Parvoscincus 3 SP Nov 2014

Zootaxa 3847 (3): 388412
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Copyright 2014 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3847.3.4
http://zoobank.org/urn:lsid:zoobank.org:pub:6F77821C-2D03-4636-92AB-1789B2383633
Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos

(Reptilia: Squamata: Scincidae), with description of three new species
CAMERON D. SILER1, CHARLES W. LINKEM2, KERRY COBB3, JESSA L. WATTERS1,
SEAN T. CUMMINGS1, ARVIN C. DIESMOS4 & RAFE M. BROWN3
1
Department of Biology and Sam Noble Oklahoma Museum of Natural History; University of Oklahoma, 2401 Chautaugua Ave., Norman, OK 73072-7029, USA. E-mails: camsiler@ou.edu, jwatters@ou.edu, sean.t.cummings-1@ou.edu
2
Department of Biology; University of Washington, Seattle, WA 98195-1800, USA. Email: cwlinkem@gmail.com
3
Biodiversity Institute and Department of Ecology and Evolutionary Biology; University of Kansas, 1345 Jayhawk Blvd, Lawrence, KS
66045-7593, USA. E-mails: cobbkerry@gmail.com, rafe@ku.edu
4
Herpetology Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos St., Manila, Philippines.
E-mail: arvin.diesmos@gmail.com
Abstract
We review the recent discovery of multiple populations of the enigmatic, semi-aquatic Sphenomorphus Group skink, Parvoscincus leucospilos Peters, and investigate the morphological and genetic diversity of isolated, allopatric populations of
this unique skink. Our investigations support the recognition of four unique evolutionary lineages distributed across Luzon
Island in the Philippines, three of which are herein described as new species (P. tikbalangi sp. nov., P. manananggalae sp.
nov., and P. duwendorum sp. nov.). All four recognized species are genetically divergent in both mitochondrial and nuclear DNA sequences, and morphologically distinct. The description of three new Luzon Island endemic species adds to
the growing body of literature suggesting that mechanisms driving the accumulation of vertebrate diversity in the Philippines may vary regionally across the archipelago.
Key words: biodiversity, conservation, cryptic diversity, endemism, lizard, Luzon Island, riparian, Southeast Asia, Sphenomorphus group
Introduction
Recent studies have revealed that our understanding of amphibian and reptile diversity in the Philippines is vastly
underestimated (Brown et al., 2013a). Phylogeny-based studies focused on species delimitation have identified and
revised numerous species complexes within the Philippines (Brown and Guttman, 2002; Brown et al., 2009, 2010;
Siler et al., 2010a,b, 2011a,b, 2012; Siler and Brown, 2010; Welton et al., 2009, 2010a,b; Linkem et al., 2010a,b;
Linkem and Brown, 2013); many of these complexes were once considered by taxonomists to be widely distributed
species (e.g., Leviton, 1963; Brown and Alcala, 1970, 1980). More recent studies, fueled with evidence
documenting high levels of genetic divergence between isolated evolutionary lineages repeatedly have found that
few endemic Philippine reptiles actually possess broad distributions spanning regionally recognized faunistic
boundaries (Brown and Diesmos, 2009; Brown et al., 2013a; Linkem and Brown, 2013). Although molecular data
have had a tremendous impact on the discovery and identification of unique biodiversity in the archipelago,
extensive survey work has resulted in the rediscovery of several enigmatic reptile species (Eutropis bontocensis
Taylor [Barley et al., 2013], Brachymeles elerae Taylor [Siler, 2010], B. pathfinderi Taylor [Siler et al., 2011c],
Hologerrhum philippinum Gnther (McLeod et al., 2011; Brown et al., 2013b), Platymantis polillensis Taylor and
P. cornutus Taylor (Siler et al., 2011c; Brown et al., 2012, 2013b). These rediscoveries have resulted in a better
understanding of the ecology and distribution of these putatively rare species, and in many cases, revealed that they
are not in fact rare, simply misunderstood or with previously unappreciated microhabitat preferences. The
rediscovery and phylogenetic assessment of the Philippine endemic, semi-aquatic skink Parvoscincus leucospilos
Peters, 1872 (Brown et al., 2000, 2013; Linkem et al., 2011; McLeod et al., 2011; Siler et al., 2011d) is exactly one
such situation.
388 Accepted by L. Grismer: 28 May 2014; published: 8 Aug. 2014
Parvoscincus leucospilos was first described by Peters (1872), who placed the species in the genus Lygosoma
Gray, section Hinulia (=Sphenomorphus Fitzinger). Subsequently, the species was transferred by Boulenger (1885)
to the genus Tropidophorus Dumril and Bibron, and was recognized as such for nearly a century until Brown and
Alcala (1980) recognized it as a member of the genus Sphenomorphus based on examinations of two available
museum specimens (ZMB 7467; BM 72.8.20.43). In 2011, a phylogenetic study of Philippine forest skinks of the
genus Sphenomorphus revised the radiation of skinks in the archipelago, and recognized five genera as independent
from Sphenomorphus (Insulasaurus Taylor, Otosaurus Gray, and Parvoscincus Ferner, Brown, and Greer,
Pinoyscincus Linkem, Diesmos, and Brown, and Tytthoscincus Linkem, Diesmos, and Brown), placing this rare
Philippine endemic species in the genus Parvoscincus (Linkem et al., 2011).
Details of the status of the type material for P. leucospilos are in need of clarification. The description by Peters
(1872) was based on material collected by Adolf Meyer on Luzon. Peters (1872) does not indicate the number of
specimens available to him at the time of description, though mention of a variable number of lateral bands
indicates that there were potentially multiple specimens. Two specimens in museums are listed as syntypes
(cotypes), ZMB 7467 and CAS 64232 (Bauer, Shea, Gnther, 2003). The CAS specimen sheds more light on the
history of this taxon. The California Academy of Sciences received this specimen as part of an exchange from
Robert Mertens at Naturmuseum Senckenberg (SMF) in 1925 under the name Tropidophorus leucospilos
Cotype, the valid name at the time based on Boulenger (1885). The combination of the transfer documents listing
this specimen as a cotype and the CAS database indicating Peters was the identifying authority demonstrates that
the CAS specimen was likely included in Peters type series. We are unable to find documentation for the transfer
of this specimen from ZMB to SMF to complete the chain of custody and cannot definitively say that it is part of
the type series. The redescription by Brown and Alcala (1980) designated ZMB 7467 as the holotype, without
comment on the status of other type material. Despite examining other specimens in the same exchange from SMF,
Walter Brown never makes reference to the CAS specimen of P. leucospilos in his publications on Philippine
lizards, indicating he may have never examined it. Based on the presence of syntypes and the lack of reference to
other types in Brown and Alcala (1980) during their designation of ZMB 7467 as the holotype, the International
Code of Zoological Nomenclature (ICZN: Article 74.5) indicates that this designation does not constitute a valid
lectotype designation. We resolve these issues as part of a redescription of P. leucospilos.
Twenty-one species are recognized in the genus Parvoscincus (P. abstrusus Linkem and Brown, P. agtorum
Linkem and Brown, P. arvindiesmosi Linkem and Brown, P. aurorus Linkem and Brown, P. banahaoensis Linkem
and Brown, P. beyeri Taylor, P. boyingi Brown, Linkem, Diesmos, Balete, Duya, and Ferner, P. decipiens
Boulenger, P. hadros Brown, Linkem, Diesmos, Balete, Duya, and Ferner, P. igorotorum Brown, Linkem, Diesmos,
Balete, Duya, and Ferner, P. jimmymcguirei Linkem and Brown, P. kitangladensis Brown, P. laterimaculatus
Brown and Alcala, P. lawtoni Brown and Alcala, P. leucospilos, P. luzonensis Boulenger, P. palaliensis Linkem and
Brown, P. palawanensis Brown and Alcala, P. sisoni Ferner, Brown, and Greer, P. steerei Stejneger, and P.
tagapayo Brown, McGuire, Ferner, and Alcala; Linkem and Brown 2013). This unique genus of lizards is
represented by both morphologically and ecologically diverse species. Body sizes in the group range from larger,
montane forest species (P. beyeri, P. boyingi, P. igorotorum, and P. hadros) to species with smaller bodies (P.
tagapayao, P. luzonensis, P. lawtoni, P. kitangladensis, P. laterimaculatus, P. steerei, and P. decipiens; Linkem et
al., 2011; Brown et al., 2010, 2013b). Unlike most species of Parvoscincus that prefer terrestrial microhabitats,
Parvoscincus leucospilos is a semi-aquatic, riparian forest species, which has only been observed active near cool,
shaded, mountain streams where it retreats under streamside rocks and forest detritus, and dives in running water
when disturbed (CDS, RMB, personal observations).
Aside from being reported to occur on Luzon Island (no specific locality was reported originally), nothing was
known of this species for more than a century, until it was rediscovered during a 1997 survey of the herpetofauna in
Aurora Province, Luzon Island (Brown et al., 2000). The collection of a single individual from the surface of leaf
litter in mature second growth forest confirmed the species presence on Luzon Island. Several recent surveys from
throughout much of Luzon have shown this species to be semi-aquatic with a preference for riparian habitat along
fast moving mountain streams (Siler et al., 2011d; McLeod et al., 2011; Brown et al. 2012, 2013b). These surveys
have resulted in the discovery of several, isolated mid-montane populations in the northern, central, and southern
Sierra Madres, the northern Cordillera Mountains and the isolated Caraballo Mountains (Fig. 1). Several of these
populations exhibit striking differences in body coloration and pigmentation pattern.
The goal of this study is to investigate the systematic relationships of isolated allopatric populations of P.
REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS
Zootaxa 3847 (3) 2014 Magnolia Press
389
leucospilos. Additionally we examine these allopatric populations for evidence of boundaries between lineage
segments (sensu de Queiroz, 1998, 1999, 2005) such that hypothesized divisions between populations might reflect
uniquely evolving, cohesive evolutionary lineages (separate species). As a result of comprehensive examination of
all recently collected specimens of P. leucospilos from central and northern Luzon Island, we revise this group and
separate isolated populations into four diagnosable and presumably separately-evolving evolutionary lineages
(species). In this paper we provide a phylogenetic analysis and the first illustrations of these taxa, fully describe
each species, and clarify taxonomic boundaries. We also provide information on each species natural history,
ecology, and geographic distribution.
FIGURE 1. Map of the Luzon Island, Philippines, showing the collection localities for the new species and P. leucospilos.
390 Zootaxa 3847 (3) 2014 Magnolia Press
SILER ET AL.
Material and methods

Field work, sample collection, and specimen preservation. We conducted fieldwork on Luzon Island in the
Philippines (Fig. 1) between 1991 and 2012. Specimens were collected during day and night searches, euthanized
with aqueous chloretone, dissected for genetic samples (liver preserved in 95% ethanol or flash frozen in liquid
nitrogen), fixed in 10% formalin and eventually (< 2 mo) transferred to 70% ethanol. Tissue samples were obtained
from nearly all samples of Parvoscincus leucospilos from throughout the range on Luzon Island. Museum
abbreviations for specimens examined or sequenced in this study follow Sabaj Perez (2013).
Taxon sampling and outgroup selection for phylogenetic analyses. Because our primary goal was to
estimate phylogenetic relationships among the various populations of Parvocincus leucospilos, we sequenced
individuals from all sampled populations. Outgroup samples were selected based on the phylogenetic relationships
in Linkem et al. (2011). We used a total of 22 ingroup samples in phylogenetic inferences.
DNA extraction, purification, and amplification. We extracted total genomic DNA from tissues using the
modified guanidine thiocyanate extraction method of Esselstyn et al. (2008). We sequenced the mitochondrial
NADH Dehydrogenase Subunit 2 (ND2) gene, and the nuclear locus prostaglandin E receptor 4 (PTGER4), for all
samples using the primers and protocols provided in Linkem et al. (2011). Detailed methods for amplification and
sequencing of the focal genes follow Linkem et al. (2011). New sequence data was deposited in GenBank
(Appendix 1).
Alignment and phylogenetic analysis. Data were aligned using MAFFT v6.717b (Katoh et al. 2005) and
concatenated using a Python script written by CWL. Partitioned Bayesian analyses were performed in MrBayes
v3.2 (Huelsenbeck and Ronquist 2001; Ronquist et al. 2012) using optimal models of evolution from jModeltest
v2.1.4 (Guindon & Gascuel, 2003; Darriba et al., 2012) for each partition (ND2 codon positions, HKY + G;
PTGER4, F81 + G). The ND2 gene was partitioned by codon position to accommodate among site rate variation
(Brandley et al. 2005). The nuclear gene PTGER4 was not partitioned due to the low level of sequence variation
between samples and rate variation could be accommodated with a gamma parameter. The final alignment thus
consisted of 1,517 characters (ND2, 1,029 characters; PTGER4, 488 characters).
The Bayesian analysis was run two times for 5 million generations, sampling every 1000th generation with
default Markov chain Monte Carlo settings. Rates of molecular evolution across partitions were unlinked and the
branch length prior was changed based on the findings of previous studies (Marshall et al. 2006; Marshall 2010).
Chain convergence of the posterior distribution was assessed using TRACER v1.5 (Rambaut and Drummond
2007) and Are We There Yet (AWTY: Wilgenbusch et al. 2004; Nylander et al. 2007). A majority-rule consensus
tree from the converged chains was calculated in MrBayes v3.2.1 and is used for interpretation of phylogenetic
relationships. We considered nodes with posterior probabilities 0.95 to be well supported (Wilcox et al., 2002;
Leach and Reeder, 2002).
Morphological data. We examined fluid-preserved specimens (Appendix 2) for variation in meristic and
mensural characters. Sex was determined by gonadal inspection, and measurements were taken to the nearest 0.1
mm with digital calipers by KC. Characters were chosen based on Brown et al. (2010), Linkem et al. (2011), and
Linkem and Brown (2013) and include: snoutvent length (SVL), axillagroin distance (AGD), tail length (TL),
total length (TotL), head length (HL), head width (HW), fore-limb length (FLL), hind limb length (HLL),
snoutforearm length (SnFa), eye diameter (ED), snout length (SNL), interorbital distance (IOD), internarial
distance (IND), auricular opening diameter (AurD), midbody scale-row count (MBSR), paravertebral scale-row
count (PVSR), ventral scale-row count (VSR), Toe-IV lamellae count (ToeIVlam), supralabial count (SL),
infralabial count (IFL), loreal count (Lo), and supraocular count (SO). In the description, ranges are followed by
mean standard deviation in parentheses.
Results
Phylogeny. Phylogenetic analyses of the molecular data resulted in posterior clade probabilities above 95% for
most nodes (Fig. 2). All analyses recover four major clades within P. leucospilos (Fig. 2). A single individual from
Ilocos Norte Province (KU 329929) is divergent from all other sampled populations with a mitochondrial
uncorrected sequence divergence from remaining samples greater than 11% (Table 1). Samples from the Laguna/
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Quezon, Camarines Norte, and Bulacan Provinces are closely related with 04.5% sequence divergence and are
over 7% divergent from all other samples (mtDNA). Individuals sequenced from Aurora and Nueva Vizcaya
Provinces are 02.5% divergent from each other and are over 4.3% divergent from all other samples (mtDNA).
Samples from the Sierra Madre Mountain province of Isabela are very closely related with < 1.5% sequence
divergence and are over 4.3% divergent from all other samples (mtDNA).
TABLE 1. Uncorrected sequence divergence for the mitochondrial (below diagonal) and nuclear genes (above diagonal)
between the four semi-aquatic Philippine skink species of the Parvoscincus leucospilos Complex. Intraspecific
mitochondrial sequence divergence shown along the diagonal in bold.
duwendorum
leucospilos
manananggalae
tikbalangi
duwendorum
0.630.85
0.851.06
0.210.64
leucospilos
11.512.5
04.5
0.821.5
0.211.06
manananggalae
11.511.8
7.68.8
02.5
0.211.03
tikbalangi
11.712.3
7.18.6
4.35.6
01.5
FIGURE 2. Partitioned Bayesian majority rule consensus estimate of molecular phylogeny from the concatenated (ND2 +
PTGER4) data of Parvoscincus. Populations within the P. leucospilos Complex are genetically unique and are diagnosed as
new species based on morphological differences. Black circles label nodes with posterior probabilities above 0.95.
Uncorrected pairwise sequence divergences are generally low within the lineages defined here as species and
high among these lineages (Table 1). Percent divergences for the mitochondrial and nuclear data show that the
monophyletic lineages defined by our phylogenetic analyses (P. leucospilos, P. duwendorum sp. nov., P. tikbalangi
sp. nov., and P. manananggalae sp. nov.) are distinguished from congeners by levels of genetic divergence on par
with those between previously defined speciesviz., P. beyeri, P. hadros, P. boyingi, P. laterimaculatus, P. sisoni,
P. tagapayo (Table 1; Fig. 2; Linkem et al., 2011). The two most closely related lineages (P. duwendorum sp. nov.
and P. manananggalae sp. nov.) are separated by 4.25.6% mitochondrial sequence divergence. Sequence
divergences among the other lineages within the P. leucospilos populations are greater than 7.1% (Table 1; Fig. 2).
Morphology. Superficially the four lineages of Parvoscincus leucospilos appear morphologically similar,
SILER ET AL.
especially in overall body size and shape, pronounced distinctiveness of the head from the nuchal region, etc;
however, upon inspection, numerous non-overlapping differences were detected in meristic, mensural, and color
pattern characters for each population, readily defining four distinct evolutionary lineages within the species
(Tables 2, 3). Variation in morphological characters (Tables 2, 3) mirrors the results observed in phylogenetic
analyses and supports the recognition of four P. leucospilos group lineages. Characters differing among these four
lineages include: total body length, head and body scale counts and patterns, and pigmentation patterns (Tables 2,
3; species accounts below), all of which are typical morphological diagnostic characters employed by taxonomists
working with this genus (Brown et al., 2010; Linkem et al., 2010a, 2011; Linkem and Brown, 2013). We observed
no intraspecific mensural or meristic differences between the sexes of any of the four species.
In summary, each lineage possesses unique and non-overlapping suites of diagnostic character states of
morphology that correspond perfectly to clades defined in phylogenetic analyses of DNA sequence data. Combined
with biogeographic evidence, and clearly separate geographical ranges, our data unequivocally support the
designation of four evolutionary lineages worthy of taxonomic recognition.
Taxonomic conclusions. Genetic and morphologically datasets support the presence of a new species from
Northwestern Luzon Island, a new species from Northeastern Luzon Island, and a new species from Central Luzon
Island (Table 1; Fig. 2). Each of the four species formerly referred to P. leucospilos are morphologically and
genetically distinct from each other and from other Parvoscincus species. Each monophyletic lineage is endemic to
geologically and biogeographically isolated regions on Luzon Island, thereby providing additional support for the
distinctiveness of each clades evolutionary history and lineage integrity.
Characters observed among our series of specimens of P. leucospilos from South-central and Southeastern
Luzon Island are in general agreement with Peters (1872) original description, and closely match Brown and
Alcalas (1980) redescription and our own inspection of the syntype deposited at the California Academy of
Sciences (CAS 64232), and include the following suite of morphological characters: (1) medium body size (SVL
5255 mm); (2) body slender; (3) tail dorsolaterally compressed; (4) loreals roughly equal in height; (5) prefrontals
in broad medial contact; (6) supralabials six or seven; (7) infralabials 69; (8) midbody scale rows 3034; (9)
paravertebral scale rows 6167; (10) Toe-IV lamellae 1517; (11) dorsal white bands 69; (12) head pigmentation
mottled tan to white; and (13) ventral body surfaces creamy white. We therefore recognize true P. leucospilos as a
species that occurs in Central and Southeastern Luzon Island (Figs. 1, 2), and recognize the three additional
lineages within this unique group each as a new species.
Taxonomic accounts
Parvoscincus leucospilos (Peters 1872)
(Figs. 35)
Lygosoma (Hinulia) leucospilos Peters, 1872:684.
Sphenomorphus leucospilos Brown and Alcala, 1980:172; Bauer et al., 1995; Linkem et al., 2011; McLeod et al., 2011.
Parvoscincus leucospilos Linkem, Diesmos & Brown, 2011; Linkem & Brown 2013 (part)
Type designation. Based on ICZN articles 74.1 and 74.7 and in accordance with recommendation 74D we
designate ZMB 7467 as the lectotype of Parvoscincus leucospilos. Specimen CAS 64232 is designated a
paralectotype.
Diagnosis. Parvoscincus leucospilos can be distinguished from congeners by the following combination of
characters: (1) body size medium (SVL 42.654.4 mm); (2) Toe-IV lamellae 1517; (3) supralabials six or seven;
(4) infralabials 69; (5) midbody scale rows 3034; (6) paravertebral scale rows 6167; (7) prefrontals in medial
contact; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation heavily mottled;
(11) upper arm pigmentation absent; (12) subcaudal pigmentation absent; (13) dorsal white spots large, welldefined; (14) dorsal white bands 913; (15) lateral body coloration bright reddish-orange; (16) tail dorsolaterally
compressed; and (17) semi-aquatic (Tables 2, 3).
Comparisons. Characters distinguishing Parvoscincus leucospilos from all species of Parvoscincus are
summarized in Tables 2 and 3. Parvoscincus leucospilos most closely resembles P. duwendorum, P.
manananggalae, and P. tikbalangi. However, P. leucospilos differs from these three taxa by having head
393
pigmentation heavily mottled (vs. moderately mottled; Figs. 4, 5), and by the absence (vs. presence) of dark
pigmentation on the upper arm surface. Parvoscincus leucospilos further differs from P. duwendorum and P.
manananggalae by having prefrontals in medial contact (vs. separated); from P. duwendorum and P. tikbalangi by
having dorsal white spots large and well-defined (vs. faint); from P. manananggalae and P. tikbalangi by the
presence of bright reddish-orange lateral coloration (vs. absence of coloration [P. manananggalae], presence,
reduced, tan [P. tikbalangi]); from P. duwendorum by having a greater number of Toe-IV lamellae (1517 vs. 12), a
greater number of paravertebral scale rows (6167 vs. 60), a greater number of midbody scale rows (3034 vs. 26),
a tendency towards a greater number of longitudinal ventral scale rows (4147 vs. 41), and fewer dorsal white
bands (913 vs. 15); from P. manananggalae by the absence of subcaudal dark pigmentation (vs. presence); and
from P. tikbalangi by having a tendency towards a greater number of paravertebral scale rows (6167 vs. 5863)
and a tendency towards a greater number of midbody scale rows (3034 vs. 2832).
Redescription (based on examination of paralectotype, CAS 64232, and 10 recently collected specimens,
Appendix 2). Details of the head scalation of an adult male (KU 313870) are shown in Figure 4, and scientific
illustrations of the dorsal and lateral views of the head of the paralectotype are shown in Figure 6. A medium sized
Parvoscincus, SVL 42.654.4 mm, with clawed, pentadactyl limbs. Head distinct from next, with enlarged jaw
adductor musculature in the temporal region. Snout pointed in dorsal aspect, rounded in lateral aspect; rostral wide
forming a nearly perpendicular margin with frontonasal, slightly rounded margin with nasals; frontonasal wider
than long, in contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in broad medial contact,
in contact with anterior and posterior loreals, frontal, frontonasal, and first supraocular, and in some specimens, in
point contact with first supraciliary; frontal greatly longer than wide, in contact with two supraoculars on right, two
on left, rounded anteriorly, pointed posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal,
in contact with supraoculars IIIV or III and IV; interparietal arrowhead-shaped; parietals in moderate to broad
medial contact, in contact with fourth supraocular, postsupraoculars, and primary and secondary temporals;
primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral
overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large.
Nasal pierced in center by large naris, surrounded anteriorly by rostral, dorsally by frontonasal, posteriorly by
anterior loreal, and ventrally by first supralabial; anterior loreal one, posterior loreal roughly equal in size;
preoculars two; supralabials six or seven; lower eyelid scaly, semi-transparent, nonscaled window absent; ear
large, moderately sunk.
Infralabials 69, decreasing in size posteriorly in series; mental small, forming straight suture with single, large
postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than ventrals.
Body elongate, cylindrical, slender, with 3034 equal sized midbody scales, limbs overlapping when
adpressed; paravertebral scales 6167, imbricate, smooth, without striations, keels or pits. Tail elongate,
dorsolaterally compressed, longer than body (TL/SVL 0.991.57 [1.26 0.18]); subcaudal scales similar to lateral
scales for basal half of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales
between pelvic region and cloaca, more elongate than ventral scales; medial precloacal scales larger.
Forelimbs smaller than hind limbs, pentadactyl; forelimb scales slightly smaller in size than body scales,
imbricate and smooth, reducing slightly in size closer to manus; lamellae becoming slightly keeled distally on each
digit; relative digit length I < V < II < III = IV; palmar scales irregular, raised, forming ventral protrusions from
palmar surface. Hind limbs small, pentadactyl; hind limb scales equal in size and shape to body scales; dorsal
scales on digits multiple. Lamellae keeled proximally and distally, flat for a few scales in between on Toe IV; Toe
IV lamellae 1517; relative digit length I < II < V < III < IV; plantar scales irregular, slightly raised.
Coloration in preservative (Figs. 3, 4). The dorsal background color is medium brown, with pronounced
mottling of various shades of brown. Three longitudinal rows of cream spots, ringed with darker brown, start just
behind the head and run the length of the body with faint light brown longitudinal stripes between. Spotting and
striping fade into a solid medium brown color approximately halfway down the tail. On the lateral surface of the
body, spots merge to become indistinct perpendicular cream stripes, with mottled medium brown between. This
pattern fades indistinctly into the solid cream present on the ventral surface. The distinctive forelimb spot is
pronounced and continues nearly to the first joint. Otherwise, the mottled pattern present on the dorsal surface of
the body continues down the dorsal surface of the fore- and hind limbs. The ventral surface of the limbs is a solid
cream, with the exception of very faint light brown mottling present on the hands and feet. The solid ventral cream
color of the body extends to about halfway down the length of the tail, then transitions to thin medium brown
SILER ET AL.
perpendicular stripes. The stripes become increasingly close together as they approach the tail tip, which is a solid
medium brown. The dorsal surface of the head is mottled the same as the dorsal surface of the body, with various
shades of brown. Three small cream spots are arranged in a triangle on the head just posterior to the eyes. A series
of distinct cream spots are visible on the labial scales, one row on the supralabial series and one on the infralabial
series. A slightly lighter brown mottling occurs just below the eye and is present between the spots. The ventral
surface of the head and body are solid cream in color.
FIGURE 3. Dorsal views of representative specimen heads and flanks, exhibiting pigmentation pattern variation among
species.
395
FIGURE 4. Photographs of heads from dorsal and lateral perspectives to show pigmentation and scale pattern variation among
species. With the exception of P. leucospilos (KU 313870), all photographs are of the holotypes of the new species.
Coloration in life (Differences from preserved specimens; Fig. 5). On the lateral surfaces of the body, the dark
brown, mottled ground coloration is replaced with a vivid reddish-orange ground color, with randomly distributed
small cream spots. The non-mottled patch on the upper forelimb proximate to the body is also a solid reddishorange color. The cream and brown mottling on the lateral surface of the head and the body anterior to the forelimb
insertion is replaced by a pale blue and brown mottled pattern.
Measurements and scale counts of paralectotype in mm. SVL 44.2; AGD 23.2; TotL 101.0; TL 56.8; HL
6.1; HW 5.4; SnFa 16.5; ED 2.2; SNL 3.7; IND 1.0; FLL 4.3; HLL 6.2; MBSR 31; PVSR 63; ToeIVlam 17; SL 7;
IFL 7; SO 4.
SILER ET AL.
Distribution, ecology and natural history. Parvoscincus leucospilos is known from south-central and
southeastern Luzon Island (Fig. 1). No mention of the specific locality or province on Luzon Island was given for
the types (Peters, 1872). Although we are certain of our identification of P. leucospilos, we are unable to pinpoint
the type locality on the basis of available specimens. Specimens are from Mt. Labo, Camarines Norte Province
(KU313870), Mt. Palali, Barangay Maddiangat, Municipality of Quezon, Nueva Vizcaya Province (KU
32580613), Barangay Kabayunan, Municipality of Dona Remedios Trinidad, Bulucan Province (KU 32938892)
and Mt. Banahao, Municipality of Tayabas, Quezon Province (TNHC 62682). The known elevational range of this
species is 200800 meters above sea level, based on Brown et al. (2013b).
Parvoscincus leucospilos occurs in primary- and secondary-growth forest habitats in riparian microhabitats;
this semi-aquatic species frequently dives into cold, rapidly running water of montane streams when disturbed. The
species is primarily found active during the day and has also been found sleeping on the edge of running water in
forest debris at night. Parvoscincus leucospilos can be found in sympatry with Parvoscincus abstrusus, P.
arvindiesmosi, P. beyeri, P. jimmymcguirei, P. laterimaculatus, and P. steerei.
We have evaluated this species against the IUCN criteria for classification, and find that it does not qualify for
Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Parvoscincus leucospilos is quite
abundant at sampled localities provided that some vegetation cover borders the streams where it has been collected
and observed. We therefore classify this species as Least Concern LC (IUCN, 2010).
Etymology. The specific epithet leucospilos apparently refers to the distinctive coloration of this species,
consisiting of prominent rows of white (from the Latin adjective leucos) spots (Latin, noun, spilos) down the
dorsal surfaces of the body.
Parvoscincus duwendorum sp. nov.

(Figs. 35)
Parvoscincus leucospilos Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2012.
Holotype. PNM 9793 (RMB Field No. 14261, formerly KU 329929), subadult, collected on 18 June 2011, on Mt.
Pao, Barangay Adams, Municipality of Adams, Ilocos Norte Province, Luzon Island, Philippines (N: 18.438, E:
120.878; WGS-84; 750 m in elevation), by RMB.
Diagnosis. Parvoscincus duwendorum can be distinguished from congeners by the following combination of
characters: (1) adult body size presumed medium (subadult SVL 33.5 mm); (2) Toe-IV lamellae 12; (3)
supralabials seven; (4) infralabials eight; (5) midbody scale rows 26; (6) paravertebral scale rows 60; (7)
prefrontals separated; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation
moderately mottled; (11) upper arm pigmentation present, patchy; (12) cloacal scale dark pigmentation absent; (13)
subcaudal pigmentation absent; (14) dorsal white spots faint; (15) dorsal white bands 15; (16) tail dorsolaterally
compressed; and (17) semi-aquatic (Tables 2, 3).
Comparisons. Characters distinguishing Parvoscincus duwendorum from all species of Parvoscincus are
summarized in Tables 2 and 3. Parvoscincus duwendorum most closely resembles P. manananggalae, P.
leucospilos, and P. tikbalangi. However, P. duwendorum differs from these three taxa by having Toe-IV lamellae 12
(vs. 17 [P. manananggalae], 1517 [P. leucospilos], 1416 [P. tikbalangi]), fewer midbody scale rows (26 vs.
3233 [P. manananggalae], 3034 [P. leucospilos], 2832 [P. tikbalangi]), a greater number of dorsal white spot
rows (15 vs. 913 [P. leucospilos], 912 [P. manananggalae, P. tikbalangi]), and the absence of precloacal scale
dark pigmentation (vs. absence or presence [P. leucospilos, P. tikbalangi], presence [P. manananggalae]).
Parvoscincus duwendorum further differs from P. leucospilos and P. manananggalae by having fewer
paravertebral scale rows (60 vs. 6167 [P. leucospilos], 6169 [P. manananggalae]); from P. manananggalae and
P. tikbalangi by having infralabials eight (vs. six or seven [P. manananggalae], seven [P. tikbalangi]); from P.
leucospilos by having head pigmentation moderately mottled (vs. heavily mottled), and the presence of dark
pigmentation on the upper arm surface (vs. absence); from P. manananggalae by having fewer longitudinal ventral
scale rows (41 vs. 4349), and the absence (vs. presence) of subcaudal dark pigmentation; from P. leucospilos and
P. tikbalangi by having prefrontals separated (vs. in medial contact [P. leucospilos], separated or in medial contact
[P. tikbalangi]); and from P. tikbalangi by having prefrontals and first supraoculars in contact (vs. separated).
397
SILER ET AL.
Locality
Mt. Banahao/Angat Dam, Luzon
Adams, Luzon
Aurora, Luzon
Isabela, Luzon
Polillo; Bicol, Luzon
Aurora, Luzon
Bicol, Luzon
Aurora, Luzon
Mt. Banahao, Luzon
Mt. Banahao, Luzon
Zambales & Bataan Mtns, Luzon
Sierra Madre, Luzon
Sierra Madre, Luzon
Northern Cordilleras, Luzon
Northern Cordilleras, Sierra Madre
Central Mindanao
S. Luzon
N. Luzon
N. Luzon
Mt. Palali, Luzon
Palawan
Panay
Widespread
Sierra Madres, Luzon
Species
leucospilos
duwendorum
manananggalae
tikbalangi
abstrusus
agtorum
arvindiesmosi
aurorus
banahaoensis
beyeri
boyingi
decipiens
hadros
igorotorum
jimmymcguirei
kitangladensis
laterimaculatus
lawtoni
luzonensis
palaliensis
palawanensis
sisoni
steerei
tagapayo
23.132.1
26.436
26.533.6
28.134.3
39.3
39.947.8
29.347.0
42.157.1
4656.3
33.844.1
51.9, 57.5
73.586.7
31.145
46.166.7
57.872.9
39.345.1
39.646.6
30.943.1
44.9
32.043.3
41.554.7
47.355.9
33.5
42.654.4
SVL
2630
2832
2426
2124
32
2729
2529
3438
3038
3137
44, 45
4547
3238
3742
3842
2832
3135
3236
39
3336
2832
3233
26
3034
MBSR
5470
5263
6168
4854
73
6573
5364
7483
6877
5872
98, 106
108111
5766
8896
88102
6266
6575
6574
71
5869
5863
6169
60
6167
PVSR
911
914
1112
1012
14
912
1015
1618
1518
1722
20
1822
1418
1921
1821
1417
1517
1619
17
1618
1416
17
12
1517
ToeIVlam
67
56
67
67
68
67
68
67
SL
67
57
78
67
79
67
69
IFL
1, 2
1, 2
+, 0
NA
+, 0
+, 0
+, 0
contact
Prefontal
Anterio
r loreals
1 & 2 forest
1 & 2 forest
Montane-forest
Montane-forest
Montane-forest
1 & 2 forest
1 & 2 forest
1 & 2 forest
1 & 2 forest
1 & 2 forest
Montane-forest
Montane-forest
1 & 2 forest
Montane-forest
Montane-forest
Montane-forest
Montane-forest
1 & 2 forest
1 & 2 forest
1 & 2 forest
Semi-aquatic
Semi-aquatic
Semi-aquatic
Semi-aquatic
preference
Microhabitat
2, A
2, D
1, C
6, A
11
size
Sample
TABLE 2. Distribution of diagnostic characters (+ present; 0 absent) among species in the genus Parvoscincus. Parvoscincus palawanensis does not have prefrontal scales. (A) Brown et al. 2010,
(B) Linkem and Brown, 2013, (C) Brown and Alcala 1980, (D) Ferner et al. 1997. Characters included for comparison are: snoutvent length (SVL), midbody scale-row count (MBSR),
paravertebral scale-row count (PVSR), Toe-IV lamellae count (ToeIVlam), supralabial count (SL), infralabial count (IFL), number of anterior loreals, contact between profrontal scales, and
microhabitat preference.
399
+
Moderately mottled
+, patchy
Faint
15
Data unavailable
8 (1)
60
41
26
7 (1)
33.5
83.5
149
3.3
10
4.0
12
12 (1)
duwendorum
(1 juv.)
Northwestern Luzon
+
Moderately mottled
+, patchy
+
+
Large, well defined
912
6 (3)
7 (2)
6169
4349
3233
7 (5)
47.355.9 (52.7 3.3)

122.9146.8 (130.8 10.8)
120168 (142 20)
4.35.5 (4.8 0.5)
810 (9 1)
5.86.7 (6.2 0.4)
1112 (12 1)
17 (5)
manananggalae
(3 m, 2 f)
East-central Luzon
Paralectotype (CAS 64232) presumed male following visual inspection of external body; authors avoided creating incision on rare type specimen.
Prefrontal contact
Prefrontal/1st supraocular contact
Head pigmentation
Upper arm dark pigmentation
Cloacal scale dark pigmentation
Subcaudal dark pigmentation
Dorsal white spots
Dorsal white bands
Lateral body coloration
IFL
PVSR
LoVent
MBSR
SL
SVL
TotL
TL/SVL
FLL
FLL/SVL
HLL
HLL/SVL
ToeIVlam
Range
leucospilos
(6 m1, 5 f)
South-central & Southeastern
Luzon
42.654.4 (50.0 3.7)
100.6126.1 (112.6 9.5)
99157 (126 18)
3.85.1 (4.5 0.4)
810 (9 0)
5.26.5 (6.0 0.4)
1113 (12 0)
15 (6)
16 (1)
17 (4)
6167
4147
3034
6 (1)
7 (10)
6 (1)
7 (6)
8 (3)
9 (1)
+
+
Heavily mottled
or +
Large, well defined

913
+, bright reddish-orange
or +
Moderately mottled
+, patchy
or +
Faint
912
+, reduced, tan
7 (8)
41.554.7 (49.0 4.6)

118.9138.5 (128.4 8.9)
138159 (150 9)
4.05.0 (4.6 0.4)
910 (9 0)
5.26.5 (6.0 0.5)
1213 (12 0)
14 (4)
15 (2)
16 (2)
5863
4144
2832
7 (8)
tikbalangi
(7 m, 1 f)
Northeastern Luzon
TABLE 3. Summary of meristic and mensural, diagnostic characters among the four semi-aquatic Philippine skink species of the Parvoscincus leucospilos Complex.. For
character definitions, please refer to Material and Methods.
FIGURE 5. Photographs in life of (A) P. leucospilos (Mt. Banahao; TNHC 62683); (B) P. leucospilos (Angat Watershed; KU
329388); (C) P. manananggalae sp. nov. (PNM 9794); and (D) P. tikbalangi sp. nov. (Holotype; PNM 9795).
Description of holotype. Details of the head scalation are shown in Figure 4. A subadult Parvoscincus, SVL
33.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor
musculature in temporal region. Snout sharply pointed in dorsal aspect, rounded in lateral aspect; rostral wide
forming a nearly perpendicular margin with nasals and frontonasal; frontonasal equally wide as long, in contact
with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals broadly separated, in contact with anterior and
posterior loreals, frontal, frontonasal, first supraciliary, and first supraocular; frontal greatly longer than wide, in
contact with two supraoculars on right, two on left, rounded anteriorly, sharply rounded posteriorly; four enlarged
supraoculars, first largest; single, large frontoparietals, in contact with supraoculars IIIV; interparietal arrowhead;
parietals in moderate medial contact, in contact with fourth supraocular, postsupraocular, and secondary temporal;
primary temporals two, ventral largest, overlapping dorsal; secondary temporals two, large, dorsal largest, ventral
overlapping dorsal; tertiary temporals two, dorsal largest, ventral overlapping dorsal; auricular opening large.
anterior loreal, and ventrally by first supralabial; anterior loreal one, posterior loreal equal in size; preoculars two;
supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent, nonscaled window absent; ear large,
moderately sunk.
Infralabials eight, decreasing in size posteriorly in series; mental small, forming straight suture with single,
large postmental and first infralabials; enlarged chin shields in three pairs; gular scales slightly smaller than
ventrals.
Body elongate, cylindrical, slender, with 26 equal sized midbody scales, limbs overlapping when adpressed;
paravertebral scales 60, imbricate, smooth, without striations, keels or pits. Tail elongate, dorsolaterally
compressed, longer than body (TL [50.0] / SVL [33.5] 1.49); subcaudal scales nondifferentiated for basal half of
tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region
and cloaca, more elongate than ventral scales; medial precloacal scales larger.
SILER ET AL.
FIGURE 6. Illustration of head of the examined Parvoscincus leucospilos paralectotype (CAS 64232) in dorsal and lateral
views. Taxonomically diagnostic head scales are labeled as follows: C, chin shield; F, frontal; FN, frontonasal; FP,
frontoparietal; IL, infralabial; IP, interparietal; L, loreal; M, mental; N, nasal; P, parietal; PF, prefrontal; PM, postmental; PO,
preocular; PSO, presubocular; PoSO, postsupraoculars; R, rostral; SC, supraciliary; SL, supralabial; SO, supraocular; T1,
primary temporal; T2, secondary temporal; and T3, tertiary temporal. Roman numerals indicate scales in the supraocular series.
Illustrations by STC and CDS.
401
imbricate and smooth, reducing slightly in size closer to manus; lamellae becoming slightly keeled distally on each
digit; relative digit length with lamellae V < I < II < III = IV; palmar scales irregular, raised, formal ventral
protrusions from palmar surface. Hind limbs small, pentadactyl; hind limb scales equal in size and shape to body
scales; dorsal scales on digits multiple. Lamellae keeled proximally and distally, flat for a few scales in between on
Toe-IV; Toe-IV lamellae 12; relative digit length with lamellae I < II < III < V < IV; plantar scales irregular, slightly
raised.
Coloration of holotype in preservative. The dorsal background color is medium brown, with tan mottling
throughout. Two parallel light brown stripes run longitudinally down the body, starting at the base of the head and
merging to one just past the start of the tail. Between these two stripes is a paravertebral row of darker brown spots.
The venter color is a solid cream, without mottling. Along the lateral surface of the body, the dorsal and ventral
color patterns merges in a scalloped pattern. There is a distinct cream spot on the forelimbs, just proximate to the
body. The dorsal fore- and hind limbs possess the same mottling pattern present on the dorsal surface of the body.
The ventral surfaces of the limbs are a solid cream color. The dorsal surface of the hands and feet are a mottled
medium brown, with the exception of the cream-colored Finger I and Finger II on the forelimbs and Toe V on the
hind limbs. The dorsal and dorsolateral surfaces of the head have a light and medium brown mottled pattern. This
coloration transitions sharply into a cream and light brown mottled pattern ventrally. The labial scales are mottled
light brown and cream. The ventral surface of the head is solid cream in color.
Color in lifeColoration in life did not differ substantially from the preserved holotype (RMB, personal
observations).
Measurements and scale counts of holotype in mm. SVL 33.5; AGD 17.5; TotL 83.5; TL 50.0; HL 4.0; HW
7.7; SnFa 12.4; ED 1.5; SNL 2.7; IND 1.3; FLL 3.3; HLL 4.0; MBSR 26; PVSR 60; ToeIVlam 12; SL 7; IFL 8; SO
4.
Distribution, ecology and natural history. Parvoscincus duwendorum is known from a single specimen
found at 750 elevation on Mt. Pao in the Ilocos Norte Province of northwestern Luzon Island (Fig. 1). Although the
species is recognized currently to occur in secondary-growth forest habitats, it is presumed the species also inhabits
suitable primary forest. This species is semi-aquatic and can be found near riparian habitat. Parvoscincus
duwendorum can be found in sympatry with Parvoscincus igorotorum, P. jimmymcguirei, and P. steerei. The new
species was also observed on Mt. Cagua at 300500 m (Brown et al 2013b) but specimens eluded biologists by
jumping in rapidly running montane streams.
At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due
to the lack of available information about its distribution and natural history. We therefore classify this species Data
Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink.
Etymology. The specific epithet is a plaural derivation of the Filipino folklore term Duwende, which is
chosen here to celebrate the countrys rich tradition of mythological forest animals and spirits. Duwendes are
believed to be little fairy-like forest creatures, such as goblins, pixies, and elves, and believed to live in trees,
termite mounds, and burrows in hillsides. The may bring bad or good fortune to humans and are often considered to
be mischievous in nature. Suggested common name: Cordillera Aquatic Skink.
Parvoscincus manananggalae sp. nov.

(Figs. 35)
Sphenomorphus leucospilos: Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al., 2000; Siler et
al., 2011d.
Holotype. PNM 9794 (RMB Field No. 10719, formerly KU 323928), adult male, collected on rocks near a rapid
flowing stream during the day on 21 June 2009, in Barangay Lipimental, Municipality of San Luis, Aurora
Province, Luzon Island, Philippines (N: 15.653; E: 121.507; WGS-84; 515 m in elevation), by RMB, CDS, L.
Welton.
Paratypes. KU 32392027, 32392930 collected 21 June 2009; CMNH 5792 (from Brown et al. 2000).
Diagnosis. Parvoscincus manananggalae can be distinguished from congeners by the following combination
of characters: (1) body size medium (SVL 47.355.9 mm); (2) Toe-IV lamellae 17; (3) supralabials seven; (4)
infralabials six or seven; (5) midbody scale rows 3233; (6) paravertebral scale rows 6169; (7) prefrontals
SILER ET AL.
separated; (8) prefrontals contact first supraocular; (9) frontoparietals fused; (10) head pigmentation moderately
mottled; (11) upper arm pigmentation present, patchy; (12) cloacal scale dark pigmentation present; (13) subcaudal
pigmentation present; (14) dorsal white spots large, well-defined; (15) dorsal white bands 912; (16) bright lateral
body coloration absent; (17) tail dorsolaterally compressed; and (18) semi-aquatic (Tables 2, 3).
Comparisons. Characters distinguishing Parvoscincus manananggalae from all species of Parvoscincus are
summarized in Tables 2 and 3. Parvoscincus manananggalae most closely resembles P. duwendorum, P.
leucospilos, and P. tikbalangi. However, P. manananggalae differs from these three taxa by the presence of
subcaudal dark pigmentation (vs. absence). Parvoscincus manananggalae further differs from P. duwendorum and
P. tikbalangi by having dorsal white spots large and well-defined (vs. faint), Toe-IV lamellae 17 (vs. 12 [P.
duwendorum], 1416 [P. tikbalangi]); from P. leucospilos and P. tikbalangi by the absence of bright lateral
coloration (vs. presence and bright reddish-orange [P. leucospilos], presence, reduced, tan [P. tikbalangi]); from P.
duwendorum by having a greater number of paravertebral scale rows (6169 vs. 60), a greater number of
longitudinal ventral scale rows (4349 vs. 41), a greater number of midbody scale rows (3233 vs. 26), dorsal
white bands 912 (vs. 15), and the presence of cloacal scale dark pigmentation (vs. absence); from P. leucospilos
by having prefrontals separated (vs. in medial contact), head pigmentation moderately mottled (vs. heavily
mottled), and the presence of dark pigmentation on the upper arm surface (vs. absence); and from P. tikbalangi by
having a tendency towards a greater number of midbody scale rows (3233 vs. 2832) and fewer paravertebral
scale rows (6169 vs. 5863), and the presence of contact between prefrontal and first supraocular scales (vs.
absence).
Description of holotype. Details of the head scalation are shown in Figure 4. An adult male Parvoscincus,
SVL 55.9 mm, with clawed, pentadactyl limbs; head distinct from neck, characterized by enlarged jaw adductor
musculature in temporal region. Snout pointed in dorsal aspect, rounded in lateral aspect; rostral wide forming a
nearly perpendicular margin with frontonasal, rounded margin with nasals; frontonasal equally wide as long, in
contact with nasals, rostral, anterior loreal, and prefrontal scales; prefrontals in point medial contact, in contact
with anterior and posterior loreals, frontal, frontonasal, first supraciliary, and first supraocular; frontal greatly
longer than wide, in contact with two supraoculars on right, two on left, rounded anteriorly, rounded posteriorly;
four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars IIIV;
interparietal arrowhead-shaped; parietals in narrow medial contact, in narrow contact with fourth supraocular, in
moderate contact with postsupraocular, and secondary temporal; primary temporals two, ventral largest,
overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals
two, dorsal largest, ventral overlapping dorsal; auricular opening large.
anterior loreal, and ventrally by first and second supralabial; anterior loreal one, narrow, sharply curved contact
with nasal, posterior loreal lower in height, 2x width of anterior loreal; preoculars two; supralabials seven, fifth
subocular; lower eyelid scaly and semi-transparent, nonscaled window absent; ear large, moderately sunk.
Infralabials seven, decreasing in size posteriorly in series; mental small, forming straight suture with single,
ventrals.
compressed, longer than body (TL [67.0] / SVL [55.9] 1.20); subcaudal scales nondifferentiated for basal half of
tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic region
and cloaca, more elongate than ventral scales; medial precloacal scales larger; left hemipene everted.
imbricate and smooth, reducing slightly in size distally; lamellae becoming slightly keeled distally on each digit;
relative digit length I < V < II < III = IV; palmar scales irregular, raised, forming ventral protrusions from palmar
surface. Hind limbs small, pentadactyl; hind limb scales equal in size and shape to body scales; dorsal scales on
digits multiple. Lamellae keeled proximally and distally, flat for a few scales in between on Toe-IV; Toe-IV
lamellae 17; relative digit length I < II < V < III < IV; plantar scales irregular, slightly raised.
Coloration of holotype in preservative. The dorsal background color is dark brown, with extensive mottling
of several shades of brown. Three dorsal longitudinal rows of cream spots start posterior to the head and continue
to the anteriormost portions of the tail, where the pattern becomes mottled more darkly brown. Lighter brown
403
mottling is present in between the three dorsal rows of spots. The dorsal mottling pattern is more broken up with
small cream spots on the lateral surface of the body, until eventually becoming a solid cream on the ventral side of
the body. Similar small cream spots are present and sporadically distributed across the dorsal surface of both sets of
limbs, continuing distal to the distinct large cream band on the forelimb, proximate to the body. The ventral
surfaces of the feet are heavily mottled brown, except for Fingers I and II and Toe V. The solid cream ventral body
coloration is broken up at the cloaca by four heavily pigmented enlarged precloacal scales. The ventral surface of
the tail is mottled medium and dark brown, except for the area just posterior to the cloaca, which is solid cream in
color. The mottled pattern on the dorsal surface of the head is a lighter brown than the dorsal surface of the body.
This mottling is slightly darker around the orbits. There are two small cream spots just anterior to the eyes. There is
a transition to a lighter mottled pattern on the lateral surface of the head starting just ventral to the eyes that
continues to the venter. The supra- and infralabial series have four distinct tan spots, surrounded on either side by
one indistinct spot. The ventral surface of the head is solid cream.
Coloration of holotype in life (Differences from preserved specimens; Fig. 5). The cream mottling and
indistinct cream spots present on the lateroventral surface of the head, body, and tail is replaced by light pale
bluish-gray coloration. Bright lateral coloration is absent in this species.
Measurements and scale counts of holotype in mm. SVL 55.9; AGD 29.3; TotL 122.9; TL 67.0; HL 12.2;
HW 7.3; SnFa 18.1; ED 2.7; SNL 4.2; IND 2.1; FLL 5.5; HLL 6.6; MBSR 32; PVSR 68; ToeIVlam 17; SL 7; IFL
7; SO 4.
Variation. Scale pigmentation was observed to vary among the examined series: dark cloacal scale
pigmentation was present (KU 323920, 323922, 323923, 323925, 323926, 32392830, 325810) or absent (KU
323921, 323924, 323927, 325807, 328808, 32581113).
Distribution, ecology and natural history. Parvoscincus manananggalae is known only from east-central
Luzon Island, in Aurora Province near the Barangay of L. Pimentel in the San Luis Mountains and from Mt. Palali
in Nueva Vizcaya Province (Fig. 1). This semi-aquatic species occurs in primary- and secondary-growth forest
habitats, and occurs in sympatry with Parvoscincus agtorum, P. palaliensis, P. steerei, and P. tagapayo.
We have evaluated this species against the IUCN criteria for classification, and find that it does not qualify for
Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Parvoscincus manananggalae has a
relatively broad geographic distribution on Luzon Island and is quite abundant at all sampled localities. We
therefore classify this species as Least Concern LC (IUCN, 2010).
Etymology. The specific epithet is a feminine noun, formed from the name Manananggal, a female, blood
sucking, vampire-like creature who flies like a bat at night to hunt humans, after separating from her lower
extremities (derived from the Tagalog word Tanggal, to separate). Manananggal can be repelled by garlic and even
killed by heavily salting her legs once she has left to hunt for the night. Suggested common name:Aurora Aquatic
Skink.
Parvoscincus tikbalangi sp. nov.

(Figs. 35)
Parvoscincus leucospilos: Linkem, Diesmos, Brown, 2011 (part); Linkem & Brown, 2013 (part); Brown et al. 2013b.
Holotype. PNM 9795 (ACD Field No. 1989, formerly KU 327785), adult male, collected on 15 February 2005, in
Sitio Apaya, Barangay Dibuluan, Municipality of San Mariano, Isabela Province, Luzon Island, Philippines (N:
17.029; E: 122.1928; WGS-84; 600 m in elevation), by ACD.
Paratypes. KU 320522, 327786 collected on 5 February 2005 in Sitio Apaya, Barangay Dibuluan,
Municipality of San Mariano, Isabela Province (same coordinates), by ACD. KU 32778796 collected on 2426
April 2005 in Barangay Del Pilar, Municipality of San Mariano, Isabela Province (N: 122.104, E: 16.8592), by
ACD.
Diagnosis. Parvoscincus tikbalangi can be distinguished from congeners by the following combination of
characters: (1) body size medium (SVL 41.554.7 mm); (2) Toe-IV lamellae 1416; (3) supralabials seven; (4)
infralabials seven; (5) midbody scale rows 2832; (6) paravertebral scale rows 5863; (7) prefrontals separated
from first supraocular; (8) frontoparietals fused; (9) head pigmentation moderately mottled; (10) upper arm
pigmentation present, patchy; (11) subcaudal pigmentation absent; (12) dorsal white spots faint; (13) dorsal white
SILER ET AL.
bands 912; (14) lateral body coloration present, tan; (15) tail dorsolaterally compressed; and (16) semi-aquatic
(Tables 2, 3).
Comparisons. Characters distinguishing Parvoscincus tikbalangi from all species of Parvoscincus are
summarized in Tables 2 and 3. Parvoscincus tikbalangi most closely resembles P. duwendorum, P. leucospilos, and
P. manananggalae, but differs from P. duwendorum by having seven infralabials (vs. eight); from P. duwendorum
and P. manananggalae by having Toe-IV lamellae 1316 (vs. 12 [P. duwendorum], 17 [P. manananggalae]); from
P. duwendorum by having a greater number of midbody scale rows (2832 vs. 26) and fewer dorsal white bands
(912 vs. 15); from P. leucospilos and P. manananggalae by having a tendency towards fewer midbody scale rows
(2832 vs. 3034 [P. leucospilos], 3233 [P. manananggalae]), fewer paravertebral scale rows (5863 vs. 6167
[P. leucospilos], 6169 [P. manananggalae]), faint dorsal white spots (vs. large and well-defined), and by the
presence of tan lateral coloration (vs. presence and bright reddish-orange [P. leucospilos], absence [P.
manananggalae]); from P. leucospilos by having seven supralabials (vs. six or seven), moderate head pigmentation
mottling (vs. heavy), the presence (vs. absence) of dark pigmentation on the upper arm surface; and from P.
manananggalae by the absence (vs. presence) of dark subcaudal pigmentation.
Description of holotype. Details of the head scalation are shown in Figure 4. An adult male Parvoscincus,
SVL 53.5 mm, with clawed, pentadactyl limbs. Head distinct from neck, characterized by enlarged jaw adductor
musculature in temporal region. Snout rounded in dorsal aspect, rounded in lateral aspect; rostral wide forming a
nearly straight margin with nasals and frontonasal; frontonasal wider than long, in contact with nasals, rostral,
anterior loreal, and prefrontal scales; prefrontals in broad medial contact, in contact with anterior and posterior
loreals, frontal, frontonasal, and first supraciliary, separated from first supraocular; frontal greatly longer than wide,
in contact with two supraoculars on right, two on left, and first supraciliary, rounded anteriorly, rounded
posteriorly; four enlarged supraoculars, first largest; single, large frontoparietal, in contact with supraoculars IIIV;
interparietal arrowhead-shaped; parietals in narrow medial contact, in narrow contact with fourth supraocular, in
moderate contact with postsupraocular, and secondary temporal; primary temporals two, ventral largest,
overlapping dorsal; secondary temporals two, large, dorsal largest, ventral overlapping dorsal; tertiary temporals
two, dorsal largest, ventral overlapping dorsal; auricular opening large.
anterior loreal, and ventrally by first and second supralabial; anterior loreal one, narrow, posterior loreal roughly
equal in size; preoculars two; supralabials seven, fifth subocular; lower eyelid scaly and semi-transparent,
nonscaled window absent; ear large, moderately sunk.
Infralabials seven, decreasing in size posteriorly in series; mental small, forming straight suture with single,
ventrals.
compressed, longer than body (TL [85.0] / SVL [53.5] 1.59); subcaudal scales equal to lateral scales for basal half
of tail, enlarged for distal half of original tail. Precloacal region with series of enlarged scales between pelvic
region and cloaca, more elongate than ventral scales; medial precloacal scales larger; hemipenes everted.
imbricate and smooth, reducing slightly in size distally; lamellae becoming slightly keeled distally on each digit;
relative digit length I < V < II < III = IV; palmar scales irregular, raised, formal ventral protrusions from palmar
surface. Hind limbs small, pentadactyl; hind limb scales equal in size and shape to body scales; dorsal scales on
digits multiple. Lamellae keeled proximally and distally, flat for a few scales in between on Toe-IV; Toe-IV
lamellae 16; relative digit length I < II < V < III < IV; plantar scales irregular, slightly raised.
Coloration of holotype in preservative. The background dorsal coloration is dark brown, with a mottled
pattern of many different shades of brown. Two faint light brown stripes run longitudinally along the length of the
body, with faint light brown spots distributed between. Rows of cream spots are distributed on the dorsolateral
surface of the body. The posterior half of the tail is solid medium brown in coloration. The lateral surfaces of the
body are heavily mottled dark brown and cream, with small cream spots distributed throughout. The limbs are
heavily mottled dark brown and cream, with the exception of a large cream spot on the upper forelimbs, proximate
to the body insertion. The ventral surface of the limbs is solid cream in color, with the exception of very faint light
brown mottling present on the hands and feet. The ventral portion of the body is solid cream in color to the tip of
405
the tail, which has a medium brown mottled pattern. The dorsal surface of the head is mottled light and medium
brown. This mottled color pattern is slightly darker between the eyes. The lateral surface of the head has a slightly
lighter brown mottled color pattern. There is a row of six distinct cream spots on the supralabials, which are paired
with another six cream spots on the infralabials.
Coloration of holotype in life (Differences from preserved specimens; Fig. 5). The cream mottled color
pattern and indistinct cream spots present on the lateral surface of the head transition into pale bluish-gray mottled
coloration. The lateroventral surfaces of the axillagroin region of the body and the tail are colored pale orangecream. Within the axillagroin region and again on the tail, the coloration transitions from more orange in
coloration anteriorly to more cream posteriorly. The surface around the upper forelimbs, proximate to the body, is
colored light orange-cream.
Measurements and scale counts of holotype in mm. SVL 53.5; AGD 26.7; TotL 138.5; TL 85.0; HL 12.2;
HW 7.2; SnFa 18.2; ED 3.1; SNL 3.8; IND 1.8; FLL 5.0; HLL 6.4; MBSR 30; PVSR 62; ToeIVlam 16; SL 7; IFL
7; SO 4.
Variation. We observed the following variation in the degree of contact between head scales: prefrontals in
medial contact (KU 320522, 32778593, 327795, 327796) or separated (KU 327794). Scale pigmentation was
observed to vary among the examined series: dark cloacal scale pigmentation was present (KU 320522, 327785,
327791) or absent (KU 32778690, 32779296).
Distribution, ecology and natural history. Parvoscincus tikbalangi is known only from northeastern Luzon
Island in the Isabela Province (Fig. 1). Samples were found in the Barangays Del Pilar and Dibuluan in the
Municipality of San Mariano, in the Sierra Madre Mountain range. This semi-aquatic species of forest skink occurs
in primary- and secondary-growth forests, in riparian microhabitats.
At this time we are unable to appropriately evaluate this species against the IUCN criteria for classification due
to the lack of available information about its distribution and natural history. We therefore classify this species Data
Deficient DD (IUCN, 2010) pending future studies on this unique semi-aquatic forest skink.
Etymology. The specific epithet is a patronym derivation of the name Tikbalang, a mythological part man,
part horse mountain forest creature from Filipino folklore. Tikbalang is said to jump down from trees to dispatch its
unsuspecting victims by decapitation. Suggested common name: Sierra Madres Aquatic Skink.
Discussion
Analyses of both loci (ND2, PTGER4) resulted in topologies with high ML bootstrap support and posterior
probabilities for four lineages representing distinct evolutionary lineages previously referred to Parvoscincus
leucospilos on Luzon Island in the northern Philippines (Fig. 2). These data, when coupled with diagnostic
morphological features, confirm the presence of at least four distinct species of semi-aquatic forest skinks in the
genus Parvoscincus, three of which we describe as new species. The four species together make up a monophyletic
complex of forest skinks specialized on low to mid-montane forest river systems; they are known only from Luzon
Island where a tendency towards within-island scincid lizard radiation prevails (Brown and Alcala, 1980; Linkem
et al., 2010b, 2011; Brown et al., 2010, 2013a).
All four species occur allopatrically on isolated mountainous regions of Luzon Island (Fig. 1). Interestingly,
the species complex spans several well-defined biogeographic regions on Luzon Island, including the Bicol
Peninsula in the southeast, the Sierra Madres Mountain Range along the northeastern coast, the isolated Caraballo
Mountains (Mt. Palali) in central Luzon, and the northernmost extreme of the Cordillera Mountain Range in the
northwest portions of the island (Fig. 1). It would not be surprising if future studies discovered additional
populations of P. leucospilos Complex skinks in other mountainous regions of Luzon Island, particularly in the
central and southern Cordillera mountain ranges, and the Zambales Mountain Range (Fig. 1).
The description of P. manananggalae, P. duwendorum, and P. tikbalangi bring the total number of species of
Parvoscincus on Luzon Island to 21 (Linkem and Brown, 2013). Notably, this diversity represents more than half
of the diversity of Philippine Sphenomorphus Group skink species (now recognized to total 38; Linkem et al.,
2011; Linkem and Brown, 2013). Over the last ten years alone, the collection of molecular data, increase in
archipelago-wide surveys, and accumulation of robust morphological datasets has led to a significantly improved
understanding of endemic scincid diversity in the Philippines (for review, see Siler et al., 2012; Linkem and
Brown, 2013; Davis et al., in press). The results of all these studies suggest one thing for certain: we most likely
SILER ET AL.
still vastly underestimate reptile diversity in this geographically dynamic island nation. Not only will future studies
undoubtedly discover and describe additional species of skinks, but, as we approach a more accurate understanding
of the countrys taxonomic diversity for various species groups, researchers will be able to begin questioning
broad-scale mechanisms of diversification (Brown et al., 2013a). For example, what are the mechanisms driving
the disparate accumulation of biodiversity in different regions of the archipelago? Considering scincid lizards
alone, why does the Luzon faunal region possess by far the highest species richness in the archipelago (Linkem and
Brown, 2013; Davis et al., in press)? Studies focused on the mechanisms driving the assembly and evolution of
vertebrate communities are poignant topics for future investigation (Brown et al., 2013a).
Acknowledgments
We thank the Protected Areas and Wildlife Bureau (PAWB) of the Philippine Department of Environment and
Natural Resources (DENR) for facilitating collecting and export permits necessary for this and related studies; we
are particularly grateful to M. Lim, C. Custodio, and A. Tagtag. Financial support for fieldwork was provided by a
Panorama Fund grant from The University of Kansas Biodiversity Institute, travel funds from The University of
Kansas Department of Ecology and Evolutionary Biology, a Madison and Lila Self Fellowship from the University
of Kansas, a Fulbright Fellowship, a Fulbright-Hayes Fellowship, NSF DEB 0804115 to CDS, and DEB 0743491,
and NSF EF-0334952 to R. Brown. Lab work was funded from NSF DEB 0910341 to CWL. ACD thanks the
Rufford Foundation for support of his Sierra Madre fieldwork. For the loans of specimens we thank J. Vindum and
D. Blackburn (California Academy of Sciences), R. Sison and J. Barnes (Philippine National Museum), J. Ferner
(Cincinnati Museum Center), A. Resetar (Field Museum of Natural History), R. Crombie and K. de Queiroz
(United States Natural History Museum), T. LaDuc (Texas Memorial Natural History Museum), J. Rosado (MCZ),
and Maklarin Lakim (Sabah Parks). For access to the Sam Noble Museum Invertebrate Paleontology Stacking
Photography Lab we thank S. Westrop and R. Burkhalter. Critical reviews of the manuscript were provided by B.
Hedges and L. Grismer.
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RMB 808
KU 325808
KU 325809
KU 325810
KU 323922
KU 323930
KU 323925
KU 323926
ACD 2423
PNM 9793 (formerly KU
329929)
RMB 3346
KU 313870
KU 329391
KU 329392
RMB 3758
TNHC 62682
KU 320522
KU 327786
KU 327793
KU 327794
KU 327795
KU 327787
KU 327788
KU 327790
KU 327791
KU 309908
Otosaurus cumingi
Parvoscincus manananggalae2
Parvoscincus decipiens
Parvoscincus duwendei1
Parvoscincus laterimaculatus
Parvoscincus leucospilos
Parvoscincus tikbalangi2
Pinoyscincus coxi coxi

Voucher
Species
Philippines, Luzon Island, Camarines Sur Province

Philippines, Luzon Island, Camarines Norte Province
Philippines, Luzon Island, Bulacan Province
Philippines, Luzon Island, Bulacan Province
Philippines, Luzon Island, Laguna Province
Philippines, Luzon Island, Laguna Province
Philippines, Luzon Island, Isabela Province
Philippines, Camiguin Sur Island, Camiguin Province
Philippines, Luzon Island, Aurora Province

Philippines, Luzon Island, Nueva Vizcaya Province
Philippines, Luzon Island, Ilocos Norte Province
Locality
KM066669
KM066672
KM066675
KM066676
KM066673
KM066674
KM066684
KM066685
KM066686
KM066687
KM066688
KM066689
KM066690
KM066691
KM066692
KM066668
KM066671
KM066655
JF498151
KM066656
KM066657
KM066658
JF498155
JF498154
KF425469
KF425470
KF425474
KM066666
KF425471
KF425472
KF425475
KF425473
GU573563
KM066677
KM066678
KM066679
KM066680
KM066681
KM066682
KM066683
KM066670
KM066659
KM066660
KM066661
KM066662
KM066663
KM066664
KM066665
JF498145
Genbank Accession Numbers

ND2
PTGER4
JF498125
KM066667
APPENDIX 1. Summary of specimens corresponding to genetic samples included in the study, general locality, and GenBank
accession number. ACD = Field collections of Arvin Diesmos, deposited at the National Museum of the Philippines; KU = University
of Kansas Natural History Museum; RMB = Field collections of Rafe Brown, deposited at the National Museum of the Philippines.
APPENDIX 2. Additional material examined.

Insulasaurus arborens: PHILIPPINES, PANAY ISLAND, Antique Province, Municipality of San Remigio, Barangay Aningalan:
KU 306805306809, 306830.
Insulasaurus traanorum: PHILIPPINES, PALAWAN ISLAND, Palawan Province, Municipality of Rizal, SW of Mt.
Mantalingajan peak: PNM 9640 (Holotype); Mt. Paray-Paray: KU 311440, 311441, 311443 (Paratypes); Mt.
Mantalingahan: KU 311423, PNM 9641 (Paratypes).
Insulasaurus victoria: PHILIPPINES, PALAWAN ISLAND, Palawan Province, Municipality of Brookes Point, Barangay
Samarinana, Mt. Mantalingahan Range: KU 309443.
Insulasaurus wrighti: PHILIPPINES, PALAWAN ISLAND, Palawan Province: FMNH 5263, 62633; Municipality of Rizal, Mt.
Paray-Paray: KU 311417311422; Mt Mantalingahan: KU 311436311439.
Otosaurus cumingi: PHILIPPINES, POLILLO ISLAND, Quezon Province, Municipality of Polillo, Barangay Pinaglubayan: KU
302923; MINDORO ISLAND, Occidental Mindoro Province, Municipality of Sablayan, Barangay Batong Buhay: KU
304067304068, 304070, 304072, 305728; LUZON ISLAND, Camarines Sur Province, Naga City, Barangay Panicuason,
Mt. Isarog: TNHC 62748; Zambales Province, Municipality of Olongapo: TNHC 62749.
Parvoscincus abstrusus: PHILIPPINES, LUZON ISLAND, Laguna Province, Municipality of Los Baos, Mt. Makiling: KU
320063, 32006570, 32658891, 330743, 330744, 331676, 33167881, PNM 9783 (Holotype); Camarines Norte
Province, Municipality of Labo, Barangay Tulay Na Lupa, Mt. Labo: KU 313859, 313861, 313864, 3138689, PNM
2087; Camarines Sur Province, Municipality of Naga City, Barangay Panicuason, Mt. Isarog: TNHC 628836; Albay
Province, Municipality of Tiwi, Barangay Banhaw, Mt. Malinao: TNHC 62887, 62888; Municipality of Malinao,
Barangay Tagoytoy, Mt. Malinao: TNHC 628968; POLILLO ISLAND, Quezon Province: KU 304073.
Parvoscincus agtorum: PHILIPPINES, LUZON ISLAND, Aurora Province, Municipality of San Luis, Barangay Lipimental:
PNM 9782.
Parvoscincus arvindiesmosi: PHILIPPINES, LUZON ISLAND, Camarines del Norte Province, Municipality of Labo, Barangay
Tulay na Lupa, Mt. Labo: KU 306560, 306561, 313866, PNM 9781 (Holotype); Quezon Province, Municipality of
Tayabas, Barangay Lalo, Mt. Banahao: PNM 8611, TNHC 62679, 62890.
Parvoscincus manananggalae: See Holotype and Referred Specimens sections.
Parvoscincus aurorus: PHILIPPINES, LUZON ISLAND, Aurora Province, Municipality Maria Aurora, Barangay Villa Aurora,
Mt. Dayap: KU 323309, 32332023, PNM 9786 (Holotype).
Parvoscincus banahaoensis: PHILIPPINES, LUZON ISLAND, Quezon Province, Municipality of Tayabas, Barangay Lalo: KU
3274267, 327635, PNM 6761, 6762, 6760, 9784 (Holotype), TNHC 6289295.
Parvoscincus beyeri: PHILIPPINES, LUZON ISLAND, Quezon Province, Municipality of Tayabas, Mt. Banahao: CAS 61183
(Holotype); Barangay Lalo, Mt. Banahao: PNM 6757, 666667, PNM 96269628, 96299631, KU 320128, 320134,
TNHC 6267374; Barangay Lalo: FMNH 2661226619, 267559, 267560.
Parvoscincus cf. beyeri: PHILIPPINES, LUZON ISLAND, Nueva Vizcaya Province, Municipality of Quezon, Barangay
Maddiangat, Caraballo Mountains, Mt. Palali: KU 308666.
Parvoscincus boyingi: PHILIPPINES, LUZON ISLAND, Zambales Province, Municipality of Masinloc, Barangay Coto, Mt.
High Peak: CMNH 365255, 365759, PNM 2301, 2302 (Holotype), 23032305, 2307, USNM 337768; Mt. Apoy:
FMNH 267561, 267664, PNM 2300.
Parvoscincus decipiens: PHILIPPINES, LUZON ISLAND, Camarines Norte Province, Municipality of Labo, Barangay Tulay Na
Lupa, Mt. Labo: KU 306559 306561, 313849, 313859313869; Camarines Sur Province, Naga City, Barangay
Panicuason, Mt. Isarog: TNHC 62883, 6288562888, 62891; Albay Province, Municipality of Malinao, Barangay
Tagoytoy, Mt. Malinao: TNHC 6289662898.
Parvoscincus duwendorum: See Holotype and Referred Specimens sections.
Parvoscincus hadros: PHILIPPINES, LUZON ISLAND, Aurora Province, Municipality of Dingalan, Mt. Mingan: KU 320132,
320133, PNM 9618 (Holotype), 9619, 9620, 9624, 9625, 9632; Municipality of San Luis, Barangay Villa Aurora, Aurora
National Park: CMNH 5772.
Parvoscincus igorotorum: PHILIPPINES, LUZON ISLAND, Kalinga Province, Municipality of Balbalan, Barangay Balbalasang,
Balbalan-Balbalasang National Park: FMNH 259448, PNM 9623 (Holotype).
Parvoscincus jimmymcguirei: PHILIPPINES, LUZON ISLAND, Cagayan Province, Municipality of Gonzaga, Barangay
Magrafil, Mt. Cagua: KU 330122, 330126, 330128, PNM 9780 (Holotype); Nueva Vizcaya Province, Municipality of
Quezon, Barangay Maddiangat: KU 325796; Bulacan Province, Municipality of Dona Remedios Trinidad, Barangay
Kabayunan: KU 329401; Ilocos Norte Province, Municipality of Adams, Barangay Adams: KU 329931, 329935, 329941,
329945, 329949.
Parvoscincus kitangladensis: PHILIPPINES, MINDANAO ISLAND, Bukidnon Province, Mt. Kitanglad Range, SE of Baunson:
CAS 191084, 250641250643 (Paratypes), 250644 (Holotype).
Parvoscincus laterimaculatus: PHILIPPINES, LUZON ISLAND, Sorsogon Province, Municipality of Bulusan, Barangay San
Roque: CAS-SU 24204 (Holotype); Albay Province, Municipality of Tabaco, Barangay Buang, Mt. Mayon National Park,
Mt. Mayon: PNM 5306; Municipality of Tiwi, Baranga Banhaw, Sitio Purok 7, Area Tamagong, Mt. Malinao: TNHC
6267662678, KU 321816, 321817; Camarines Norte Province, Municipality of San Lorenzo Ruis, Mt. Labo Range:
PNM 2119, 2128, FMNH 27021, 27022; Camarines Sur Province, Municipality of Naga City, Barangay Panicuason, Mt.
411
Isarog National Park, Mt. Isarog: CAS 191800, PNM 2311 (formerly USNM 318342; Holotype of S. knollmanae), 9673,
TNHC 62675, USNM 318341, 318343, 318344; MARINDUQUE ISLAND, Marinduque Province, Municipality of Boac:
LSUMZ 5258952591.
Parvoscincus lawtoni: PHILIPPINES, LUZON ISLAND: FMNH 177674; Bicol Peninsula: FMNH 270525270527; Nueva
Vizcaya Province, Municipality of Quezon, Barangay Maddiangat, Mt. Palali: KU 308643, 308652, 308661, 308662,
308668, 308686; Kalinga Province, Municipality of Balbalan: CAS 6150161502, 61504.
Parvoscincus leucospilos: PHILIPPINES, LUZON ISLAND: CAS 64232 (Paralectotype); Quezon Province, Municipality of
Tayabas, Barangay Lalo, Mt. Banahao: TNHC 6268062683, PNM 96369639; Camarines Norte Province, Municipality
of Labo, Barangay Tulay Na Lupa, Mt. Labo: KU 313870, 32938892.
Parvoscincus luzonensis: PHILIPPINES, LUZON ISLAND, Highlands of Kipanto: CAS-SU 24147; Mountain Province,
Municipality of Bauko, Barrio Sinta: USNM 51276062; Kalinga Province, Municipality of Balbalan, Balbalasang:
FMNH 258990.
Parvoscincus palaliensis: PHILIPPINES, LUZON ISLAND, Nueva Vizcaya Province, Municipality of Quezon, Barangay
Maddiangat, Mt. Palali: KU 308651, 30869092, PNM 9785 (Holotype).
Parvoscincus palawanensis: PHILIPPINES, PALAWAN ISLAND, Palawan Province, Municipality of Puerto Princesa, Barangay
Iwahig, Malabo Peak: CAS-SU 23122 (Holotype), 91905 (Paratype).
Parvoscincus tikbalangi: See Holotype and Referred Specimens sections.
Parvoscincus sisoni: PHILIPPINES, PANAY ISLAND, Antique Province, Municipality of Culasi, Barangay Alojipan, Hanggud
Tubig, Mt. Madja-as: ACD 002 (deposited at PNM), CAS 193110, CMNH 37973799 (Paratypes), PNM 2308
(Holotype), 2309.
Parvoscincus steerei: PHILIPPINES, LUZON ISLAND, Municipality of Masinloc, Barangay Coto, south slope, Mt. Apoy:
CMNH 436167; PNM 245764; CEBU ISLAND, Cebu Province, Cebu City, Taptap barrio: CAS 139099, 13910107,
139111139112.
Parvoscincus tagapayo: PHILIPPINES, LUZON ISLAND, Aurora Province, Municipality of San Luis, Barangay Villa Aurora,
Aurora National Park, Kabatangan River drainage: PNM 5767 (Holotype), 5766, 5668, CMNH 5631, 5632 (Paratypes);
Municipality of Maria Aurora, Barangay Villa Aurora: CMNH 5633 (Paratype); Nueva Vizcaya Province, Municipality of
Quezon, Barangay Maddiangat, Mt. Palali: KU 325818325825.
Pinoyscincus abdictus: PHILIPPINES, POLILLO ISLAND, Quezon Province, Municipality of Polillo, Barangay Pinaglubayan:
KU 302911302922; LUBANG ISLAND, Occidental Mindoro Province, Municipality of Lubang, Barangay Vigo: KU
304041304049; BABUYAN CLARO ISLAND, Cagayan Province, Municipality of Calayan, Barangay Babuyan Claro: KU
304787304790.
Pinoyscincus coxi: PHILIPPINES, MINDANAO ISLAND, Davao Province: FMNH 5257151573; Agusan del Sur Province,
Agusan Valley, Bunawan: CAS 6204462046.
Pinoyscincus jagori: PHILIPPINES, PANAY ISLAND, Antique Island, Municipality of San Remigio, Barangay Aningalan: KU
306805306809; DINAGAT ISLAND, Dinagat Province, Municipality of Loreto, Barangay San Juan: KU 306535306544.
Pinoyscincus llanosi: PHILIPPINES, LEYTE ISLAND, Leyte Province, Municipality of Baybay, Barangay Pilim: KU 311269,
311270; SAMAR ISLAND, Eastern Samar Province, Municipality of Taft, Barangay San Rafael: KU 310322310324,
310786310795, 310840, 310856, 310857.
Pinoyscincus mindanensis: PHILIPPINES, MINDANAO ISLAND, Agusan Del Norte Province: CAS 133291, 133322; DINAGAT
ISLAND, Dinagat Province, Municipality of Loreto, Barangay Santiago: KU 310135; SAMAR ISLAND, Eastern Samar
Province, Municipality of Taft, Barangay San Rafael: KU 310809.
Sphenomorphus acutus: PHILIPPINES, SAMAR ISLAND, Eastern Samar Province, Municipality of Taft, Barangay San Rafael:
KU 310818; MINDANAO ISLAND, Agusan del Sur Province, Municipality of San Francisco, Barangay Bagasan II, Mt.
Magdiwata: KU 319961319964.
Sphenomorphus diwata: PHILIPPINES, MINDANAO ISLAND, Agusan del Norte Province, Diwata Mountains: CAS 2478
(Holotype), 1331415, KU 320129 320131, PNM 96339635.
Sphenomorphus fasciatus: PHILIPPINES, MINDANAO ISLAND, Davao City: FMNH 5262352627; Zamboanga City Province,
Municipality of Pasonanca, Barangay Pasanonca, Pasonanca Natural Park: KU 315061315066; LEYTE ISLAND, Leyte
Province, Municipality of Baybay, Pilim, San Vicente: KU 311252, 311253; SAMAR ISLAND, Eastern Samar Province,
Municipality of Taft, Barangay San Rafael: KU 310808; Western Samar Province, San Jose de Buan, Barangay Poblacion:
KU 310807; DINAGAT ISLAND, Dinagat Province, Municipality of Loreto, Barangay San Juan: KU 310093.
Sphenomorphus variegatus: PHILIPPINES, MINDANAO ISLAND, Davao City: FMNH 52617, 52627, 52630, CAMIGUIN SUR
ISLAND, Camiguin Province, Municipality of Mambajao: Barangay Pandan, KU 309899309907;
Tytthoscincus atrigularis: PHILIPPINES, MINDANAO ISLAND, Zamboanga City Province, Municipality of Pasonanca,
Barangay Pasanonca; Sitio Canucutan; Tumaga River, Pasonanca Natural Park: KU 305031305060; Davao City
Province, Municipality of Malalag, Kibaualan: CAS-SU 28789, 28791, 28794, 28795; BASILAN ISLAND, Basilan
Province, Municipality of Abung-abung: CAS 60242, 60244, 60246, 60247.
Tytthoscincus biparietalis: PHILIPPINES, JOLO ISLAND, Jolo Province, Municipality of Jolo: CAS 60703, 60704, 60706-08,
60711, 60714, 60718.
SILER ET AL.

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Zootaxa 3847 (3): 388412

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)

Taxonomic revision of the semi-aquatic skink Parvoscincus leucospilos

388 Accepted by L. Grismer: 28 May 2014; published: 8 Aug. 2014

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

390 Zootaxa 3847 (3) 2014 Magnolia Press

Material and methods

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

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REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

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REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

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Parvoscincus duwendorum sp. nov.

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

398 Zootaxa 3847 (3) 2014 Magnolia Press

Mt. Banahao/Angat Dam, Luzon

Polillo; Bicol, Luzon

Mt. Banahao, Luzon

Mt. Banahao, Luzon

Zambales & Bataan Mtns, Luzon

Sierra Madre, Luzon

Sierra Madre, Luzon

Northern Cordilleras, Luzon

Northern Cordilleras, Sierra Madre

Mt. Palali, Luzon

Sierra Madres, Luzon

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

47.355.9 (52.7 3.3)

Large, well defined

41.554.7 (49.0 4.6)

400 Zootaxa 3847 (3) 2014 Magnolia Press

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

Parvoscincus manananggalae sp. nov.

402 Zootaxa 3847 (3) 2014 Magnolia Press

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press

Parvoscincus tikbalangi sp. nov.

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REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

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REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

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Cordillera Mountain Range. Check List, 8, 469490.

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REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

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Philippines, Luzon Island, Camarines Sur Province

Philippines, Luzon Island, Aurora Province

Genbank Accession Numbers

APPENDIX 2. Additional material examined.

REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Zootaxa 3847 (3) 2014 Magnolia Press