The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands

Abstract We summarize all available amphibian and reptile species distribution data from the northeast Mindanao faunal region, including small islands associated with this subcenter of endemic vertebrate biodiversity. Together with all publicly available historical information from biodiversity repositories, we present new data from several major herpetological surveys, including recently conducted inventories on four major mountains of northeast Mindanao, and adjacent islands of Camiguin Sur, Dinagat, and Siargao. We present species accounts for all taxa, comment on unresolved taxonomic problems, and provide revisions to outdated IUCN conservation status assessments in cases where our new data significantly alter earlier classification status summaries. Together, our comprehensive analysis of this fauna suggests that the greater Mindanao faunal region possesses distinct subcenters of amphibian and reptile species diversity, and that until this area is revisited and its fauna and actually studied, with on-the-ground field work including targeted surveys of species distributions coupled to the study their natural history, our understanding of the diversity and conservation status of southern Philippine herpetological fauna will remain incomplete. Nevertheless, the northeast Mindanao geographical area (Caraga Region) appears to have the highest herpetological species diversity (at least 126 species) of any comparably-sized Philippine faunal subregion.


Introduction
Recent efforts to conduct comprehensive herpetological surveys of the various islands of the Philippines have provided near-complete estimation of the amphibian and reptile diversity and endemism of several islands, mountain ranges, or other conspicuous geographical subcenters of diversity in the northern reaches of the archipelago ). These efforts have focused on the northern Babuyan and Batanes islands (Oliveros et al. 2011), the large northern island of Luzon (McLeod et al. 2011;Siler et al. 2011a;Brown et al. 1996Brown et al. , 2000aBrown et al. , 2012aBrown et al. , 2013aDevan-Song and Brown 2012), and central island groups (Brown and Alcala 1961, 1964, 1986Siler et al. 2012a), but little recent activity has documented other parts of the archipelago, especially the western island of Palawan, the Sulu Archipelago, and portions of the large southern island of Mindanao (Leviton 1963;Brown and Alcala 1970a;Peterson et al. 2008;Jones and Kennedy 2008;). Mindanao supports high levels of herpetological diversity (Taylor 1920a(Taylor ,b, 1921(Taylor , 1922aBrown and Alcala 1970;Brown et al. 2013a;Diesmos et al. 2015) and considerable endemism, despite its close proximity to the larger landmasses of Sundaland (Borneo, Sumatra, and Java) and Wallacea (Sulawesi; Fig. 1). The current lack of information relating to Mindanao's herpetological diversity, can be attributed to several recent decades of increasing bureaucratic and security-related logistical obstacles to research, combined with earlier, possibly incorrect impression of some biologists that its herpetofauna is reasonably well known (Taylor 1920a(Taylor , 1921(Taylor , 1922aInger 1954;Brown and Alcala 1970, 1978Ross and Lazell 1990).
The perception that southern portions of the Philippine archipelago are sufficiently studied and/or reasonably understood may derive in part from proximity-based expectations of faunal similarity between Mindanao, Sulu, and Palawan versus the islands of Sundaland and Wallacea (Boulenger 1920, Inger 1954, Leviton 1963a, Brown and Alcala 1970. Alternatively, some of this perception stems from the fact that Taylor worked extensively in the region, lived on Mindanao for over two years, collected more than 2,000 specimens from 1912-1915, and returned to survey sites in southern portions of the Philippines periodically until 1924 (Taylor 1975;Duellman 2015). He published widely on southern Philippine herpetofauna (Taylor 1917a(Taylor ,b, 1918a(Taylor ,b, 1919(Taylor , 1920a(Taylor ,b, 1921(Taylor , 1922a(Taylor ,b,c,d, 1923(Taylor , 1928(Taylor , 1960(Taylor , 1965(Taylor , 1975, with his own collections forming the basis for much of this work (Taylor 1975). With this impressive body of work as an early 20 th Century starting point, later workers may have been motivated to focus elsewhere in their efforts to characterize new faunas (i.e., central and northern portions of the archipelago ;Inger 1954;Leviton 1963a;Brown and Alcala 1970). As a result, few modern systematic treatments have revisited the southern Philippine endemics with modern collections-based technologies (fresh specimens,  Table 1. Colored contours correspond to elevation (elevation key). Dinagat Island (Site 13) is often presumed to possess the same herpetofauna as Siargao (the small island to its southeast), whereas the fauna of Camiguin Sur Island (Sites 9-12) has been assumed to be somewhat distinct from that of Mindanao. Neither of these generalizations has been examine critically Alcala 1967, 1970;Ross and Lazell 1991;Nuñeza and Galorio 2015). Province names are abbreviated with unique letter combinations used in text and tables (provincial map key), and numbered sites are assigned to province (surveyed sites key) for clarity. integrative techniques, ecological information, life history studies, acoustic data, genetic resources, or molecular data); in fact, many recent attempts at archipelago-wide revisions of Philippine fauna have had no alternative but to base decisions regarding species diversity (e.g., Diesmos et al. 2015) and conservation status (IUCN 2016) on Taylor's original specimens, collected over a century ago (Leviton 1964a(Leviton ,b,c,d, 1965a(Leviton ,b, 1967(Leviton , 1968Brown and Alcala 1978, 1994Brown and Guttman 2002;Siler et al. 2011b).
Our prevailing perspective on patterns of endemism and subdivision of terrestrial biodiversity in the archipelago includes a general acceptance of a model of diversification based on late Pleistocene sea level oscillations and the generation of periodic land connectivity; this has been termed the Pleistocene Aggregate Island Complex (PAIC) paradigm (Inger 1954;Voris 2000;Brown et al. 2013b). According to this model for understanding patterns of diversity ) and processes of diversification Esselstyn and Brown 2009;Oaks et al. 2013;Brown et al. 2013b) the Mindanao PAIC is one large biogeographically significant sub-province within the archipelago, with Mindanao proper having been connected to the neighboring islands (Dinagat, Siargao, Samar, Leyte, Bohol, Biliran, etc.) of today by land bridges during the Pleistocene (Taylor 1928;Inger 1954;Voris 2000;). Because of this perspective, herpetologists have assumed that when sea levels were low during the Pleistocene, the emergence of land bridges allowed faunal exchanges between connected landmasses, producing highly endemic faunas within PAICs Diesmos 2001, 2009) and archipelago-wide diversity generated by prevailing species boundaries between PAICs. Augmenting the PAIC paradigm, appreciation for recent geological events, such as accretion of paleoisland precursors along identified suture zones (Yumul et al. 2003(Yumul et al. , 2009, and repeated bouts of colonization (Siler et al. 2011b;Brown et al. 2013b;Brown and Siler 2013;Justiniano et al. 2015;Brown 2015) are assumed to contribute to the evolutionary process of diversification via within-and among-PAIC isolation. Habitat fragmentation, ecological gradients, and barriers in ecological suitability are assumed to have contributed as well, although these ideas are less studied potentialities for future research programs (Steppan et al. 2003;Esselstyn and Brown 2009;Brown et al. 2010a;Esselstyn et al. 2011;Roberts et al. 2011;Sanguila et al. 2011;Gonzales et al. 2014;Hosner et al. 2014). Together, these geological, geographic and faunal distribution patterns contribute to the Philippine archipelago's recognition as a "model" archipelago for the study of evolutionary processes of biotic diversification . How is it, then, that the herpetofauna from the southern portion of this model island archipelago remains so poorly understood, with the many of Mindanao's endemic species absent in modern molecular phylogenetic studies, and fundamental phylogeographic studies of even its most common species sorely lacking (but see McGuire and Kiew 2001;Evans et al. 2003;Sanguila et al. 2011;Siler et al. 2011b;Brown and Siler 2013).
Mindanao was formed by the accretion of the island-arc related to the eastern-central block and the western continental Zamboanga peninsula block, separated by the active Sindangan-Cotabato-Daguma lineament (Yumul et al. 2003). The eastern portion of the island was previously divided into the high mountains of the Pacific Cordillera from the Central Cordillera, separated by the Agusan-Davao trough (Yumul 2004(Yumul , 2009). Many of the large isolated mountains in central Mindanao are separated from each other by substantial stretches of low-lying areas (Hall 1998) that were formed as a result of collision, accretion and subduction events over the past ten million years (Hall 1996(Hall , 1997Yumul et al. 2003Yumul et al. , 2009). This highly dynamic geological history suggests the possibility of faunal limits to dispersal within early paleoislands, a mechanism that might possibly have contributed to the diversification of the first amphibian and reptile lineages that colonized Mindanao (Brown and Alcala 1970;Brown and Guttman 2002;Evans et al. 2003;Sanguila et al. 2011;Brown et al. 2013b; Barley et al. 2014).
Previous studies have described the possibility of an "island-hopping" mode of dispersal across paleoislands to explain colonization of the eastern Philippine island arc (Diamond and Gilpin 1983;Brown and Guttman 2002;Jones and Kennedy 2008;Oliveros and Moyle 2010;Roberts et al. 2011;Sanguila et al. 2011;Brown and Siler 2013;Gonzales et al. 2014); many of these conceived west-to-east dispersal/diffusion route of colonization (Brown and Guttman 2002;Evans et al. 2003;) from the western continental Zamboanga peninsula, as evidenced by the series of island formations distributed across the southern island (Hall 2002;Yumul 2004;Gonzales et al. 2014).
Because of obstacles to biologists' access to parts of Mindanao, our current understanding of the island's herpetofauna comes in large part from the historical works of Boulenger (1882Boulenger ( , 1884Boulenger ( , 1920, van Kampen (1923), Smith (1930Smith ( , 1935, as well as the synthetic works of Taylor (1920aTaylor ( ,b, 1921Taylor ( , 1922a, Leviton (1963a), Alcala (1970, 1978, 19780), and a few, scattered and site-specific studies (e.g., Smith 1993a,b;Ross and Lazell 1990). These have been subsidized by recent inventories from the eastern (Delima et al. 2006(Delima et al. , 2007Ates et al. 2009;Relox et al. 2011), central , and western (Nuñeza et al. 2009) portions of the island. Each are welcomed additions but most have been constrained in scope and lacking biogeographical context.
In this paper, we take what we hope will be a first step towards ameliorating Mindanao's herpetological information shortage, by initiating the second study in a series of attempts towards a comprehensive review of the herpetofauna of the island. In this paper, we focus on the regional diversity and endemism of amphibians and reptiles from one subcenter (northeast Mindanao) of the biogeographically distinct Mindanao PAIC. We present species accounts using data from our own intensive herpetological surveys of northeast Mindanao and its adjacent islands, and provide notes on each species' microhabitats and natural history. To provide a biogeographical synthesis, we include historical museum records from all accessible biodiversity repositories. The anticipated result will be a new opportunity to review numerous unresolved taxonomic problems, provide a new standardized reference for species distributional data, a much needed biogeographical reconsideration, and a platform from which biodiversity specialists can undertake revisions of the conservation status of the poorly understood herpetofauna of the southern Philippines.

Materials and methods
We surveyed amphibian and reptile diversity at four major sites in Surigao del Norte, Surigao del Sur, Misamis Oriental, Agusan del Norte and Agusan del Sur provinces (Camiguin Sur, Dinagat and Siargao islands; Table 1) using standardized sampling techniques (Heyer et al. 1994) and specimen collection and preservation methodology (Brown et al. 2000a(Brown et al. , 2012aASIH 2004;Siler et al. 2011a;Simmons 2015). Our most recent surveys (March-April, 2009;June-August, 2012) involved intensive elevational transects on Mts. Balatukan and Lumot, Misamis Oriental Province and Mts. Magdiwata and Hilong-hilong, Agusan del Norte Province (Fig. 1). Surveys were conducted in early mornings, mid-day, afternoons, and evenings by experienced teams of four to eight individuals, sampling a wide variety of habitat types within each general study location. Habitats included dry forest on ridges, moist ravines, forest trails at all elevations, dry intermittent streambeds, small streams, seaps and swampy areas, large rivers, forest gaps and edges, and grassy open areas. Investigators at each sampling location made extensive surveys of each area (on foot) to ascertain habitat types and then visited each at varying times of the day. Nocturnal searches (1800-2400 hr) were conducted at each habitat type, within each sampling site, on dry and rainy nights. By concentrating field survey efforts to span the end of the dry season and the beginning of the rainy season (June-August) we were able to assure that each habitat type at each location was sampled under differing atmospheric conditions. Sampling Locations. Data presented here include results of our own surveys (Table 2) and a variety of pre-existing collections, both intensive and incidental, from major Philippine and U.S. Museum collections (see acknowledgements). In addition to extensive collections housed at the California Academy of Sciences (CAS: fieldwork of E. H. Taylor, W. C. Brown, ACA, and colleagues), the University of Kansas (KU) and the National Museum of the Philippines (PNM; field work of RMB, ACD, MBS, KAC, and CDS), we summarize historical records from the Smithsonian (USNM), the Carnegie (CM), and Harvard (MCZ), originating from Misamis Oriental (northern Mindanao Region), Agusan del Norte, Agusan del Sur, Surigao del Norte, Surigao del Sur provinces (Caraga Region), Dinagat, and Siargao records (previously summarized by Ross and Lazell 1990) and Camiguin Province (Camiguin Sur Island). To be as comprehensive as possible in our treatment of northeast Mindanao Island, we include all of these records here, with the caveat that methods of surveying herpetological communities most likely differed among collection efforts and locations.

Results
We document at least 126 species of amphibians and reptiles from northeast Mindanao and adjacent islands, including 40 species of frogs, one species of caecilian, 49 species of lizards, 35 species of snakes, one species of freshwater turtle, and one species of crocodile (Table 2). This diversity represents approximately 36% percent of the total Table 1. Northeast Mindanao faunal region sites included in this study (where herpetological specimens have been collected and/or observations have been recorded).
Numbered sites correspond to Figure 1 (map; note that some sites are included under a single number in Figure 1 in cases of close proximity) and coordinates and elevation are included when available. * = sites georeferenced for this study; ** = combined extremely proximate sites into one set of coordinates.  (Taylor, 1915) H/N N N LC Cyrtodactylus annulatus (Taylor, 1915) Brown et al. 2008;Diesmos and Brown 2011;Brown and Stuart 2012;Diesmos et al. 2015) and 85-90% of the taxa reported here are endemic to the Philippines. We provide accounts for each species, provide notes on their habitat and natural history, and draw attention to many unresolved taxonomic problems (involving ~40% of the species included) relevant to particular taxa. We also comment on the conservation status of individual species when our new data suggest that existing conservation status assessments (IUCN 2010(IUCN , 2016Diesmos et al. 2014Diesmos et al. , 2015 are out of date or require revision (Siler et al. 2011a;McLeod et al. 2011;Brown et al. 2012aBrown et al. , 2013a. In cases of taxonomic uncertainty involving frogs of the family Ceratobatrachidae, we refer to undescribed species by informal place-holder nicknames derived for distinctive characteristics (phenotype, habitat, calls;Brown et al. 2012a) and species identification numbers, following Brown et al. (2015)

Amphibia Family Bufonidae
Ansonia muelleri (Boulenger, 1887) This species (Fig. 2) is widespread across the island of Mindanao and is the sister species to A. mcgregori from the southern tip of the Zamboanga peninsula (Matsui et al. 2010;Sanguila et al. 2011). With its highly specific torrent-dwelling larval habitat requirements (Inger 1960(Inger , 1985, this species is limited to montane habitats (and lowlands immediately adjacent to mountains) with high gradient stream flow. As a result, it is widespread but distributed patchily throughout Mindanao in strips of geographic ranges surrounding major mountain ranges (Inger 1954(Inger , 1992(Inger , 2005Duellman and Trueb 1994;Matsui et al. 2010). In suitable habitat this species was observed in large numbers and was often the most commonly observed amphibian in an immediate area. Adults were typically collected in splash zones near rapids or waterfalls although they were also found some distance from water, especially after a period of rainfall. A recent phylogeographic study identified five genetically divergent and geographically structured haplotype groups in central and eastern Mindanao. Sanguila et al. (2011) argued that, at a minimum, these lineages should be recognized as evolutionarily significant units (ESUs) for conservation purposes, and that they may represent distinct species. Two of these ESUs are represented in this study and morphological differences were observed between one form found at Mts. Hilong-hilong and Magdiwata and another morphotype present at Mts. Balatukan and Lumot. Little can be made of this species current conservation status ("Vulnerable:" B1ab(iii); IUCN 2016) because no actual studies of its population status have ever confirmed the "populations trend decreasing" assessment and because its "severely fragmented" distribution appears to be the result of its natural, larval habitat requirements. However, if Sanguila et al. (2011) data can be interpreted as evidence for multiple cryptic species contained within A. muelleri, then each of these five divergent lineages would have much more limited distributions, perhaps qualifying each for some level of threat category. In the absence of firm conclusions either way, and until taxonomic studies are undertaken to confirm or refute this possibility, A. muelleri should be considered "Data Deficient" (DD; IUCN 2010; Diesmos et al. 2014

Pelophryne brevipes (Peters, 1867)
As currently understood, this species (Fig. 3) inhabits an improbably disjunct distribution that includes Mindanao and Basilan Islands (type locality: Zamboanga), and parts of the Malay Peninsula, Sumatra, Mentawi Islands and Natuna Islands (Inger 1954(Inger , 1999. Considered "Least Concern" (LC) by IUCN (2016), this uncommon and patchily distributed small species can be found calling (with soft "peeping" vocalization) on the surfaces of shrubs and understory trees in the vicinity of running water. It appears to tolerate moderate levels of forest disturbance and has been found on ornamental plants around buildings on forest edges (RMB personal observations). We suspect that this species eventually will be recognized as a Philippines endemic, and that populations outside of the country will be referred to other species. At that time, reconsideration of its conservation status will be required.

Rhinella marina (Linnaeus, 1758)
We frequently observe this common introduced species in the vicinity of human habitations and in agricultural areas on Mindanao. It may have originally been introduced to the Philippines during the industrial revolution and the major sugar cane agricultural production boom on the central Philippine island of Negros (Alcala 1957; Brown and Alcala 1970), and has now become widespread throughout the country Diesmos et al. 2006Diesmos et al. , 2015. We observed this species around human populations at most sites, but did not encounter it in forested areas. We collected a single specimen at low elevation on Mt. Lumot. Sites and specimens: MO 6: KU 333803.

Family Ceratobatrachidae
Platymantis sp. 34: P. cf. corrugatus (Duméril, 1853) Platymantis cf. corrugatus (Fig. 4) is a commonly encountered widespread species on the Mindanao PAIC. Populations on these islands are morphologically and acoustically distinguishable from Luzon and Visayan PAIC lineages and molecular studies are underway to determine if these slight differences could constitute the basis for taxonomic recognition. Although the call on all three PAICs is similar, the lineage on the Mindanao PAIC ("sp. 34;" Brown et al. 2015) have a different preferred calling habit (calling from exposed areas, versus under leaf litter on Luzon) and daily pattern of activity (calling all evening versus strictly crepuscular; RMB personal observations.). Platymantis corrugatus is classified by the IUCN as "Least Concern" (LC; IUCN 2016).

Platymantis guentheri (Boulenger, 1884)
As currently understood, this common, widespread species (Fig. 5) is now known from seven Philippine islands throughout the Mindanao PAIC and is known from at least six of Mindanao's major mountain ranges (AmphibiaWeb 2013); Although P. guentheri has been considered "Vulnerable" (VU: B1ab(iii); IUCN 2016) on the basis if an "extent of occurrence less than 20,000 square km'" and distribution "extremely fragmented distribution" with populations trends inferred to be declining on the basis of "continuing decline in the extent and quality of its habitat" (IUCN 2016), continued treatment of this species as under threat of extinction is no longer tractable. Our now extensive surveys throughout Bohol, Samar, Leyte, Dinagat and Mindanao indicate that this very widespread species persists in second growth, degraded, and fragmented habitat patches and is be highly abundant and commonly encountered (and easily identified on the basis of its frequency sweep advertisement call), given sufficient precipitation and any form of vegetation pre- A B sent. We therefore downgrade this species to "Near Threatened" (NT; IUCN 2010; Diesmos et al. 2014) and emphasize that we do not anticipate circumstances that could result in re-elevating of this species to a higher threat category in the foreseeable future.

Platymantis cf. guentheri sp. 2
We collected a single specimen of a morphologically distinctive arboreal Platymantis and high elevation on Mt. Lumot. Molecular studies are under way to determine the genetic affinities of the single specimen. Site and specimens: MO 2: KU 334329.

Platymantis rabori Brown, Alcala & Diesmos, 1998
Considered "Vulnerable" (VU; B1ab (iii); IUCN 2016) for the same reasons as P. guentheri (above), Platymantis rabori ( Fig. 7) is relatively widespread on Bohol, Leyte, Samar, and Mindanao (AmphibiaWeb 2013), but is much less commonly encountered and locally considerably less abundant. This species is easily identified by its distinctive morphology (relatively large body size, widely expanded terminal disks of the fingers) and slowly repeated pulsed call, similar to other species of the P. guentheri complex (Brown et al. 1997) and it usually calls high in the forest canopy, which may explain why it is seldom collected and rare in collections. This suspected new species of Platymantis ( Fig. 9) has now been recorded at numerous sites on Mindanao, Bohol, Samar and Leyte islands and is readily identified by its small body size and unique advertisement call, consisting of a long chain of clicking pulses. Preliminary phylogenetic analyses of DNA sequence data suggest that these populations constitute a distinct lineage that is divergent from other Mindanao PAIC species. Sites and specimens: AS 5: KU 319508-12, KU 319403-17.

Platymantis sp. 38: "Cliff loud"
We collected this morphologically and acoustically distinctive undescribed species of Platymantis ( Fig. 10)  Two specimens of an acoustically unique Platymantis species (Fig. 11) were collected by ACD, MBS and party on Mt. Balatukan. The advertisement call of this population is so distinct that we cannot ally it with any known Platymantis.

Family Dicroglossidae
Fejervarya moodiei (Taylor, 1920) This endemic species, formerly considered conspecific with F. cancrivora but now afforded the status of an endemic Philippine species, due to its genetic distinctiveness (Kurniawan et al. 2010(Kurniawan et al. , 2011, is common in coastal areas of NE Mindanao, brackish water swamps and mangroves, and river mouth estuarine areas. CAS specimens were collected in coastal areas, along riverbanks, four decades ago; an assessment of this species status in these heavily populated areas would be advisable. This species' conservation status is "Data Deficient" (DD; IUCN 2016). As lowland and coastal habitats throughout the country continue to be invaded by the introduced species Hoplobatrachus rugulosus and Kaloula pulchra (Diesmos et al. 2006, these voracious generalist species replace entire populations of native frogs (RMB, ACD personal observation), either via competition for resources or direct predation on native taxa. We suspect this widespread Philippine endemic may soon become threatened with extinction; accordingly we recommend field-based surveys targeting conservation status assessment.

Fejervarya vittigera (Wiegmann, 1824)
The first endemic Philippine species known to science, F. vittigera (Fig. 12) inhabits low elevation aquatic habitats and is often found in streams, drainage ditches and flooded rice fields. It is easily identified at a distance by its loud "honking" advertisement call and aggregation in large choruses. This species conservation status is currently "Least Concern" (LC; IUCN 2016) but we revise it to "Data Deficient" (DD; IUCN 2010 to reflect the paucity of available data on its actual confirmed (versus presumed; IUCN 2016) distribution and the degree to which it may be threatened by invasive species. Like F. moodiei, this is another species that is quickly replaced when Hoplobatrachus rugulosus and Kaloula pulchra invade its low elevation riparian habitat (RMB, ACD personal observation; Diesmos et al. 2006Diesmos et al. , 2014Diesmos et al. , 2015.

Limnonectes diuatus (Brown & Alcala, 1977)
Recent collections of this species (Fig. 13) at high elevations on Mts. Balatukan and Lumot and on Dinagat Island make it clear that, as presently understood, this species is considerably more widespread than previously thought. Previously recorded in the Diwata range (Brown and Alcala 1977) and Mt. Kitanglad (Bukidnon Province; based on a single, tentatively identified specimen in FMNH; ACD personal observation), this species is most likely widely distributed on montane formations throughout much of northern and central Mindanao. This species' conservation status has been arbitrarily listed since 2004 as "Vulnerable" (IUCN 2016) based solely on its previously assessed range of less than 20,000 square kilometers and possessing a "severely fragmented" distribution with "continuing decline in the extent and quality of its forest habitat," (IUCN 2016) none of which has actually been determined with accompanying field based data. Now that is it known to inhabit a much wider geographic range (data presented here), we adopt the suggested revision of Diesmos et al. (2014) who downgraded this species status to "Near Threatened" (NT: IUCN 2010). Given that there have, in fact, been no actual field studies of habitat fragmentation in its actual range and that we know nothing of the extent or quality of its required habitat, we fail to see how this species' status could be elevated to a higher level, unless drastic land use changes at high elevations were to occur. Many of the known areas of occurrence are now protected areas (Mts. Hilong-hilong, Kitanglad) or proposed protected areas (e.g., Mt. Lumot), so this species may well be reasonably well protected for the foreseeable future.
Sites and specimens:

Limnonectes leytensis (Boetger, 1893)
The Leyte Swamp Frog, L. leytensis (Fig. 14), is widely distributed on the Mindanao, Visayan, and Romblon PAICs and is frequently observed in swamps or marshes, but also along small streams or other bodies of water. We collected specimens on grassy banks of streams on Dinagat Island, and in marshy areas of Bunawan. Its conservation status is "Least Concern" (LC; IUCN 2016).
Sites Limnonectes cf. magnus (Stejneger, 1910) Stejneger (1910) described Limnonectes magnus from specimens collected at high elevation on Mt. Apo. Over the century, this name was widely applied to all large bodied fanged frogs of the Mindanao PAIC (Samar, Leyte, Bohol; Brown and Alcala 1970;Alcala and Brown 1998). Evans et al. (2003) demonstrated the genetic distinctiveness of the high elevation Mt. Apo species, which is considerably divergent from the widespread low elevation species. Thus, the taxonomic status of the widespread, low elevation species remains unresolved with respect to real L. magnus at its type locality. Limnonectes magnus has been treated as "Near Threatened" because the species is hunted for its meat. However, the degree to which this species actually is "over-harvested" has never been properly ascertained (contrary to IUCN 2016) and there have been no studies whatsoever of its status on Mt. Apo. It remains possible that the widespread low elevation form ( Fig. 15) should be downgraded to "Least Concern," whereas the high elevation Mt. Apo populations may actually be range-restricted and worthy of higher conservation status. Until taxonomic studies have properly been undertaken, surveys for genetic variation are conducted throughout its range, and populations at high elevations have been studied, we argue that this species should be treated as "Data Deficient" (DD; IUCN 2016) and that field-base conservation efforts be focused on the Mt. Apo population.
Sites and specimens:   Limnonectes parvus (Taylor, 1920) Commonly encountered and locally abundant in central, southern, and western Mindanao, this species (Fig. 16) is conspicuously absent at most NE Mindanao localities and has not been recorded on Bohol, Leyte, Dinagat, Siargao, or Samar. Considered "Vulnerable" (VU; IUCN 2016: B1ab (iii) because of is supposedly small (< 20,000 square kilometers), its "severely fragmented distribution" and "continued decline in quality and extent of Mindanao's forests, this species is now known from Bunawan in NE Mindano (the origin of our specimens), throughout much of south-central Mindanao, and all the way west to Zamboanga and Basilan. Given that no studies of its distribution have ever actually been conducted, it appears widespread, commonly encountered, and locally abundant wherever present, we consider a revision of its conservation status warranted. We take issue with the IUCN (2016) characterization of is range as "severely fragmented" and we follow Diesmos et al. (2014) in their downgrading of this species to "Near Threatened" (NT) using IUCN status assessment criteria (IUCN 2010). Our acceptance of this proposal (a downgrade from VU to NT, and not Least Concern) is based on the observation that the species appears somewhat dependent upon vegetation cover (but second growth and nursery forests appear sufficient to sustain large populations; RMB, ACD, MBS personal observation).

Occidozyga laevis (Günther, 1859)
Yellow-bellied Puddle Frogs are widespread, aquatic, non-endemic species found in a wide variety of freshwater habitats from streams, rivers, swamps, and flooded rice fields at low elevation, to cascading mountain streams in montane environments. This species' conservation status is "Least Concern" (

Leptobrachium lumadorum Brown, Siler, Diesmos & Alcala, 2009
Widespread throughout Mindanao and Basilan, but not Dinagat, Siargao, Leyte, Samar or Bohol ), this species (Fig. 17) occurs at naturally low abun-  dances in a wide variety of habitats provided that some vegetation persists. We have encountered this species in agricultural areas adjacent to second growth, tree nurseries, along riparian habitats in lowland forest, and up to 1,500 m on Mts. Magdiwata, Hilong-hilong, Balatukan, and Lumot. Previously unassessed, we categorize this species as "Least Concern" (LC) using IUCN conservation status assessment criteria (IUCN 2010 Megophrys stejnegeri (Taylor, 1920) This species conservation status has been listed as "Vulnerable" (VU; IUCN 2016). Since 2004, this classification is no longer tenable given new information on its extremely widespread distribution (throughout all islands of the Mindanao PAIC), its wide ecological tolerance of disturbance, and the fact that it is commonly encountered and locally abundant. For the same reasons we have suggested downgrading Platymantis güntheri, Limnonectes parvus and L. diuatus, we similarly propose a downgrade of Megophrys stejnegeri (Fig. 18) to "Near Threatened" (NT; IUCN 2010) and emphasize that we can foresee no circumstances that conceivably would result in a higher level of threat category being assigned to this species in the near future. Sites

Chaperina fusca Mocquard, 1892
Known only from Mindanao, Jolo, and Palawan in the Philippines, but also from Peninsular Malaysia, Thailand and central Borneo, this species (Fig. 19) inhabits a wide geographic range, but is characterized by a patchy and unpredictable distribution. We commonly find this species in moderately sized (10-25 individuals), tightly clustered choruses surrounding stream-side pools (preferred breeding habitat) in rock impressions. The advertisement call of this species is a series of high frequency, quiet "peeps" and it is currently classified as "Least Concern" (LC; IUCN 2016); that status should be revisited following molecular studies aimed at determining the number of evolutionary lineages in this widespread, but poorly understood species.

Kalophrynus sinensis Peters, 1867
Until recently, Kalophrynus pleurostigma Tschudi, 1838 ( Fig. 20) was considered a widespread species from the Philippines, Indonesia, Singapore, Malaysia, Thailand and Myanmar (AmphibiaWeb 2013). Zug (2015) clarified the status of the Philippine population and resurrected K. sinensis-the oldest available name, Calophrynus pleurostigma var. Sinensis Peters (1867)-for the Philippine lineage. In doing so, he considered all Philippine populations to be conspecific, and thus, considered Boettger's 1897 Calophrynus acutirostris (type loc.: Samar Island) and Stejneger's (1908) Kalophrynus stellatus (Basilan Island) to be junior synonyms of K. sinensis. If future genetic studies determine that significant geographic structure exists to warrant the recognition of allopatric islands populations, additional names may need to be resurrected from the synonymy of K. sinensis (Zug 2015). Kalophrynus sinensis frequently is encountered in the rainy season, calling while floating in temporary pools or water filled cavities in a variety of habitats of varying levels of disturbance. It is classified as

Kaloula conjuncta meridionalis Inger, 1954
The curious, patchy, and unpredictable distribution of the Mindanao PAIC representative of the Kaloula conjuncta group (Inger 1954;Alcala and Brown 1998) leads us to think that this "subspecies" (Fig. 21) has a distinctly different natural history than the other taxa in this group (e.g., K. c. conjuncta from Luzon and K. c. negrosensis from the western Visayan islands of Panay, and Negros). In a recent phylogenetic analysis, Blackburn et al. (2013) demonstrated the monophyletic and genetic distinctiveness of  each of the subspecies of Kaloula conjuncta, suggesting that each may warrant specific status. If this action is followed by taxonomists, the conservation status of K. c. meridionalis will need to be independently assessed. Kaloula conjuncta currently is considered "Least Concern" (LC; IUCN 2016) and its subspecies have not been assessed individually. Frogs of the Kaloula conjuncta Complex emerges at the start of the rainy season, forming large breeding aggregations. At other times of the year they virtually undetectable (RMB personal observation), which will be a challenge for future conservation status assessments; we consider this taxon to be a distinct evolutionary lineage, worthy of species rank; as such we classify it as Data Deficient (DD; IUCN 2010).

Kaloula picta (Duméril & Bibron, 1841)
Distributed throughout the Philippines and formerly quite commonly encountered in dense aggregations in rice fields and temporary bodies of water in the rainy season, Kaloula picta ( Fig. 22) has been considered "Least Concern" (LC; IUCN 2016). We consider Kaloula picta to be "Data Deficient" (DD; IUCN 2010), owing to lack of recent field observations confirming the species' actual (versus presumed; IUCN 2016) distribution and the extent to which it may be threatened by invasive species. Unevaluated threats to this species may include competition and/or direct predation from recently introduced invasive species K. pulchra and H. rugulosus (Diesmos et al. 2006. Blackburn et al. (2013) demonstrated the genetic uniformity of this species throughout the Philippines, which would tend to refute Inger's (1954) suggestion that regional morphological differentiation within K. picta might eventually warrant taxonomic partitioning.

Kaloula sp. (undescribed)
An undescribed species of forest cavity-dwelling (tree hole) Kaloula has been documented on Leyte, Samar islands (Blackburn et al. 2013), and northeast Mindanao (Fig. 23). Distictive "honking" vocalizations were heard during our 2012 survey of Mt. Hilonghilong (RMB, unpublished data) but the source of these apparently species-specific calls was never documented because of the sheer, canyon-like terrain at Eye Falls. Nevertheless, we are certain of this new species' identity, having traced its distinctive advertisement call to individual calling males on numerous occasions on Samar and Leyte, and so inclusion of this undescribed species in the this report is clearly advisable at this time.
Sites and specimens: none.

Oreophryne cf. nana Brown & Alcala, 1967
Described from Camiguin Sur Island (Brown and Alcala 1967), O. nana reportedly differs from O. anulata (from Mt. Apo) on the basis of the absence of subarticular tubercles on the hand. Now that specimens of Oreophryne have been collected from numerous high and low elevation sites throughout Mindanao, a comprehensive appraisal of the genetic and morphological variation in this group is overdue. Our impression is that there are several sites where the degree of distinctiveness of the subarticular tubercles varies and that this character may require additional study. For now, we refer all our Northeast Mindanao Oreophryne to "O. cf. nana" with the caveat that we are not at all certain of this identification. This species (Fig. 24) is listed as "Data Deficient" (DD; IUCN 2016) due to taxonomic and species distribution uncertainty and as a taxonomic reappraisal becomes available, the status for the Camiguin Sur Island and northeast Mindanao populations will need to be revisited. Specimens from our Mt. Lumot expedition were positive for chytrid fungus Diesmos et al. 2012Diesmos et al. , 2015, which leaves us with concerns for the long-term status of these populations.

Family Ranidae
Pulchrana grandocula (Taylor, 1920) This species (Fig. 25) is widespread, commonly encountered at high abundances, and distributed throughout the Mindanao PAIC islands (Brown and Guttman 2002;Brown and Siler 2013). Classified as "Least Concern" (LC; IUCN 2016) the species can be found in a variety of disturbed habitats and is distributed across much of the elevational relief of Mindanao. Males form choruses around pools in streams and rivers and call when water levels are relatively low; females have been observed in these same riparian habitats, but also relatively far away from water (100 m or more), sometimes perching in low branches of understory trees, and occasionally aggregating in shallow caves formed by overhanging stream banks (RMB personal observation). The discovery of a new, morphologically similar, and exceedingly rare stream frog species that had previously been confused with Hylarana grandocula Brown 2015) leads us to speculate that mountains of northeast Mindanao may also harbor undocumented populations of this second Mindanao Hylarana taxon. Oliver et al. (2015) recently published a phylogeny for many members of the African, Papuan, and southeast Asian members of the genus Hylarana and recognized "Pulchrana" as the available name corresponding to the Hylarana signata complex (sensu Brown and Guttman 2002;Brown and Siler 2013). Although this action is arbitrary and unnecessary and no justification for a maximally atomized classification was provided (Wiens et al. 2009;Poe 2013;Brown et al. 2015) we adopt the most recently published name. Sanguirana albotuberculata (Inger, 1954) Previously considered a subspecies of "Rana everetti" (Inger 1954), the treatment of the Samar-Leyte-northeast Mindanao population of slender stream frog as a full species, distinct from the southwest Mindanao population (topotypic Sanguirana everetti) is an arrangement that was first postulated on the basis of morphometric data (Brown et al. 2000bFuiten et al. 2011), and has now been confirmed with genetic data (Brown et al. 2016). When reproductively active (July-August on Mt. Lumot) this species (Fig. 26) can be exceedingly common and locally abundant in riparian habitats, and usually perches in streamside vegetation, on steep banks, or large boulders, several meters from water; we have observed individuals as high as 4-5 m in riverbank trees. Lacking vocal sacs, Sanguirana albotuberculata males call with a slow, dull, pulsed rattle vocalization (the apparent advertisement call). Females also vocalize with a series of rapid "squeaks" and "whistles" delivered in a series of descending frequency notes (RMB personal observation) as has been reported two related species, S. luzonensis (Brown et al. 2000) and the lineage from the West Visayan islands' (Brown et al. in review). Previously considered "Data Deficient" (DD; IUCN 2016), this species has now been recorded at a sufficient number of localities that we can evaluate it against IUCN (2010) conservation status criteria. Sanguirana albotuberculata does not qualify for elevated threat status; we consider it "Least Concern" (LC; Diesmos and Brown 2011;Diesmos et al. 2014

Staurois natator (Günther, 1858)
Common throughout the Mindanao faunal region, Staurois natator (Fig. 27) is a frequently observed component of most amphibian communities of the southern Phil-

Family Rhacophoridae
Theloderma (Nyctixalus) spinosum (Taylor, 1920) This somewhat rarely encountered species (Fig. 28) is known from Basilan, Leyte, Samar, Mindanao, and Bohol. Taylor speculated that its apparent "rarity" might be due to the species' arboreal microhabitat preferences; he reported finding individuals underneath leaf litter inside a tree cavity (Taylor 1920a). We suspect that this species, which does not form choruses, is rare in collections because of the difficulty of localizing and tracking its soft, tonal advertisement call, and so we do not recommend elevating conservation status (sensu IUCN 2016) of this species on the basis of the frequency with which it has been collected historically in faunal surveys. We typically locate individuals on shrubs and saplings after triangulating (with multiple field collectors) on the source of its recognizable, but soft whistling call. Theloderma spinosum calls with a quiet, high frequency, pure tone chirp or a rapid series of quiet, high frequency, ascending chirps. Based on our observations this species seems to rely on primary and mature secondary forest and has been collected along elevational ranges of 300-1,100 m (Brown and Alcala 1994;Alcala and Brown 1998). On the basis of a molecular phylogenetic analysis Poyarkov et al. (2015), recommended that Nyctixalus be considered a subgenus of Theloderma. This species was previously classified as "Vulnerable" (VU; IUCN 2016). It has recently been reassessed ) on the basis of new data and qualifies only for "Near Threatened" (NT; IUCN 2010; Diesmos et al. 2014Diesmos et al. , 2015.

Philautus acutirostris (Peters, 1867)
This small shrub frog (Fig. 29) is one of the most conspicuous, locally abundant, and commonly recorded montane forest frogs-provided that field workers recognize its distinctive advertisement call (a rattle, with decremental pulse rate), conduct surveys on nights following heavy rains, and search its preferred microhabitat (small leaf perches, including undersides of leaves, 0.3-1.0 m above forest floor). It is known from Jolo and Basilan islands of the Sulu Archipelago, and throughout the entirety of Mindanao Island from the extreme southwestern Zamboanga City area (RMB, ACD, CDS, and MBS personal observations) to the mountains of the northeast (Brown and Alcala 1994). As noted by Diesmos et al. (2014), the species IUCN conservation status ("Vulnerable:" B1ab(iii); IUCN 2016) is incorrect because no actual population research has ever confirmed the "populations trend decreasing due to continuing decline in the extent and quality of its forest habitat" justification (IUCN 2016). We found this species to be quite common, in dense aggregations, even in highly disturbed matrices of regenerating second growth and shifting agriculture on the lower slopes of Mts. Balatukan, Magdiwata, Hilong-hilong, and Lumot. Characterizing this species' range as "severely fragmented" also appears to be erroneous because it is generally a mid-high-montage species (and, thus, has a naturally discontinuous distribution on isolated mountains). Philatus acutirostris was screened extensively for fungal; pathogens in 2010-2012 and we detected no signs of infection by chytrid fungus.
Sites and specimens:

Philautus poecilius Brown & Alcala, 1994
This high elevation Mindanao endemic shrub frog (Fig. 30) calls from a variety of perches in primary forests above 700 or 800 meters. It has been recorded from forests of eastern Mindanao (Brown and Alcala 1994;Plaza and Sanguila 2015) to the more central high elevation forests of Mt. Lumot to Mt. Malingdang of western Mindanao (Nuñeza et al. 2010). This is a species (in contrast to P. acutirostris and P. worcesteri), which actually does appear to be limited to relatively smaller areas of occurrence towards the higher elevation reaches of montane habitats. Although we emphasize that a naturally discontinuous geographical distribution does not constitute a "severely fragmented" range (IUCN 2016), that this species occurs in multiple protected areas, and that additional montane localities are certain to be added to the species geographical range once the mountains of Mindanao are properly surveyed, we follow  and hold downgrading of Philautus poecilius's conservation status in abeyance because available data do suggest that this species is a high elevation specialist, with a distribution limited to original forest. At one high elevation site (Mt. Lumot) where we found this species, we detected high prevalence and levels of chytrid infection in other species of treefrogs; Philautus poecilius, however, was negative for chytrid.

Philautus surrufus Brown & Alcala, 1994
Described originally from Dapitan Peak (10 km from Masawan, Misamis Occidental Province), this species (Fig. 31) is also known from another site in the same province (Mt. Malingdang; Nuñeza et al. 2010), from Mt. Kitanglad (IUCN, 2016) and now from our work on Mt. Lumot (Misamis Oriental Province), constituting a substantial extension of its range, to the east. When Diesmos et al. (2014) downgraded from "Endangered" (EN; IUCN 2016) to "Vulnerable" (VU; IUCN 2016), they found that the species no longer qualified for the higher threat category principally on the basis of its much wider area of occurrence, and also the fact that it appears to tolerate at least some levels of disturbance to its preferred upper montane forest habitat.

Philautus surdus (Peters, 1863)
The presence of Philautus surdus (Fig. 32) on Mindanao is attributed exclusively to specimens from several sites in Agusan Del Norte province. Elsewhere in the Philippines, this species has been recorded from Bohol, Polillo, Catanduanes, and throughout Luzon (Brown and Alcala 1994). Considered "Least Concern" (LC; IUCN 2016) because of its wide distribution across multiple islands and including many protected areas. However, we emphasize that Philippine Philautus are notoriously difficult to identify (because of their similar size, variable appearance, and because most species exhibit color pattern polymorphism and have similar "crunch" mating calls), and we would not be surprised if future molecular data revealed the presence of multiple species masquerading within this oddly widely distributed species.
Sites and specimens:  Brown et al. (1998) resurrected Philautus worcesteri from the synonomy of Platymantis guentheri (where it was placed by Inger [1954]). Once recognized as the distinctive large-bodied Philautus of Mindanao, it was recorded at numerous montane sites throughout the island (Brown and Alcala 1994) and since has been documented at a variety of sites in the island's northeast and southwest. This species (Fig. 33) no longer qualifies for a listing of "Vulnerable" (VU; IUCN 2016) because (1) it is no known from a much greater area of occurrence than when it was originally assessed (IUCN 2004), (2) it is known from many lower elevation areas (originally considered restricted to 800-2,000 m), and (3) because it has recently been recorded at several highly disturbed agro-forest, lower-elevation sites (Zamboanga City area, lower slopes of Mts. Lumot and Hilong-hilong). We therefore classify it as "Near Threatened" (NT; IUCN, 2010) and we urge field workers to focus on its distinctive advertisement call (a loud, "crunch," sounding to the human ear like "Yaak!," repeated two or three times) when conducting future surveys. We suspect that this species has, in some past studies, not been detected because it perches higher and calls less frequently under drier atmospheric conditions. We screened this species for chytrid fungus at multiple sites between 2010 and 2012 and no infection was detected.
Sites and specimens:

Polypedates leucomystax (Gravenhorst, 1829)
Common and distributed throughout the archipelago, Polypedates leucomystax (Fig. 34) is a species that persists well in disturbed habitats, is ubiquitous in agricultural areas,

Kurixalus appendiculatus (Günther, 1858)
Previously classified as a species of Rhacophorus, K. appendiculatus (Fig. 35) is distributed across much of the archipelago's eastern island arc (Mindanao, Dinagat, Samar, Leyte, and northern Luzon), but is conspicuously absent on the intervening Bicol Peninsula (Brown and Alcala 1994). In a recent phylogeographic analysis of samples from throughout this range, Gonzales et al. (2014) suggested a novel island colonization scenario and provided some evidence to suggest that K. appendiculatus in the Philippines may be composed of several independent evolutionary lineage, which eventually may be recognized as separate species. Based on this information and the unresolved taxonomic status of the three major Philippine lineages, Diesmos et al. (2014) recommend classification of Philippine populations of K. appendiculatus as "Data Deficient" (DD; IUCN 2016). The resolution of the status of these populations is an urgent conservation priority for the immediate future. This is a swamp, ephemeral pool, and stagnant water specialist species, which most likely lays its eggs in mud; the tadpoles are unknown.

Rhacophorus bimaculatus (Peters, 1867)
This common tree frog (Fig. 36) inhabits overhanging understory vegetation surrounding rapidly cascading streams in lower-to mid-montane forests. Its distinctive advertisement call is a single brief, high frequency, shrill chirp-and can be heard over the sound of waterfalls (its favored microhabitat). Previously considered uncommon, this species is now appreciated for its very specific microhabitat preference, wherein it can  (Gonzales et al. 2014), and the predictability with which it can be found, now that its habitat is known and can be purposefully surveyed .
Sites and specimens:

Rhacophorus pardalis (Günther, 1858)
This swamp-and ephemeral pond-breeding species (Fig. 37), as presently conceived, is distributed throughout the archipelago (Brown and Alcala 1994;Alcala and Brown 1998;Gaulke 2011). Outside of the country, populations referred to the same species have been reported from Peninsular Thailand, Borneo, and Sumatra (Manthey and Grossman 1997). Patchily distributed, this species is most often encountered perched in vegetation above stagnant pools such as feral pig wallows, and nearly any water-filled in original or even highly disturbed forest (RMB personal observation). Because of its wide distribution, this species is considered "Least Concern" (LC; IUCN 2016).

Family Ichthyophiidae
Ichthyophis mindanaoensis (Taylor, 1960) A single specimen putatively identified as I. mindanaoensis (Fig. 38) has been collected at Barangay Bayugan II, Municipality of San Francisco, Mt. Magdiwata. Collected in dry soil beneath a log several hundred meters from a small stream, this specimen appears to be a juvenile male. The taxonomy and extent of occurrence of I. mindanaoensis is somewhat uncertain. Other specimens have been reported from Bukidnon (central Mindanao), Mt. Malindang (western Mindanao; most likely misidentified exemplars of I. glandulosus, a close relative known from Basilan Island and the southern Zamboanga Peninsula), Davao City Province (eastern Mindanao) and South Cotabato Province (southern Mindanao; Taylor 1920aTaylor , 1960Taylor , 1965Inger 1954. A much-needed effort to reexamine and reconsider the range of variation exhibited in traditional morphologically diagnostic characters (Taylor 1960(Taylor , 1965) would now be possible, given the accumulation of specimens from throughout the island. Given the current uncertainty, however, concerning its distribution taxonomic status, this species is considered by the IUCN to be "Data Deficient" (DD; IUCN 2016).
This species (Fig. 39) was found asleep at night in shrubs and saplings 2-4 meters above the ground in secondary forest and along forest edges in agricultural areas. Individuals were especially common at site MO 6 at the edge of primary forest abutting a river. This species has a widespread distribution that includes Northeast Mindanao, Camiguin Sur, Dinagat and Siargao islands. At present, with the taxonomy of this group confused and unassessed using molecular data (Hallermann 2005), the conservation status of "B. cristatella" remains unassessed (IUCN 2016). If this taxon is as widespread as currently conceived (Hallermann 2005), its extremely broad distribu- tion throughout the Philippines and surrounding Sundaic landmasses, plus its constant presence in a wide variety of forested and unforested habitats, would qualify it for "Least Concern" (LC) using IUCN conservation status assessment criteria (IUCN 2010). We recommend treating this species as "Data Deficient" (DD; IUCN 2016) until a formal taxonomic review using both morphological and genetic data, can be performed. We suspect Philippine populations are not conspecific with the lineage at the type locality (Java Island, Indonesia); although Taylor named Bronchocela marmorata from northern Luzon, that name most likely would not apply to the distinctive population on the Mindanao PAIC (see also : Grismer et al. , 2016.

Draco bimaculatus (Günther, 1864)
This species occurs throughout the Mindanao and Sulu faunal regions (McGuire and Alcala 2000). We encountered D. bimaculatus (Fig. 40) in both primary and secondary forests as well as at the edges of coconut plantations immediately adjacent to forest; our

Draco cyanopterus Peters, 1867
This species was quite common in coconut plantations and, like Draco bimaculatus, was found far from forest in a few instances. This species is classified by the IUCN as "Least Concern" (LC; IUCN 2016) and its population size may be increasing due to the expansion of Mindanao's ubiquitous coastal coconut plantations. Sites

Draco mindanensis Stejneger, 1908
This species (Fig. 41) is rarely encountered in the wild and the extent of its occurrence is poorly known; it has been documented from Mindanao, Leyte, Samar, Dinagat and Siargao isands (Taylor 1922a;Ross and Lazell 1990;Smith 1993a;McGuire and Alcala 2000;Realubit et al. 2015). Specimens have only been observed in primary and mature secondary growth forest habitats, active during the day on trunks, usually quite high above the ground (5-10 m above the ground; RMB personal observation). A true forest obligate, this phylogenetically distinct (McGuire and Kiew 2001) species qualifies for formal recognition as a "Vulnerable" (VU; IUCN 2010, 2016) conservation status taxon. It is noteworthy that biologists have always considered this species to be rare (for review, see McGuire and Alcala 2000) and almost a century ago, following several years of near-continuous fieldwork in heavily forested central Mindanao, Taylor (1922a) had only collected a few specimens.

Gonocephalus cf. interruptus
Gonocephalus interruptus is the only species of Gonocephalus in the Philippines with a type specimen bearing specific locality data indicating that it was collected on Mindanao Island (although where on Mindanao is unclear). Precise type locality data are unavailable for G. semperi and G. sophiae, species originally reported only to have been originally collected from "The Philippines." Although recent taxonomists and biogeographers have referred to specimens from Mindoro, Mindanao, and Bohol to G. semperi and others from Luzon, Mindanao and the western Visayas (Negros, Panay, Masbate) to G. sophiae, both species' original descriptions were not accompanied by specific locality data. Thus, detailed comparisons of the name-bearing type specimens to fresh material from known localities will be necessary to definitively determine the proper application of these names to Philippine populations. Based on the crest morphology of our specimens and comparison to original illustrations (Boulenger 1885;Taylor 1922aTaylor , 1923 and their known provenance (Mindanao), we treat this taxon as most likely representative of G. interruptus (Fig. 43). We concede that additional phylogeny-based taxonomic review will be required to confirm this designation and properly assign this name to the phenotypically most similar population, following examination of the name-bearing type (Boulenger 1885

Hydrosaurus pustulatus Eschsholtz, 1829
An inhabitant of lowland riparian corridors, coastal forests, and mangroves (Smith 1993a, we encountered H. pustulatus (Fig. 44) along rivers at lower elevation sites ranging from 170-420 m above sea level. Specimens were collected at night while sleeping in rock crevices as well as during the day when this species is active. Due to the rapid and continuing decline and fragmentation of the habitat upon which it depends, this species has been classified as having a "Vulnerable" conservation status (VU; IUCN 2016). A recent phylogeographic study supports the monophyly of Philip-  pine H. pustulatus, demonstrates that H. amboinensis does not occur in the Philippines, and indicates that there are six well structured haplotype groups across the archipelago, although given the data used in the study, not all haplotype groups corresponded with unique geographic ranges . Specimens from Camiguin Sur and Dinagat island populations were recovered in several of these clades. Genetic samples from Northeast Mindanao were not available at the time of this study, and so additional research will be necessary to determine the relationship of the Northeast Mindanao populations to other populations of H. pustulatus in the Philippines, and whether or not they constitute their own Evolutionarily Significant Unit (ESU) for conservation.
Sites and specimens:

Dibamus cf. leucurus Taylor, 1915
We collected one specimen (Fig. 45) at site MO 6 on Mount Lumot; this individual matches Taylor's description of the holotype of D. argenteus, which he collected from Butuan, Agusan Del Norte, Mindanao Island. We suspect that the southern Philippine population currently referred to P. leucurus is diagnosable as a distinct species, but this will need to be verified with a comparison to specimens from Sumatra, Indonesia (the type locality for D. leucurus). To our knowledge our new specimen is now the only representative of the genus from Northeast Mindanao, as Taylor's type specimen was destroyed during WWII (Brown and Alcala 1980). Other documented localities include Basilan Island and Camiguin Sur (CAS specimens). Our specimen was found beneath a pile of discarded coconut husks within a coconut plantation, mixed with secondary growth forest. So little is known about these secretive lizards that we are unable to comment on their conservation status and regard them as "Data Deficient" (DD; IUCN 2016).

Family Gekkonidae
Cyrtodactylus agusanensis (Taylor, 1915) Formerly part of a species complex known from Mindanao, Leyte, Dinagat and Siargao islands (Taylor 1922a, Brown and Alcala 1978, Ross and Lazell 1990, the distri- bution of this species (Fig. 46) has since been restricted to Eastern Mindanao following taxonomic revision (Welton et al. 2010). Consistent with observations reported by Brown and Alcala (1978), Smith (1993a), and Welton et al. (2010), we collected this species among boulders on the banks of streams. We also found them to be common on vegetation adjacent to streams as well as albeit less common on vegetation well away from water. One individual was collected from inside a cave at site 12. This species is currently listed as "Least Concern" (LC; IUCN 2016).
Sites and specimens:  (Taylor, 1915) Occurring in sympatry with C. agusanensis, C. annulatus (Fig. 47) can be identified by its smaller size and lower non-overlapping range of paravertebral scales (Brown and Alcala 1978;Welton et al. 2009). Cyrtodactylus annulatus has been reported from a number of localities across the archipelago including Camiguin Sur Island and in the northeastern regions of Mindanao Island. We frequently encountered these geckos on

Cyrtodactylus mamanwa Welton, Siler, Linkem, Diesmos & Brown, 2010
Formerly recognized as part of the C. agusanensis Complex, the Dinagat Island clade is now referred to as a unique Philippine endemic species, C. mamanwa (Fig. 48), as a result of an analysis of molecular and morphological evidence and a comprehensive taxonomic review (Welton et al. 2010). This species was collected among large boulders and logs on the bank of a stream at night. Future surveys may reveal additional populations on Siargao Island. This species does not qualify for any elevated threat categories and we classify it as "Least Concern" (LC; IUCN 2010).

Gehyra mutilata (Wiegmann, 1834)
This species of house gecko is quite common throughout the Philippines, and regularly encountered in residential habitats (in poorly lighted areas, in contrast to most house geckos such as Hemidactylus platyurus and H. frenatus, which are most frequently encountered below lights) mixed and disturbed forest edge habitats. To date, no phylogenetic studies have focused on understanding population genetic structure or patterns among the widespread species of house geckos in the Philippines (G. mutilata, Hemidactylus platyurus, or H. frenatus). Owing to its widespread distribution, this species does not qualify for any elevated threat categories and is classified as "Least Concern" (LC; IUCN 2010). Sites

Gekko cf. mindorensis
A recent phylogeographic (Siler et al. 2012c investigation identified eight genetically divergent and geographically structured clades within this Philippine endemic species, confirming previous speculation (Ferner et al. 2001) that many Philippine populations constitute a cryptic species complex. In a follow-up study, Siler et al. (2014b) recognized G. kikuchii as a member of the G. mindorensis Complex, with a distribution that included populations from the islands of Luzon (northern Philippines) and Lanyu (Taiwan). Because these changes render G. mindorensis paraphyletic, Siler et al. (2014b) suggest the recognition of G. mindorensis as a monophyletic clade that includes populations from the Mindoro PAIC, the Bicol Peninsula and Catanduanes Island of the Luzon PAIC, the Visayan PAIC, and Bohol and Camiguin Sur islands of the Mindanao PAIC. Following these studies, we recognize populations of Camiguin Sur (Fig. 49) and Panglao as G. cf. mindorensis in anticipation of taxonomic resolution of this complex and diverse group of Philippine endemic geckos. Gekko mindorensis is classified as "Least Concern" (LC; IUCN 2016) but should now be considered "Data Deficient" (DD; IUCN 2010) until a taxonomic assessment of species diversity can be performed.

Gekko gecko (Linnaeus, 1758)
This species occurs throughout Southeast Asia and has been recorded at most well surveyed sites throughout the Philippines with the exception of the Batanes and Babuyan Island Groups Alcala 1970, 1978;Siler et al. 2012). We frequently observed this species on manmade structures, in well-lighted areas (Brown et al. 2012a or in nearby trees such as plantations. Its conservation status is "Least Concern" (LC; IUCN 2016). Sites and specimens: C 12: KU 304145-46; C 15: KU 309880-82; KU 310360; MO 3: KU 334015-16.

Gekko monarchus (Shlegel, 1836)
Gekko monarchus although reported from a few localities in the oceanic portions of the Philippines (Taylor 1922a;Brown and Alcala 1978) has most frequently and convincingly been reported from Palawan. Recent molecular analyses of Gekko samples from throughout the country (Siler et al. 2012 have failed to find true G. monarchus at any Philippine localities beyond Palawan, leading us to consider that the species may be extremely rare on the Mindanao PAIC or may be attributed to the oceanic (non-Palawan) portion of the archipelago in error.

Hemidactylus platyurus (Schneider, 1792)
Like its congener, H. platyurus is understudied and often overlooked in faunal inventories because it is a constant presence in and around human habitations. Although it may warrant its current IUCN conservation status ("Least Concern"; IUCN 2016), this categorization remains to be critically evaluated.

Hemiphyllodactylus cf. typus Bleeker, 1860
In response to Zug's (2010) consideration of Hemiphyllodactylus typus as a widespread, "low diversity" and invariant taxon, Grismer et al. (2013) showed deep phylogenetic structure within H. "typus" suggesting the existence of many distinct species, several of which subsequently have been named. The taxonomic status of the Mindanao population(s) have not yet been studied, so we refer to them as H. cf. typus in hopes that a thorough review of their status (at multiple sites throughout the island) will soon be undertaken. The conservation status of H. typus formally is "Least Concern" (LC; IUCN 2016) although pending taxonomic revision may show this species to be much more range restricted. Similarly, Philippine populations most likely will be partitioned into a number of species (Grismer et al. 2013) suggesting that like G. mindorensis, all Hemiphyllodactylus populations in the Philippines should now be considered "Data Deficient" (DD; IUCN 2010) until taxonomic reassessment in available.

Lepidodactylus aureolineatus Taylor, 1915
This Mindanao PAIC endemic is a species that historically has been heavily collected (possibly indicating its stable and widespread population status; Brown and Alcala 1978) but which has been conspicuously absent in recent surveys. Considered "Least Concern" (LC; IUCN 2016), this species may be common at low elevations and thus, was frequently encountered in late 1900s surveys, but absent in recent studies targeting montane habitats. A focused investigation of its distribution and taxonomic status would be highly desired.

Lepidodactylus labialis (Peters, 1864)
This previously considered common and "Least Concern" (LC; IUCN 2016) species has not been recorded in any of our recent surveys, even those focused near the type locality (Mt. Hilong-hilong. Siler et al. (2014c) demonstrated that this species had previously been incorrectly placed in the genus Pseudogekko (Brown and Alcala 1978), and actually possesses diagnostic characters that ally it with Lepidodactylus. This species has not been observed since its original collection and should now be designated "Data Deficient" (DD; IUCN 2010) until an assessment in the form of a field survey at the type locality can be undertaken. It is possible that Lepidodactylus labialis could be a range-restricted microendemic species and a major conservation concern.
Sites and specimens:

Pseudogekko pungkaypinit Siler et al., 2014
This rare forest-obligate species (Fig. 50) is a member of an endemic genus of geckos in the Philippines, and formerly recognized as a member of the P. compresicorpus Complex Alcala 1978, Siler et al. 2014c, d). Recently, the species complex was revised using genetic and morphological data, restricting the range of true P. compresicorpus to the Luzon PAIC, and recognizing three new species within the species complex (Siler et al. 2014c,d). The longer-bodied species found in central and northeastern Mindanao is now recognized as P. pungkaypinit, and at this time does not qualify for elevated or threatened conservation status (LC; IUCN 2016).

Brachymeles tiboliorum Siler, Jones, Diesmos, Diesmos & Brown, 2012
One juvenile of undetermined sex (KU 326109) collected in Initao National Park, Barangay Initao, Municipality of Tubigan, Misamis Oriental Province in May 2001 by ACD. This specimen was tentatively identified as B. tiboliorum during a recent review of the B. gracilis complex . Brachymeles tiboliorum is otherwise restricted to southern Mindanao, which makes this single specimen record somewhat tentative. It is conceivable that, with the collection of additional specimens from northern Mindanao, this population may be recognized in future studies as a distinct species. Although this species has not yet been assessed (IUCN 2016) it is quite clear that it should be regarded as "Data Deficient" (DD; IUCN 2016) Sites and specimens: MO: KU 326109.

Brachymeles hilong Brown & Rabor, 1967
Like Brachymeles orientalis, this species (Fig. 51) of slender skink is quite common in Caraga Region of northeast Mindanao; it formerly was classified as a subspecies of the "widespread" Brachymeles gracilis group (Brown and Alcala 1980) (Smith 1993a;Siler et al. 2012b). Individuals are often found in residential, disturbed, and secondarygrowth forest at low elevations. They are commonly observed within rotting logs, in leaf litter, in the root networks of tree buttresses, as well as beneath rotting piles of coconuts . The species has been found to qualify for a conservation status of "Near Threatened" (NT; IUCN 2016; Siler et al. 2012a,b

Brachymeles orientalis Brown & Rabor, 1967
This common species of semi-fossorial slender skink (Fig. 52) can be found throughout the eastern and northern islands of the Mindanao PAIC, as well as Bohol, Leyte, and  Camiguin Sur islands (Brown and Alcala 1980;Smith 1993a;Siler et al. 2011b) and Samar Island (specimens in KU). The species can often be found in sympatry with B. hilong on northeastern Mindanao, B. samad on Leyte and Samar islands, and B. vulcani on Camiguin Sur Island . Individuals are often found in residential, disturbed, and secondary-growth forest at low elevations. They are commonly observed in soil and forest floor detritus around tree buttresses and under nearly any form of ground cover in coconut palm plantations adjacent to forest . Because of its wide geographical distribution and tolerance of disturbance, this species does not qualify for any elevated threat categories and is classified here as "Least

Eutropis multicarinata (Gray, 1845)
The southern Philippine common sunskink traditionally known as Eutropis multicarinata multicarinata (Fig. 53) has been documented on most of the islands of the Mindanao PAIC (Brown and Alcala 1980;Smith 1993a). Although the type locality of this species was designated simply "The Philippines," Barley et al. (2013) matched the type material phenotype to populations from Mindanao, Leyte, and Dinagat islands. Two additional lineages in the large E. multicarinata Group clade were identified in the southern Philippines: one from southwest Mindanao ("Clade A;" Barley et al. 2013) and another from Mindanao, Bohol, Dinagat and Siargao ("Clade E;" Barley et al. 2013. Although the type locality of this species was designated simply AS 2: USNM 2293 of associated genetic samples in older collections) under the name Eutropis multicarinata pending a taxonomic revision that will be published elsewhere (Barley et al. unpublished data). Given the taxonomic uncertainty and morphologically cryptic nature of these populations, this taxon must be considered "Data Deficient" (DD; IUCN 2016).
Sites Eutropis cf. multicarinata Barley et al. (2013) demonstrated the genetic distinctiveness of some individuals of "Eutropis multicarinata" at selected populations on Mindanao, Bohol, and Dinagat (and possibly and Siargao) islands ("Clade E;" Barley et al. 2013 (Brown & Alcala, 1980) This relatively common Philippine sunskink has been reported as occurring throughout the southern portion of the archipelago (Brown and Alcala 1980), including Palawan and even northern Borneo. Some populations from the northern portion of the archipelago have been described as a distinct, but closely related species, E. cumingi. When naming E. indeprensa, Brown and Alcala (1980) restricted the type locality to Mindoro, which made clear Barley et al.'s (2013) assignment of the name-bearing types to a definite population on Mindoro (Barley at al. 2013). Brown and Alcala (1980) reported the species from Mindanao, Leyte, Samar, "Camiguin" (presumably Camiguin Sur Island), Palawan, Negros and Cebu, but Barley et al. (2013), using a multilocus molecular analysis showed that these populations do not form a monophyletic clade, and that this species is restricted to Mindoro Island and Borneo-a highly unusual and disjunct distribution if true. Barley et al.'s (2013) highly divergent "Clade C" (which is not closely related to true, type-locality E. indeprensa) contained samples from Dinagat, Mindanao, Samar, Panay, Cebu, and the Bicol Peninsula of Luzon (see Barley et al. 2013: Figure 3) and we find it likely that these eventually will be recognized as a distinct species. Eutropis indeprensa species has not been assessed for conservation status but may qualify for an elevated level of threat should additional studies confirm a Mindoro-only distribution in the Philippines (IUCN 2010); for now it is best considered "Data Deficient" (DD; IUCN 2010).

Eutropis multifasciata (Kuhl, 1820)
This geographically widespread habitat generalist species (Brown and Alcala 1980;Grismer 2011) has been recorded and observed throughout the Philippines, most of the western landmasses of the Indo-Australian archipelago, southwest Asian and north-ward into Indochina. Related to E. grandis (syn. E. macropthalma) and E. rudis , this species (Fig. 54) clearly possesses the dispersal capacity to overcome marine barriers to colonization and yet, at a much finer scale, has been shown to possess the genomic signature of habitat fragmentation in the Philippines (Barley et al. 2015). A truly widespread species, E. multifasciata does not qualify for any elevated level of threat and must be considered "Least Concern" (DD; IUCN 2016). Sites

Lamprolepis smaragdina philippinica (Mertens, 1928)
This species is widely distributed throughout the archipelago and locally abundant in coastal and agricultural areas (Brown and Alcala 1980;Smith 1993a). We typically encountered L. smaragdina (Fig. 55) on coconut trees in plantations, as well as developed areas in close proximity to agricultural areas. Due to its tolerance for human disturbance and its ubiquitous distribution, this species is considered "Least Concern" (LC; IUCN 2016). A recent phylogeographic study  identified the presence of numerous highly divergent evolutionary lineages in the Philippines, suggesting that reassessment of conservation status should be attempted once taxonomic reconsideration of this group has been undertaken. Sites

Lipinia auriculata herrei (Taylor, 1922)
Taylor (1917b) described the species Siaphos (currently Lipinia) auriculata on the basis of a few specimens from northern Negros and he later described L. kempi from Mindoro (Taylor 1919) and L. herrei from Polillo Island (Taylor 1922a). At the time of their review of Philippine scincid lizards, Brown and Alcala (1980) remarked on the close phenotypic similarity of the original specimens, and therefore considered them to be closely related and recognizable only at the level of subspecies (Brown and Alcala 1980). Interestingly, although few rare forest Philippine reptile taxa had been recorded to possess wide ranges encompassing the Luzon, Mindanao and Mindoro faunal regions (Taylor 1928;Leviton 1963;Brown and Alcala 1970), Brown and Alcala (1980) assigned one specimen from Bohol and two specimens from northeast Mindanao to the subspecies L. a. herrei on the basis of a single character in common: their shared, distinctly paired (i.e., unfused) frontoparietal scales. Remarking on this oddly disjunct distribution, the authors did however explicitly state that they anticipated the species would eventually be recorded at interven-  ing localities on the islands of Leyte and Samar (Brown and Alcala 1980:90). Lipinia auriculata has been classified by the IUCN (2016) to be of "Least Concern." Sites and specimens: AN 3: CAS 133700, CAS 133778.

Lipinia pulchella pulchella Gray, 1845
Gray (1845) described Lipinia pulchella (the type species for the genus; Fig. 56) on the basis of a single specimen from Mindanao Island; Taylor (1917b) subsequently confirmed the species on Negros Island (= L. p. taylori Brown and Alcala 1956) and Brown and Alcala (1980)

Lipinia quadrivittata (Peters, 1867)
Peters (1867) described "Lygosoma" (=Lipinia) quadrivittata from three Mindanao Island specimens, and the species subsequently has been reported from Bohol, Leyte, Negros, Palawan, Siquijor, Cebu and Camiguin Sur islands (Taylor 1922a;Brown and Alcala 1980). The Mindanao specimens with habitat data were collected from under the moss on tree trunks at 800+ m above sea level and one was collected from an arboreal fern (Brown and Alcala 1980). This species has not been assessed for conservation status (IUCN 2016) and we consider it "Data Deficient" (DD; IUCN 2016) given the absence of any other information on its distribution and status. Although L. quadrivittata appears to have a wide distribution on numerous Philippine islands, we hesitate to classify it as "Least Concern" on the arbitrary basis of distribution alone. On Negros, Brown and Alcala (1980) noted that this species was predictably observed on trees in swamps. Given that swamp ecosystems are severely imperiled throughout the archipelago, plus the fact that this habitat is quite different than that noted for Mindanao (montae forest), we suspect that more L. quadrivittata, as currently conceived, may be a complex of more than one species. The species' anomalous distribution pattern (three Philippine faunal regions + Borneo) reinforces our suspicion that more than one species is involved. Lipinia quadrivittata should be the subject of a taxonomic review, which should immediately be followed by conservation status assessment.

Otosaurus cumingi Gray, 1845
Otosaurus cumingi, the largest Philippine Sphenomorphus-Group lizard of the family Scincidae is a monotypic taxon, which was recently elevated to the level of a distinct genus (previous recognized as a member of the genus Sphenomorphus; Brown and Alcala 1980) by Linkem et al. (2011). Not closely related to other Philippine Sphenomorphus-Group lizards, this large-bodied taxon is part of an unresolved polytomy involving the genus Otosaurus the Philippine clade, and the Australian clade (Linkem et al. 2011). The single species of Otosaurus (Philippine O. cumingi) is widespread throughout the archipelago, except Palawan (Taylor 1922a;Brown et al. 1980;Brown et al. 2012aBrown et al. , 2013. This widespread species is considered "Least Concern" (LC; IUCN 2010).

Parvoscincus kitangladensis (Brown, 1995)
Described from high elevation forested plateaus of Bukidnon Province, central Mindanao (Brown 1995), this small forest species (Fig. 57) is now also known from eastern and northeastern Mindanao. Parvoscincus kitangladensis is most closely related to the miniature Luzon forest species (e.g., P. lawtoni, and possibly P. luzonensis) but is phenotypically similar to members of the P. decipiens complex and other small bodied taxa from Luzon (i.e. P. laterimaculatus, P. leucospilos; Linkem et al. 2011;Siler et al. 2014e) and even Palawan (e.g., Insulasaurus victoria, I. traanorum; Linkem et al. 2010aLinkem et al. , 2011. Parvoscincus steerei (Stejneger, 1908) Members of the large, geographically structured, genetically diverse taxon Parvoscincus steerei are widespread throughout the oceanic islands of the Philippines (Brown and Alcala 1980) and exhibit some color variation, but very little variability in external morphology (scalation; Linkem et al. 2011;RMB personal observation). Frequently encountered from disturbed second growth and agro-ecosystem matrices, this species extends up to mid-elevation original forests (700-900 m) and is common in leaf litter, under woody forest debris, and in forest detritus surrounding roots and buttresses of trees. This widespread species, as presently conceived, is considered "Least Concern" (LC; IUCN 2010, 2016) but genetic evidence suggest that P. steerei is a species complex composed of numerous evolutionary lineages; these may warrant individual conservation assessment with future investigation.

Pinoyscincus jagori jagori (Peters, 1864)
Unlike the taxonomically confused and highly polyphyletic "species" P. abdictus and P. coxi, P. jagori fall into a single monophyletic clade (Linkem et al. , 2011. Known from eastern Mindanao, Dinagat and Siargao, populations keying out to P. jagori constitute four distinct, genetically divergent lineages (Linkem et al. , 2011: three of these, are identified as P. jagori jagori but nested within them is the population identified as P. jagori grandis (from the western Visayan islands; Brown and Alcala 1980). These large bodied skinks are common throughout disturbed and forested areas from near sea level to 500 or 600 m (RMB personal observation). This common species is considered "Least Concern" (LC; IUCN 2010IUCN , 2016.

Pinoyscincus mindanensis (Taylor, 1915)
This unusual small-bodied species (Fig. 59) was recently placed in the new genus Pinoyscincus (Linkem et al. 2011) together with former members of the Sphenomorphus abdictusjagori-coxi assemblage (Taylor 1922a;Brown and Alcala 1980), in recognition of its close relationship with this large clade including numerous species, named subspecies, and morphologically cryptic but genetically distinct lineages . Unlike most other members of this group, P. mindanensis is a small bodied, montane forest species, whereas other taxa in this genus are large-bodied habitat generalists most frequently encountered at lower elevations (RMB personal observation). The other outlier in this group is P. llanosi, a medium-sized, low elevation riparian habitat specialist most frequently encountered on Samar and Leyte islands (but possibly present in northeast Mindanao) that is unique in that it retreats into running water when disturbed. Previously considered "Near Threatened" (NT; IUCN 2010), this species' conservation status assessment is outdated and in need of reconsideration given the new distributional data presented here. Sites

"Sphenomorphus" acutus (Peters, 1864)
An ecologically unique and morphologically highly distinctive species, "Sphenomorhus acutus" was placed in the large clade of Philippine skinks (genera Pinoyscincus and Parvoscincus), but with low support for its phylogenetic position (Linkem et al. 2011). In recognition of its uncertain phylogenetic affinities and its extremely distinctive morphology, the authors elected to leave this taxon and S. diwata (its sister species) incertae sedis, and thus temporarily remaining in Sphenomorphus, pending further investigation (Linkem et al. 2011). This species has most frequently been encountered in recent years during nocturnal surveys, in which it has been located asleep in suspended coils of dry wild banana and abaca leaves, hanging 1-2 m above the forest floor (in original forest; RMB personal observations). It has been reported active in arboreal substrates (understory tree branches) during the day (Brown and Alcala 1980) and has been observed retreating into running forest streams when pursued by field workers (RMB personal communication with J. Fernandez). Despite this species' enigmatic ecology and natural history, its widespread distribution and generalist habitat characterization render it "Least Concern" (LC; IUCN 2010IUCN , 2016.

Sphenomorphus fasciatus (Gray, 1845)
Unrelated to other Philippine scincids, S. fasciatus (Fig. 60) is a morphologically distinctive species that is most closely related to Solomon Island taxa (e.g., S. cranei, S. solomonis, S. concinatus) and appears to be a single-species lineage that resulted in only one Philippine taxon (Linkem et al. 2011). Found throughout the islands of the Mindanao faunal region, this species is common in low elevation disturbed, second growth, and original coastal forests (Brown and Alcala 1980;Smith 1993a). This common species is considered "Least Concern" (LC; IUCN 2016). Sites

Sphenomorphus variegatus (Peters, 1867)
Unrelated to other Philippine scincid lizards, S. variegatus (Fig. 61) is a member of a clade consisting of non-Philippine species S. maculatus, S. indicus, S. cyanolaemus, S. sabanus, and S. multisquamatus (Linkem et al. 2011). The single Philippine representative of this clade, S. variegatus is a commonly encountered low elevation species wherever mature second growth or first growth forest is present on Mindanao (Taylor 1922a;Brown and Alcala 1980;Smith 1993a). This common widespread species' con-

Tropidophorus misaminius Stejneger, 1908
This Philippine endemic species (Fig. 62) is restricted to Basilan, Camiguin Sur, Dinagat, and Mindanao (type locality: Mt. Malindang, western Mindanao). During the day, we collected specimens on stream banks as well as in the forest where they were under logs or out in the open on the forest floor. At night, we found them sleeping among rocks and logs on stream banks. This species would often flee to the water in the presence of humans, demonstrating a remarkable adaptation for a semiaquatic lifestyle. Our observations of this species were restricted to primary and secondary growth forests, leading us to believe it is not tolerant of degraded habitat. However (Beukema 2006) reports this species as being able to tolerate some degree of habitat disturbance. This species is currently listed as "Least Concern" (LC; IUCN 2016). Sites

Tropidophorus partelloi Stejneger, 1910
Also a Philippine endemic, T. partelloi is restricted to the islands of Mindanao and Siargao (Taylor 1922a;Brown and Alcala 1980;Ross and Lazell 1990) with an apparently wide yet patchy distribution that does not include areas impacted by anthropogenic disturbance (Smith 1993a). At one site (AN 12) we found this species in sympatry with T. misaminius. We collected this species both on the banks of streams and in the forest well away from water. In the forest we collected specimens from under logs or from the forest floor. On the banks of streams this species was found both out in the open as well as asleep among rocks and logs. Similarly to T. misaminius, this species will flee to the water when disturbed. This species is currently listed as "Least Concern" (LC; IUCN 2016).

Varanus cumingi Martin, 1839
This Mindanao faunal region endemic monitor lizard recently was distinguished, at the level of full species, from its closest known relative Varanus samarensis, an endemic species native to Bohol, Samar and Leyte islands, just north of Mindanao proper (Welton et al. , 2014a; these two species apparently diverged as long as 2 million years before present (1.5-2.5; Welton et al. 2014b). Varanus cumingi (Fig.  63) is an extremely common, widespread species and its conservation classification is "Least Concern" (LC; IUCN 2016). Our specimen from Mt. Lumot was salvaged from a villager who had killed the animal as a pest (it had raided his chicken coop on several recent occasions, in search of eggs and newly hatched chicks). We salvaged the decomposing specimen after several days; it has been prepared as an osteological specimen, making it the only known skeleton of this species in existence.

Reptilia: Snakes Family Colubridae
Ahaetulla prasina preoccularis (Taylor, 1912) Ahaetulla prasina is a common, widely distributed snake encountered in a variety of habitats throughout the Philippines (Leviton 1963a(Leviton , 1967. We frequently encounter this species at night, sleeping on branches of bushes and saplings in low elevation, selectively logged forest, primary forest, or even secondary growth or along edges of agricultural areas. We noted exceptionally high densities of this species at low elevation, in riparian habitats on Mt. Hilong-hilong. It is considered "Least Concern" (LC; IUCN 2016) for conservation purposes. Sites and specimens: AN 10-KU 334411-21; AN 11: KU 334407-09; AN 13: KU 334410; D 5: KU 310162; MO 6: KU 334422-30; AN 3: CAS 133398.

Boiga cynodon (Boie, 1827)
Leviton (1970a) described B. cynodon as widely distributed in Southeast Asia and the Philippines, as part of the enormous range of this highly variable species. We collected one specimen on Dinagat Island and assume that it occurs in riparian habitats throughout the Caraga region and all of northeast Mindanao. Although it is classified as "Least Concern" (LC; IUCN 2016), we anticipate that eventual taxonomic studies may find this species to be a complex of evolutionary lineages, which will require individual evaluation in the future.

Calamaria lumbricoidea H. Boie in F. Boie, 1827
Calamaria lumbricoidea is frequently encountered throughout the Mindanao PAIC islands and we collected specimens from sea level to 1,200 m on several mountains of northeast Mindanao. It is widespread in Southeast Asia (type locality: Java). Its color pattern suggests adaptation for prey avoidance via a coral snake mimicry system, and its overall similarity to banded forms of Hemibungarus and Calliophus is striking (Fig.  65). This species is classified as "Least Concern" (LC; IUCN 2016).

Chrysopelea paradisi Boie, 1827
This widespread, non-endangered ("Least Concern;" IUCN 2016) species has been encountered on Dinagat Island several times, but only once in northeast Mindanao proper. This common "flying snake" has been reported to us on numerous occasions in the Caraga region by farmers and workers in coconut palm plantations who witness their directed aerial descent as a frequent locomotor mode (Dudley et al. 2007) during active foraging during the day. This species is infrequently encountered sleeping in vegetation 2-4 meters above the ground (in contrast to species of Dendrelaphis and Ahaetulla), suggesting its preferred retreat is higher in the forest canopy (or coconut A B palms) than biologists had suspected previously (Leviton 1964d). This species is classified as "Least Concern" (LC; IUCN 2016).

Coelognathus erythrurus erythrurus (Duméril, Bibron & Duméril, 1854)
Composed of four currently recognized subspecies, this Philippine rat snake has three distinctive phenotypes in the Philippines, corresponding to named taxa: Coelognathus erythrurus manillensis from Luzon (Brown et al. 1996(Brown et al. , 2000McLeod et al. 2011;Devan-song and Brown 2012), Coelognathus erythrurus psephenoura from the West Visayan, central islands of Negros, Cebu, Panay and possibly the Romblon Island Complex (Leviton 1979;Ferner et al. 2001;Gaulke 2011;Siler et al. 2012a), and Coelognathus erythrurus erythrurus, a form originally described from Java, Indonesia, but also reported on various islands of the Mindanao PAIC (Leviton 1979), including Mindanao, Camiguin Sur, Samar, and here: Dinagat (new island record). This complex, suspected to be composed of multiple evolutionary lineages deserving a specific rank, has not been evaluated by IUCN (2016) for conservation purposes.
Sites and specimens: AS 5: KU 319988; C 1: CAS-SUR 28272; C 14: KU 309924; D 4: KU 305168, KU 305648. Leviton, 1967 Reported first by Taylor (1923), Leviton (1967) noted an apparent east-west hiatus in scalation across central Mindanao and recognized Cyclocorus nuchalis nuchalis (Zamboanga Peninsula's southern most populations, and Basilan Island) for the western form and Cyclocorus nuchalis taylori (Fig. 66) for Mindanao's eastern population. Since being reported by Smith (1993b) and , we have also collected the species recently on Camiguin Sur, and Dinagat islands in addition to populations on Siargao Island (Ross and Lazell 1990), and farther north on Samar and Leyte islands. Considered "Least Concern" (LC; IUCN 2016), primarily due to its wide Philippine distribution, the taxonomic and conservation status of the variable populations will need to be evaluated with a molecular phylogenetic analysis.

Dendrelaphis marenae Vogel & Van Rooijen, 2008
An extremely common, widespread Philippine arboreal snake, D. marenae was collected by us in Agusan, Mindanao and previously has been reported throughout the Philippines (e.g., Leviton 1968; Gaulke 2011;Siler et al. , 2012Brown et al. 2012aBrown et al. , 2013Devan-song and Brown 2012). It has been documented previously in the northeast Mindanao faunal region by Ross and Lazell (1990;Siargao Island) and was recently recognized as a Philippine and Sulawesi form by Vogel and van Rooijen (2008) who split the eastern form (D. marenae: Sulawesi and the Philippines) from the western nominate population (D. pictus: Sunda shelf and Asian mainland). This species has not been assessed for conservation status but its wide distribution would lead is to consider it "Least Concern" (LC; IUCN 2010).

Dendrelaphis philippinensis (Günther, 1879)
Previously considered part of a widespread polytypic species complex (Leviton 1968), referred to Dendrelaphis caudolineatus, the name Dendrelaphis philippinensis recently was resurrected by van Rooijen and Vogel (2012) and applied to southern Philippine populations. The holotype specimen of D. philippinensis was collected from northern Mindanao (Günther 1879), indicating that this application is most likely correct. The biogeographically anomalous distribution of the populations currently referred to D. philippinensis (van Rooijen and Vogel 2012)-including Mindanao PAIC landmasses, but also Cebu Island (the West Visayan faunal region), the Bicol Peninsula and Polillo islands (the Luzon faunal region; )-will require independent verification via a molecular test of these revised species boundaries. This species currently has not been assessed for conservation status, but its wide distribution, if correct, would lead as to consider it "Least Concern" (LC; IUCN 2010). Sites and specimens: AN 11: KU 334483; C 6: CAS-SUR 26328; C 14: KU 310367; D 4: KU 310379.

Lycodon dumerilii (Boulenger, 1893)
This frequently encountered northeast Mindanao faunal region endemic has been reported from Mindanao and Samar (Taylor 1922e;Brown and Alcala 1970), and Leviton restricted the unspecified "Philippines" type locality of Boulenger (1893) to Surigao, northeast Mindanao (Leviton 1965b). Lanza (1999) described an apparently related species (L. ferroni) from Samar, and the phylogenetic distinctiveness of the various, somewhat variable (RMB personal observations). Northeast Mindanao faunal region populations (Fig. 69) has not yet been examined in detail ). This species is ranked "Least Concern" (LC; IUCN 2016).

Stegonotus muelleri Duméril, Bibron & Duméril, 1854
Presumably because of the common species epithet, this species has been erroneously confused in taxonomic literature with Lycodon muelleri a species from Luzon, Polillo, and presumably Catanduanes and Marinduque islands . Stegonotus muelleri (Fig. 71), in fact is a large (1.5-2.0 m body length) rat snake that is unique among Mindanao PAIC herpetofauna in that it is a true Papuan biogeographical element, akin to the frog genus Oreophryne and extends no farther into the Philippines  than the Mindanao faunal region (Inger 1954(Inger , 1967(Inger , 1999Leviton 1963a;Brown and Alcala 1970). We have collected this common species throughout northeast Mindanao and Dinagat islands despite the fact that a decade ago, it qualified as "Near Threatened" (NT;IUCN 2010IUCN , 2016; the status of this common widespread species needs to be reconsidered now that there are new distributional data are available. We consider the species to be "Least Concern" (LC; IUCN 2010) given all of our data presented here and those not yet published from Leyte and Samar islands. Sites and specimens: AN 11: KU 334773-74; AS 5: KU 320001-04; D 5: KU 310178; MO 6: KU 334775.

Rhabdophis auriculata auriculata (Günther, 1858)
A widespread and exceptionally abundant species Rhabdophis auriculata (Fig. 72) is another species with taxonomic differentiation across central Mindanao (Günther 1858;Taylor 1922e;Leviton 1963a). Leviton (1970b)   Mindanao have noted this species (e.g. Smith 1993b; , and we have collected it on the forest floor and riparian habitats in disturbed areas at numerous sites on Mindanao and Dinagat (Ross and Lazell 1990). This species is not considered threatened (LC; IUCN 2016).

Family Elapidae
Calliophis philippina (Günther, 1864) This species (Fig. 75) is widespread in the Mindanao PAIC islands of Mindanao, Samar and Leyte (Leviton 1963b;Leviton et al. 2014) and is known from Agusan del Norte and Misamis Occidental provinces in the northeast, as well as Zamboanga Province in the west, and Davao and Lanao del Sur provinces in the south. Previously considered a subspecies of the widespread polytypic Calliophis intestinalis (which is considered "Least Concern" by IUCN [2016]) the revised taxonomy, considering Mindanao PAIC populations as a separate species has not yet been assessed for conservation purposes. Nevertheless, we consider this to be a reasonably common species in areas with any kind of forest cover and we have encountered specimens in many of our recent surveys in Agusan Del Norte and Agusan Del Sur provinces (Mindanao Island), and Camiguin Sur Island, as well as on recent surveys in extreme western Mindanao. Based on its wide distribution and lack of any specific threats, we consider this species to be "Least Concern" (IUCN 2010).
Sites and specimens:

Calliophis sp.
We collected single specimen of a new species of elapid snake, tentatively assigned to the genus Calliophis, at the Barangay Santiago, Sitio Cambinlia (Sudlon), Municipality of Loreto, Dinagat Island. The specimen is so distinct as to initially defy identification to genus (Leviton et al. 2014). Possibly more closely related to the red-headed blue coral snake (Calliophis bivirgata) than to the sympatric and common striped coral snake Calliophis philippina (Eric Smith, personal communication), the new species will formally be named to science in another publication. A previous molecular phylogenetic analysis of Philippine "coral snakes" of the genus Hemibungarus determined those Philippine elapids were unrelated to true coral snakes and, rather, more closely related to cobras (Castoe et al. 2007). A phylogenetic analysis of DNA sequence data will likewise be necessary to determine the phylogenetic affinities of this unusual new species (E. Smith and RMB unpublished data). Without any information on the distribution of this new species, we hold conservation status assessment in abeyance until more data are available.

Naja samarensis Peters, 1861
This venomous cobra species is known throughout Mindanao, Bohol, Samar, Leyte, Dinagat, and Camiguin Sur islands (Taylor 1922e;Leviton 1963aLeviton , 1964bWüster and Thorpe 1990;Smith 1993b;Leviton et al. 2014). Our specimen from Barangay Pandan, Municipality of Mambajao (Camiguin Sur Island) was active during the day on the edge forest and coconut palm plantations. Like all recognizably venomous snakes, Samar cobras are heavily persecuted by humans whenever the two species come in contact; however, N. samarensis is known from a number of protected areas, and clearly can tolerate anthropogenic modification of its habitat; it is considered "Least Concern" (LC; IUCN 2016) despite the fact that all cobras are included in Appendix II of CITES (IUCN 2016) due to their commercial harvest for skin (leather) and traditional medicine trades. Sites and specimens: AN 7: CAS 15340; C 14: KU 309923; C 7: CAS-SUR 26322; D 1: USNM 229324. Leviton et al. (2014) list multiple sea snakes form the coastal areas of Mindanao, including Hydrophis atricepts and Laticauda laticaudata. We were able to locate credible, identified specimens of only a single species from northeast Mindanao, although we do not discount sight records of earlier workers (Taylor 1922d;Leviton 1963;Brown and Alcala 1970).

Hydrophis platyurus (Linnaeus, 1766)
This extremely widely distributed sea snake is most likely found throughout the Philippines (Leviton et al. 2014), and outside of the country as far as Hawaii and the western coastal United States. The two specimens from Surigao Del Norte are hemipene preparations, taken by J. C. Thompson (a U. S. Marines office, stationed in the Philippines in the early 1900s) from voucher specimens deposited then in the University of Santo Tomas (Manila) collection. No information is available regarding the condition of these voucher specimens in the UST collection, and we have not been granted access to this historically important Manila-based collection (Taylor 1920a(Taylor ,b, 1922aBrown and Alcala 1978). This species has been assessed by the IUCN as "Least Concern" with populations stable (IUCN 2016). The status of sea snakes in the waters surrounding Mindanao has been very poorly studied and almost nothing is known of species diversity in the southwest Philippines (Taylor 1922e;Uetz and Hošek 2015).

Psammodynastes pulverulentus (Boie, 1827)
This non-endemic, very widespread species has not been assessed by IUCN (2010IUCN ( , 2016 but clearly is not threatened with extinction due to its widespread distribution across many Southeast Asian countries and landmasses, along with its obvious tolerance of disturbance. We frequently find adults on the ground foraging, and juveniles in the branches (sleeping at night) of very small saplings. This species is not considered endangered at any elevated threat level (LC; IUCN 2016). Sites

Aplopeltura boa (Boie, 1828)
This widely distributed slug-eating snake (Fig. 77) is most frequently encountered in the vicinity of rivers and streams where it feeds exclusively on arboreal slugs and snails that are abundant following rain. It is considered "Least Concern" by IUCN (2016) due chiefly to its wide geographical distribution across many Southeast Asian landmasses.

Malayopython reticulatus (Schneider, 1801)
Caraga region reticulated pythons are quite common in various habitats, including agricultural plantations, and even residential areas-localities where humans have subsidized their prey base by inadvertently increasing rodent populations. We received numerous reports of pythons from residents around disturbed forests patches in the foothills of Mt. Hilong-hilong, Mt. Balatukan, Mt. Magdiwata, and Mt. Lumot. Hunted for meat (Gaulke 1998), M. reticulatus is also exploited in other parts of its range for leather. The conservation status of natural populations of Philippine pythons is unclear; this species has not been assessed by the IUCN (2016), but it most likely will not require additional measures of protection unless it becomes commercially exploited in the area. We collected one specimen in a coconut palm plantation at night, at the base (170 m) of Mt. Hilong-hilong.

Ramphotyphlops cf. cumingii (Gray, 1845)
Owing to the near complete lack of information on the distribution of Ramphotyphlops cumingii (Gray 1845;Taylor 1922e;McDowell 1974;McDiarmid et al. 1999), plus wholesale taxonomic confusion surrounding species boundaries in these secretive taxa (McDiarmid et al. 1999;Wynn et al. 2015), Ramphotyphlops cumingi is considered "Data Deficient," as are most species of Philippine typhlopid snakes. Imperfectly matching published accounts of the diagnostic scalation in R. cumingii, our specimen from low elevation foothills of Mt. Hilong-hilong (coconut palm plantations) may represent yet another undescribed form.

Malayotyphlops sp.
Collected in 1971 above 700 m on the west side of Mt. Hilong-hilong on approach from the Municipality of Cabadbaran, CAS 133604 is the only known specimen of an undescribed species. Despite our return to the general area in 2010 and 2012, no additional specimens have been secured. In previous assessments, this specimen has been identified as "Typhlops luzonensis" Taylor  Trimeresurus flavomaculatus (Fig. 78) is an exceedingly common pit viper, encountered in a variety of habitats on Luzon, Catanduanes, and Polillo (Taylor 1919;McLeod et al. 2011;Siler et al. 2011a;Brown et al. 2012aBrown et al. , 2013bDevan-song and Brown 2012). On Mindanao PAIC islands, Trimeresurus cf. flavomaculatus is widespread and but somewhat common (Leviton 1964a;Leviton et al. 2014), prone to be encountered in mature forest, and with a distinctive color pattern. It is entirely possible that these observations may be simple inter-populational differences not indicative of specieslevel diagnostic characteristics. This certainly has been the case for Trimeresurus flavomaculatus halieus, a subspecies (sensu Leviton 1964a) originally described from Polillo Island (Taylor 1919) but which has been shown to be a mere color pattern variant, frequently found on Luzon (Brown et al. 2000Devan-song and Brown 2012;Leviton et al. 2014). On the other hand, previously recognized subspecies Trimeresurus flavomaculatus mcgregori (now recognized as T. mcgregori, a Batanes-Babuyan archipelago endemic; Leviton et al. 2014), and Trimeresurus flavomaculatus schultzei (a Palawan endemic) have been elevated to the level of full species (Brown et al. 2000;Gumprecht et al. 2004;Oliveros et al. 2011;Leviton et al. 2014), leading us to consider a molecular test of species potential species boundaries in Luzon PAIC versus Mindanao PAIC Trimeresurus flavomaculatus populations highly desirable. Given its  C be separate evolutionary lineages. Fischer (1885) described. Trimeresurus schadenbergi from Mindanao (currently a synonym of T. flavomaculatus). This would seem to be the available name for the Mindanao faunal region lineage, should it be deemed worthy of specific rank. According to David and Ineich (1999), Leviton's (1964a) restriction of the type locality of T. flavomaculatus to Luzon may not be valid, thus necessitating that a full reconsideration of phenotypic variation in name-bearing types (or literature descriptions, given that some type specimens were destroyed in WWII; Brown and Alcala 1978;Uetz and Hosek 2015) be undertaken in coordination with any effort to distinguish these putative taxa using molecular data. Sites and specimens: AN 11: KU 334432-33; AN 12: KU 334434-35; AS 5: KU 320006-07; D 5: KU 310378; MO 2: KU 334436.

Tropidolaemus subannulatus (Gray, 1842)
This common and widespread Luzon and West Visayan pit viper (Fig. 79) is frequently encountered in coastal to mid-montane elevation forested areas (Taylor 1919Leviton 1964a;Brown et al. 2012aBrown et al. , 2013aSiler et al. 2011a;Devan-song and Brown 2012). Vogel et al. (2007) revised the Tropidolaemus wagleri complex and, in restricting T. wagleri to non-Philippine landmasses, resurrected Gray's (1842) T. subannulatus for Philippine populations on numerous islands, including northeast Mindanao populations. The authors noted the "highly problematic" nature of these populations, with respect to their diagnosis from conspecific T. philippensis (below) and we concur: although some specimens clearly meet the definition of Gray's T. subannulatus, others from the same or nearby populations are definitively T. philippensis. The status of these two possibly distinct and sympatric lineages needs to be addressed with molecular data and a full appraisal of variation in the name-bearing type specimens (Vogel et al. 2007). This species is considered "Least Concern" (LC; IUCN 2016).

Tropidolaemus philippensis (Gray, 1842)
As discussed above, Vogel et al. (2007) recently resurrected Gray's (1842) Tropidolaemus philippensis (Fig. 80); both taxa are exemplified by holotypes bearing locality data no more specific than "The Philippines." Although Taylor (1919 recognized them both, Leviton (1964a) placed both in the synonymy of T. wagleri (Vogel et al. 2007;Leviton et al. 2014). That action, no longer tenable, has been revised to recognize all three species (and others) with the unresolved problem of a lack of clarifica- tion between T. subannulatus and T. philippensis (Vogel et al. 2007). With reference to clearly T. philippensis phenotypes from Samar and Leyte (e.g., KU 338863, 311290, 326317, from Leyte Island), we do not hesitate to identify a subset of our available specimens to T. philippensis. This species' conservation status has not been assessed.

Cuora amboinensis (Riche in Daudin, 1802)
This common geomydid freshwater turtle species (Fig. 81) has been documented by Diesmos et al. (2008) throughout the country. It is considered widely distributed and composed of two poorly differentiated subspecies, which may or may not be valid (the nominate form, C. a. amboinensis is reported to occur throughout the eastern Philippines, including northeast Mindanao). Our records all originate from Dinagat Island where it is common on forest edges and agricultural areas and streams in secondary, disturbed forest. Although the taxonomy of this group is unresolved and despite its  wide distribution throughout the Philippines, and adjacent landmasses of the western Australasian archipelago and adjacent mainland, this species is considered "Vulnerable" to extinction (VU; IUCN 2016) in recognition of the fact that freshwater turtles are so heavily exploited throughout Asia (Lehr and Holloway 2003;Schoppe 2008). Sites and specimens: D 2: KU 305165, KU 305167, KU 305557-58; D 5: KU 310179.

Crocodylus porosus Schneider, 1801
This species, a saltwater and estuarine crocodile, is found distributed from Southeast Asia up to northern Australia (Iskandar 2000;Lewis et al. 2013). In the Philippines, Indo-pacific crocodiles are found throughout the archipelago and in north-central Mindanao; their presence previously has been documented inland at Agusan Marsh. The species inhabits inland lakes, swamps and marshes, as well as coastal brackish waters and tidal sections of rivers; terrestrial nest sites and basking areas are frequently observed along many Mindanao freshwater bodies of water. This species has an IUCN status of "Least Concern" (LC; IUCN 2016).

Discussion
Our collective knowledge of the herpetological diversity of Mindanao Island (and, in general, the southern Philippines) relies on important mid-to late-19 th Century historical European museum collections and a few recent surveys, mostly concentrated at one region: northeast Mindanao (e.g., collections of E. H. Taylor, W. C. Brown and ACA; Plaza and Sanguila 2015), with scattered, limited-scope reports from the western (Nuñeza et al. 2010), central , and southeastern (Smith 1993a,b;Delima et al. 2006Delima et al. , 2007 corners of the island. The historical survey works of Taylor (1919Taylor ( , 1920aTaylor ( ,b 1921Taylor ( , 1922aTaylor ( ,b, 1923Taylor ( , 1960Taylor ( , 1965) based on his extensive survey work (1912)(1913)(1914)(1915) from the north-central portion of the island (Taylor 1975) resulted in records of approximately 100 species, which provided the first picture of Mindanao's herpetological diversity (Dickerson 1928;Taylor 1928Taylor , 1975. After this seminal work, the only subsequent summaries of Mindanao fauna were those of revisionary taxonomists (Inger 1954;Leviton 1963aLeviton -c, 1964aLeviton -c, 1965aLeviton ,b, 1967Leviton , 1968Leviton , 1970aLeviton ,b, 1979Leviton , 1983) and the biogeographical reviews of Brown and Alcala (1970, 1975, 1994, which relied primarily on Taylor's specimens and augmented his records with all that was known of the island at the time (ancillary collections). Since Taylor's work, the majority of our understanding of the herpetofaunal diversity of Mindanao (at least in terms of published literature and data in the public domain) has been augmented by excursions concentrated on specific sites. These include the records of 14-41 reptile species (Amoroso 2000;Beukema 2010) and 25 amphibian species records from central Mindanao (Delima et al. 2006), 34 species from southeastern portions of the island (Delima et al. 2007;Ates et al. 2009), 27 species reported from Mt. Hilong-hilong (Plaza and Sanguila 2015), 26-59 species recorded in the northwestern reaches of Mindanao (Tabaranza et al. 2001;Nuñeza et al. 2010), and 80-90 species recorded on the extreme western part of the island (Brown et al. unpublished data). Despite a paucity of information, the existence of high species diversity on Mindanao has been cited to rival other well-studied areas of the northern portions of the archipelago, where newly compiled levels of species diversity estimates range upwards of 100 species in some areas (Brown et al. 2012a. Because of our continued uncertainty regarding the true herpetological biodiversity of the southern Philippines, we strongly urge continued survey work aimed at providing accurate estimates of species richness in other subcenters of high-diversity within the Mindanao faunal region. Updated and continued assessment of the conservation status of Mindanao species Diesmos et al. 2014Diesmos et al. , 2015 is, likewise, urgently needed. Using new approaches of concentrated herpetological sampling efforts and repeat visits to several historical sites, we continue to ask: (1) how diverse is Mindanao's herpetological fauna? (2) Are there differences among sites that would suggest ecological and evolutionary processes behind observed patterns? (3) Is the continued treatment of the Mindanao PAIC's northern islands (e.g. Samar, Leyte) as simple faunal subsets of Mindanao an over simplification? (4) Just how different is the poorly studied fauna of western (Zamboanga del Norte, Zamboanga Sibugay, and Misamis Occidental provinces) and southern Mindanao (Maguindanao, Lanao del Norte, Lanao del Sur, Sultan Kudarat, South Cotabato, and Sarangani provinces) from the now moderately well-known central and northeastern sub-centers of diversity? (5) Do small islands, off the coast of Mindanao harbor endemic species and unrecognized diversity? These and other question should be the focus of a newly invigorated research program focused on the Mindanao faunal region.
To augment the record provided by published literatutre and publically served museum data, we surveyed isolated islands (Camiguin Sur and Dinagat) and mountain ranges spanning several provinces (Misamis Oriental, Agusan del Norte, Agusan del Sur, Surigao del Norte and Surigao del Sur) of northeastern Mindanao (Table 1). Based on our newly synthesized data (Table 2), we have confirmed the occurrence of numerous historically recorded species, and highlight many new distribution records on Dinagat Island, Agusan del Sur, and Misamis Oriental provinces. Our results suggest high herpetofaunal species diversity (approximately 35% of the total Philippine herpetofauna) in northeast Mindanao and adjacent small islands. We fully anticipate that this estimate of species diversity will continue to increase with future survey efforts targeting many inaccessible and/or poorly sampled intervening sites (Fig. 1) for which data currently are unavailable.
High estimates of regional herpetofaunal diversity in the Philippine archipelago have been confirmed in well-studied biogeographically distinct island groups such as Luzon (Brown and Stuart 2012;Brown et al. 2012aBrown et al. , 2013a. Understanding of the processes that have led to high species diversity in part relies on increased quality of geological inference (Hall 1996(Hall , 2002Yumul et al. 2009), an understanding of the routes of colonization of the archipelago (Brown and Guttman 2002;Brown and Siler 2013), and mechanisms associated with in situ speciation (i.e., geographical and/or marine barriers, ecological gradients, and habitat heterogeneity (Welton et al. 2009Brown et al. 2013b). Our new appreciation of Mindanao as an aggregate terrain, formed from recent accretion events of paleoisland precursors as well as subduction events associated with the uplift of previously submerged crustal platforms (Hall 1998;Yumul 2003Yumul , 2009), leads us to expect that these same mechanisms, inferred in islands to the north, may have likewise contributed to Mindanao's within-island provincial diversity in the south (McGuire and Alcala 2000; Sanguila et al. 2011;Brown and Siler 2013;Brown 2015) From our surveys, we note new Philippine records for native but non-endemic widespread species (those with a portion of their distributions outside the Philippines). These include Pelophryne brevipes (Bufonidae), Chaperina fusca (Microhylidae), Polypedates leucomystax, Rhacophorus pardalis, and Kurixalus appendiculatus (Rhacophoridae). These species' distribution data supplement our understanding of the eastern island arc colonization route (Sulu archipelago-Mindanao-Leyte-Samar-Luzon) as the hypothesized entryway to explain the distribution of herpetofaunal species on Mindanao (Brown and Guttman 2002;Gonzales et al. 2014;Brown and Siler 2014), and to other parts of the archipelago today (review: Brown et al. 2013b). Our new species distribution records from the islands of Camiguin Sur and Dinagat (e.g., Cyrtodactylus mamanwa, Eutropis cf. indeprensa, E. cf. multicarinata, Brachymeles vulcani, Calliophis sp.) re-affirm the prediction of faunal colonization from Mindanao to these ephemerally isolated islands via over water dispersal or colonization along land bridges (Inger 1954;). Both, apparently, have contributed to the processes of isolation, divergence, and speciation (Welton et al. 2010;Linkem et al. 2011;Siler et al. 2012b;Barley et al. 2013).
Although we did not explicitly analyze archipelago-wide distributional data, our new survey data (consisting of intensive elevational transects sampling, covering differing habitat types) corroborate the expectation of regional herpetofaunal diversity of Mindanao (Taylor 1928;Inger 1954;Leviton 1963a;Brown and Alcala 1970;Ross and Lazell 1990;Brown et al. 2013b;Gonzales et al. 2014). Recent surveys have uncovered numerous new species on the island (Brown, Diesmos, Sanguila, Siler unpublished data), which will supplement our understanding of species distributions in relation to other biogeographically distinct island groups of the archipelago Siler et al. 2011a;Barley et al. 2013). Our results not only confirmed additional species distribution records (Table 2), but also provided an opportunity for us to identify many herpetofaunal species complexes (27% of the total species included), which will require taxonomic work in the near future (e.g., Sanguila et al. 2011;Barley et al. 2013;Brown and Siler 2014;Brown 2015). Recent studies aimed at resolving species complexes have identified cryptic genetic diversity and important evolutionary units thereby clarifying these species' taxonomy and conservation status (e.g., Welton et al. 2009;2010;Sanguila et al. 2011;. Such studies have confirmed the importance of using of multiple sources of data for a sound understanding of species complexes (Brown and Guttman 2002;Grismer et al. 2013;Welton et al. 2013aWelton et al. , 2013bBrown et al. 2014;Siler et al. 2014a-d).
In this study, we have taken a simple, but important first step, aimed at identifying similar issues on Mindanao. Future work necessarily will involve the resolution of species complexes with taxonomic clarifications, via species descriptions and revisionary work. This will provide an important source of data for use in future studies focused on ecology, conservation, and diversity of many Mindanao amphibian and reptile species complexes.
Our current review of Mindanao amphibians and reptiles also serves as a template to address broader conservation issues. Hosting a large percentage of the remaining forested areas in the archipelago, the forests of Mindanao are threatened by commercial (legal) and illegal logging (deforestation), land conversion for agricultural and commercial purposes (Conservation International 2014;Brown 2015), and recently, the outbreak of an infectious disease that threatens the long term survival of amphibian populations (Swei et al. 2011;Brown et al. 2012b;Diesmos et al. 2012). The first two factors occur in the west and south, and also heavily in the east, while the threat of chytridiomycosis is now established in north-central portions of the island. An additional major threat comes in the form of introduced and invasive frog species (Diesmos et al. 2006, which appear to be having negative effects (extirpation of native species) along the island's coasts and lowlands (MBS, KC, RMB personal observations) The entire Mindanao landscape, particularly lower elevation forests, has been heavily impacted by widespread deforestation, and climate change has become a particular concern for some Mindanao forest and shrub frogs (Alcala et al. 2012). Aside from anthropogenic factors that threaten Mindanao's terrestrial biodiversity, one issue requiring attention from conservationists is a wholesale dearth of herpetological species' occurrence data Diesmos et al. 2014Diesmos et al. , 2015 throughout Mindanao. Few comprehensive, repeat-visit, site-based surveys are available for the island. Most recent published accounts have been derived from single-visit site-based studies (Amoroso 2000;Tabaranza et al. 2001;Delima et al. 2006Delima et al. , 2007Ates et al. 2009;Beukema 2010;Nuñeza et al. 2010), which we necessarily view as incomplete (see Brown et al. 2012aBrown et al. , 2013a. Although conservation status assessment efforts attempt to address these factors in current assessments (IUCN 2016; still badly out of date despite recent comprehensive revisions ] and based on the Global Amphibian Assessment of 2004), we have revised and supplemented IUCN conservation status of numerous species with new information from field surveys, plus all available occurrence data in the public domain  Table 2). Based on our new distributional records and natural history information for relevant species, we downgrade (or support the Diesmos et al.'s [2014] downgrading of eight species to lower threat categories (Platymantis guentheri, Limnonectes diuatus, L. parvus, Megophrys stejnegeri, Theloderma spinosum, Philautus acutirostris, Philautus worcesteri, Philautus surrufus, and Rhacophorus bimaculatus). In addition to providing novel assessments for previously unassesed taxa (Leptobrachium lumadorum, Bronchocela sp., Cyrtodactylus mamanwa, Brachymeles tiboliorum, Sphenomorphus variegatus, Lipinia quadrivittata, Dendrelaphis philippinensis, Calliophis philippina, and Psammodynastes pulverulentus), another predominant trend to our assessments have been the conversion of species of primarily "Least Concern" status to "Data Deficient" (IUCN 2010) due to recent demonstration of taxonomic uncertainty, a lack of information regarding species boundaries, and/or a lack of sufficient species occurrence or natural history data with which to make informed assessments.. Thirty-two of the total 126 species (~20%) recorded in this study require immediate systematic revision before informed conservation status assessments can be undertaken ( Table 2). Most of these are cases where we have evidence that a widespread species (sensu Taylor [1928], Inger [1954], Leviton [1963a], Brown and Alcala [1970]) actually is composed of several more range-restricted taxa. We anticipate that in many cases, these more geographically limited putative species will be associated with serious conservation concerns.
Our newly revised conservation reassessment of Mindanao species is based on novel species distribution data, combined with all available historical data (Table 2; see also Diesmos et al. 2014Diesmos et al. , 2015. The new survey data summarized here involved actual field surveys and, as such, should greatly inform existing (IUCN 2016) conservation assessments. Because the available IUCN (2016) assessments were compiled more than a decade ago, were based on secondary sources such as inferred forest cover, and indiscriminately applied and often incorrect assumptions about the natural history of many taxa (e.g., the degree to which species are, or are not, dependent on primary forest), they are now out of date and becoming incrementally ineffectual with time (Brown et al. 2012aDiesmos et al. 2014). Fortunately, the process of conservation status reassessment is underway ) and continuing, in an effort to stay current with each major faunal summary. Even in the face of Philippine central government efforts to establish the country's own, independent red list of endangered species (ACD unpublished data), staying current with IUCN (2016) conservation status assesments will be an ongoing challenge for future generations of conservation biologists if the country is to interact with neighboring nations, exchange internationallyrelevant information on conservation priorities, or cooperate with global wildlife trade monitoring, law enforcement, and/or conservation agencies (e.g., CITES, USF&W, IUCN, Conservation International).
We have attempted to provide a summary of all available information on the distribution of amphibians and reptiles of northeast Mindanao. This work both presents a state-of-the-art picture of the region's herpetofauna, but also identifies areas now sorely in need of additional survey work. Prioritization of remaining areas for field-based surveys (pockets of Data Deficient species like those of western Mindanao and the Sulu Archipelago) has not been been critically evaluated in past conservation assessments (IUCN 2016). Although some groups have been the subject of comprehensive taxonomic review, which has then allowed for subsequent conservation status assessment Brown 2015;Siler et al. 2011bSiler et al. , 2012bSiler et al. , 2014c, an analysis of suspected cryptic species diversity, and unrecognized (unprotected) new taxa is sorely lacking for approximately 20% of Mindanao's fauna. Great progress could be made with increased field-based survey work across all of central, southern, and western Mindanao. This mandate identifies a clear and urgent focus for ameliorating the lack of herpetological biodiversity information from the southern Philippines, and promotes the development of effective conservation management plans, tailored to the needs and challenges of individual protected areas and/or biodiversity hotspots, charged with conserving the remarkable herpetological biodiversity of Mindanao Island.
the Philippine Department of Environment and Natural Resources (DENR), for their continued support of our field research program. We thank R. G. Dahonog and R. M. Tawantawan (DENR Region X), R. Igot and M. Mendoza (DENR Region XIII) and DENR CENRO Cabadbaran City for logistical support. Our recent field surveys were conducted under a Gratuitous Permit to Collect Biological Specimens (GP) permit nos. 201, 210 and 212, provided by the Department of Environment and Natural Resources (DENR) Biodiversity Monitoring Bureau (BMB). In particular, we appreciate the efforts of our partners in fieldwork: J. Plaza, M.G. Medrano, J. Fernandez, W. Bulalaco, J. Cantil, N. Antoque, M. Tuto, B. Paulson, K. Ingenloff, M. Janra, E. DiBlasi, M. Pabillore, R. Edma, and V. Yngente. K. Andam and B. Pascual provided invaluable logistical support on Mts. Balatukan and Lumot. During the Mt. Lumot surveys, we especially appreciated the enthusiastic efforts of our dedicated field assistant, Baba, who secured several important species records (e.g., Dibamus) included here. We thank A. Barley, C. Linkem, L. Welton, and J. Vindum for helpful discussion of lizard taxonomy and access to specimen data and we are grateful to Aaron Bauer, Maren Gaulke and Indraneil Das for critical reviews of the manuscript. During initial stages of manuscript preparation MBS was supported by a Fulbright Senior Advanced Research Grant; this paper represents and her work towards this collaboration constitutes contribution No. 1 from the FSUU Biodiversity Informatics and Research Center.