parvoscincus 3 sp nov 2014

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

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN

1175-5334

(online edition)Copyright 2014 Magnolia Press

Zootaxa3847 (3): 388412www.mapress.com/zootaxa/

Article

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 skinkParvoscincus 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

1Department of Biology and Sam Noble Oklahoma Museum of Natural History; University of Oklahoma, 2401 Chautaugua Ave., Nor-

man, OK 73072-7029, USA. E-mails: [email protected], [email protected], [email protected] of Biology; University of Washington, Seattle, WA 98195-1800, USA. Email: [email protected] Institute and Department of Ecology and Evolutionary Biology; University of Kansas, 1345 Jayhawk Blvd, Lawrence, KS

66045-7593, USA. E-mails: [email protected], [email protected] Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos St., Manila, Philippines.

E-mail: [email protected]

Abstract

We review the recent discovery of multiple populations of the enigmatic, semi-aquatic Sphenomorphus Group skink,Par-

voscincus 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., andP. duwendorumsp. nov.). All four recognized species are genetically divergent in both mitochondrial and nu-

clear 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 Philip-

pines may vary regionally across the archipelago.

Key words: biodiversity, conservation, cryptic diversity, endemism, lizard, Luzon Island, riparian, Southeast Asia, Sphe-

nomorphusgroup

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 thatfew 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 philippinumGnther (McLeod et al., 2011; Brown et al., 2013b),Platymantis polillensis Taylor and

P. cornutusTaylor (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 skinkParvoscincus leucospilos

Peters, 1872 (Brown et al., 2000, 2013; Linkem et al., 2011; McLeod et al., 2011; Siler et al., 2011d) is exactly onesuch situation.


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Zootaxa3847 (3) 2014 Magnolia Press 389REVISION OF PHILIPPINE SEMI-AQUATIC SKINKS

Parvoscincus leucospiloswas first described by Peters (1872), who placed the species in the genusLygosoma

Gray, sectionHinulia(=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 Sphenomorphusbased 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 Sphenomorphusrevised 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,

PinoyscincusLinkem, Diesmos, and Brown,and Tytthoscincus Linkem, Diesmos, and Brown), placing this rare

Philippine endemic species in the genusParvoscincus(Linkem et al., 2011).

Details of the status of the type material forP. leucospilosare 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 ofP. leucospilos.

Twenty-one species are recognized in the genus Parvoscincus(P. abstrusus Linkem and Brown, P. agtorum

Linkem and Brown,P. arvindiesmosiLinkem and Brown,P. aurorusLinkem and Brown,P. banahaoensisLinkem

and Brown, P. beyeri Taylor, P. boyingi Brown, Linkem, Diesmos, Balete, Duya, and Ferner, P. decipiens

Boulenger,P. hadrosBrown, Linkem, Diesmos, Balete, Duya, and Ferner,P. igorotorumBrown, 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. palaliensisLinkem 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, andP. decipiens; Linkem et

al., 2011; Brown et al., 2010, 2013b). Unlike most species of Parvoscincus that prefer terrestrial microhabitats,

Parvoscincus leucospilosis 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.


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SILERET AL.390 Zootaxa3847 (3) 2014Magnolia Press

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 ofP. leucospilosfrom 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 andP. leucospilos.


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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 1000 th 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 withinP. 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.

FIGURE 2.Partitioned Bayesian majority rule consensus estimate of molecular phylogeny from the concatenated (ND2 +PTGER4) data ofParvoscincus. Populations within the P. leucospilosComplex are genetically unique and are diagnosed asnew 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. duwendorumsp. nov.,P. tikbalangi

sp. nov.,andP. 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. duwendorumsp. nov.

and P. manananggalae sp. nov.) are separated by 4.25.6% mitochondrial sequence divergence. Sequence

divergences among the other lineages within theP. leucospilos populations are greater than 7.1% (Table 1; Fig. 2).Morphology. Superficially the four lineages of Parvoscincus leucospilos appear morphologically similar,

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


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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 fourP. leucospilosgrouplineages. 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 trueP. 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) leucospilosPeters, 1872:684.Sphenomorphus leucospilosBrown and Alcala, 1980:172; Bauer et al., 1995; Linkem et al., 2011; McLeod et al., 2011.Parvoscincus leucospilosLinkem, Diesmos & Brown, 2011; Linkem & Brown 2013 (part)

Type designation. Based on ICZN articles 74.1 and 74.7 and in accordance with recommendation 74D wedesignate 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, well-

defined; (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


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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.

manananggalaeby having prefrontals in medial contact (vs. separated); fromP. duwendorum andP. tikbalangiby

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]); fromP. 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); fromP. manananggalae by the absence of subcaudal dark pigmentation (vs. presence); and

fromP. tikbalangiby 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


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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 amongspecies.


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FIGURE 4. Photographs of heads from dorsal and lateral perspectives to show pigmentation and scale pattern variation amongspecies. With the exception ofP. 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 reddish-

orange 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.


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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 ofP. 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, andP. 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 leucospilosLinkem, 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 duwendorumcan 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,andP. 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]); fromP. 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); fromP. manananggalaeby 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 fromP. tikbalangiby having prefrontals and first supraoculars in contact (vs. separated).


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12/25


TABLE

3.Summaryofmeristicandmensural,diagnosticcharactersamongthe

foursemi-aquaticPhilippineskinkspec

iesoftheParvoscincusleucospilosCom

plex..For

characterdefinitions,pleaserefertoMa

terialandMethods.

leucospilos

(6m

1

,5f)

duw

endorum

(1juv.)

mananangg

alae

(3m,2f)

tikbalangi

(7m,1f)

Range

South-central&Southeastern

Luzon

No

rthwesternLuzon

East-centralLuzon

NortheasternLuzon

SVL

42.654.4(50.03.7)

33.5

47.355.9(

52.73.3)

41.554.7(49.0

4.6)

TotL

100.6126.1(112.69.5)

83.5

122.9146.8(130.810.8)

118.9138.5(128.48.9)

TL/SVL

99157(12618)

149

120168(1

4220)

138159(150

9)

FLL

3.85.1(4.50.4)

3.3

4.35.5(4.80.5)

4.05.0(4.60

.4)

FLL/SVL

810(90)

10

810(91

)

910(90)

HLL

5.26.5(6.00.4)

4.0

5.86.7(6.20.4)

5.26.5(6.00

.5)

HLL/SVL

1113(120)

12

1112(121)

1213(120)

ToeIVlam

15(6)

16(1)

17(4)

12

(1)

17(5)

14(4)

15(2)

16(2)

PVSR

6167

60

6169

5863

LoVent

4147

41

4349

4144

MBSR

3034

26

3233

2832

SL

6(1)

7(10)

7(1)

7(5)

7(8)

IFL

6(1)

7(6)

8(3)

9(1)

8(1)

6(3)

7(2)

7(8)

Prefrontalcontact

+

or+

Prefrontal/1stsupraocularcontact

+

+

+

Headpigmentation

Heavilymottled

Mo

deratelymottled

Moderately

mottled

Moderatelymottled

Upperarmdarkpigmentation

+,patchy

+,patchy

+,patchy

Cloacalscaledarkpigmentation

or+

+

or+

Subcaudaldarkpigmentation

+

Dorsalwhitespots

Large,welldefined

Faint

Large,well

defined

Faint

Dorsalwhitebands

913

15

912

912

Lateralbodycoloration

+,brightreddish-orange

Dataunavailable

+,reduced,tan

1

Paralectotype(CAS64232)presumedmalefollowingvisualinspectionofexternalbody;authorsavoidedcreatingincisiononraretypespecimen.


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FIGURE 5. Photographs in life of (A)P. leucospilos(Mt. Banahao; TNHC 62683); (B)P. leucospilos(Angat Watershed; KU329388); (C)P. manananggalae sp. nov.(PNM 9794); and (D)P. tikbalangisp. 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 adductormusculature 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.


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FIGURE 6. Illustration of head of the examined Parvoscincus leucospilos paralectotype (CAS 64232) in dorsal and lateralviews. 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; T 1,

primary temporal; T2, secondary temporal; and T3, tertiary temporal. Roman numerals indicate scales in the supraocular series.Illustrations by STC and CDS.


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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 withParvoscincus igorotorum, P. jimmymcguirei, andP. 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 etal., 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


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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 distinguishingParvoscincusmanananggalae from all species of Parvoscincusare

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 fromP. 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]); fromP.

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 fromP. 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


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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 withParvoscincus agtorum, P. palaliensis, P. steerei, andP. 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 manananggalaehas 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 armpigmentation present, patchy; (11) subcaudal pigmentation absent; (12) dorsal white spots faint; (13) dorsal white


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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 resemblesP. duwendorum,P. leucospilos,and

P. manananggalae, but differs fromP. duwendorum by having seven infralabials (vs. eight); fromP. duwendorum

andP. 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); fromP. leucospilosandP. manananggalaeby 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]); fromP. leucospilosby 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


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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 orange-

cream. 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 skinkoccurs

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

genusParvoscincus, 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 BicolPeninsula 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. leucospilosComplex 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 ofP. manananggalae,P. duwendorum, andP. tikbalangibring 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 andBrown, 2013; Davis et al., in press). The results of all these studies suggest one thing for certain: we most likely


20/25


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 theRufford 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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