5
supralabial (in the case of the presence of seven supralabials);
subocular size: length on right side of head 0.6–1.6 (1.0 ± 0.3, n = 6) and 0.8–1.8 (1.1 ± 0.3, n = 7) on left side,
height 0.6–1.7 (0.9 ± 0.4, n = 6) on right and 0.6–1.9 (1.0 ± 0.4, n = 7) on left side of head; 40–48 (43.1 ± 2.8, n =
7) alternating, light ventral blotches, two ventrals wide at midbody, three ventrals wide at midbody in a single
specimen; light blotches on lateral surfaces of prefrontals might be fused into a bar running across the snout; light
‘X’-shaped marking on ventral surface of head might be dissolved into a reticulated pattern.
Etymology. The specific epithet subocularis is a compound adjective of sub (Latin: ‘under,’ ‘beneath’) and
ocularis (Latin: ‘pertaining to the eye’), referring to the presence of a subocular scale in the new species.
Distribution and natural history. The new species is only known from Grabag on the south coast of
Purworejo Regency, Central Java Province, Java, Indonesia (Fig. 7). The type locality in the South Central Java
basin area is enclosed by mountain ranges to the north, west, and east, which include active volcanoes (Darman &
Sidi 2000).
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FIGURE 7. Distribution map of Cylindrophis subocularis sp. nov. and C. ruffus sensu lato on Java, Indonesia. The black
triangle marks the type locality of C. subocularis sp. nov. at Grabag. The white circles with letters identify localities of
examined specimens of C. ruffus sensu lato, including (a) Jakarta (Batavia), (b) Bogor (Buitenzorg), (c) Sukabumi
(Soekaboemi), (d) Indramayu (Indramajoe), (e) Cirebon (Cheribon), (f) Kagok, Tegal, (g) Pekalongan, (h) Semarang
(Samarang), (i) Rembang, (j) Kediri, (k) Surabaya (Surabaja, Soerabaja), (l) Mount Arjuno (Ardjoeno), (m) Malang (Malary/
Malang?), and (n) Tengger Mountains. Note that not all locality data of museum vouchers provided necessarily correspond to
towns and their environs but may refer to district names at the time of specimen collection. Locality names in parentheses refer
to historical names provided on museum labels or in museum catalogues. Base map modified from Wikipedia © Sadalmelik /
Wikimedia Commons / CC-BY-SA-3.0 by Sven Mecke.
During the geological history of Sundaland, Java was connected to the islands of Borneo and Sumatra (Voris
2000; Sathiamurthy & Voris 2006; Wilting et al. 2012), and according to Natus (2005) many elements of the
Javanese terrestrial vertebrate fauna descended from Bornean and Sumatran lineages that migrated to Java during
or even before the Pleistocene and Holocene. Natus (2005) also identified eight endemism centers for terrestrial
vertebrates in Java (Natus 2005: Fig. 4.22), which can be divided into two major groups: the lowlands in the
northwest (immediately adjacent to Sumatra) and the eastern parts of Java, and the highlands of the Neogene-
Quaternary volcanic arc that stretches longitudinally through the centre of Java. The South Central Java basin,
however, has long been isolated to the north by the central volcanic chain (based on the maps presented in
Sathiamurthy & Voris 2006) that may have largely prevented immigration events to the south, leading to vicariant
evolution. Although the range of Cylindrophis subocularis is probably not restricted to Grabag, it may indeed
exhibit a relatively limited distribution in the South Central Java basin and therefore should be regarded as a
regional endemic.
Based on the lifestyle of congeneric species, we assume that Cylindrophis subocularis is semifossorial and
preys mainly on elongate vertebrates (e.g., fishes, caecilians, skinks, and snakes: Schmidt 1928; Taylor 1965;
Pauwels et al. 2000; Kupfer et al. 2003; pers. obs.), which are subdued by constriction (Greene 1983). Both the
limited distribution and the secretive semifossorial lifestyle of C. subocularis may explain its apparent rarity in
museum collections.
One specimen of the new species (RMNH.RENA 8958) contains eggs covered by a thin membrane. An
incision into the membrane of one of the largest eggs (length 26.8 mm, width 13.3 mm) revealed the presence of an
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embryo (approximately at developmental stage 26−27, following Zehr 1962). We believe that this observation
confirms that Cylindrophis subocularis is a viviparous species (sensu Blackburn 1994), with viviparity being the
reproductive strategy for most, if not all, Cylindrophis species (de Rooij 1917; Smith 1943; McDowell 1975;
Blackburn 1985; Brischoux et al. 2011). We also found one specimen of the closely related C. ruffus from Java
(NMW 21558.6) that contains fully developed embryos. No further information is available on the biology of C.
Remarks. While we discovered six of the seven type specimens of Cylindrophis subocularis in the collection
of the RMNH, all of which were collected by Felix Kopstein (1893−1939) and accompanied by precise collection
locality data, a single specimen was found in the collection of the NMW. For this specimen (NMW 21559.1) the
collection locality is limited to “Java,” but the specimen label lists Felix Kopstein as the collector of the specimen.
Based on specimen labels in the RMNH, Kopstein collected Cylindrophis specimens at other localities in Java,
such as at “Indramajoe” (Indramayu, on the north coast of Central Java). We have examined these, as well as 113
additional Javanese specimens, and all lack a subocular scale and have the prefrontal usually in broad contact with
the orbit. We believe that NMW 21559.1 is part of the series Kopstein collected on the south coast of Central Java,
but deposited mostly in Leiden, with the single specimen deposited in the Vienna collection
. We discovered an
additional specimen of C. subocularis in the Berlin collection (ZMB 53459). In the absence of a listed collection
locality and collector’s name, we chose not to include this specimen in our type series.
Two specimens (RMNH.RENA 47931–32, formerly RMNH.RENA 8785.80–81) from the same original jar
(jar number 8785) as the holotype (RMNH.RENA 8785, formerly RMNH.RENA 8785.51) and supposedly also
collected at Grabag, are not conspecific with Cylindrophis subocularis. In the original catalogue of the
herpetological section of the RMNH, we found the following entry:
De fles [8785] bevat nu 3 ex, zij zijn bewerkt door E.M.J. Jaspars en door hem voorzien van de nrs. 51,
80, 81. Mogelijk zijn de nrs 80 en 81 door bewerker bij vergissing in deze fles ondergebracht en zijn zij
afkomstig van Buitenzorg [Bogor], Java.
[The jar [8785] now contains three specimens; they were examined by E.M.J. Jaspars and labeled with the
numbers 51, 80, 81. Potentially, the numbers 80 and 81 have been misplaced in the jar by the researcher and
they may have originated in Buitenzorg [Bogor], Java.]
We agree with the catalogue entry that RMNH.RENA 47931–32 (formerly RMNH.RENA 8785.80–81) were
most likely misplaced in the jar; these specimens strongly resemble Cylindrophis ruffus from Bogor (n = 9) in
having no subocular and the prefrontal in broad contact with the orbit, PrefO/ED = 0.42 and 0.47 respectively (vs.
subocular present and prefrontal in narrow contact with or separated from the orbit in C. subocularis, PrefO/ED =
0.0–0.27, 0.11 ± 0.11, n = 8). An additional specimen (RMNH.RENA 11255), with greatly damaged anterior head
scalation, but lacking a subocular scale, was supposedly also collected at the type locality of C. subocularis. Due to
the consistent presence of a subocular scale in the Grabag population, we have reasonable grounds to believe that
RMNH.RENA 11255 is also not conspecific with the new species. We believe that RMNH.RENA 11255 was most
likely also misplaced or erroneously labeled, as was the case with RMNH.RENA 47931–32.
Discussion and outlook
Species of Cylindrophis have generally been described from small series of specimens collected at remote localities
(e.g., Roux 1911; Boulenger 1920; Stuebing 1994; Smith & Sidik 1998) or, especially in the early days of
taxonomy, were described using insufficient or unsuitable characters (e.g., Laurenti 1768; Wagler 1828–1833).
Taking into account the distribution of the morphologically variable taxon Cylindrophis ruffus sensu lato (Java,
Borneo, Sumatra, Singapore and Peninsular Malaysia), which heretofore had been considered even more widely
10. It is perhaps incongruous that an Austrian naturalist with ties to the NMW would not deposit a majority of specimens at what was essentially his
home institution (without formal ties). It is possible that Kopstein had designs on an appointment at the RMNH, and he perhaps sent a significant
number of specimens there to court favor. Unfortunatel y for Kopstein, he died before his appointment might have become reality (Marinus
Hoogmoed, in l itt.).
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distributed, it appears that the diversity of Cylindrophis in general, and of forms hidden under the name C. ruffus in
particular, is still significantly underestimated. While C. ruffus has long been identified as a species complex in
need of a thorough and comprehensive revision, including the designation of a neotype (Mecke et al., in prep.), we
feel it necessary to caution against taxonomic studies of such historically difficult taxa without a solid basis of
comparative material, without a wide range of characteristics used, and when personally unfamiliar with relevant
specimens. While a general aim of these studies is to achieve greater taxonomic stability, the example of C. mirzae
shows that, even with the best intentions, a small data set may yield an unsatisfactory result.
Cylindrophis subocularis is superficially similar to other forms currently referred to as C. ruffus sensu lato. It
is, however, ‘inconspicuously conspicuous,’ because it is easily diagnosed by its unique pholidotic characters: the
presence of a subocular and the prefrontal in narrow contact with or separated from the orbit. The former character
has been considered of broad taxonomic importance in snake systematics and has readily been used to identify
distinct species (e.g., Schätti 1987; Dowling & Price 1988
; O’Shea 1998, 1999; Murphy et al. 2005). We are
confident that the subocular scale in C. subocularis represents a true, distinctly differentiated scale and not an
aberrant horizontal division of the 4
or 5
supralabial (in specimens with six or seven supralabials respectively). In
contrast to developmental aberrations in head scales, which usually occur only on one side of the head, the
subocular occurs bilaterally in all specimens in precisely the same position below the orbit. This convincingly
demonstrates that the occurrence of a subocular scale in the genus Cylindrophis is a stable character found only in
a single, probably isolated population and does not represent a sporadic aberration found across the genus.
Moreover, the scale is always of the same rectangular shape and is clearly independent of the supralabial below it.
In one specimen (RMNH.RENA 11263), the subocular is fused with the postocular on the right side of the head,
but still clearly separated from the supralabial, which supports the concept of this scale as an independent,
bilaterally occurring pholidotic character. During our examination of Cylindrophis specimens from the entire range
of the genus (451 specimens), we found ten specimens (2.2 %) with aberrant head scale conditions, of which seven
(70 %) were unilateral anomalies of bilaterally occurring scales and three (30 %) were aberrant divisions or fusions
of azygous head scales. Unilateral anomalies of bilaterally occurring scales included deformations and were never
found to occur in a single population with any specific frequency.
Cylindrophis subocularis is one of several poorly known species with a rather restricted area of distribution,
and in that it is similar to C. aruensis, C. boulengeri, C. engkariensis, C. isolepis, and C. yamdena. As outlined
above, the new species is only known from eight specimens collected almost 80 years ago, six of which were
evidently collected at a single locality in southern Java. Although it appears to be generally accepted that the
Javanese herpetofauna is relatively well studied compared to the herpetofaunas of the other Greater Sunda Islands
(e.g., Teynié et al. 2010), we argue that historic and recent research has mostly been conducted along the north
coast and the western and eastern parts of the island. Hence, species diversity for the whole of Java may still be
underestimated. The recent discovery of new bent-toed gecko species (genus Cyrtodactylus) in Java (Riyanto et al.
2014, 2015; Hartmann & Mecke et al., 2016) indicates that new species, some of which have a rather limited area
of distribution, are still being identified.
It is uncertain at this time whether Cylindrophis subocularis exhibits a wider distribution than the single
collection locality would indicate, or is truly a localized endemic. Herpetological surveys of southern coastal
localities in Java are required to investigate the taxon’s distribution and population size, and to assess any potential
threats that may impact its conservation status. It may be noted that Central Java has little remaining forest, and that
the long history of deforestation and intensification of agriculture along the south-central coast potentially led to
local species extinctions in the region (Whitten et al. 1996). As the almost 80-year-old type series of C. subocularis
is unsuitable to obtain molecular data, it would be desirable to obtain fresh tissue samples for molecular genetic
approaches to investigate its phylogenetic affinities, especially in relation to C. ruffus sensu lato.
During our work with specimens of Cylindrophis, we have progressively been able to recognize morphological
and ontological patterns in these snakes that would not be recognizable when working with only a few selected
specimens, let alone only type specimens. Detailed revisions of the C. ruffus and the C. melanotus complexes,
including the description of new species, are ongoing and will be published elsewhere (Kieckbusch et al. & Mecke
et al., in prep.).
11. Dowling & Price (1988) called suboculars “lorilabial scales.”
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For the loans of specimens and for access to collections under their care during our visits, we are very grateful to
Esther Dondorp (RMNH); Georg Gassner, Silke Schweiger, and Heinz Grillitsch (NMW); and Frank Tillack and
Mark-Oliver Rödel (ZMB). Further important material for comparison was provided by Lauren Vonnahme, David
A. Dickey, David A. Kizirian, and Christopher J. Raxworthy (AMNH); Andreas Schmitz (MHNG); Markus Auer
and Raffael Ernst (MTKD); Stefan T. Hertwig (NHM); Urs Wüest and Denis Vallan (NMB); Linda Acker and
Gunther Köhler (SMF); Jakob Hallermann (ZMH); and Kelvin Lim (ZRC). We are very grateful to Harry W.
Greene (Cornell University, Ithaca, USA), John C. Murphy (FMHN), Marinus S. Hoogmoed (MPEG), Roy W.
McDiarmid (USNM), George R. Zug (USNM), Frank Tillack, Gernot Vogel, and an anonymous reviewer for their
helpful comments on earlier versions of the manuscript. SM thanks Glenn Shea (AM) for fruitful discussions
regarding scale characteristics and pholidotic aberrations. We thank Britta Döring (Philipps-Universität Marburg,
Germany) for contributing to data collection. Many thanks to Felix Mader who prepared the drawings in Figs. 4 &
6, and to Gunther Köhler who provided the photograph on which the illustration in Fig. 6 is based. We further thank
Linda Acker, Aaron M. Bauer (Villanova University, Villanova, USA), Esther Dondorp, Georg Gassner, Heinz
Grillitsch, Gunther Köhler, Silke Schweiger, and Frank Tillack for providing some of the literature cited below.
This study was supported by an AMNH collection study grant to SM. This paper is contribution No. 19 from the
Tropical Research Initiative at Victor Valley College.
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APPENDIX. Specimens examined for comparison.
Cylindrophis burmanus.—Myanmar: Kachin State: Bhamo: NMB-REPT 479, NMW 21552.3–4, ZMB 11619, ZMH R06256;
Rakhine State: “Aracan”: MTKD 14867.
Cylindrophis boulengeri.—Indonesia: Maluku Province: Wetar Island: without precise locality data: RMNH.RENA
5529A.168, 5529B.169; Ilwaki: Wetar Island, SMF 16996 (holotype).
Cylindrophis engkariensis.—Malaysia: Sarawak (Borneo): Second Division, Lubok Antu District, Lanjak-Entimau,
headwaters of the Engkari River, Nanga Segerak: ZRC 2.3398 (holotype).
Cylindrophis isolepis.—Indonesia: South Sulawesi Province: Jampea Island: RMNH.RENA 11269A.171, 11269B.72.
Cylindrophis jodiae.—Malaysia: Kedah State: NMW 39624.2; Penang: NMW 21570.1, 21570.4. Thailand: no precise locality
data: NMW 21556.4, ZMH R09798–99, R09801–02, MTKD 24126–27, SMF 16987, 16991, ZMB 30205, 52611;
Bangkok Province: Bangkok: MHNG 1335.17, NMW 21561, 21562.1−4, 21563.1−2, 21564.1−11, SMF 58675, 58679,
61903, 64838, ZMB 4394, 4545, 58428, ZMH R09794, ZRC 2.4583; Chiang Mai Province: MTKD 39216; Dangrek
Mountains: Phu Khi (Pu-Kin, Don-Rek): NMW 21556.2, 21569.1−3; Don Pia Fei Mountains: NMW 21565.1−6,
21566.1−6; Muang Pou Vieng (Pu Wieng): NMW 21567.1−2; Phang Nga Province: Khaolak-Luk National Park: ZMB
55188; Phetchaburi Province: Puek Tian: NMW 21569.1–3; Saraburi Province: Saraburi: MHNG 1471.30, MHNG
1530.9. Vietnam: no precise locality data: NMBE 1015768−69; Ho-Chi-Minh Province: Ho-Chi-Minh City: NMBE
1015764–66, ZMB 31123, 50774; “South Vietnam”: MHNG 1325.30, 1551.18–20.
Cylindrophis lineatus.—Singapore (in error): AMNH R-12872.
Cylindrophis maculatus.—Sri Lanka (occasionally labeled as “Ceylon”): without specific localities: MHNG 762.65, 1199.44,
2745.34, MTKD D14873–76, NMW 21574.1–5, NMW 21575.1–2, RMNH.RENA 160–63, SMF 16995, ZMB 1456,
18550, 18551.A–B, 24125, 49460, 77698, ZMH R09785, R09792, R09795–96. Central Province, Kandy District,
Peradenyia: ZMB 31506. Sabaragamuwa Province: Kitulgala: MHNG 2156.29; Ratnapura: MHNG 2156.30. Weste rn
Province: near Colombo: MHNG 1199.30–32.
Cylindrophis melanotus.—Indonesia: North Maluku Province: Bacan Island: SMF 16975; Halmahera: ZMB 34313 (holotype
of Cylindrophis heinrichi Ahl, 1933)
; Sanana Island (Soela-Sanana): RMNH.RENA 5104.176. Central Sulawesi
Province: Poso: ZMA.RENA 11453.117–19; Lake Wawontoa: ZMB 62929. South Sulawesi Province: Lake Tempe:
ZMA.RENA 11464.116; Makale: RMNH.RENA 11274.88; “Patmmang” (possibly Ujung Pandang, today’s Makassar):
NMW 21571.1–3. North Sulawesi Province: Lake Moat: ZMB 50020; Manado: RMNH.RENA 19.82, 173.18B, 174.18A,
5459.41–42; without precise locality data: RMNH.RENA 5461.34–40, ZMA.RENA 11451.112–15. Southeast Sulawesi
Province: Buton Island, Bau Bau: RMNH.RENA 11265.87; Kolaka: RMNH.RENA 11276.89. Mainland Sulawesi
(occasionally labeled as “Celebes”): without precise locality data: RMNH.RENA 17.83–84, 17.86, ZMA.RENA
11459.120, ZMB 1450, 4049 (potential holotype of Tortrix rufa var. celebica Schlegel, 1844).
Cylindrophis opisthorhodus.—Indonesia: East Nusa Tenggara Province: Flores Island: SMF 23301, ZMB 33787. West Nusa
Tenggara Province: Lombok Island: SMF 23299, ZMA.RENA 12135, 14082; Sumbawa Island: SMF 23300.
Cylindrophis ruffus sensu lato.—Indonesia: without precise locality data: ZMH R09749, R09786, R09793, R09797. “East
coast of Borneo”: RMNH.RENA 3924.15–17. “Java”: MHNG 2745.35–38, MTKD D5614–15, D7071, D14868–72,
NMW 13835–36, 21558.1, 21558.3, 21558.6, 21558.8, 21559.2–14, NMBE 1015767, RMNH.RENA 1.65–68, 46, 47927–
28, SMF 16976–78, 16981–82, 16984–86, 16990, ZMA.RENA 10495, 11452.145, 11467.151–53, 14460, ZMB 1455,
4908, 13129, 29696. “South Java”: ZMB 14443, 58433. “Sumatra”: NMW 21550.4–5. Aceh Province (Atje), Sumatra:
NMW 21550.2. Bangka-Belitung Islands Province: Bangka Island: ZMA.RENA 10487, 23068, 23070; Belitung Island:
ZMA.RENA 11471.177–79. Central Java Province: Kagok, Tegal: ZMA.RENA 11455.155; Pekalongan: ZMA.RENA
11468.157; Rembang: RMNH.RENA 11252.105; Semarang (Samarang): RMNH.RENA 5.60–61, ZMA.RENA
11461.158, ZMB 14351, 58429–30. Central Kalimantan Province (Borneo): Muara Teweh: NMW 21554.6. East Java
Province: without precise locality data: RMNH.RENA6928.52–55; Kediri: ZMA.RENA 11462.159, 11454.146–50;
Malang (Malary): NMW 21558.4–5; Mount Arjuno (Ardjoeno): RMNH.RENA 11260.108–09, 11261.93–94; Surabaya
(Surabaja, Soerabaja): RMNH.RENA 5791.49, 5999.58–59, 11251, 11252.105, ZMA.RENA 11457.154; Tengger
Mountains: NMB-REPT 471–73. Jakarta Province (Java): Jakarta (historically: Batavia): MTKD D14750, NMB-REPT
20441. North Sumatra Province: Langkat: RMNH.RENA 6349.25–26; Tanah Merah, Bindjey Estate: ZMH R09751–52.
Riau Province (Sumatra): Rantau Island: RMNH.RENA 8185.13; Sungai Lala: ZMH R09787. South Sumatra Province:
Tanjung Enim: ZMA.RENA 11458.126. Sultanate of Deli (Sumatra): NMW 21550.1, 21550.3, 21568.1–6, RMNH.RENA
6968.27–33, ZMA.RENA 10490, 11463.125, 11465.127, 11466.124. Sultanate of Serdang (Sumatra): ZMA.RENA
11460.123. West Java Province: Bogor (historically: Buitenzorg): NMB-REPT 462–70, RMNH.RENA 11256.110,
11258.92, 11272.98, SMF 16979–80, 16992–94, ZMB 20525; Cirebon (Cheribon): ZMA.RENA 11469.129–33;
Indramayu (Indramajoe): RMNH.RENA 8956.56, 8972.62–64; Itjabe: MHNG 676.67; Sukabumi (Soekaboemi):
ZMA.RENA 11456.156. West Kalimantan Province (Borneo): Badau: NMW 21554.5; Landak: ZMA.RENA 10488,
23064; Pontianak: RMNH.RENA 8234.2–3, 8264.5–6, 8264.8–11, 8264.14. Malaysia: Johor State: no precise locality
data: AMNH R-12873; Johor Bahru: ZRC 2.3009–10. Kelantan State: Kuala Lebir: ZRC 2.3011. Penang State: no precise
locality data: NMW 21570.2–3; Sarawak (Borneo): Baram: NMW 21554.1; Sungai Tangap, Niah: AMNH R-111923.
Singapore: no precise locality data: ZMH R09788–89, ZRC 2.3017–20, ZRC 2.3021, ZRC 2.3023, ZRC 2.6907; Bukit
Timah Road: ZRC 2.3022; Sembawang: Naval Base: ZRC 2.3029.
... Asian pipesnakes comprise a single genus (Cylindrophis Wagler, 1828) in the monotypic family Cylindrophiidae. There are 14 currently recognized species distributed in Sri Lanka and Southeast Asia (Kieckbusch et al. 2016(Kieckbusch et al. , 2018Uetz et al. 2020). These snakes are generally secretive but can be encountered in forests, wetlands, canals, flooded rice fields, and even suburban gardens and inside houses (Wall 1925;Smith 1943;Das 2015;. ...
... The other publications lack any voucher information. Kieckbusch et al. (2016) restricted C. ruffus, a taxon for which tissue samples appear unavailable, to parts of Maritime Southeast Asia (mainland Malaysia, Singapore, and the Greater Sunda Islands). This indicates that the sequence data for "C. ...
... for subsequent studies remains problematic due to the secretive nature and potential rarity of many of the relevant species, some of which are known from only very few (and sometimes historical) specimens with genetic samples unavailable (e.g., McDowell 1975;Stuebing & Goh 1993;Stuebing 1994;Gower et al. 2005;Kieckbusch et al. 2016Kieckbusch et al. , 2018. As a result, intrageneric relationships for Cylindrophis have recently been based on morphological characters. ...
Full-text available
Cylindrophis is a genus of secretive, semi-fossorial, non-venomous snakes comprising 14 species, characterized by a generally cylindrical body, uniform scales (with barely enlarged ventrals), and vestiges of pelvic and limb bones, the latter terminating in a claw lateral to the vent. We reconstructed a concatenated molecular phylogeny of seven taxa of Cylindrophis taxa based on one nuclear (R35) and two mitochondrial (16S, ND2) genes. Analyses recovered the Sri Lankan endemic C. maculatus as sister to all other sampled Cylindrophis. The mainland Southeast Asian species C. burmanus and C. jodiae form successive sister lineages to a monophyletic Wallacean island group containing C. boulengeri, C. isolepis, and C. yamdena. We also describe a new species of Cylindrophis, morphologically similar to C. burmanus, from Kachin State in northern Myanmar. Cylindrophis slowinskii sp. nov. is distinguished from all congeners by the following combination of characters: 19 dorsal scale rows at midbody, 216-220 ventrals, eight subcaudals, a dark venter with > 60 very narrow diffuse pale blotches, and a pale bar running along the posterior border of the prefrontals. In our phylogeny, the new species is strongly supported as the sister species of C. burmanus. It is the 15th currently recognised species in the genus, and the fourth from mainland Southeast Asia.
... Research on amphibians and reptiles in Java that had been conducted, e.g., , Riyanto and Kurniati (2014), Riyanto et al. (2015), Hartmann et al. (2016), Kieckbusch et al. (2016), Hamidy et al. (2018), Cahyadi and Arifin (2019) and Riyanto et al. (2019). They have demonstrated that herpetofauna diversity in the region is still underestimated. ...
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Megantara EN, Jauhan J, Shanida SS, Husodo T, Fauzi DA, Hendrawan R, Wulandari I, Yuansah. 2022. Herpetofauna distribution in different land cover types of West Java, Indonesia. Biodiversitas 23: 2990-2999. Herpetofauna is very sensitive and can be used as a biodiversity indicator. Herpetofauna can experience environmental stress due to agricultural activities, tourism, and other disturbances that result in habitat loss. Herpetofauna can be found on various land covers, such as natural forests and human-modified land. This study revealed species associated with natural forests, human-modified land, and both. This study aims to investigate: 1) herpetofauna distribution based on land cover types and 2) the disturbance that threatens the herpetofauna habitat in West Java. Visual Encounter Survey combination with the Auditory Encounter Survey, was applied in this study. The result of this study, the species were found in the natural forest (53 species), human-modified land (63 species), and crater (2 species). Thirty-nine species were found both in the natural forest and human-modified land. The habitat disturbance potentially threatens the species, such as land clearing (it will affect the microclimate of land cover), roadkill, infrastructure development, and tourism activities.
... The region experiences high anthropogenic impacts both from infrastructure and tourism activities. Research on amphi-bians and reptiles in Java that had been conducted, e.g., ; Riyanto and Kurniati (2014); Riyanto et al. (2015); Hartmann et al. (2016); Kieckbusch et al. (2016); Hamidy et al. (2018); Riyanto et al. (2019b); and Cahyadi and Arifin (2019) had revealed that amphibian and reptile diversity in the region is still underestimated. Therefore, it is essential to show the diversity of herpetofauna species to support conservation efforts. ...
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Erawan TS, Jauhan J, Husodo T, Wulandari I, Fauzi DA, Megantara EN, Shanida SS. 2021. Herpetofauna diversity and distribution based on the elevational range in West Java, Indonesia. Biodiversitas 22: 4308-4319. The herpetofauna discovery in West Java is still low and has not yet described the herpetofauna community's species diversity entirely and thoroughly in West Java. It experiences high anthropogenic impacts both from infrastructure and tourism activities. If left further, herpetofauna species will become extinct locally because of anthropogenic factors. Therefore, it is essential to reveal the diversity and distribution of herpetofauna species. This study aimed to show diversity and distribution based on the elevational range in West Java. Visual Encounter Survey in combination with the Auditory Encounter Survey was applied in this study. We found 74 species where the Colubridae family were found most commonly in the study sites. Of 74 species, four species have high conservation status, including Rhacophorus reinwardtii, Ophiophagus hannah, Varanus salvator, and Malayopython reticulatus. Besides, ten species were found to be endemic to Java and Java-Bali. Based on elevation, herpetofauna is mainly located at elevations < 1000 m asl. The higher the height, the fewer the number of species found. In Amphibians, the highest number of species tend to be found at 400-1200 m asl and 1400-1700 m asl, while reptiles tend to be located at 400-900 m asl. Keywords: Auditory Encounter Survey, West Java
... Our remeasurement of tail length brings the body proportions in line with the expected parameters. It is also important to ensure that all type material is known as such and, perhaps, to allow more light to be shed on the important collections made by the Commissie, parts of which were apparently dispersed to other important European collections of the day (e.g., Kieckbusch et al. 2016;Mecke et al. 2016a). It is noteworthy that, despite a good match in length measurements with RMNH 325B, Müller's count of 200 ventral and 70 subcaudal scales is not a count found in any of the three extant specimens of S. lividus, which may indicate that a fourth specimen existed at some point in the past. ...
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We report on the discovery of a third, male specimen of Stegonotus lividus in the collection of the Muséum National d'Histoire Naturelle in Paris, France, and demonstrate that it is not only a member of the original type series but the only one of the three syntypes, whose morphology was detailed in the original description. We herein identify it as a paralectotype. In their description of S. lividus, Duméril et al. (1854) attributed authorship of the name to the German zoologist Salomon Müller, whose work was never published. By the rules of zoological nomenclature, author attribution solely via an unpublished manuscript is inadmissible, and the species is therefore properly listed as Stegonotus lividus (Duméril et al., 1854). The recent discovery of Müller's handwritten manuscript, along with an unpublished drawing of one of these snakes by the Dutch artist Pieter van Oort, allows a better assessment of color and pattern for a species that remains known from only three preserved vouchers, as well as improved differentiation from other taxa occurring in the Lesser Sundas and Moluccas.
... Photographs of these species in life are not likely to be forthcoming in the near future, and even some of their genus allocations are speculative. Of course, digitization of data on the world's natural history museum collections is an ongoing process and we anticipate that some of these species may be "rediscovered" (and that undescribed species still lurk in undigitized, unexamined museum jars; e.g., Kaiser et al., 2020;Kieckbusch et al., 2016;O'Shea et al., 2020). ...
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The secretive behavior and life history of snakes makes studying their biology, distribution, and the epidemiology of venomous snakebite challenging. One of the most useful, most versatile, and easiest to collect types of biological data are photographs, particularly those that are connected with geographic location and date-time metadata. Photos verify occurrence records, provide data on phenotypes and ecology, and are often used to illustrate new species descriptions, field guides and identification keys, as well as in training humans and computer vision algorithms to identify snakes. We scoured eleven online and two offline sources of snake photos in an attempt to collect as many photos of as many snake species as possible, and attempt to explain some of the inter-species variation in photograph quantity among global regions and taxonomic groups, and with regard to medical importance, human population density, and range size. We collected a total of 725,565 photos—between 1 and 48,696 photos of 3098 of the world's 3879 snake species (79.9%), leaving 781 “most wanted” species with no photos (20.1% of all currently-described species as of the December 2020 release of The Reptile Database). We provide a list of most wanted species sortable by family, continent, authority, and medical importance, and encourage snake photographers worldwide to submit photos and associated metadata, particularly of “missing” species, to the most permanent and useful online archives: The Reptile Database, iNaturalist, and HerpMapper.
... Recent publications have either considered the Singapore type locality to be in error (e.g., Stuebing 1991: 327;Wallach et al. 2014: 204;Kieckbusch et al. 2016: 25;) or that the correct distribution of C. lineatus should be Borneo (e.g., Stuebing et al. 2014: 63;Kieckbusch et al. 2016;10;Bernstein et al. 2020: 556). It would therefore appear that C. lineatus is endemic to the island of Borneo and that the action of Smedley (1923: 11, 12) and de Haas (1950: 526) in correcting the type locality of Cylindrophis lineatus Dennys, 1880b from Singapore to Borneo is justified (Recommendation 76A.2 of the Code; ICZN 1999: 87). ...
The snake genus Cylindrophis Wagler, 1828 belongs to the monogeneric family Cylindrophiidae comprising 15 species distributed predominately throughout SE Asia, with one extralimital species occurring in Sri Lanka (Bernstein et al. 2020: 535). Cylindrophis lineatus is a rare species known from only eight museum specimens (discussed herein), and a photograph of one live individual from Kuching, Sarawak, East Malaysia (Stuebing et al. 2014: 63). Despite being originally described with Singapore as the type-locality (Blanford 1881: 217, 218), it is currently understood that C. lineatus is endemic to western Sarawak, East Malaysia (Stuebing et al. 2014: 63). Wallach et al. (2014: 204) stated that C. lineatus is also found in Kalimantan, but did not provide any references. This appears to have been followed by Bernstein et al. (2020: 537), who provide a map indicating C. lineatus occurrences in Kalimantan. The original description of Cylindrophis lineatus is conventionally cited as Blanford (1881: 217, 218, pl. 20). Herein, we demonstrate that the authorship and date of publication of this taxon should correctly be Cylindrophis lineatus Dennys, 1880b, and discuss that the type locality should be changed to “Borneo”.
... Gunung Muria is separated from Sumatera by ~ 580 km, Kalimantan by ~350 ROCK GECKO FROM JAVA, INDONESIA Zootaxa 4608 (1) © 2019 Magnolia Press · 171 km, and Belitung Island (the type locality of C. purnamai) by ~470 km. The African-Asian Frog genus Chiromantis also occurs in Java (Riyanto & Kurniati 2014;Wostl et al. 2017), as do other recently described reptiles and amphibians Hartmann et al. 2016;Kieckbusch et al. 2016;Wostl et al. 2017;Hamidy et al. 2018). These novelties have shown Java to be a more herpetologically interesting place than heretofore believed, even though it was believed to have been relatively well studied during the long history of exploration during the Dutch colonial period. ...
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We describe a new species of rock gecko of the genus Cnemaspis from Java, Indonesia, representing the first record of the genus for this Island. The new species was collected from the southern slopes of Gunung Muria, a dormant volcano in Central Java. The new species is easily distinguished from all congeners by having a maximum SVL of 58.1 mm in males and 56.9 mm in females; a pair of sharp conical tubercle clusters on the occiput; a warty bridge on the nuchal loop, extending from the upper tympanum and curving to the nape; dorsal tubercles not linearly arranged; 18–20 paravertebral tubercles; postmentals separated by one scale; gular, pectoral and abdominal scales, ventral scales of fore- and hindlimbs, and subcaudal scales keeled; no tubercles on lower flank; precloacal and femoral pores absent; enlarged submetacarpal scales present on the first digit of the manus; 38–40 ventral scales; 31–35 lamellae under fourth toe; two postcloacal tubercles on each side; enlarged median subcaudal scales row present; caudal tubercles encircling tail; and a sexually dimorphic ventral color pattern, with males having a yellow belly and females white and the ventral surface of the tail in males yellow proximally changing to white at mid-length, whereas in females, alternating black and white rings completely encircle the tail, which is black distally.
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An inventory of herpetofauna species from western part of Nusa Kambangan Island, Central Java, Indonesia, is presented. There are 43 herpetofauna species reported (16 amphibians and 27 reptiles). This study confirmed new distribution record and list some of threatened species. In light of the imminent human disturbances on Nusa Kambangan Island, a conservation plan is urgently needed.
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West Java Province has the largest population amongst others in Java, and therefore land conversion rate in the region is increasing. Approximately 40% of forest areas in West Java has been converted between 1990–2015. As a consequence, the number of bi­odiversity in the region is decreasing, including amphibians and rep­tiles. These groups play an important role in the food chain of an ecosystem, and are very sensitive to environmental changes. How­ever, comprehensive research on amphibian and reptile species in West Java is suboptimal. Visual Encounter Survey has performed in seven districts in West Java for one month and has recorded 26 amphibian species and 27 reptile species. These species were in­cluding Javan endemic species (for example: Fejervarya iskandari, Huia masonii, Limnonectes microdiscus, Megophrys montana, and Microhyla achatina), introduced species (Calotes versicolor), and species with a new distribution record (Leptophryne borbonica and Kalophrynus minusculus). In addition, cryptic species (Genus Lep­tophryne and Cyrtodactylus), which are interesting for further stud­ies, were observed. This study has demonstrated that the potential of a comprehensive study of amphibian and reptile species in West Java is great. Apart from this, conserving the biodiversity in the region also challenging due to the high rate in land conversion
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A new species of the gekkonid lizard genus Cyrtodactylus Gray, 1827 is described from Klakah, Lumajang Regency, Jawa Timur Province, Java, Indonesia. Cyrtodactylus klakahensis sp. nov. can be distinguished from all other congeners by the presence of (1) a deep precloacal groove in males, (2) three rows of enlarged precloaco-femoral scales, of which the third row bears 37–38 pores in males, (3) three or four rows of enlarged scales between precloacofemoral scale rows and cloaca, forming distinct chevrons, (4) raised and strongly keeled dorsal tubercles in 15–19 rows at midbody, (5) an indistinct lateral fold, (6) 17–20 subdigital lamellae under 4th toe, and (7) the absence of transversely enlarged subcaudal scales.Cyrtodactylus klakahensis sp. nov. is only the third bent-toed gecko species described from Java, indicating that the diversity of this genus in Java has been neglected in the past. Furthermore, we confirm that C. fumosus (Müller, 1895) is a species that possesses a precloacal groove in males and may be restricted to northern Sulawesi. The species is defined by a single female type specimen (holotype;NMB-REPT 2662). Specimens in museum collections catalogued as C. fumosus from localities elsewhere are misidentified and likely represent undescribed species.
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Cyrtodactylus petani sp. nov. is a new species of Bent-toed Gecko from Java, Indonesia that had been masquerading under the name C. fumosus (Müller, 1895). The new species is differentiated from C. fumosus and all its Sundaland congeners by having the following combination of morphological characters: a maximum SVL of 57.2 mm; nine or ten supralabials; seven or eight infralabials; strongly tuberculate body and limbs; 20-25 paravertebral tubercles; 30-35 ventral scales; enlarged precloacal scales; enlarged femoral scales; 17-18 subdigital lamellae on the fourth toe; 31-35 continuous precloacal and femoral pores in males, pores absent in females; no precloacal groove; no enlarged median subcaudals; tubercles on anterior portion of tail; no reticulated pattern on top of head; a blotched dorsal pattern; and no paired, dark, semi-lunar shaped blotches on the nape.
A new species of Cylindrophis closely related to Cylindrophis ruffus is described from Yamdena, Tanimbar Archipelago, Indonesia.
From the existing literature and data from museum specimens an overview is presented of all currently known terrestrial and semi-aquatic snakes of the Lesser Sunda Islands, in the Wallacean area of Indonesia. In total, twenty-nine species are known to inhabit the area. Of these eight are endemic to the area: Boiga hoeseli, Coelognathus subradiatus, Dendrelaphis inornatus, Stegonotus florensis, Cylindrophis opisthorhodus, Broghammerus timoriensis, Liasis mackloti and Typhlops schmutzi. Insular endemism is only found at the subspecific level, including Liasis mackloti dunni (Wetar), Liasis mackloti savuensis (Sawu), Ramphotyphlops polygrammicus brongersmai (Sumba), Ramphotyphlops polygrammicus elberti (Lombok) and Ramphotyphlops polygrammicus florensis (Flores). Such endemism may be due to the relatively young geological age of the Lesser Sunda Islands and that the snake fauna is still underestimated. Taxonomy of the genus Cylindrophis, the species Coelognathus subradiatus, Dendrelaphis inornatus, Cryptelytrops insularis, and the fve subspecies of Ramphotyphlops polygrammicus need to be reviewed. Ecological studies are urgently required to establish if the species Broghammerus timoriensis and Liasis mackloti savuensis are endangered and which conservation measures should be taken.