PRIMARY RESEARCH PAPER | Philippine Journal of Systematic Biology DOI 10.26757/pjsb2020b14012 urn:lsid:zoobank.org:pub:0938ACCF-60DF- 45D1-B207-0827356E97A7

A new species of Sun Skink (Reptilia: Scincidae: Eutropis) from the Zamboanga Peninsula, southwestern Mindanao Island, Philippines Anthony J. Barley1*, Marites B. Sanguila2 and Rafe M. Brown3

Abstract

We describe a new species of lizard in the genus Eutropis Fitzinger 1843 from the southwestern tip of the Zamboanga Peninsula on the western part of Mindanao Island, Philippines. The new species is related to Eutropis rugifera, which is a secretive, forest-adapted skink that ranges widely outside the Philippines from the western extent of its distribution on Nicobar Island (the type locality) through southern Peninsular Malaysia, Sumatra and the Mentawai islands, Borneo, Java, and as far east as Bali Island. The discovery of a new, morphologically distinct, and genetically highly divergent Sun Skink lineage in the low elevation forests of Zamboanga Peninsula creates a puzzling disjunct geographic distribution (E. rugifera has not been reported from the Sulu Archipelago). The new species is estimated to have diverged ~10–16 mya from E. rugifera, from which it appears to have an extralimital and isolated distribution. Considering the dynamic geological history and ancient continental origin of the Zamboanga Peninsula, colonization by the new species may have been facilitated by pre-Pleistocene overseas long-distance dispersal, saltatory range expansion and subsequent contraction/extinction in the Sulu Archipelago, and/or possibly paleotransport on the ancient crustal fragment of Zamboanga. The new species is known only from Zamboanga City’s primary surface water supply catchment at the lowest elevations inside the boundaries of Pasonanca Natural Park, despite the fact that there have been historical surveys of herpetological diversity at multiple sites to the northeast (Zamboanga, western Mindanao) and to the southwest (Sulu Archipelago). The new species, thus, may be limited to just the tip of the Zamboanga Peninsula, possibly rendering Pasonanca’s low elevation forests its most critical habitat resource for long term persistence and survival of the species.

Keywords: IUCN Red List, Palawan microcontinent block, Pasonanca Natural Park, Sulu Archipelago, Surface catchment watershed biodiversity

Introduction other groups of tropical vertebrates, preliminary genetic studies suggest that cryptic diversity is likely present within populations The genus Eutropis is a moderate size group of scincid of many species (Ota et al. 2001; Mausfeld and Schmitz 2003; lizards that are widespread and abundantly distributed across Barley et al. 2013), and in recent years, new species have been Asia (Mausfeld and Schmidt 2003; Grismer 2011; Uetz et al. described from across Asia (Mausfeld and Böhme 2002; Das et 2020). Also known as Sun Skinks, these lizards historically al. 2008; Batuwita 2016). Much of this overlooked diversity can have constituted a major taxonomic conundrum. As in many be ascribed to the historical taxonomic focus on identifying lineages using external morphological characters (which 1* School of Life Sciences, University of Hawai‘i, Honolulu, HI 96822, frequently vary little among populations), as well as the group’s USA; broad distribution across southern Asia, including many 2 Biodiversity Informatics and Research Center, Father Saturnino Urios countries and landmasses that are politically divided and University, San Francisco Street, 8600 Butuan City, Philippines logistically difficult for field biologists to access (Barley et al. 3 Biodiversity Institute and Department of Ecology and Evolutionary 2015a; Amarasinghe et al. 2017). Biology, University of Kansas, Lawrence, KS 66045, USA Although genetic studies are providing novel insights into sun skink diversity, this new source of information is also *Corresponding email: [email protected] challenging to interpret from a taxonomic perspective. Taxonomic resolution in many sun skink lineages is hampered Date Submitted: 6 July 2020 by data limitations. For example, many widespread species lack Date Accepted: 1 February 2021

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Barley et al.: A new species of Eutropis from the southern Philippines robust population-level sampling (but see Barley et al. 2015a), and the genetic information that is available is often limited to one, or a small number of loci, which is widely recognized as being insufficient for reliably identifying species boundaries (Irwin 2002; Funk and Omland 2003; Hinojosa et al. 2019). Additionally, many species occur in naturally fragmented habitats, such as archipelagos, where population differentiation is expected to be high, although these genetic differences will often not be representative of a speciation event. Thus, reliance on morphological information will often lead to taxonomic conclusions that underestimate species diversity, whereas reliance on genetic information may lead researchers to overestimate species-level diversity in these types of systems (e.g., Barley et al. 2013). Combining both sources of data with information about ecology and geographic distributions can help mitigate these sources of taxonomic bias in sun skinks, and potentially aid in the recognition of the total diversity of evolutionary lineages (species), whether or not they can be Figure 1. Map showing sampling localities within the geographic readily identified on the basis of external morphology (Barley et distribution of Eutropis alcalai sp. nov. from the Zamboanga Peninsula al. 2020). Of particular interest is the Philippine Archipelago, (green dots), and the much broader range of its closest relative, E. rugifera (type locality: Nicobar Island; red dots). Inset: known records widely recognized as a global biodiversity hotspot where for E. alcalai sp. nov. within Pasonanca Natural Park (southern point species diversity is thought to be substantially underestimated in 0.5 km above intake area, Barangay Pasonanca [type locality]; northern nearly all major terrestrial vertebrate groups (Brown and point Cabo Negro Outpost area, Barangay Tolosa). Diesmos 2009; Brown et al. 2013). Sun skink species diversity, first comprehensively reviewed in the Philippines 40 years ago current location 10–6 mya (Yumul et al. 2009a; Hall 1998, by W.C. Brown and A.C. Alcala (Brown and Alcala 1980; see 2002; Blackburn et al. 2010). Similar patterns have been also Taylor 1922) was nearly tripled in a single lineage of documented in the southwestern Philippines, in particular the Philippine Eutropis by a recent taxonomic revision (Barley et Sulu Archipelago and Mindanao Island, which have been al. 2020) and biogeographic studies suggest that the Philippines interpreted as receiving substantial proportions of their reptile was likely invaded from the Sunda Shelf by Eutropis 2–4 faunas by overland colonization from the Sunda Shelf (Taylor independent times (Barley et al. 2015a). 1928; Inger 1954; Leviton 1964; Brown and Alcala 1970). Biogeographic patterns in island systems are shaped by Finally, in addition to dispersal corridors via land bridges (Inger their geologic history, with older and more isolated islands 1954; Diamond and Gilpin 1983), the ‘species-pump’ action of typically having a higher proportion of endemic species (Evans oscillating sea levels may have contributed to regional et al. 2003; Siler et al. 2012; Brown et al. 2016). In Southeast population differentiation within the oceanic portions of the Asia, historical sea-level fluctuations have played a major role Philippines (Brown and Diesmos 2009), although this determining terrestrial diversity patterns, where dry land mechanism may only explain a portion of in situ species connections that appeared among islands when sea levels diversification in the endemic terrestrial vertebrate clades dropped to 120 m below present during the last glacial (Esselstyn and Brown 2009; Siler et al. 2010; Setiadi et al. 2011; maximum (Miller et al. 2005) connected populations and Blackburn et al. 2013; Oaks et al. 2013, 2019; review: Brown et facilitated dispersal. For example, this led some species on al. 2013). Palawan Island of the southwestern Philippines to be more The “five-keeled” or “rough-scaled” Sun Skink [Eutropis closely allied to those on Sundaland than the remainder of the rugifera (Stoliczka 1870)] occurs widely across Southeast Asia, Philippines which remained isolated (Inger 1954; Heaney its distribution encompassing Peninsular Malaysia south of the 1985). Other groups appear to have arrived on Palawan via Isthmus of Kra, virtually all large islands on the Sunda Shelf, dispersal from the north (Esselstyn et al. 2010) or possibly as a and the Nicobar Islands (the species’ type locality). In 2015, a result of paleotransport on the Palawan Microcontinent Block related population was first reported from the extreme (Yumul et al. 2009a; Siler et al. 2012; Brown et al. 2016) which southwestern Philippines (Barley et al. 2015a). Similar to other rifted from continental Asia 40–35 mya and arrived at its members of the genus, E. rugifera is a generalist insectivore

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Barley et al.: A new species of Eutropis from the southern Philippines found in interior, lowland, primary and mature secondary locality. However, this dataset was not combinable with the tropical forests (Karin et al. 2017; Amarasinghe et al. 2017). former E. rugifera dataset because it lacked overlapping Nicobar Island and newly discovered Philippine populations taxonomic sampling. are, thus, the biogeographically disjunct extreme western and For each dataset we performed a concatenated Bayesian eastern limits of the group’s documented range (Fig. 1). phylogenetic analysis of the data using MrBayes 3.2.6 (Ronquist In a range-wide treatment of the Eutropis rugifera group, et al. 2012), selecting models of molecular evolution (from Karin et al. (2017) interpreted ~6 mya (divergence time among the 20 commonly implemented in MrBayes) and the estimated through a relaxed-clock phylogenetic analysis optimal partitioning strategy using Bayesian information calibrated with a typical reptile mtDNA substitution rate), criterion in PartitionFinder v2.1.1 (Lanfear et al. 2014). We ran divergences among populations of E. rugifera from Sundaland MrBayes analyses (consisting of two runs with eight chains landmasses (Sumatra, Malay Peninsula, Mentawai islands and each) for 50 million generations, sampling every 5,000 Borneo) as evidence that environmental barriers likely persisted generations, and assessed convergence using the average through the Pleistocene despite the existence of land-bridges. standard deviation of split frequencies and potential scale These xeric environmental barriers may have maintained and reduction factor diagnostics in MrBayes, and by ensuring that all reinforced population structure (by inhibiting dispersal and gene parameters had effective sample sizes >1,000 using Tracer v1.7 flow) in forest obligate species, which formerly were thought to (Rambaut et al. 2018). Topological convergence was assessed have been unified (via corridors for dispersal) upon exposure of using the R package RWTY (Warren et al. 2017). We rooted the the Sunda Shelf during Pleistocene sea-level reductions (Inger phylogenetic tree from the genus-wide analysis according to 1954; Voris 2000; Bird et al. 2005; Cannon et al. 2009). Barley et al. (2015a), whereas the two analyses focused on the Previous work also clearly demonstrated that the significant E. rugifera complex were rooted with E. multifasciata (Kuhl evolutionary divergence between the Zamboanga population 1820) (not shown). and the rest of the species complex likely has resulted in a speciation event (Barley et al. 2015a; Karin et al. 2017). In this Morphological Data paper, we formally recognize this lineage as a highly distinctive We compared specimens from our Philippine collections to new Sun Skink from the low elevation rainforests of Pasonanca Eutropis rugifera specimens from landmasses of the Sunda Natural Park on the outskirts of Zamboanga City. Shelf (Sumatra, Borneo), and we relied heavily on the recent study of Amarasinghe et al. (2017) who reported on specimens Materials and Methods from the type locality (Camorta, Nicobar Island, northwest of Sumatra; Fig. 1), redescribed the holotype of E. rugifera in Genetic Data and Phylogeny Estimation detail, reviewed the taxonomic history of the species and its We assembled several datasets using genetic data available synonyms (Euprepes percarinatus Peters 1871 & E. p. var. from GenBank to evaluate the evolutionary relationships of borneensis Peters 1871 [Java], Mabuia rubricollis Bartlet 1895 populations assigned to E. rugifera with respect to other [Borneo], Mabuia quinquecarinata De Rooij 1915 [Sumatra]), Southeast Asian Eutropis. To summarize what is currently and compared morphological variation among specimens from known about the broadscale phylogenetic relationships of the disjunct populations located on various Sundaic landmasses Eutropis, we first combined one mitochondrial and nine nuclear throughout the range of this widely distributed, yet poorly genes (see Barley et al. 2015a) with the available 12s and 16s phenotypically characterized species (Chan-ard et al. 1999; data. For this analysis, we selected a single individual per Malkmus et al. 2002; Das 2004; Chan et al. 2010; Grismer 2011; species, though we also created several chimeric sequences in Venugopal 2010; Karin et al. 2017). We assembled our dataset order to maximize phylogenetic resolution if we were confident from newly collected specimens, historical museum specimens that the sequences were drawn from the same lineages (based (see Specimens Examined section) and the literature mentioned on preliminary analyses of the individual gene trees and current above. Fluid-preserved specimens we examined are housed at taxonomic knowledge; Table 1). To evaluate the genetic the University of Kansas Biodiversity Institute (KU), The Field relationships among E. rugifera populations, we combined data Museum (FMNH), the Smithsonian Institution (USNM), the for the ND2 mitochondrial gene and the MC1R, BRCA1, and Museum of Vertebrate Zoology (MVZ) at the University of RAG1 nuclear genes from Barley et al. (2015a) and Karin et al. California, Berkeley, the Museum Zoologicum Bogoriense (2017). We reconstructed an additional E. rugifera dataset for (MZB) and the National Museum of the Philippines (PNM); the ribosomal RNA 16S mitochondrial gene (Amarasinghe et al. Institutional museum codes follow Sabaj (2019). 2017) because a single individual was sampled from the type

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We used Mitutoyo digital calipers (RMB measured the taxon E. greeri based on examination of photos of the specimens, in mm, ± 0.1 mm) and a Leica Wild MZ12 specimen (Kanishka Ukuwela, personal communication). microscope with installed micrometer, to measure mensural Additionally, our data suggest the ‘E. indeprensa’ of Karin et al. characters (from the left side of the body) and take meristic data (2016) is likely to be a specimen of E. cumingi (Brown & Alcala (scale counts), following definitions of Amarasinghe et al. 1980), as the individual they sequenced is from Lubang (the (2017) with a few modifications (see below) to clarify species to which that population is assigned; Barley et al. 2020). ambiguities; these modifications with respect to head length, We follow the unified, general concept of species as separately paravertebrals, and ventrals allow for direct comparisons with evolving metapopulations when interpreting the taxonomic Amarasinghe et al. (2017), and also comparisons to other, implications of the results (De Queiroz 1998, 2007). earlier works (Brown and Alcala 1980; Greer and Nussbaum 2000; Das 2004; Grismer 2011). Measurements included snout– Genetic Data and Phylogeny Estimation vent length (SVL), and the lengths of the forearm, femur, tibia, The twelve-gene species-level phylogenetic analysis of foot, head, snout, and 4th toe. Because of our difficulty reliably Eutropis (Fig. 2a) suggests that the E. rugifera lineage is identifying the posterior edge of the mandible (Amarasinghe et evolutionarily divergent from other Southeast Asian Eutropis al. [2017] used “posterior edge of the mandible to the tip of the and indicates weak support for a sister relationship with E. snout” as head length), we also measured and report what we tytleri (Theobald 1868), a poorly known endemic from the feel is a more repeatable head length definition: from the Andaman Islands (Chandramouli and Amarasinghe 2020). The anterior edge of the auricular opening to the tip of the snout. We four-gene phylogeographic analysis shows that the Philippine also measured head width, horizontal eye (orbit) diameter, populations are genetically divergent from Malaysian and auricular opening diameter, axilla-groin distance, eye–auricular Indonesian populations of E. rugifera at both mitochondrial and opening distance, and eye–nostril distance. We counted nuclear loci (Fig. 2b). Phylogenetic analyses of the 16s data supralabial and infralabial scales from the upper and lower jaw show that the Malaysian and Indonesian E. rugifera populations symphysis to rostral and mental scales, respectively. To are minimally divergent from the type locality populations on facilitate comparisons with other published work, we reported Nicobar Island (Fig. 2c). Although these data also suggest a ventrals and dorsals using the various methods used by different close relationship between the Nicobar and the Sumatran E. investigators (Brown and Alcala 1980; Das 2004; Grismer rugifera populations (a result also seen in other co-distributed 2011; Amarasinghe et al. 2017); thus, we counted and reported species), the remaining phylogeographic relationships within ventrals from the postmental to the precloacals and also this lineage are poorly resolved. We do not use degrees of between the midpoints of limb insertions. We counted genetic distance among populations to diagnose the new species paravertebral scales between nuchals (included) to the region described below. However, we emphasize that the genetic posterior to the thigh immediately dorsal to the opening of the divergence (~17–18% uncorrected p-distance for ND2 mtDNA) cloaca (counted in a straight line immediately left of the and ancient date estimated for the divergence of the Zamboanga middorsal line) and also along the spine (the middorsal-most lineage from its common ancestor with E. rugifera (10–16 mya) scale row) between the dorsal midpoints of limb insertion. We provide overwhelming support for the two lineages being counted subdigital lamellae underneath the 4th toe from the first recognized as distinct species (Barley et al. 2015; Karin et al. proximally differentiated (enlarged) scale to the distal-most 2017). lamella before the claw. Sex was determined by everted hemipenes or ventral tail dissection (males) or presence of Morphological Data oviducts and embryos (females). For standardization of color Our examination of our small series of specimens from descriptions of the new species, we incorporated the numbered Pasonanca Natural Park (on the outskirts of Zamboanga City, at color scheme from Köhler’s (2012) standardized color guide. the SW tip of Zamboanga Peninsula, Mindanao Island) identified unambiguous, discretely variable (i.e., non- Results overlapping ranges) character values, in multiple meristic and color pattern character states between the Zamboanga lineage We update two taxonomic identification errors to avoid and all other populations of Eutropis rugifera from throughout continued confusion from previous phylogenetic studies. the Sundaic landmasses (Borneo, Bali, Java, Sumatra, Mentawai Batuwita (2016) suggested that the ‘Eutropis madaraszi’ of Islands, Bawean Island, the Malay Peninsula, and Nicobar Mausfeld et al. (2003) actually corresponds to E. austini Island; Table 2). We use only fixed character state differences to Batuwita 2016, however, we tentatively assign the sequence to diagnose the new species, but several other narrowly

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– – – –

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MC1R

KJ574603 KJ574594 KJ574548 KJ574633 KJ574549 KJ574551 KJ574608 KJ574555 KJ574559 KJ574565 KJ574561 KJ574601 KJ574575 KJ574606 KJ574581 KJ574583 KJ574629 KJ574590 KJ574587 KJ574580 KJ574612 KJ574613 KJ574571 KJ574615 KJ574617 KJ574620 KJ574567 KJ574578

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KF235210 KF235153 KF235219 KF235169 KF235162 KF235208 KF235183 KF235215 KF235190 KF235191 KF235189 KF235175 KF235171 KF235187

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FOXP2

KJ574474 KJ574457 KJ574495 KJ574451 KJ574453 KJ574454 KJ574455 KJ574464 KJ574488 KJ574470 KJ574468 KJ574476 KJ574493 KJ574479 KJ574481 KJ574484

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KJ574772 KJ574756 KJ574790 KJ574759 KJ574763 KJ574785 KJ574768 KJ574787 KJ574774 KJ574777 KJ574779 KJ574782

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KJ574988 KJ575003 KJ574979 KJ574999 KJ574984 KJ574990 KJ574994 KJ574997

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Summary of dataof used Genbank inSummary genus

borealis Species alcalai allapallensis ashwamedhi greeri beddomei bibronii bontocensis caraga carinata clivicola cumingi cuprea dissimilis gubataas indeprensa islamaliit lapulapu longicaudata macropthalma 1 macularia cf. 2 macularia cf. macularia multicarinata multifasciata nagarjunensis palauensis quadricarinata rudis rugifera sahulinghang- ganan sibalom trivittata tytleri Table 1. Table

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Barley et al.: A new species of Eutropis from the southern Philippines

Table 2. Distribution of diagnostic character states in selected charac- ters among Eutropis alcalai, new species, and E. rugifera, its unam- biguous most closely-related species. See Barley et al. (2020) for diagnoses distinguishing the new species from all other Philippine species of Sun Skinks in the genus Eutropis. “PM” = postmental scale; “infra2” = second infralabial scale; see text for further explanation of characters. E. alcalai, new Character E. rugifera species PM–infra2 contact – + 4th toe lamellae 29–31 21–26 Differentiated 2 medially enlarged 6 equivalent precloacals scales

Infralabials 7 6 3rd & 4th finger lengths equivalent 4th longer than 3rd

Dorsal color pattern Very dark grayish- Chestnut–brown, brown, faint, bright yellow incomplete stripes stripes Figure 2. (a) Consensus tree from Bayesian phylogenetic analysis of Chin & infralabials White Yellow-orange twelve genes, summarizing current knowledge of phylogenetic relationships among Eutropis species and placement of E. rugifera/E. Fernandez, C.D. Siler, L.J. Welton and J.W. Phenix, in low alcalai sp. nov. lineage. (b) Consensus tree from phylogenetic analysis elevation climax rainforest (semi-open canopy, thin layer of dry of four genes (ND2, BRCA1, RAG1, MC1R) illustrating divergence leaf litter, 160 m above sea level) 10 m from the bank of the among Philippine populations and E. rugifera populations from the Tumaga River, 0.5–0.75 km upstream from Intake Area, Sunda Shelf. (c) 16s gene tree illustrating relationships between E. rugifera individual sampled from the type locality (Nicobar Island) and Pasonanca Natural Park, Barangay Pasonanca, Zamboanga City, three populations from Indonesia. Circles represents nodes where SW tip of Zamboanga Peninsula, western Mindanao Island, posterior probability ≥0.99. Scale bar units are in number of Philippines (6.9922333°N, 122.0604333°E). substitutions per site. Paratypes.—KU 321834 (RMB 11651), adult male, collected 10 overlapping tendencies in character state distributions are August, 2009, RMB and C. Infante, in mid-elevation secondary, emphasized in the holotype description discussed below. The regenerating rainforest (semi-open canopy, thin layer of dry leaf new species is also clearly differentiated, both morphologically litter, 550 m above sea level) Cabo Negro Outpost, Sitio Santa and biogeographically, from the populations from which the Clara, Pasonanca Natural Park, Barangay Tolosa, Zamboanga currently recognized E. rugifera junior synonyms are derived City (7.0941°N, 122.0876°E) ; KU 321833 (RMB 11580), (which closely match the diagnostic characters of E. rugifera; juvenile of undetermined sex, 12 August, 2009, RMB and C. Table 2; see Amarasinghe et al. 2017). Infante.

Taxonomic Account Diagnosis and comparisons (see also Table 2).—Eutropis Eutropis alcalai sp. nov. alcalai sp. nov. can be distinguished from its closest relative, E. Eutropis rugifera Barley et al. 2015:296. rugifera, by its smoother head scales, with light embossing only, and keels limited to posterior margins of parietals, temporals LSID: registration: urn:lsid:zoobank.org:pub:0938ACCF-60DF- and nuchals (vs. rugose to strongly keeled in E. rugifera), by 45D1-B207-0827356E97A7 absence (vs. presence) of contact between the postmental and Nomenclatural act: urn:lsid:zoobank.org:act:31AE197D-516D- second infralabial, by the presence of 29–32 (vs. 21–26) 49C5-9387-1FD8FB4C8197 subdigital lamellae under the 4th toe, by the presence of seven

infralabials (six in E. rugifera), the presence of differentiation in Holotype.—PNM 9878 (formerly KU 315013; Field No. RMB the precloacal scales series (two, medial precloacals enlarged, 9241), adult male, collected 15 July, 2008, by RMB, J. B. transversely expanded and bordered on either side by two

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Barley et al.: A new species of Eutropis from the southern Philippines

undifferentiated scales) in E. alcalai sp. nov. (vs. six equal sized scales in E. rugifera), by equivalent length of 3rd and 4th fingers (vs. 4th finger longer than 3rd), and by its white chin and infralabial scales (vs. yellow to orange in E. rugifera).

Description of holotype.—The holotype is an adult male, with hemipenes partially everted, in excellent condition, with a midventral incision (portions of the liver removed and preserved separately in 100% ethanol for genetic material) with an apparent caudal autotomy scar 10 mm posterior to the vent (Fig 3b), followed by a largely regenerated tail (Figs 3a, 4b); snout– vent length 70.1 mm, tail length (regenerated) 78.6 mm. Body robust, relatively wide and slightly compressed, limbs moderately developed, overlapping when adpressed, Figure 3. Eutropis alcalai sp. nov. (holotype PNM 9878; formerly KU hindlimbs conspicuously longer than forelimbs; head small and 315013; adult male, SVL 70.1 mm), photographed before preservation short (its length 42.5% of axilla–groin distance when measured in dorsal (a) and ventral (b) views. snout–mandible, 54.7% when measured snout–auricular opening); head narrow (its width 85.5% of length when measured snout–mandible; 66.3% of length when measured snout–auricular opening); head width 16.8% SVL; head barely distinct from neck, triangular in dorsal aspect, compressed in lateral view with snout rounded; head length 19.6% (snout– mandible) and 25.3% (snout–auricular opening) of SVL, 42.6%, 54.8% of axilla–groin distance, respectively. Head scales smooth and, except for posterior-most supraoculars which are lightly striated but unkeeled; rostral wider than high, contacting frontonasal; frontonasal as wide as high; prefrontals large, not in medial contact but broadly contacting first supraocular and anterior point of frontal; frontal diamond-shaped, with elongate posterior point extension, not contacting first supraocular but in broad contact with second supraocular and frontoparietals; supraoculars four, second largest; frontoparietals in broad medial contact, and also contacting 2nd, 3rd, and 4th supraoculars anteriorly and parietals and minute interparietal, posteriorly; interparietal without a parietal eyespot; parietals in medial contact behind interparietal, for distance equivalent to length of interparietal itself; parietals large, strongly keeled on posterior margins, separated anteriorly by interparietal, contacting 4th supraocular; nuchals in single pair, strongly keeled. Lateral head scales lightly embossed; nostril pierces small, trapezoidal nasal at its center (nostril completely enclosed); nasals widely separated, contacting rostral anteriorly, supranasals dorsally, anterior loreal posteriorly, first Figure 4. Eutropis alcalai sp. nov. (holotype PNM 9878; formerly KU supralabial ventrally; nasal makes point contact with second 315013; adult male, SVL 70.1 mm), photographed in life: (a) close-up supralabial on right, but not left; postnasals tall, curving forward of anterior body region; (b) full body; collected from Pasonanca slightly above nasal, frontonasal and prefrontal; loreal single, Natural Park, Barangay Pasonanca, on the outskirts of Zamboanga rd th City, Zamboanga Peninsula, western Mindanao Island. Photos: RMB. large, broader than high, contacting 3 and 4 supralabials ventrally, and prefrontals dorsally, but not contacting 1st supraocular; second loreal in contact posteriorly with first

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Barley et al.: A new species of Eutropis from the southern Philippines supraciliary, minute scales in preocular position (enlarged equivalent to above vent); 22 rows between midpoints of dorsal preoculars absent), and first of two enlarged suboculars, limb insertions; 43 ventral scales from postmental to vent (23 separated from remainder of subocular series (3 unkeeled scales between midpoints of limb insertions); two greatly enlarged, posterior to orbit) by greatly enlarged 6th supralabial; eye large, laterally expanded precloacals, overlapped on either side by two orbit diameter 26.1% head length when measured snout-to- undifferentiated scales in lateral precloacal position. mandible (20.2% snout-to-auricular opening); supraciliaries six, Scales of dorsal distal portions of limbs same size as first slightly larger than others; enlarged pretemporals two, ventral limb scales; keels continue outer surface of radial keeled, both in contact with parietals; supralabials 7, 6th largest, segment of forelimbs (inner forearm scales smooth), before subocular, second and third longer than others; one transitioning to smooth scales on dorsal surface of hands and postsupralabial, keeled; one primary temporal, keeled; fingers; keels continuously distributed through tibial segments secondary temporals two, lower much larger than upper, and of hindlimbs, all the way to dorsal surfaces of foot; relative contacting parietal; tertiary temporals three, keeled, length of fingers: IV = III > II > V > I; dorsal surfaces of fingers approximately equal size, uppermost contacting parietals and with one row of smooth scales; proximal ends (one or two nuchals. Lower eyelid translucent, scaled, lacking transparent phalangeal elements) with additional row; subdigital lamellae unscaled window; lower eyelid scales tall, columnar; smooth, slightly transversely expanded, widest at base of each infralabials seven, first two are equivalent in size, then digit; each finger terminating in moderately recurved claw; progressively increasing their width until sixth which is palmar surface of hands with non-imbricate, smooth, convexly approximately twice width of the first, followed by a smaller 7th rounded, juxtaposed; single enlarged, raised scale present on infralabial scale, same size as 1st and second infralabials; mental posterolateral edge of palm; relative length of toes: IV > III > V twice as wide as deep; single large postmental in contact with > II > I. scales on dorsal surfaces of toes variably scaled, with first infralabial; first pair of enlarged chin shields in contact, two rows of keeled scaled on proximal two phalanges of 2nd following postmental and contacting 1st and 2nd infralabials; through 5th toe, and proximal first toe, and all distal dorsal toe second row of chin shields separated by a single median scale, surfaces covered by single rows of smooth scale; subdigital contacting 2nd and 3rd infralabials dorsally; subsequent chin lamellae of toes moderately transversely expanded, smooth, shields and gular scales equivalent in size, not transitioning to numbering 29 under 4th toe; subdigital lamellae conspicuously larger ventral body scales until sternum; auricular opening more expanded and widest (~1.5–2 times wider) at phalangeal small, punctiform, round, its diameter 44.4% eye diameter, articulations; toes terminating in large, recurved claws; plantar tympanum deeply recessed; two small lobules present on surface of foot with non-imbricate, convexly rounded, ventral edge of auricular opening. juxtaposed scales; four differentiated, large subcylindrical scales Dorsal and lateral body scales, and those of proximal half on the heel. of limbs (humeral segment of forelimbs and femoral portions of Tail cylindrical at base, not original, with autotomy scar hindlimbs) strongly keeled, imbricate, without apical pits. Keels and subsequent regenerated regrowth to approximate length prominent from posterior half of parietal scales, nuchals, and equivalent of SVL, approximately 2/3 anticipated original tail remaining body; parietals and nuchals each with 8–10 keels; length; regenerated portion of tail much more slender than nape and body scales with 4–6 prominent keels (mucros) per inferred original tail width (Fig. 3a), median subcaudal row scale, for full length of body; most middorsal scales each with transversely expanded, wider than subcaudals anterior to four strong keels, flanked on either side with weak keels; keels autotomy scar (Fig. 3b). arranged in longitudinal rows, end to end, from posterior margin of each scale to anterior margin of next; lateral keeled scales Measurements of holotype (in mm).—SVL 70.1; forearm length transition slightly to less-keeled scales at ventrolateral body 19.1; femur length 13.4; tibia length 13.1; foot length 16.4; head regions, and eventually to smooth midventrals; first two rows of length 13.8; snout length 7.4; length of 4th toe 11.6; head width ventrals (at ventrolateral region) slightly keeled; posterior 11.8; eye diameter 3.1; auricular opening 1.9; axilla–groin margin of dorsal and lateral scales more strongly keeled than distance 32.5; eye–tympanum distance 5.6; eye–nostril distance anterior margins of each scale, producing keeled projections 4.3. (mucros), posteriorly from each scale, giving posterior edge of each scale a crenulate margin, due to projecting keel points Coloration in life.— Dorsal surfaces of head, neck and anterior (mucros); ventrals smooth (with striae), same size as dorsals trunk dark fawn (Köhler’s [2012] color 258) with cinnamon and lateral body scales; transverse scale rows around midbody highlights (255, 260) on neck and glaucous (289) to grayish 28; paravertebrals in 33 rows from parietals to base of tail (point horn (268) on snout; posterior trunk surfaces darker, glaucous

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Barley et al.: A new species of Eutropis from the southern Philippines dark brown (291) to burnt umber (48); dorsum with three interrupted series of longitudinal, body-length stripes; similar Coloration in preservative.—As with most species of Eutropis, but distinct dorsolateral stripes on each side; on anterior our specimens of Eutropis alcalai sp. nov. have demonstrably portions of body, longitudinal stripes are orange-rufus (56) to faded, and lost brighter coloration and contrasting colors in dark salmon (59), and these fade to less vibrant sayal (41) to preservative, even when kept (in this instance, for more than a mikado brown (42) by posterior trunk regions, where they are decade) in complete darkness at constant temperatures and in barely ascertainable from background dorsal pelvic region ethanol concentration. In preservative, the holotype is brownish- coloration; in anterior portions of trunk, distinctiveness of the olive (276) with dark neutral gray (299) dorsal and dorsolateral five dorsal, longitudinal stripes is heightened by increased longitudinal body stripes, offset by jet black single scales and contrasting jet black (300 ocelli patterning, with each black spots, each tightly juxtaposed with beige (254) markings; dorsal marking tightly coupled to with bright chamois (84) to straw surfaces of limbs are not noticeably different from dorsal body yellow (53) markings (Figs. 3a, 4b), some of which are confined surfaces; ventral surfaces cream white (52) with pearl gray (262) to single troughs between individual scale keels. laterally, and light lilac (221) scale margins on ventrolateral Sutures surrounding the outer perimeter of each scale on transitions with flanks; ventral surfaces of limbs, palmar surface dorsal and lateral head surfaces demarcated with jet black of hand, and plantar surface of foot cream white (52) with pale- pigment (Fig4a); supraciliaries and circumorbital minute scales to light (296, 297) gray subdigital lamellae of digits. bright straw yellow (53); lateral nuchal region, axilla-groin region (flanks), and lateral caudal surfaces (anterior to Variation.—A summary of variation in the type series is autotomy; Fig. 4b) olive- (265) to smoke-gray (256); presented in Table 3. KU 321834 has a substantial portion of its regenerated dorsal and lateral tail surfaces jet black (300); original tail, indicating that when it has not been autotomized aggregation of black-and-chamois markings present above and regenerated, the tail typically tapers gradually. In forelimb insertion (Fig 4a) but not hindlimb; a sharp light preservative, dorsal surfaces of one paratype differs from the grayish-brown above–white below ventrolateral transition holotype by being raw umber (280; paratype KU 321834) present throughout head, neck, and trunk surfaces (Fig. 3b, 4a); whereas ventral surfaces of both paratypes is pale mauve (204) forelimbs and hind-limbs generally darker ground color, amber with individual scale margins marked by dark blue gray (194); (51) on forelimbs, dark grayish brown (284) on hindlimbs, with both paratypes (KU 321833 and 321834) both have ventral distinct jet black (300) margins among individual dorsal limb sternal regions cream white (52), and KU 321834’s original scales (Fig4a); dorsal surfaces of hand and foot similar to limbs; subcaudal coloration is pale buff (1). dorsal surfaces of digits distinctly lighter, with brighter pearl gray (290) highlights transversely alternating with sepia black Distribution.—This species is only known from Pasonanca (286) coloration distally and terminating with Payne’s gray Natural Park near Zamboanga City at the southwestern tip of the (293) claws. Zamboanga Peninsula. A closely related species (E. rugifera) Ventral body color bright smokey white (261) to pale buff occurs on Borneo (among other islands of the Sunda Region), (1) under head and throat, darkening and becoming more and although neither species has been collected from the Sulu yellowish (light buff 2) by posterior ventrum and distinctly Archipelago, one of them may occur on its larger islands. yellowish (pale horn 11) posterior to yellowish (light ocher 13) cloacal opening (overlying precloacal scales); margins between Habitat and natural history.—No detailed, field based natural ventral scale rows very light pale bluish gray (287) giving history studies of the new species have been performed but appearance of seven or eight faintly, smokey strips running given its conservation significance and its unique biogeographic longitudinally, the length of the body (Fig. 3b), especially status as an endemic species of extreme southwestern Mindanao, evident on chin and throat; tail posterior to autotomy scar pale ecological studies of habitat use and requirements, behavior, bluish gray (287) to pale neutral (296), with increasingly dense activity, and diet would all be very informative. Although little aggregation of black pigment condensing by tail’s black is known about the natural history of E. alcalai sp. nov., it subcaudal tip coloration; palmar surfaces of palm, plantar clearly shares some aspects of its biology with E. rugifera in surfaces of feet, and proximal subdigital portions of digits light being “uncommon” and “forest adapted” (Karin et al. 2017; neutral gray (297), fading progressively to jet black (300) by Amarasinghe et al. 2017), which constitute a challenge for terminal phalanges and terminal claws of digits (Figs. 3b, 4a). future ecological studies. Other inferences that are likely safe to Iris surrounding circular pupil flesh ocher orange (57) to light extrapolate from E. rugifera include our assumptions that E. pratt’s rufus (71). alcalai sp. nov. is diurnal, a generalist insectivore, most likely

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Barley et al.: A new species of Eutropis from the southern Philippines

Table 3. Variation in the type series of Eutropis alcalai, new species. See text for further explanation of multiple ways head length, paravertebrals, and ventrals were recorded.

Specimen: PNM 9878 KU 321834 KU 321839

(holotype) (paratype) (paratype)

Sex male male juvenile

SVL 70.1 64.4 46.5

Tail length 78.6 regenerated 27.8 autotomized 36.0 autotomized

Forearm length 19.1 17.6 11.8

Femur length 13.4 12 7.9

Tibia length 13.1 11.45 7.8

Foot length 16.1 14.7 11.8

Head length (mandible) 13.8 12.9 9.5

Head length (auricular) 17.8 14.7 11.5

Snout length 7.4 6.8 5.8

4th toe length 11.6 10.7 6.5

Head width 11.8 11.3 8.2

Eye diameter 3.6 3.1 2.2

Auricular opening 1.6 1.4 1

Axilla–groin distance 32.5 30 22.2

Eye–auricular opening 5.6 4.7 4.1

Eye-nostril 4.3 4.5 4.2

Ventrals: total 43 45 43

Ventrals: limb insertions 23 24 23

Dorsals: total 22 23 23

Dorsals: limb insertions 33 34 34

Midbody scales 29 29 30

4th tow lamellae 29 32 29

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Barley et al.: A new species of Eutropis from the southern Philippines reliant on cool forest floor arthropods found in the leaf litter and Etymology.—We take great pleasure in naming this distinctive herbaceous layer of vegetation, in lowland primary forest and/or new species for our colleague Angel C. Alcala, in recognition of mature secondary forests (Das 2004; Grismer 2011; his numerous foundational contributions to the natural history, Amarasinghe et al. 2017). In the protected lowland original systematics, ecology, and conservation of Philippine lizards of forests of Pasonanca, RMB and MBS observed the new species the family Scincidae (Alcala and Brown 1966, 1967; Alcala moving through sun spots on the otherwise relatively dark, 1970; Brown and Alcala 1956, 1961, 1963a,b, 1980, 1986). humid forest floor (07:00–10:00 am; Fig. 5). In addition to the Suggested common name: “Alcala’s Quinque-carinate (Five- holotype and paratype localities, several specimens were keeled) Sun Skink,” or “Alcala’s Rough-scaled Sun Skink.” observed on the forest floor near the river at 6.9922333° N, 122.0604333°E. In several instances, individuals were Discussion observed as they darted rapidly under leaf litter when disturbed by approaching people, and another took refuge under bark and The Zamboanga Peninsula of Western Mindanao in the other woody debris when pursued. When captured, the new Philippines has an enigmatic geologic history that has been species exhibited anti-predatory behaviors/adaptations: it bit subject to debate (Lambeck and Chappell 2001; Hall 2002; while twisting its body in an “alligator roll” maneuver, Yumul et al. 2003; Padrones et al. 2017). It appears to have had defecated, and easily lost scales when grasped (RMB and MBS, an ancient geologic origin much further west, closer to the personal observation). Sunda Shelf, before this exposed, originally continental crust fragment moved eastward and accreted to the remaining Mindanao Island landmass (Hall 2009; Padrones et al. 2017). Given the estimated age of the divergence of E. alcalai sp. nov., it is conceivable that these geological processes potentially contributed to, or facilitated, colonization of the southern Philippines from the Sunda Shelf, as has been hypothesized in other species (Clouse et al. 2011; Brown et. al. 2013). Genetic data suggest that E. alcalai sp. nov. diverged from populations of E. rugifera on the Sunda Shelf more than 10 mya (Karin et al. 2017), providing extensive time for these lineages to embark on independent evolutionary trajectories (Gavrilets 2000; Coyne and Orr 2004). Divergence among the remaining populations of E. rugifera in Sundaland are much shallower and consistent with them being intraspecific, albeit highly structured (Karin et al. 2017). The close relationship between the type locality E. rugifera populations on Nicobar Island and the Sumatran populations illustrate a biogeographic relationship that has been identified in other species as well (Mani 1974; Das 1999; Teynié et al. 2010; Ganeshaiah et al. 2019). Our phylogenetic results also highlight a novel sister relationship between E. rugifera/E. alcalai sp. nov. and E. tytleri, a result that is biogeographically compelling given that E. tytleri is endemic to the Andaman Islands which are thought to have shared a similar geological history to the Nicobar Islands (Ripley and Beehler 1989).

Eutropis Taxonomic Diversity Eutropis taxonomy remains in a state of flux. Genetic data are lacking for several taxa and, thus, assessing the taxonomic Figure 5. Characteristics of vegetation in low elevation primary forest status of rare or poorly known, seldomly collected Eutropis of Pasonanca Natural Park, at the type locality of Eutropis alcalai sp. species remains a challenge. Genetic material is not available in nov. Photo: RMB. publicly accessible natural history collections for E. andamanensis (M.A. Smith 1935), E. chapaensis (Bourret

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Barley et al.: A new species of Eutropis from the southern Philippines

1937), E. darevskii (Bobrov 1992), E. floweri (Taylor 1950), E. could be explained by insufficient sampling or non- gansi (Das 1991), E. innotata (Blanford 1870), E. madaraszi comprehensive field surveys, which may be needed to (Méhelÿ 1897), E. quadratilobus (Bauer & R. Günther 1992), adequately characterize terrestrial vertebrate faunal assemblages E. tammana Das, da Silva & Austin 2008 and E. englei (Taylor (Brown et al. 1996, 2000, 2012; Devan-song and Brown 2012; 1925) due to the fact that many of these species have not been Oliveros et al. 2011; Siler et al. 2011; Sanguila et al. 2016). encountered by biologists since their original discovery (but Nevertheless, the estimated divergence of E. alcalai sp. nov. note the recent discovery of the latter by Pitogo et al. 2020). from E. rugifera appears to have been ancient (16–10 mya; Other species such as E. dattaroyi, E. lankae, E. lewisi, and E. Karin et al. 2017 ) and, as such, the establishment of an resetarii were recently described or elevated primarily on the evolutionary lineage in the Zamboanga Peninsula (itself an basis of morphological characters, without reference to genetic ancient crustal fragment, and part of the geological component information. Genetic data for the remaining lineages are largely known as the Palawan Microcontinent Block; Dimalanta et al. limited to a handful of genes used to assess phylogenetic 2009; Yumul et al. 2009b; Zamoras et al. 2004, 2008; Padrones relationships (Fig. 2; Supporting Information). Only in the et al. 2017) could be related to a seldom-considered, but exceedingly common species, like E. multifasciata, has increasingly invoked mode of faunal transport—paleotransport sufficient sampling accumulated to allow for fine-scale via movement of crustal fragments of continental origin examination of population-level genomic variation (Barley et al. (Blackburn et al. 2010; Siler et al. 2012; Brown et al. 2016). 2015b) though phylogeographic studies of widespread Eutropis Distinguishing between this hypothesis and that of long-distance species through international collaboration are beginning to lead overwater dispersal versus island-hopping through the Sulu to a much clearer understanding of Eutropis diversity (Barley et Archipelago faunal corridor/filter zone (Inger 1954; Diamond al. 2015a; Datta-Roy et al. 2015; Karin et al. 2017). and Gilpin 1983; Brown et al. 2013a; Brown 2016) is beyond Following resolution of many of the systematic problems the scope of the presently available data, but presents an in the Philippine radiation, another major taxonomic challenge intriguing challenge for future biogeographic inquiry in the within this group is in the E. macularia complex. The type southern Philippines. We interpret the presence of a largely low- locality of E. macularia (Blyth 1853) was restricted to elevation, forest obligate, endemic scincid lizard, with the Bangladesh by Taylor and Elbel (1958) and the complex entirety of its geographical distribution limited to Pasonanca includes the species E. austini Batuwita 2016, E. clivicola Natural Park, at the end of the Zamboanga Peninsula, as a high- (Inger et al. 1984), E. allapallensis (K.P. Schmidt 1926) E. value target species for conservation. Such species convey the greeri, E. madaraszi and E. tammanna (Fig. 2a). Barley et al. supreme importance of Pasonanca’s low-elevation climax (2015a) demonstrated that populations from the rest of rainforests and formal protected areas as a shining example of mainland Asia are not closely related to populations from India the value of forested surface catchment watersheds for the near the type locality (Rangpur) and that these mainland preservation of Philippine terrestrial vertebrate biodiversity. populations represent at least two species based on the genetic data (a result that is congruent with the karyotype work of Ota IUCN Species Status of Eutropis alcalai sp. nov. et al. [2001] on the populations from Thailand). How this Eutropis alcalai sp. nov. should be classified under the taxonomic information relates to the three subspecies of E. IUCN Red List Status of Data Deficient (IUCN 2012). The macularia described by Taylor and Elbel (1958) is unknown. known localities for this new species fall within a very small Additionally, because the data presented in the original range, which could be interpreted as “range restricted” and a description of E. tammanna (Das et al. 2008) are not publicly higher threat status. However, there have been no sustained available on GenBank, it is unclear how this species is related to efforts aimed at detecting this species in the region over multiple other lineages in the complex that have been identified in years, and no data exist on abundance or population trends. It is subsequent, more comprehensive studies. possible that the species has a broader distribution in the Zamboanga Peninsula and possibly western Mindanao or the Biogeographical and conservation significance Sulu Archipelago (in particular on Basilan Island). Eutropis Although it is tempting to interpret the apparent absence of alcalai sp. nov. also occurs in one of the most threatened habitat a Eutropis rugifera complex species (E. alcalai sp. nov. or E. types in the Philippines: lowland dipterocarp forest (Kummer rugifera) in the Sulu Archipelago (Fig. 1) or farther up the 1992; Bankoff 2007). Therefore, the species may be benefited Zamboanga Peninsula and western mainland Mindanao, we by landscape/ecosystem-level conservation initiatives, although hesitate to do so for the simple reason that negative species it is crucial that efforts are made to assess this species occurrence data are fraught with assumptions and pitfalls, which conservation status in the future with additional empirical data.

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Although widespread, E. rugifera is rarely found in disturbed and Brown 2012; Barley et al. 2020). Higher elevation areas (Grismer 2011), which may suggest it is relatively populations in the Cordillera Mountains are genetically intolerant of anthropogenic impacts compared to other Eutropis divergent from the other populations on Luzon, and the species. Based on our current knowledge, it seems likely the investigation of mid-elevation contact zones would be valuable same is true of E. alcalai. If the new species is indeed relictual to understand patterns of gene flow. Recent surveys by RMB and largely restricted to the watershed reservation in Pasonanca and AJB have found this species to be common in parts of Natural Park (~12,000 ha), it would likely qualify under a Luzon, including in highly disturbed habitats (e.g., on the threatened category on the IUCN Red List. Ramon Magsaysay Technological University Botolan Campus). We assign them a status of Least Concern. IUCN Species Status of other Philippine Eutropis Eutropis cuprea.—Eutropis cuprea is a poorly known In light of a recent taxonomic revision (Barley et al. 2020), species, currently having only been documented from a single we also provide IUCN Red List assessments for all other locality in southwestern Mindanao (Barley et al. 2020). The Philippine Eutropis here. region is logistically challenging to work and this species is Eutropis bontocensis.—Based on Taylor’s original difficult to distinguish from the more widespread species E. description (Taylor 1923), E. bontocensis was initially thought caraga that also occurs in the region, so it is possible that further to be restricted to high elevation habitats and endemic to the survey work would demonstrate that the species has a larger Cordillera Mountains in northern Luzon. However, recent range in southwestern Mindanao than is currently known. research has demonstrated that it also occurs on numerous small However, the type locality (Tampakan) is also the site of a large islands north of Luzon (Oliveros et al. 2011; Barley et al. 2013). and controversial mining project that was recently extended. We These habitats have experienced considerably less assign this species status to be Data Deficient at present, anthropogenic impacts than others in the region. Historical however, if it is indeed endemic to this small region of populations in Baguio have likely declined or been extirpated Mindanao (and particularly if it is sensitive to anthropogenic due to urbanization, however, surveys by AJB in 2012 found disturbance), the species likely deserves to be elevated to a this species to be common along road cuts near Bontoc, threatened status (in this case “Near Threatened”). suggesting this species tolerates disturbance well and probably Eutropis gubataas.—Eutropis gubataas is known from remains abundant in many areas. Thus, this species should be northern Luzon and the Babuyan Islands (Barley et al 2020). considered Least Concern at present. While the region is relatively large, its distribution is relatively Eutropis borealis.—Eutropis borealis is broadly poorly understood, particularly on Luzon where it has been distributed across the northern and central Philippine Islands, documented only at several, scattered, upland localities. Survey with populations in the Visayan Islands being somewhat work by RMB suggests it is substantially less abundant than the divergent from the northern Philippine populations (Barley et al. morphologically similar and sympatric congener E. borealis. It 2020). Surveys over the past decade by RMB and AJB have appears to be at least somewhat tolerant of disturbance (having found this species to be abundant in many low elevation been collected in agricultural and residential areas near forests) habitats and to tolerate disturbance well (e.g., they are and occurs in at least some protected areas in northeastern commonly found in agriculture and residential areas), so we Luzon (e.g., Aurora Memorial National Park). Thus, we assign this species a status of Least Concern. tentatively assign this species a status of Least Concern, but Eutropis caraga.—Eutropis caraga is broadly distributed suggest that additional survey data would be valuable to across the southern Philippine Islands with populations from understanding this species and any conservation management Siargao and Dinagat being somewhat genetically divergent from considerations. populations across the rest of the species range (Barley et al. Eutropis indeprensa.—The distribution of E. indeprensa 2020). Recent survey work by RMB has found the species to be was recently restricted to the islands of Mindoro and common across of range of elevations and to tolerate northeastern Borneo (Barley et al. 2020). Given that, one would disturbance well (having been reported from agricultural and also expect this species to occur on Palawan, though only the residential areas near forest), so we consider this species to have morphologically similar congener E. sahulinghangganan has a status of Least Concern. been documented from the island. Recent collections of the Eutropis cumingi.— Eutropis cumingi is broadly species are primarily from low elevation, secondary growth distributed across Luzon Island, as well as numerous small forest. Further work is needed to understand population trends islands in the northern Philippines including Lubang, and the and tolerance to disturbance, though we tentatively assign the Batanes and Babuyan Islands (Brown et al. 1996; Devon-song species a status of Least Concern.

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Barley et al.: A new species of Eutropis from the southern Philippines

Eutropis islamaliit.—Eutropis islamaliit is an intriguing Virtually nothing is known about its natural history, population species from a biogeographic perspective. It is known from trends, the extent of its distribution, or tolerance to disturbance. relatively few specimens from scattered localities across We assign the species a status of Data Deficient. It occurs in at Semirara, Lubang, and Samar Islands in the Philippines, and least one protected area (Sibalom Natural Park), though surveys Turtle Island, in Sabah, Malaysia (Barley et al. 2020). The fact are needed to determine if the species occurs more broadly that the species is that broadly distributed, but known from so across the island of Panay, or if it is restricted to the limited area few specimens is surprising, with the majority of specimens from which it is currently known. having been collected in primary and secondary growth forest. Perhaps it is more of a habitat specialist or relictual species Acknowledgements (given its presence on several small offshore islands) than is currently appreciated, athough it is worth noting that recent In addition to our field companions J. B. Fernandez, C. D. surveys by RMB on Samar found populations that were Siler, C. Infante, C. Oliveros, J. Menegay, A. C. Diesmos, and J. sympatric with multiple other morphologically similar species W. Phenix, who we thank for their company and good humor, of Eutropis. Further work is needed to understand population we thank Angel C. Alcala for his many years of support, trends, abundance, and tolerance to disturbance; we tentatively guidance, and (together with the late Walter C. Brown), for assign this species a status of Least Concern, primarily because serving as an inspiration for our team—by setting an example of its apparently broad distribution. with their career-long, ‘role model’ of an international academic Eutropis lapulapu.—This species was recently described collaboration. We thank the staff and management of the from populations that had previously been assigned to E. Zamboanga City Water District, the Pasonanca Natural Park indeprensa (Barley et al. 2020). The species has a broad Protected Area Management Board, Golden Buddha Security, distribution across numerous islands in the central and southern and DENR Region IX office for logistical support, for assistance Philippines where it occurs in a broad range of habitats and in processing regional export permits, and for hosting our visits appears to tolerate disturbance well (having been collected in to Zamboanga; special thanks are due to M. Dela Cruz, A. agricultural and residential areas). Therefore, we assign E. Baysa, A. Adorable, D. Melana, M. Layson, E. Callanta, T. lapulapu a status of Least Concern. Gadon, M. Tee, G. Fernandez, L. Alejo, E. Toribio, L. Sorsone, Eutropis multicarinata.—This species distribution was L. Narag, A. Rojas and A. Braulio. We thank K. Ukuwela for recently restricted to encompass populations in northeastern providing taxonomic advice. We acknowledge financial support Mindanao, Samar, Leyte, and Dinagat Islands (Barley et al. for field work (US National Science Foundation Grant, DEB 2020). It appears to inhabit a range of habitats in the region, 0743491, to RMB) and central Manila DENR and the though it has long been confused with the recently described, Biodiversity Management Bureau for granting our Gratuitous more widespread species E. caraga, which is morphologically Permits to Collect biological specimens (2007–2012); live very similar. Data are needed on abundances and tolerance to animal handling protocols were approved by KU’s Institutional disturbance, although here we assign E. multicarinata a status Animal Use and Care Committee (IACUC AUS 185-05 to of Least Concern. RMB). Phylogenetic analyses were performed using the Eutropis sahulinghangganan—Eutropis sahulinghangga- CIPRES Science Gateway. nan appears to be endemic to the island of Palawan (Barley et Literature Cited al. 2020), although we would expect it to eventually be recorded on satellite islets including the Balabac Island group (to the Alcala, A.C., 1970. Notes on the population biology of the lizard south), as well as Busuanga, Coron, and Culion (to Palawan’s Mabuya multicarinata. The Philippine Biota, 3: 591–611. north). Collections of the species are limited, but it is reported Alcala, A.C. & W.C. Brown, 1966. Thermal relations of two from a variety of low and higher elevation forest types on tropical island lizards on Negros Island, Philippine Islands. Palawan Island proper. Although information is needed on Copeia, 1966: 593–594. population abundances and tolerance to dispersal, these habitats Alcala, A.C. & W.C. Brown, 1967. Population ecology of the are relatively abundant on the island (which is considered a tropical scincoid lizard Emoia atrocostata, in the protected area), and therefore we assign it a status of Least Philippines. Copeia, 1967: 596–604. Concern. Amarasinghe A.A.T, N.A. Poyarkov Jr., P.D. Campbell, S. Leo, Eutropis sibalom.—Eutropis sibalom is known only from J. Supriatna & J. Hallermann, 2017. Systematics of Eutropis several specimens collected from low elevation, secondary rugifera (Stoliczka, 1870) (Squamata: Scincidae) including growth forest in southwestern Panay Island (Barley et al. 2020). the redescription of the holotype. Zootaxa, 4272: 103–118.

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