[Herpetology • 2026] Bronchocela carestiai & B. mallarii • A Comprehensive, Archipelago-Wide Taxonomic Reassessment of the Genus Bronchocela (Squamata: Agamidae) in the Philippines, with Descriptions of Two New Species

 

(A) Bronchocela marmorata from Luzon,
(B, C) B. philippina from Mindanao,
 (D) Bronchocela carestiai from Mindoro, and
(E) B. mallarii from Palawan. 

 

Supsup, Pitogo, Sanguila, Hallermann, Denzer, Wood & Brown, 2026

Herpetological Monographs. 40 (1),

Photographs by Rafe M. Brown, Christian E. Supsup, and Kier Mitchel E. Pitogo.

 Abstract  

One challenge for species delimitation involves closely related species, characterized by both intra- and interspecifically variable morphologies, for which genetic information is not yet available. One such example is exhibited by agamid lizards of the genus Bronchocela. This group of conspicuously vibrant green lizards is widely distributed in Southeast Asia, with 15 recognized species. However, the geographic distributions of several named taxa have remained unclear because of the imprecise application of names in taxonomic summaries and faunal checklists. For more than a century, many authors have noted that a few species, often considered “widespread,” persist throughout nearly the entire range of the genus. This is particularly true for populations identified as Bronchocela cristatella sensu lato and, to a lesser extent, B. marmorata in the Philippines. In this study, we reassess the systematics and taxonomy of Bronchocela populations in the Philippine archipelago. Using multiple lines of evidence from molecular and morphological data, we uncover four distinct lineages of Philippine Bronchocela, all of which we interpret as being distantly related to B. cristatella sensu lato from Malaysia and Indonesia. We argue that these lineages should now be interpreted as Philippine endemic species. Given the strong congruence in molecular and morphological analyses, we now consider populations from the Luzon faunal region (including Catanduanes, Marinduque, Lubang, and Polillo Islands), Romblon (Tablas, Romblon, and Sibuyan Islands), and the West Visayas (Negros, Cebu, and Panay Islands) as B. marmorata. The nomenclaturally available name, B. philippina, is now designated to the unique lineage from the Mindanao faunal region. Additionally, we recognize two new, distinctive species from the Mindoro (B. carestiai) and Palawan (B. mallarii) faunal regions. Overall, our findings demonstrate how conspicuous and widely distributed species in Southeast Asia remain poorly understood, overlooked, and taxonomically unresolved. Hence, the taxonomic resolution of Bronchocela populations in the island archipelagos of Southeast Asia will greatly benefit from the urgent priority of determining which landmass the name “B. cristatella” applies.

KEYWORDS: Bronchocela carestiai sp. nov, Bronchocela marmorata, Bronchocela mallarii sp. nov, Bronchocela philippina, Southeast Asia, taxonomy, Widespread species

In life photographs of Bronchocela marmorata from Luzon (A), B. philippina from Mindanao (B, C),
B. carestiai from Mindoro (D), and B. mallarii from Palawan (E).

Photographs were taken by Rafe M. Brown, Christian E. Supsup, and Kier Mitchel E. Pitogo.

Bronchocela carestiai sp. nov.

 Bronchocela mallarii sp. nov. 

Christian E. Supsup, Kier Mitchel E. Pitogo, Marites B. Sanguila, Jakob Hallermann, Wolfgang Denzer, Perry L. Wood Jr. and Rafe M. Brown. 2026. A Comprehensive, Archipelago-Wide Taxonomic Reassessment of the Genus Bronchocela (Squamata: Agamidae) in the Philippines, with Descriptions of Two New Species. Herpetological Monographs. 40 (1), 1-44, (20 July 2026) DOI: doi.org/10.1655/HERPMONOGRAPHS-D-24-00003

https://bioone.org/journals/herpetological-monographs/volume-40/issue-1/HERPMONOGRAPHS-D-24-00003/A-Comprehensive-Archipelago-Wide-Taxonomic-Reassessment-of-the-Genus-Bronchocela/10.1655/HERPMONOGRAPHS-D-24-00003.full

[Herpetology • 2021] Eutropis alcalai • A New Species of Sun Skink (Scincidae: Eutropis) from the Zamboanga Peninsula, southwestern Mindanao Island, Philippines

Eutropis alcalai  

Barley, Sanguila & Brown, 2021

Alcala’s Rough-scaled Sun Skink ||  facebook.com/PhilippineSystematists 

Abstract 

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

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

Eutropis alcalai sp. nov. (holotype PNM 9878; formerly KU 315013; adult male, SVL 70.1 mm), photographed in life:
(a) close-up of anterior body region; (b) full body;
collected from Pasonanca Natural Park, Barangay Pasonanca, on the outskirts of Zamboanga City, Zamboanga Peninsula, western Mindanao Island.

Photos: Rafe M. Brown.

 

Eutropis alcalai sp. nov. 

Diagnosis.— Eutropis alcalai sp. nov. can be distinguished from its closest relative, E. rugifera, by its smoother head scales, with light embossing only, and keels limited to posterior margins of parietals, temporals and nuchals (vs. rugose to strongly keeled in E. rugifera), by absence (vs. presence) of contact between the postmental and second infralabial, by the presence of 29–32 (vs. 21–26) subdigital lamellae under the 4th toe, by the presence of seven infralabials (six in E. rugifera), the presence of differentiation in the precloacal scales series (two, medial precloacals enlarged, transversely expanded and bordered on either side by two undifferentiated scales) in E. alcalai sp. nov. (vs. six equal sized scales in E. rugifera), by equivalent length of 3rd and 4th fingers (vs. 4th finger longer than 3rd), and by its white chin and infralabial scales (vs. yellow to orange in E. rugifera).

Distribution.— This species is only known from Pasonanca Natural Park near Zamboanga City at the southwestern tip of the Zamboanga Peninsula. A closely related species (E. rugifera) occurs on Borneo (among other islands of the Sunda Region), and although neither species has been collected from the Sulu Archipelago, one of them may occur on its larger islands.

Etymology.— We take great pleasure in naming this distinctive new species for our colleague Angel C. Alcala, in recognition of his numerous foundational contributions to the natural history, systematics, ecology, and conservation of Philippine lizards of the family Scincidae (Alcala and Brown 1966, 1967; Alcala 1970; Brown and Alcala 1956, 1961, 1963a,b, 1980, 1986). Suggested common name: “Alcala’s Quinque-carinate (Five-keeled) Sun Skink,” or “Alcala’s Rough-scaled Sun Skink.”

Anthony J. Barley, Marites B. Sanguila and Rafe M. Brown. 2021. A New Species of Sun Skink (Reptilia: Scincidae: Eutropis) from the Zamboanga Peninsula, southwestern Mindanao Island, Philippines. Philippine Journal of Systematic Biology. DOI: 10.26757/pjsb2020b14012

   asbp.org.ph/wp-content/uploads/2021/05/PJSB-ACA-2020-012.pdf

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[Herpetology • 2016] Brachymeles dalawangdaliri • Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Squamata: Scincidae) III: A New Species from Tablas Island

Tablas Slender Skink |  Brachymeles dalawangdaliri  

 Davis, Geheber, Watters, Penrod, Feller, Ashford, Kouri, Nguyen, Shauberger, Sheatsley, Winfrey, Wong, Sanguila, Brown & Siler, 2016.  

 DOI: 10.11646/zootaxa.4132.1.3  

Abstract

Studies of the diversity of Philippine amphibians and reptiles have resulted in the continued description of cryptic species. Species formerly thought to range across multiple recognized faunal regions are now considered to be assemblages of multiple unique species, each restricted to a single faunal region. This pattern continues to hold true when considering Philippine skinks of the genus Brachymeles. Recent studies have resulted in the description of numerous unique species with many exhibiting various degrees of digit loss or limb reduction, as well as suggesting that unique lineages are still present in the B. bonitae Complex. In this paper, we describe a new species of fossorial skink within this species complex from Tablas Island based on collections made nearly 50 years ago. Although no genetic data are available for the new species, examinations of morphological data (qualitative traits, meristic counts, and mensural measurements) support its distinction from all other members of the genus. Brachymeles dalawangdaliri sp. nov. is differentiated from other members of the genus based on a suite of unique phenotypic characteristics, including a small body size (SVL 66.0–80.9 mm), bidactyl fore-limbs, digitless, unidactyl, or bidactyl hind limbs, a high number of presacral vertebrae (49), the absence of auricular openings, and distinct dorsal head scale patterns. The description of the new species increases the diversity of endemic vertebrates recognized to occur in the Romblon Island Group in the central Philippines.

Keywords: Reptilia, biodiversity, endemism, faunal region, fossoriality, limb reduction, non-pentadactyl, pentadactyl, Romblon Island Group

Taxonomic account

Brachymeles dalawangdaliri sp. nov.

Brachymeles bonitae Duméril & Bibron 1839; Taylor 1917; Brown 1956:5; Brown & Rabor 1967:526; Brown & Alcala 1970; Brown & Alcala 1980:20; Davis et al. 2014; Geheber et al. 2016; Siler et al. 2016. 

Diagnosis. Following recent taxonomic revisions of Brachymeles (Siler et al. 2011; Davis et al. 2014) the new species is assigned to the B. bonitae Complex based on the following suite of morphological characters: (1) limbs present, (2) non-pentadactyl, (3) fore-limbs with 0–3 fingers. (4) hind limbs with 0–2 toes, (5) paravertebral scale rows ≥ 91, (6) presacral vertebrae 47–53, (7) supraoculars four, (8) enlarged, differentiated nuchals present, (9) longitudinal rows of dark spots around the body absent, and (10) auricular opening absent.

Brachymeles dalawangdaliri sp. nov. can be distinguished from congeners by the following combination of characters: (1) body size small (SVL 66.0–80.9 mm), (2) fore-limbs bidactyl, (3) hind limbs digitless, unidactyl, or bidactyl, (4) limb length short, (5) supralabials six or seven, (6) infralabials six or seven, (7) supraciliaries five, (8) supraoculars four, (9) midbody scale rows 24 or 25, (10) axilla–groin scale rows 80–83, (11) paravertebral scale rows 97–101, (12) prefrontal contact absent, (13) frontoparietal contact present or absent, (14) enlarged chin shields in two pairs, (15) nuchals enlarged, (16) auricular opening absent, (17) presacral vertebrae 49, and (18) uniform body color (Tables 1, 2).

Etymology. The name of the new species presented in this paper is derived from the Tagalog (Filipino) terms ‘dalawa,’ meaning two, and ‘daliri,’ meaning finger, referencing one of the defining characteristics of the new species, the two digits on the fore-limbs. Suggested common name: Tablas Slender Skink.

Distribution. Brachymeles dalawangdaliri sp. nov. is known only from Tablas Island, Romblon Island Group, Philippines (Fig. 1).

 Drew R. Davis, Aaron D. Geheber, Jessa L. Watters, Michelle L. Penrod, Kathryn D. Feller, Alissa Ashford, Josh Kouri, Daniel Nguyen, Kathryn Shauberger, Kyra Sheatsley, Claire Winfrey, Rachel Wong, Marites B. Sanguila, Rafe M. Brown and Cameron D. Siler. 2016.
 Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) III: A New Species from Tablas Island.
Zootaxa. 4132(1); 30-43.  DOI: 10.11646/zootaxa.4132.1.3

[Herpetology • 2016] Brachymeles ilocandia • Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Squamata: Scincidae) II: A New Species from the northern Philippines

Ilokano Slender Skink |  Brachymeles ilocandia 

Siler, Davis, Freitas, Huron, Geheber, Watters, Penrod, Papeș, Amrein, Anwar, Cooper, Hein, Manning, Patel, Pinaroc, Diesmos, Diesmos, Oliveros & Brown. 2016.    DOI: 10.11646/zootaxa.4132.1.2 

Abstract

We describe a new digitless scincid lizard of the genus Brachymeles from northern Luzon and Camiguin Norte islands in the Philippines. This species belongs to the Brachymeles bonitae Complex, and both molecular and morphological data confirm that this species is distinct from all other congeners. Formerly considered to be a single widespread species, this group of species has been the focus of recent systematic reviews. Here we describe a new species in the B. bonitae Complex, recognized currently to constitute five species. Brachymeles ilocandia sp. nov. is the second digitless and the seventeenth non-pentadactyl species in genus. The description of this species brings the total number of species in the genus to 40, and provides new insight into unique distribution patterns of species of the northern Philippines.

Keywords: Reptilia, Babuyan Island Group, biodiversity, ecological niche modeling, endemism, faunal region, fossorial, Luzon Island, non-pentadactyl, pentadactyl

Brachymeles ilocandia sp. nov.

Brachymeles bonitae Duméril & Bibron 1839; Taylor 1917; Brown 1956:5; Brown & Rabor 1967:526; Brown & Alcala 1970; Brown & Alcala 1980:20; Davis et al. 2014; Geheber et al. 2016.

Diagnosis. Following recent taxonomic revisions of Brachymeles (Siler et al. 2011; Davis et al. 2014; Geheber et al. 2016) the new species is assigned to the B. bonitae Complex based on the following suite of morphological characters: (1) limbs present, (2) non-pentadactyl, (3) fore-limbs with 0–3 fingers, (4) hind limbs with 0–2 toes, (5) paravertebral scale rows ≥ 91, (6) presacral vertebrae 47–53, (7) supraoculars four, (8) enlarged, differentiated nuchals present, (9) longitudinal rows of dark spots around the body absent, and (10) auricular opening absent.

Brachymeles ilocandia sp. nov. can be distinguished from congeners by the following combination of characters: (1) body size small (SVL 65.7–77.6 mm), (2) limbs digitless, (3) limb length short, (4) supralabials six, (5) infralabials five or six, (6) supraciliaries five, (7) supraoculars four, (8) midbody scale rows 22–24, (9) axilla–groin scale rows 80–82, (10) paravertebral scale rows 97–100, (11) mental/first infralabial fusion present or absent, (12) prefrontal contact absent or in point contact, (13) frontoparietal contact present, (14) enlarged chin shields in three pairs, (15) nuchals enlarged, (16) auricular opening absent, (17) presacral vertebrae 50–53, and (18) uniform body color (Tables 1, 2).

Distribution, ecology and natural history. Brachymeles ilocandia sp. nov. is known only from northern Luzon and Camiguin Norte islands (Fig. 1A). The new species likely once occurred in low- to mid-elevation primary forest habitats; however, all recent observations of this species have occurred in secondary growth forest habitats. In contrast to the other members of the B. bonitae Complex, this species appears to be quite common in secondary growth forest fragments throughout the northern Philippines.  Brachymeles ilocandia sp. nov. is found in parts of its distribution with B. bicolor, B. kadwa, and B. boulengeri. Other species of Brachymeles known to occur in the Luzon PAIC include B. bicolandia, B. bonitae, B. brevidactylus, B. cobos, B. elerae, B. isangdaliri, B. kadwa, B. lukbani, B. makusog, B. minimus, B. muntingkamay, and B. wrighti (Davis et al. 2014). 

We have evaluated this species against the International Union for Conservation of Nature (IUCN) criteria for classification and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Not only does B. ilocandia sp. nov. occur on more than one island, but also the species appears common in secondary growth and disturbed habitats throughout its recognized distribution. Therefore, we recommend that this species be classified as Least Concern, LC (IUCN 2015).

Etymology. The specific epithet is chosen in reference to the biogeographically and culturally distinct homeland, “Ilocandia”, of the Ilokano people of the northern Philippines, the third largest ethnolinguistic group in the country. Including the Babuyan Island Group north of Luzon Island, Ilocandia stretches from the western coast of northern Luzon, across the Cagayan Valley, to parts of central Luzon and the boundaries of Aurora Province in the east. The region is home to many endemic vertebrates, diverse geographic landscape, unique local cuisine, and rich cultural traditions. Suggested common name: Ilokano Slender Skink.

Cameron D. Siler, Drew R. Davis, Elyse S. Freitas, Nicholas A. Huron, Aaron D. Geheber, Jessa L. Watters, Michelle L. Penrod, Monica Papeș, Andrew Amrein, Alyssa Anwar, Dontae Cooper, Tucker Hein, Annalisa Manning, Neeral Patel, Lauren Pinaroc, Arvin C. Diesmos, Mae L. Diesmos, Carl H. Oliveros, and Rafe M. Brown. 2016.
Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) II: A New Species from the northern Philippines.
Zootaxa. 4132(1); DOI: 10.11646/zootaxa.4132.1.2

[Herpetology • 2016] Brachymeles ligtas • Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Squamata: Scincidae) I: A New Species from Lubang Island

Lubang Slender Skink |  Brachymeles ligtas

  Geheber, Davis, Watters,  Penrod, Feller, Davey, Ellsworth, Flanagan, Heitz, Moore, Nguyen, Roberts, Sutton, Sanguila, Linkem, Brown & Siler, 2016

DOI: 10.11646/zootaxa.4132.1.1 

Abstract

A new species of slender skink is described from the Philippines. The species is endemic to Lubang Island, and is assigned to the Brachymeles bonitae Complex based on phenotypic and genetic data. Specimens were collected from Lubang Island between 1991 and 2012, and were examined based on morphological data (qualitative traits, meristic counts, and mensural measurements). Published genetic sequence data from phylogenetic studies of the genus reveal the new species to be highly divergent from congeners. Brachymeles ligtas sp. nov. is differentiated from other members of the genus based on a number of distinct morphological features, including small body size (SVL 60.7–79.6 mm), bidactyl fore-limbs, digitless hind limbs, high number of presacral vertebrae (50), and the absence of auricular openings. Additionally, the new species has diagnostic, distinct dorsal head scale patterns. This new species becomes the only member of the genus known to occur on the deep-ocean island of Lubang.

Keywords: Reptilia, biodiversity, endemism, fossorial, limb reduction, non-pentadactyl, pentadactyl, Philippines

FIGURE 3. Photograph of holotype in life of Brachymeles ligtas sp. nov. (PNM 9818). Note: Individual is about to shead, resulting in lighter scale coloration.

Photograph taken by Cameron D. Siler 

DOI: 10.11646/zootaxa.4132.1.1

Etymology. The specific epithet is derived from the Tagalog (Filipino) term "nakaligtas," meaning "survivor" and "ligtas," meaning "salvation." We name this species in honor of the people of Lubang Island who endured nearly three decades of violence and guerrilla warfare, from 1945 to 1974, led by the Imperial Japanese Army intelligence officer Hiroo Onoda, and four Japanese soldiers. After being driven into the jungle of Lubang Island by allied forces near the end of World War II, Onoda resisted surrender for 29 years believing the war was not yet over. Onoda would finally surrender in 1974, allowing the communities of Lubang to move on from the hardships faced during this time period, including the loss of over 30 lives and injuries to dozens more. Suggested common name: Lubang Slender Skink.

Distribution. Brachymeles ligtas sp. nov. is currently known only from Lubang Island (Fig. 1) and we expect it is found on several smaller nearby landmasses.

Natural history. Brachymeles ligtas sp. nov. likely once occurred in low- to mid-elevation primary forest habitats. As most primary forest on Lubang Island has been destroyed, the recent observations of this species have occurred in secondary growth forest habitats. In contrast to the other members of the B. bonitae Complex, this species appears to be relatively common in secondary growth forest fragments on the island. To date, no other congeners have been documented on Lubang Island. 

We have evaluated this species against the International Union for Conservation of Nature (IUCN) criteria for classification and find that it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Although B. ligtas sp. nov. is known from a single island only, the species appears relatively common in secondary growth forest on Lubang, and until additional data are presented to support otherwise, we classify this species as Least Concern (LC; IUCN 2015).

 Aaron D. Geheber, Drew R. Davis, Jessa L. Watters,  Michelle L. Penrod, Kathryn D. Feller, Conner S. Davey, Elyse D. Ellsworth, Rachel L. Flanagan, Brendan B Heitz, Tana Moore, Marie D.C. Nguyen, Austyn Roberts, John Sutton, Marites B. Sanguila, Charles W. Linkem, Rafe M. Brown and Cameron D. Siler. 2016. Additions to Philippine Slender Skinks of the Brachymeles bonitae Complex (Reptilia: Squamata: Scincidae) I: A New Species from Lubang Island. Zootaxa. 4132(1); 1-14. DOI: 10.11646/zootaxa.4132.1.1

[Herpetology • 2012] Did Gekko Geckos Ride the Palawan Raft to the Philippines?

Abstract

Aim: We examine the genetic diversity within the lizard genus Gekko in the Philippine islands to understand the role of geography and geological history in shaping species diversity in this group. We test multiple biogeographical hypotheses of species relationships, including the recently proposed Palawan Ark Hypothesis.

Location: Southeast Asia and the Philippines.

Methods: Samples of all island endemic and widespread Philippine Gekko species were collected and sequenced for one mitochondrial gene (NADH dehydrogenase subunit 2) and one nuclear gene (phosducin). We used maximum likelihood and Bayesian phylogenetic methods to derive the phylogeny. Divergence time analyses were used to estimate the time tree of Philippine Gekko in order to test biogeographical predictions of species relationships. The phylogenetic trees from the posterior distribution of the Bayesian analyses were used for testing biogeographical hypotheses. Haplotype networks were created for the widespread species Gekko mindorensis to explore genetic variation within recently divergent clades.

Results: Both maximum likelihood and Bayesian phylogenetic analyses indicated that Philippine Gekko species are a diverse clade with a long history in the archipelago. Ancestral range reconstruction and divergence time analyses suggest a Palawan microcontinental origin for this clade, coinciding with Palawan’s separation from Asia beginning 30 Ma, with subsequent diversification in the oceanic Philippine islands. The widespread species G. mindorensis and G. monarchus diversified in the late Miocene/early Pliocene and are potentially complexes of numerous undescribed species.

Main conclusions: 

The view of the Philippine islands as a ‘fringing archipelago’ does not explain the pattern of species diversity in the genus Gekko. Philippine Gekko species have diversified within the archipelago over millions of years of isolation, forming a large diverse group of endemic species. Furthermore, the Philippine radiation of gekkonid lizards demonstrates biogeographical patterns most consistent with stochastic colonization followed by in situ diversification. Our results reveal the need to consider deeper time geological processes and their potential role in the evolution of some Philippine terrestrial organisms.

Keywords: Biogeography; Buruanga Peninsula; diversification; Gekko; islands; lizards; Palawan Ark Hypothesis; Philippines; rafting; Southeast Asia

Cameron D. Siler, Jamie R. Oaks, Luke J. Welton, Charles W. Linkem, John C. Swab, Arvin C. Diesmos and Rafe M. Brown. 2012. Did Geckos Ride the Palawan Raft to the Philippines? Journal of Biogeography. 39(7); 1217–1234. DOI: 10.1111/j.1365-2699.2011.02680.x

http://www.nhm.ku.edu/rbrown/Siler.etal.2012.Gekko.J.Biogeog.pdf