Table of Contents

by Peter K. L. Ng and Jose Christopher E. Mendoza

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14001

Abstract (Primary Research Paper)

Date Posted (Final Published Version) :August 3, 2020

Abstract

The poorly known Philippine freshwater crab, Sundathelphusa picta (von Martens, 1868) from Luzon Island is re-described and re-illustrated, using type material as well as other specimens sampled from near its type locality. Two similar congeners from Luzon, S. uva sp. nov. and S. angelito sp. nov., from the provinces of Bataan and Rizal, respectively, are described as new. These three species are united by their relatively small size, rounded and dome-shaped carapaces, proportionately short ambulatory legs, and stout male first gonopods. They are distinguished from each other by a suite of morphological characters, particularly of the carapace, male pleon and gonopods.

KEYWORDS: Decapoda, Sundathelphusa uva, Sundathelphusa angelito, taxonomy, Bataan, Bicol, Rizal

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by Camila G. Meneses, Cameron D. Siler, Juan Carlos T. Gonzalez, Perry L. Wood, Jr., and Rafe M. Brown

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14002

Abstract (Primary Research Paper)

Date Posted (Final Published Version) :August 10, 2020

Abstract

We report on the first molecular estimates of phylogenetic relationships of Brachymeles dalawangdaliri (Scincidae) and Pseudogekko isapa (Gekkonidae), and present new data on phenotypic variation in these two poorly known taxa, endemic to the Romblon Island Group of the central Philippines. Because both species were recently described on the basis of few, relatively older, museum specimens collected in the early 1970s (when preservation of genetic material was not yet standard practice in biodiversity field inventories), neither taxon has ever been included in modern molecular phylogenetic analyses. Likewise, because the original type series for each species consisted of only a few specimens, biologists have been unable to assess standard morphological variation in either taxon, or statistically assess the importance of characters contributing to their diagnoses and identification. Here we ameliorate both historical shortfalls. First, our new genetic data allowed us to perform novel molecular phylogenetic analyses aimed at elucidating the evolutionary relationships of these lineages; secondly, with population level phenotypic data, from the first statistical sample collected for either species, and including adults of both sexes. We reaffirm the distinctiveness of both named taxa as valid species, amend their diagnoses to facilitate the recognition of both, distinguish them from congeners, and consider the biogeographic affinities of both lineages. Our contribution emphasizes the conservation significance of Sibuyan Island’s Mt. Guiting-Guiting Natural Park, the diverse, idiosyncratic biogeographic histories of its variably-assembled, highly endemic reptile fauna, and the critical importance of multiple, repeated, survey–resurvey studies for understanding forest community species composition and the evolutionary history of Philippine biodiversity.

KEYWORDS: biodiversity, endemism, forest geckos, faunal region, fossoriality, limb reduction

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by Emerson Y. Sy, Sabine Schoppe, Mae Lowe L. Diesmos, Theresa Mundita S. Lim, and Arvin C. Diesmos

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14003

Abstract (Primary Research Paper)

Date Posted (Final Published Version) :August 11, 2020

Abstract

The Philippine or Palawan Forest Turtle Siebenrockiella leytensis is the only endemic turtle known to occur in the Philippines. It was assessed as Critically Endangered in 2000 and has been considered as one of the world’s top 25 most endangered turtles since 2003. The species is accorded protection nationally by the Wildlife Protection and Conservation Act of 2001 and its international commercial trade is regulated by the Convention on International Trade in Endangered Species (CITES). However, the publication of its rediscovery in 2004 triggered unrelenting poaching and trafficking for the pet trade nationally and internationally. With the aim of quantifying the extent of poaching and to provide insight on the trade dynamics, we analyzed seizure records from 2004–2018 and conducted physical and online market surveys in 2017–2018. Twenty-three (23) seizure incidents involving 4,723 Philippine Forest Turtles were recorded in the last 15 years. Based on an online survey, we estimated that an additional 1,200 Philippine Forest Turtles were smuggled and illegally sold in China in 2015. The majority of the 74 live individuals exported legally from the Philippines were likely sourced illegally from the wild and declared fraudulently as captive bred by exporters to obtain CITES permits. While habitat loss or degradation is a major threat, the illegal pet trade remains the most important factor threatening the survival of the Philippine Forest Turtles in the wild.

KEYWORDS: chelonian, CITES, pet trade, trafficking, wildlife laundering

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by Abner A. Bucol, Rainier I. Manalo, Angel C. Alcala, and Paulina S. Aspilla

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14004

Abstract (Primary Research Paper)

Date Posted (Final Published Version): November 2, 2020

Abstract

Crocodilians have been assumed to influence aquatic primary productivity and fishery yield. However, strong empirical evidence to support such claims is lacking. The long-standing assumption first hypothesized by Fittkau (1970), is that local fisheries (secondary productivity) in areas inhabited by crocodilians would be expected to improve. We tested this hypothesis at two locations in the Philippines, inhabited by the Philippine Crocodile (Crocodylus mindorensis) in Paghungawan Marsh in Siargao Island Protected Landscape & Seascape (SIPLAS), Jaboy, Pilar, Surigao Del Norte, and the Indo-Pacific Crocodile (Crocodylus porosus) in the Rio Tuba River, Bataraza, southern Palawan Island. Water chemistry parameters, with emphasis on nutrient (nitrate and phosphate) levels, were determined using using standard protocols. Catch-per-Unit Effort (CPUE) of gillnets in sites with crocodiles was compared with corresponding control sites without crocodiles. CPUE was higher in areas inhabited by crocodiles, but appeared not to be directly influenced by nutrient levels. Increased fish catches in areas inhabited by crocodiles might be attributed to several factors, such as reduced fishing pressure due to the presence of crocodiles which discouraged the local fishermen to fish intensively. Overall, while fish catch was higher in areas inhabited by crocodiles, it is too early to attribute this to the nutrient output from crocodiles due to several confounding factors.

KEYWORDS: estuarine, fish catch, freshwater, nutrient

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by Cameron D. Siler, Elyse S. Freitas, Jennifer A. Sheridan, Stephanie N. Maguire, Drew R. Davis, Jessa L. Watters, Kai Wang, Arvin C. Diesmos, and Rafe M. Brown

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14005

Abstract (Primary Research Paper)

Date Posted (Final Published Version): November 16, 2020

Abstract

The diversity of Philippine amphibians and reptiles has increased over the last few decades, in part due to re-evaluation of species formerly believed to be widespread. Many of these investigations of widespread species have uncovered multiple closely related cryptic lineages comprising species complexes, each restricted to individual Pleistocene Aggregate Island Complexes (PAICs). One group in particular for which widespread cryptic diversity has been common is the clade of Philippine skinks of the genus Brachymeles. Recent phylogenetic studies of the formerly recognized widespread species Brachymeles bonitae have indicated that this species is actually a complex distributed across several major PAICs and smaller island groups in the central and northern Philippines, with numerous species that exhibit an array of digit loss and limb reduction patterns. Despite the recent revisions to the B. bonitae species complex, studies suggest that unique cryptic lineages still exist within this group. In this paper, we resurrect the species Brachymeles burksi Taylor 1917, for a lineage of non-pentadactyl, semi-fossorial skink from Mindoro and Marinduque islands. First described in 1917, B. burksi was synonymized with B. bonitae in 1956, and has rarely been reconsidered since. Evaluation of genetic and morphological data (qualitative traits, meristic counts, and mensural measurements), and comparison of recently-obtained specimens to Taylor’s original description support this species’ recognition, as does its insular distribution on isolated islands in the central portions of the archipelago. Morphologically, B. burksi is differentiated from other members of the genus based on a suite of unique phenotypic characteristics, including a small body size, digitless limbs, a high number of presacral vertebrae, the absence of auricular openings, and discrete (non-overlapping) meristic scale counts. The recognition of this central Philippine species further increases the diversity of non-pentadactyl members of the B. bonitae complex, and reinforces the biogeographic uniqueness of the Mindoro faunal region.

KEYWORDS: biodiversity, endemism, faunal region, fossoriality, limb reduction

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by Jeffrey L. Weinell, Alan E. Leviton, and Rafe M. Brown

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14006

Abstract (Primary Research Paper)

Date Posted (Final Published Version): February 8, 2021

Abstract

We describe a new species of reed snake of the genus Calamaria Boie 1827, from Mindoro Island, Philippines. The new species differs from all other species of Calamaria by having the following combination of characters: a high number of subcaudal scale pairs (> 40 in males, > 30 females) and ventrals + subcaudals (> 205 in males, > 210 in females); mental scale not contacting chin shields; dorsal surface of head, body, and tail uniformly dark brown; and ventral surface of body (extending to include part or all of first longitudinal row of dorsals) uniformly pale (yellow or white in life). The new species is likely most closely related to Calamaria schlegeli Duméril, Bibron, and Duméril 1854, which also has a high number of subcaudal scales compared to other Calamaria species. The new species is the second Calamaria species known from Mindoro Island and the eighth known from the Philippines, and its presumed distant relationship from other Philippine Calamaria suggests an additional colonization of the Philippines by this genus from continental Asia.

KEYWORDS: biodiversity, biogeography, Calamaria alcalai new species, Serpentes, Squamata, systematics

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by Kin Onn Chan, Sabine Schoppe, Edmund Leo B. Rico and Rafe M. Brown

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14007

Abstract (Primary Research Paper)

Date Posted (Final Published Version): March 22, 2021

Abstract

Focusing on the phylogenetic relationships of puddle frog populations spanning the biogeographic interface between Sundaland (Borneo) and the Philippines, we demonstrate, for the first time, a widespread geographic pattern involving the existence of multiple divergent and co-distributed (sympatric) evolutionary lineages, most of which are not each other’s closest relatives, and all of which we interpret as probable distinct species. This pattern of co-occurrence in the form of pairs of ecologically distinct puddle frog forms (dyads), prevails throughout northern Borneo, Palawan, Tawi-Tawi, the Sulu Archipelago, and western Mindanao (Zamboanga). Previously obscured by outdated taxonomy and logistical, legal, and security obstacles to field-based natural history studies, this pattern has remained hidden from biogeographers and amphibian biologists by an uncontested proposal that Philippine Occidozyga laevis is a single, “widespread,” and “highly variable” species. In this paper we use an integrative synthesis of new genetic data, organismal phenotypic data, historical literature reports, and ecological observations to elucidate an interesting and potentially widespread pattern of puddle frog species coexistence at the Sundaland–Philippine biogeographic interface. Calling attention to this pattern opens promising possibilities for future research aimed at understanding the scope of this dyads pattern, and whether it extends to the more northern reaches of the Philippines. On either side of Huxley’s and Wallace’s lines, data suggest that the majority of puddle frog dyads at a given locality are not each other’s closest relatives (are more distantly related, or non-monophyletic) and, thus, assembled ecologically, likely coexisting now as a result of their ecological tendencies toward distinct microhabitats (warmer stagnant pools in open areas, versus cool, flowing streams enclosed in forest). If these pairs of species types are determined to be the geographic norm among the more isolated, central, and northern, Philippine faunas, an obvious question will be whether they have evolved into dual ecological forms, possibly in response to ecological opportunity and/or reduced competition.

KEYWORDS: biogeography, taxonomy, microhabitat, cryptic species

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by Alan T. White

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14008

Abstract (Primary Research Paper)

Date Posted (Final Published Version): March 22, 2021

Abstract

This review shares lessons learned from the establishment of early marine protected areas (MPAs) in the Philippines about the need to establish baseline information, do systematic monitoring of the status of the marine environment, and to progress towards more integrated forms of management that involve key stakeholders in coastal areas. The tendency for human society to change its perception of what is “normal” through the phenomena of “shifting baselines” is pointed to as a reason why more concerted action is not taken to stop the downward trends of Philippine coastal resources and environment. The small MPAs of Apo, Sumilon and Olango Islands as well as the large Tubbataha Reefs Natural Park, are cited as examples of how the establishment of baselines and the implementation of effective monitoring over time for both biophysical and governance parameters, has been instrumental in maintaining and improving the quality of the marine environment and bringing benefits to people. The development of integrated coastal management and coastal resource management programs within local government units is explained as a way of harnessing local institutions to lead the way towards improved management and stewardship of coastal resources and provide tangible benefits to coastal communities. And, the role of national government is highlighted as a facilitator and a source of technical support to local governments in the implementation of marine conservation and coastal resources management. Finally, the significant influence of Dr. Angel Alcala in marine conservation in the Philippines is noted through his research and related conservation efforts for small-island and fisheries management and his mode of sharing results with local communities and governments so that they could learn from their own mistakes and successes and become better stewards of their resources.

KEYWORDS: Apo, community, coral reefs, Sumilon, tourism, Tubbataha

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by Indraneil Das and Genevieve V.A. Gee

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14009

Abstract (Short Communication)

Date Posted (Final Published Version): April 14, 2021

Abstract

In this essay, we commemorate the zoological and herpetological contributions of Angel Chua Alcala, with a review of stamps and pictorial cancellations on herpetological themes from the Philippines. Between 1982 and 2017, a total of 79 such stamps, stamp sheetlets, and undenominated tabs, depicting amphibians and reptiles have been officially issued by the postal administration of the country, all but one within its commemorative stamp releases. Species featured are those of ecotourism importance, in addition to threatened or endemic taxa, although stylized as well as non-local species too have featured on stamps produced by the country.

KEYWORDS: Philippines, philately, stamps, postmarks, amphibians, reptiles

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by Harvey B. Lillywhite

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14010

Abstract (Primary Research Paper)

Date Posted (Final Published Version): April 16, 2021

Abstract

Biodiversity and the function of tropical shallow-water marine environments are threatened by numerous anthropogenic factors, especially climate change, overharvesting of resources, and destruction of habitat. Marine snakes are important components of coastal shallow-water systems and should be considered as indicators of the health of coastal ecosystems such as mangroves. Acrochordid snakes (Acrochordidae: Acrochordus) represent a highly distinct evolutionary lineage with unusual adaptations to shallow water habitats and importance to biodiversity of tropical coastal regions. One of three congeneric species, Acrochordus granulatus (file snake), is an interesting and common inhabitant of coastal estuaries and mangroves in the Philippines. This paper reviews unusual attributes of A. granulatus and provides a perspective for its conservation in coastal habitats. Morphological, physiological, and behavioral characters of this snake are specialized for life in shallow-water marine environments such as mangroves. Unusual and specialized features confer abilities for prolonged submergence and include low metabolic rate, large capacity for oxygen storage, cutaneous gas exchange, nearly complete utilization of oxygen stores during aerobic submergence, intracardiac and cutaneous shunts for regulating blood flow, and reclusive behavior. Fresh water is required for water balance, and file snakes are dependent on rainfall in many habitats where they drink from freshwater lenses formed by precipitation on the surfaces of marine water. File snakes feed largely on fishes and are candidates as bio-indicators of the health of shallow-water coastal habitats. Attention should be given to threatening insults on coastal environments including climate change, habitat destruction, harvesting of resources, and other factors in need of research, monitoring, and plans for abatement. Importantly, conservation can be promoted by educating people about the docile behavior, unusual traits, and interesting ecology of A. granulatus.

KEYWORDS: mangrove, shallow water, Acrochordidae, little file snake, conservation physiology, ecophysiology

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by Danilo S. Balete†, Lawrence R. Heaney , and Eric A. Rickart

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14011

Abstract (Primary Research Paper)

Date Posted (Final Published Version): February 14, 2021

Abstract

Small mammal communities that occur in habitats on volcanic soil substrates have been extensively studied on Luzon Island, but those that occur in forest over limestone are poorly known and have not been directly compared to those over volcanic soils. We conducted field surveys of small mammals in forest over limestone from ca. 100 m to 590 m elevation in the vicinity of Callao Cave, and in adjacent lowland dipterocarp forest over volcanic soil from 490 m to 900 m, near the location of prior surveys from 1300 m to1550 m on Mt. Cetaceo, an extinct volcanic peak in the northern Sierra Madre range. Despite moderately heavy disturbance to the habitats over karst (limestone) and moderate disturbance to forest over volcanic soils, we found native small mammals overall at levels of species richness and abundance similar to what we have documented elsewhere on Luzon over the same elevational range. Non-native mammals were present at all localities in the karstic habitat but were absent in all types of forest over volcanic soils, even in areas recovering from prior disturbance. Although non-natives were moderately common in karstic areas, they rarely were more common than native species, and non-native species were no more successful at invading the disturbed karstic habitat than the native species were at persisting there. The most abundant small mammal in dipterocarp forest over volcanic soil (Apomys sierra) was absent in karstic localities, despite occurring in adjacent areas at overlapping elevation. Overall, the difference between small mammals in karst and lowland dipterocarp forest was mainly due to species composition rather than total abundance. Comparisons with data from a prior study on the upper slopes of Mt. Cetaceo showed that total native species abundance was highest in montane and mossy forest, typically about three times higher than in lowland dipterocarp forest. We confirmed the current presence of one species, Apomys microdon, reported as a fossil from Callao Cave, but the apparent absence of one other, Batomys sp.; both were from deposits dated as ca. 65,000 BP. We also summarize information about large mammals in the study areas. Further study of mammals in the distinctive forest over limestone is clearly needed.

KEYWORDS: biodiversity, biogeography, Cagayan Valley, disturbed forest, elevation, fossils, Muridae, Sierra Madre, Soricidae

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by Anthony J. Barley, Marites B. Sanguila, and Rafe M. Brown

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14012

Abstract (Primary Research Paper)

Date Posted (Final Published Version): April 14, 2021

Abstract

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

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

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by Arne Redsted Rasmussen, Anders Hay-Schmidt, Farnis Boneka, Morten E. Allentoft, Kate Laura Sanders, and Johan Elmberg

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14013

Abstract (Primary Research Paper)

Date Posted (Final Published Version): April 25, 2021

Abstract

Shallow tropical marine ecosystems are under great anthropogenic pressure due to habitat destruction, overfishing, shrimping, climate change, and tourism. This is an issue of global concern as such environments hold a tremendous biodiversity much of which remains to be described. The present situation urgently calls for time- and resource-efficient methods to identify and delineate the most valuable remaining areas and to set up priorities for their management and conservation. Using indicator species can be a way to accomplish this goal. In this paper we evaluate whether viviparous sea snakes can serve as bioindicators for other rare or cryptic tropical marine fauna. Based on seven generally acknowledged criteria for bioindicators, we argue that using viviparous sea snakes as bioindicators can help monitoring marine habitats to gauge the effects of climate change, habitat change and loss, decline in biodiversity and other anthropogenic changes. However, to maximize their efficacy as bioindicators, deeper knowledge about viviparous sea snakes natural history is urgently needed. Topics for expanded research programs include the taxonomy of some groups, their breeding and feeding biology, habitat selection and their geographical distribution. Despite these gaps in our understanding, we argue that viviparous sea snakes can be utilized as bioindicators of marine ecosystem health.

KEYWORDS: anthropogenic changes, conservation, herpetology, marine habitat, monitoring

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by Terrence M. Gosliner

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14014

Abstract (Primary Research Paper)

Date Posted (Final Published Version): July 12, 2021

Abstract

The Verde Island Passage of the Philippines is renowned for its rich marine biodiversity and heterobranch mollusks are demonstrable models of that species richness of the region. Chromodorid nudibranchs represent a particularly rich taxon, with species of Chromodoris being one of the most iconic and abundant taxa in shallow water coral reef environments. Despite being one of the best-documented clades of nudibranch mollusks, recent work has shown that numerous cryptic and pseudocryptic species are abundant in the waters of the Coral Triangle region. This paper reviews the species richness and distribution of Chromodoris species in the Philippines and provides a description of a new species of Chromodoris from the region. Chromodoris alcalai Gosliner, n. sp. is named to honor Dr. Angel Alcala’s 90th birthday and his pioneering contributions to preserving the marine biodiversity of the Philippines. This species is most similar externally to C. dianae Gosliner and Behrens, 1998, with which it was erroneous lumped. Distinctness of molecular data, external morphology and coloration, and internal anatomical features clearly distinguish these two species.

KEYWORDS: systematics, Nudibranchia, coral reefs, Indo-Pacific, Coral Triangle

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by John C. Murphy and Harold K. Voris

Philippine Journal of Systematic Biology, Volume 14, Issue 2, 2020, DOI: 10.26757/pjsb2020b14015

Abstract (Primary Research Paper)

Date Posted (Final Published Version): July 19, 2021

Abstract

Homalopsid snakes are monophyletic and contain two major subclades: a fangless clade and rear-fanged clade. They are distributed in South Asia, Australasia, and the Western Pacific. The fangless clade is restricted to the eastern Indonesian Archipelago and the island of Sumatra and is poorly known in terms of its natural history. Molecular data support the eastern Indonesian fangless endemic genus Brachyorrhos as the sister to the rear-fang clade. Here we recognize the identity of the Brachyorrhos population from the island of Morotai as B. wallacei and describe a new species of dwarf Brachyorrhos from the island of Seram, Malukus, Indonesia. The new species can be distinguished from all congeners by a lower number of ventral scales, the presence of a preocular scale and a loreal scale, as well as its exceptionally diminutive size. The new species is a candidate for the smallest alethinophidian snake. The three fangless genera, Brachyorrhos, Calamophis, and Karnsophis, have been suggested to form a clade of homalopsid snakes restricted to the Indonesian Archipelago, and we discuss their biogeography.

KEYWORDS: biogeography, Calamophis, Homalopsidae, Karnsophis, small snakes

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