BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research
libraries, and research funders in the common goal of maximizing access to critical research.
Taxonomic Reappraisal of the Northeast Mindanao Stream Frog, Sanguirana
albotuberculata (Inger 1954), Validation of Rana mearnsi, Stejneger 1905, and
Description of a New Species from the Central Philippines
Author(s): Rafe M. Brown, Allyson Prue, Chan Kin Onn, Maren Gaulke, Marites B. Sanguila, and
Cameron D. Siler
Source: Herpetological Monographs, 31(1):182-203.
Published By: The Herpetologists' League
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and
environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published
by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries
or rights and permissions requests should be directed to the individual publisher as copyright holder.
Herpetological Monographs, 31, 2017, 182–203
Ó2017 by The Herpetologists’ League, Inc.
Taxonomic Reappraisal of the Northeast Mindanao Stream Frog, Sanguirana albotuberculata
(Inger 1954), Validation of Rana mearnsi, Stejneger 1905, and Description of a New Species
from the Central Philippines
RAFE M. BROWN
,CHAN KIN ONN
,MARITES B. SANGUILA
,AND CAMERON D. SILER
Department of Ecology and Evolutionary Biology and Biodiversity Institute, University of Kansas, Lawrence, KS 66045-7561, USA
Haskell Indian Nations University, Lawrence, KS 66045, USA
GeoBio Center LMU, Richard-Wagner-Strasse 10, 80333 M ¨
Biodiversity Informatics and Research Center, Father Saturnino Urios University, San Francisco Street, 8600, Butuan City, Philippines
Department of Biology and Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK 73072-7029, USA
ABSTRACT: With a published multilocus phylogenetic analysis as our guide, we use new data from the external phenotype and genetically
defined distributions of evolutionary lineages to resolve species boundaries associated with the southwest Mindanao Stream Frogs, Sanguirana
everetti (Boulenger 1882), its junior synonym, Rana mearnsi, Stejneger 1905, and the northeast Mindanao Stream Frogs, Sanguirana
albotuberculata (Inger 1954). Consideration of relationships, distributions, type localities, phenotypic data, and type specimens clearly indicates
that the names R. mearnsi and S. albotuberculata refer to the same lineage, and we recognize the oldest available name (Sanguirana mearnsi) for
this species. We also define the central Philippine lineage (from Negros, Masbate, and Panay islands) as a distinct new species. Long confused
with S. everetti, the new taxon is readily diagnosed and biogeographically restricted to the West Visayan faunal region. The new multilocus
estimate of phylogeny and our multivariate analysis of morphological variation demonstrate that the new species is closely related and
phenotypically most similar to northern Philippine Sanguirana luzonensis, to the exclusion of S. everetti, the southern species with which it
previously was confused. Morphological characters distinguishing the new species include body size, the absence of infracloacal tubercles, the
presence of smooth dorsal skin without dorsolateral folds or dermal asperities, its degree of sexual size dimorphism, uniquely stratified flank
coloration, bright white subarticular tubercles, bold pectoral patches, dark transverse bars on the limbs, and various body proportions.
Recognition of this new species further emphasizes the predictable nature of island bank-structured endemism in the Philippines and
demonstrates that the country’s vertebrate diversity remains underestimated. The new species is relatively rare, patchily distributed and, with so
little natural forest remaining in the central Philippines, it constitutes an immediate conservation concern. Management of this problem will
require continued, field-based collection of data on the new species, distribution, local abundance, population trends, natural history,
reproductive biology, and larval ecology—most of which currently is unknown.
Key words: Anuran biodiversity; Biogeography; Cascade Frogs; Endemicity; Ranidae; Sanguirana acai sp. nov.; Slender Stream Frogs
PHILIPPINE amphibian diversity currently consists of 112
species, with most (~85%) of these taxa endemic to the
archipelago (Brown 2007; Diesmos and Brown 2011; Die-
smos et al. 2014, 2015). Rates of species discovery in the
archipelago show no signs of slowing (Brown et al. 2008,
2013a; Brown and Stuart 2012; Diesmos et al. 2015), and
nearly 30% of the country’s amphibian fauna have been
discovered and described in the last 2 decades (Diesmos and
Brown 2011; Brown 2015; Diesmos et al. 2015). Unfortu-
nately, more than a third of the archipelago’s species have
been found to qualify for formal threatened status at some
level (Diesmos et al. 2014; IUCN 2015).
Endemic ranoid frogs are particularly diverse with at least
11 species of Limnonectes and 2 species of Occidozyga
(Dicroglossidae; Taylor 1920, 1922; Evans et al. 2003; Siler
et al. 2009; Setiadi et al. 2011), 32 or more Platymantis
(Ceratobatrachidae; Siler et al. 2010; Brown et al. 2015a,b), 2
species of Staurois (Ariﬁn et al. 2011), and 13 native species
of ranids (Brown 2007; Diesmos et al. 2015). Excluding
introduced species such as Hoplobatrachus rugulosus,
Hylarana erythraea, and Lithobates catesbeianus (Diesmos
et al. 2006, 2015; Brown 2007), Philippine ranids are divided
into three genera: Pulchrana with ﬁve species (Brown and
Guttman 2002; Brown and Siler 2013; Brown 2015),
‘‘Amnirana,’’ consisting of one nonendemic native species
(Inger 1954, 1999; Brown and Alcala 1970; Oliver et al. 2015;
Diesmos et al. 2015; Chan and Brown 2017), and
Sanguirana, containing seven species formerly referred to
the Rana everetti Complex (Brown et al. 2000a; Brown 2007;
Fuiten et al. 2011; Brown et al. 2016). Philippine Pulchrana
and Sanguirana are found on most major islands of the
archipelago and are distributed in accordance with geolog-
ically deﬁned (Inger 1954; Voris 2000) biogeographic regions
known as Pleistocene Aggregate Island Complexes (PAICs;
Brown and Diesmos 2002, 2009; Brown et al. 2013a), with
most PAICs possessing at least one widespread species
(Inger 1999; Brown et al. 2000a, 2016; Brown and Siler
2013) and the largest islands (Luzon and Mindanao)
supporting 2–4 species, with distributions structured geo-
graphically and/or along elevational gradients (Taylor 1922;
Brown et al. 2000a, 2016; Brown 2015; Fuiten et al. 2011).
Boulenger (1882) described Rana everetti from an
unspeciﬁed type locality of ‘‘Zamboanga’’ (an elongate
peninsula of western Mindanao Island, southern Philippines;
Fig. 1A), and more than 2 decades later Stejneger (1905)
named Rana mearnsi from the mountains of eastern
Mindanao. Taylor (1920) recognized both of these taxa and
named a third Mindanao Stream Frog, Rana dubita, from
Bunawan, east-central Mindanao. Particularly important for
the current study, in the same work he also referred some of
his own specimens from an allopatric population on southern
Negros Island (Fig. 1A) to Rana mearnsi.
When Inger (1954) later synonymized Rana mearnsi and
R. dubita with R. everetti, he characterized the Negros
Island population as conspeciﬁc with the Mindanao popu-
lation. He conceived of R. everetti as a polytypic taxon
containing three subspecies: R. e. everetti Boulenger 1882,
R. e. luzonensis Boulenger 1896, and R. e. albotuberculata
Inger 1954. The West Visayan populations (Negros, Mas-
bate, and Panay islands) have resided in synonymy with
Sanguirana everetti ever since (Brown and Alcala 1970;
Sison et al. 1995; Ferner et al. 2000; Brown et al. 2000a,b,
2016). This arrangement has persisted, but without explicit
scrutiny of its underlying assumptions, and despite the fact
that under the prevailing biogeographic framework (Brown
and Alcala 1970; Brown and Diesmos 2009; Brown et al.
2013a), such a distribution is highly anomalous because it
spans widely allopatric yet restricted geographic regions on
multiple PAICs (Brown and Guttman 2002; Brown and
Diesmos 2009; Brown and Siler 2013).
Meanwhile, recent taxonomic works have recognized all of
Inger’s former subspecies as full species (Brown et al. 2000a;
Brown 2007; Diesmos et al. 2015), resurrected Taylor’s
(1922) Rana igorota (Brown et al. 2000a), and described two
additional species, S. tipanan (Brown et al. 2000a) and S.
aurantipunctata (Fuiten et al. 2011). Most recently, follow-
ing higher-level phylogenetic analyses (Wiens et al. 2009),
Fuiten et al. (2011) expanded and augmented the deﬁnition
of the genus Sanguirana (Dubois 1992; Brown et al. 2000a)
to include the Palawan Wood Frogs S. sanguinea (Boettger
1893). This Palawan PAIC endemic had previously been
considered a Papuan-derived Philippine faunal element
(Inger 1954; Dubois 1992); the morphological and biogeo-
graphic distinctiveness of S. sanguinea most likely led to this
view never being challenged by anuran taxonomists (Inger
1954; Fuiten et al. 2011). Recent phylogenetic analyses
(Brown et al. 2016) demonstrate that S. sanguinea is actually
the ﬁrst-diverging lineage in a Palawan–Ark-facilitated
biogeographic diversiﬁcation scenario (Blackburn et al.
2010; Siler et al. 2012), suggesting that Sanguirana ﬁrst
diversiﬁed on the isolated Palawan Micro-continental Block
(Zamoros et al. 2008; Yumul et al. 2009a; Aurelio et al. 2013)
before undergoing range expansion via overseas dispersal
after colonization of multiple oceanic Philippine landmasses
(Brown et al. 2016; Chan and Brown 2017).
The genus Sanguirana now consists of species with largely
allopatric distributions including S. albotuberculata from
Leyte, Samar, and eastern Mindanao islands (Sanguila et al.
2016); S. aurantipunctata from a few sites in the mountains
of central Luzon Island (Fuiten et al. 2011); S. everetti from
southwestern Mindanao Island (Inger 1954); S. sp. (‘‘ S. cf.
everetti’’) from Negros, Masbate, and Panay islands (Sison et
al. 1995; Ferner et al. 2000; Gaulke 2011); S. igorota from
the Cordillera Mountain Range of Luzon Island (Taylor
FIG. 1.—(A) Distributions of the eight species of the Philippine endemic genus Sanguirana with vouchered localities indicated with symbols colored when
corresponding to genetic samples; white in cases where no genetic data are available; m ¼type locality of Rana mearnsi (Baganga River); a ¼type locality of
Rana everetti albotuberculata (Cabalian); Pleistocene Aggregate Island Complexes (PAICs; Brown and Diesmos 2009) indicated with incremental gray
shading (key). (B) Multilocus Bayesian phylogenetic estimate of evolutionary relationships in the genus Sanguirana (from Brown et al. 2016); black dots at
nodes indicate strongly supported clades (likelihood bootstraps 70%; posterior probabilities 0.95); gray node moderately supported (,70% / .0.90);
symbols at branch tips correspond to those plotted on map (A) and question marks at tree tips indicate populations of uncertain taxonomic status. See text
and Brown et al. (2016) for additional details. A color version of this ﬁgure is available online.
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
1920); S. luzonensis from throughout most islands of the
Luzon PAIC (Brown et al. 2000a, 2016); S. sanguinea from
the Palawan PAIC landmasses (Boulenger 1894; Inger 1954;
Brown 2007); and S. tipanan from the Sierra Madre
Mountain Range, of Luzon Island (Brown et al. 2000a;
Fuiten et al. 2011; Fig. 1).
Molecular phylogenetic analyses strongly support the
monophyly of the clade and conﬁrm the inclusion of S.
sanguinea as part of this genus (Bossuyt et al. 2006; Stuart
2008; Wiens et al. 2009; Holder et al. 2010). However, a
recent multilocus phylogenetic study demonstrated that the
West Visayan populations form a highly divergent, well
supported clade unrelated to Sanguirana everetti, precluding
their continued identiﬁcation as that taxon and necessitating
this study (Brown et al. 2016).
In this paper we reconsider the taxonomic status of the
lineage from the northeast Mindanao faunal region, S.
albotuberculata (Inger 1954), in light of genetically veriﬁed
species distributions (Brown et al. 2016), relevant type
localities, phenotypic variation, and examination of the
name-bearing type specimens of these taxa. We ﬁnd the
substitution of S. mearnsi (Stejneger 1905) for S. albotu-
berculata (Inger 1954) advisable at this time and we place
the latter in synonymy with the former. We also revisit the
issue of the biogeographically anomalous West Visayan
(Negros, Masbate, and Panay islands) populations of
‘‘Sanguirana everetti’’ and ﬁnd character-based morphomet-
ric, biogeographic, and genetic evidence for the recognition
of a new central Philippine endemic species.
MATERIALS AND METHODS
Morphological Character Differences
Specimens of all species of the genus Sanguirana were
examined (Appendix; museum institutional codes/acronyms
follow Sabaj 2016) and data from types were incorporated
into deﬁnitions and diagnoses presented here. Specimens
were examined for the presence–absence of diagnostic
morphological character states including color pattern, body
proportions, nuptial pad shape, digital characters, dermal
asperities, infracloacal tubercles, dermal ﬂanges along limbs,
and raised dorsolateral ridges (Taylor 1920; Inger 1954;
Brown et al. 2000a; Fuiten et al., 2011).
Sex was determined by body size (for mature females),
the presence–absence of conspicuous secondary sexual
characteristics (nuptial pads in males; male Sanguirana lack
vocal sacs), and/or by gonadal inspection in the case of
specimens of intermediate sizes.
Frog vocalizations were recorded with an analog tape
recorder (Sony WM DC6 Professional Walkman) with a
directional microphone (Sennheiser ME80 condenser mi-
crophone, equipped with K3U power module). Calls were
recorded at distances of 1–3 m after which ambient and
cloacal temperatures were collected. Calls were digitized and
analyzed with Raven Pro v1.5 (Bioacoustics Research Group,
Cornell Lab of Ornithology, Ithaca, NY) software set on
default spectrogram parameters (256 samples and 50%
overlap). We examined oscillograms (waveforms), audio-
spectrograms (sonograms), and results of the Fast Fourier
Transformation (FFT; power spectrum) for a series of
spectral and temporal call characteristics following Brown
and Guttman (2002) and Brown and Gonzales (2007). Calls
are archived at the Cornell Laboratory of Ornithology
Macaulay Library (ML) under ML digital media catalog
numbers 224181 and 224348.
Analyses of Continuously Varying Phenotypic Variation
To examine Sanguirana populations for lineage-based
structure in continuously varying morphometric characters,
we supplemented published morphological and mensural
data for the genus Sanguirana (Brown et al. 2000a; Fuiten et
al. 2011) with new data from all species and exhaustive
sampling of Sanguirana albotuberculata from northeast
Mindanao, Leyte, and Samar islands and S. cf. everetti from
Negros, Masbate, and Panay, islands (Appendix). We
included all named Sanguirana from the oceanic Philippine
islands, the West Visayan population of ‘‘ S. everetti’’ (the
new species), and excluded only S. sanguinea from Palawan
Island, a species shown to be highly morphologically distinct
from congeners (Inger 1954; Brown et al. 2000a; Fuiten et al.
2011). We treated S. luzonensis as two putative operational
taxonomic units (OTUs) on the basis of observed variation in
phylogenetic analysis resulting from mitochondrial (mtDNA)
and nuclear (nDNA) DNA datasets (Brown et al. 2016). In
our previous study (Brown et al. 2016), we observed
moderately supported incongruence between mtDNA and
nDNA datasets, suggesting that S. igorota and S. tipanan
may be nested within S. luzonensis, with some northern
Luzon Island populations of S. luzonensis sister to a ([S.
igorota,S. tipanan] southern S. luzonensis) clade. Thus,
given that we did not ﬁnd strong support for the monophyly
of all S. luzonensis populations, and the possibility that some
northern S. luzonensis populations could be a distinct
evolutionary lineage (but see Brown et al. 2016, for
discussion of other possibilities), we designated the northern
and southern Luzon populations as two OTUs for our
analysis of continuously varying morphometric variation.
We collected data for the following 19 mensural
characters following the character deﬁnitions of Brown et
al. (2000a) and Fuiten et al. (2011): snout–vent length (SVL),
head, and snout lengths; head width, interorbital and
internarial distances; eye and tympanic annulus diameters;
lengths of forearm, femur, tibia, tarsus, foot, and hand,
Finger I, Finger III, and Toe IV; Finger III and Toe IV
terminal disk widths; and nuptial pad length. All measure-
ments (taken by AP and RMB only to reduce intermeasurer
bias; Lee 1982, 1990; Hayek et al. 2001) were measured to
the nearest 0.1 mm (with digital calipers and stereomicro-
scope when necessary) from sexually mature adult males;
data were excluded to minimize the impact of allometric
ontogenetic variation (juveniles) and due to insufﬁcient
sample sizes among all groups (females).
Prior to analyses, measurements were corrected for
differences in ontogenetic composition (Thorpe 1983a) using
the following allometric equation: X
), where X
is the adjusted value of the morpho-
metric variable and Xis the original value; SVL ¼snout–vent
¼overall mean SVL length; and bis the
within-OTU coefﬁcient of the linear regression of each
original character value (X) against SVL (following Thorpe
1975, 1983b; Turan 1999; Chan et al. 2013). Based on the
values of b, a subset of 14 informative morphometric
184 Herpetological Monographs 31, 2017
characters were selected for inclusion in subsequent
analyses. These characters include SVL, head and snout
lengths, head width, tympanic annulus diameter, forearm,
femur, tibia, tarsus, foot, and hand width, Toe IV, Finger III
disc widths, and nuptial pad length. Separately conducted
Shapiro–Wilk tests indicated violations of the assumptions of
normality for SVL, head width, and lengths of snout, tibia,
tarsus, hand, nuptial pad length, and Toe IV disc width (P.
0.05), and Levene’s tests of homogeneity of variance
indicated most were heteroscedastic. Accordingly, we log-
transformed all data and then conﬁrmed that they satisﬁed
assumptions of normality and homoscedasticity before
performing subsequent multivariate analyses and analyses
of variance (ANOVAs) with post hoc Tukey tests (or Tukey–
Kramer tests in cases of unequal sample sizes) to identify
individual character differences among means of our seven
A principal component analysis (PCA) was performed to
ﬁnd the best low-dimensional representation of morpholog-
ical variation in the data and to further determine whether
continuous morphological variation could form the basis of
statistically detectable group structure. Principal compo-
nents with eigenvalues of 1.0 or higher were retained in
accordance to Kaiser’s criterion (Kaiser 1960). To further
characterize clustering and distance in morphospace, a
discriminant analysis of principal components (DAPC) was
performed for all congeners to ﬁnd the linear combinations
of morphological variables that have the largest between-
group variance and the smallest within-group variance. The
DAPC relies on data transformation using PCA as a prior
step to discriminant analysis (DA), ensuring that variables
included in the DA are uncorrelated and number fewer than
the sample size (Jombart et al. 2010). All analyses were
implemented and visualized using the statistical software
environment R v3.1.2 (R Core Team 2015). The DAPC
analysis was performed using the R package adegenet v2.0.0
We refer to the recently published study of Brown et al.
(2016) which included sampling from 161 individuals from
throughout the Philippine archipelago (47 localities) and
specimens of all currently recognized species of the genus
Sanguirana (Fig. 1B). That study included a concentrated
analysis of 6098 nucleotide positions from two mitochondrial
gene regions and six nuclear loci, and standard phylogenetic
analyses using likelihood (ML) and Bayesian (BA) methods.
Details of polymerase chain reaction temperature regimes,
manufacturer laboratory protocols, inference of nucleotide
substitution models, partitioning strategy, and details of
phylogenetic analyses are provided in Brown et al. (2016).
For simplicity, because ML and BA analyses produced
identical topological estimates, we summarize here just the
Bayesian estimate of phylogeny and posterior probabilities of
nodal support. All sequences are deposited in GenBank
(Brown et al. 2016: Supplemental Appendix).
We embrace the General Lineage Concept of species (de
Queiroz 1998, 1999) as the logical extension of the
Evolutionary Species Concept (Simpson 1961; Wiley 1978),
which has been articulated in a manner (de Queiroz 2005,
2007) that is particularly consistent with our deﬁnition of this
new species. A species is the most inclusive lineage segment
(ancestor–descendant series of metapopulations) identiﬁed
as distinct from other such lineages, within which there is
evidence of reproductive cohesion, for which we can infer a
unique evolutionary history and predict an independent
future evolutionary trajectory or ‘‘ fate’’ (Wiley 1978; Frost
and Hillis 1990; Brown and Diesmos 2002). We recognize as
distinct evolutionary lineages those ancestor–descendent
population segments that are (1) sympatric or parapatric
(occur on the same landmass) but with discrete, diagnostic,
phenotypic, and/or ecological character state differences,
and a genetic evidence of lineage cohesion (inferred absence
of reticulation or gene ﬂow with other sympatric congeners)
and, thus, lineages for which the hypothesis of conspeciﬁcity
can be rejected; or those that are (2) allopatric or
geographically isolated (i.e., as insular or PAIC endemic
lineages and, thus, demonstrably unique evolutionary
entities) and morphologically, ecologically, and/or genetically
For the purpose of recognizing the noncontroversial
evolutionary lineages of the Mindanao PAIC (Brown et al.
2000a, 2016), criterion (1), for example, is applicable: the
northeast Mindanao, Leyte, and Samar islands’ (Fig. 1)
lineage was recognized originally (described as Rana mearnsi
[Stejneger 1905]; see below), and later was redescribed
(Inger 1954; as Rana e. albotuberculata) with an accompa-
nying analysis of intraspeciﬁc mensural and meristic data.
This lineage originally was diagnosed as part of a polytypic
taxon (Inger 1954), and later redeﬁned as an evolutionary
species (Brown et al. 2000a) distinct from the parapatric
southwest Mindanao Island Sanguirana everetti (see Brown
et al. 2000a, 2016, for evolutionary species deﬁnition,
illustration of diagnostic characters, phylogeny, and biogeo-
graphical inference). Likewise, for the purpose of the new
species recognized here, criterion (2) is clearly applicable,
and the recognition of the new species is not surprising
because it represents a distinct evolutionary lineage on a
separate geological Pleistocene island bank platform and is
noncontroversial in that most widespread Philippine verte-
brate groups possess distinct species on separate PAICs
(Brown et al. 2000a, 2013a, 2016; Brown and Diesmos 2002,
Deﬁnition of Sanguirana and Assignment of Taxa
We follow the Fuiten et al. (2011) deﬁnition of the genus
Sanguirana, and we place taxa in this genus based on
phylogenetic evidence (Brown et al. 2016) and possession of
diagnostic character states. Members of the genus can be
distinguished from all other Philippine ranids (Inger 1954;
Diesmos et al. 2015) by the following combination of shared
characters: (1) thin, elongate body; (2) extremely expanded
terminal digital disks with circummarginal grooves; (3)
elongate nuptial pad, covering nearly entire medial portion
of the ﬁrst digit (Finger II), present (most species) or absent
(S. sanguinea); (4) absence of vocal sacs; (5) posterior
abdomen coarsely glandular; and (6) absence of humeral
glands (Boulenger 1882; Inger 1954; Taylor 1920; Brown et
al. 2000a; Fuiten et al. 2011).
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
Taxonomic Reappraisal of Sanguirana albotuberculata
(Inger 1954) and Rana mearnsi Stejneger 1905
In considering the status of West Visayan faunal region
species, we clarify the boundary between populations now
referred to Sanguirana everetti (a taxon now restricted to
western Mindanao Island; Fig. 1A; Brown et al. 2016: Fig. 1)
versus its sister species S. albotuberculata (Inger 1954;
Brown et al. 2000a) of Leyte, Samar, and eastern Mindanao
islands (Diesmos et al. 2015) and the unnamed evolutionary
lineage of the West Visayan islands. Confusion has resulted
from Taylor’s (1920) referral of the Negros population to
Rana mearnsi Stejneger 1905, combined with Inger’s
placement of Rana mearnsi in synonymy with R. everetti
everetti Boulenger 1882. Additionally, in the same work,
Inger (1954) named the northeast Mindanao faunal region
lineage as a new subspecies, Rana everetti albotuberculata.
We assume that the combination of these actions has
resulted in a historical delay in what might otherwise have
been a natural reconsideration of priority with regard to
available names for the eastern Mindanao, Leyte, and Samar
As a result of the Brown et al. (2016) phylogenetic study,
we have no doubt that Sanguirana mearnsi Stejneger 1905
has priority over, and is thus the valid name that must be
substituted for (a nomen substitutum), the northeast Mind-
anao PAIC species currently referred to as S. albotubercu-
lata (Inger 1954; Brown et al. 2000a; Diesmos et al. 2015).
We base this name substitution on the chronological order of
relevant publications and because several lines of evidence
indicate the names Rana mearnsi Stejneger 1905 and R.
everetti albotuberculata Inger 1954 refer to the same
First, the distribution of the species from the northeast
Mindanao PAIC is now very well documented (Fig. 1A;
Brown et al. 2016: Fig. 1; Sanguila et al. 2016), with
genetically conﬁrmed identities of fresh samples from
northern and central Samar Island, at numerous sites
through Leyte Island, and from sites along the northeast
coastal mountains of eastern Mindanao to the southeast
corner of the island (Brown et al. 2016: Fig. 1). Second,
the type locality of Rana mearnsi Stejneger 1905 (Baganga
River, Eastern Mindanao; .300 m above sea level;
Stejneger 1905; Cochran 1961) falls without any uncer-
tainty within this geographical span of genetically con-
ﬁrmed localities (Fig. 1A). Third, the distributions of S.
everetti (southwest Mindanao) and ‘‘S. albotuberculata’’
(¼S. mearnsi) are now well circumscribed, conﬁrmed
with documented genetic sampling, and demonstrably do
not overlap (Fig. 1A). Fourth, the Rana mearnsi Stejneger
1905 holotype (USNM 35258) is indistinguishable mor-
phologically from similarly sized ‘‘S. albotuberculata.’’
Finally, we note that Stejneger’s (1905) original descrip-
tion mentions character states used by Inger (1954) to
diagnose R. e. albotuberculata from R. e. everetti (distinct,
ﬂeshy glandular dorsolateral folds, prominent ‘‘pustules’’
[termed ‘‘asperities’’ in Inger 1954; see Brown et al.
Admittedly, the poor state of preservation of the Rana
mearnsi holotype (USNM 35258; preserved in blackberry
brandy, brittle, and broken into multiple pieces; as
originally reported by Stejneger 1905) now prevents
evaluation of some previously emphasized character states
(Taylor 1920; Inger 1954; Brown et al. 2000a). These
include the distribution of pustules/asperities on lateral
surfaces of the head, the shape of the nuptial pad,
morphometric variation, and live color or dorsum, thick
dorsolateral folds, and infracloacal tubercles (Inger 1954;
Brown et al. 2000a). However, all other evidence points to
a single hypothesis.
In summary, despite the absence of genetic material
from the exact type locality, the multiple lines of evidence
discussed above, plus examination of the relevant name-
bearing types, convinces us that Rana mearnsi Stejneger
1905 has priority over Rana everetti albotuberculata Inger
1954 and that Sanguirana mearnsi (Stejneger 1905) is the
ﬁrst available, valid name to be applied correctly to
populations of the Stream Frogs (Fig. 2) from the
northeast Mindanao PAIC (Leyte, Samar, eastern Mind-
anao, and most likely Bohol islands). In addition to Rana
everetti albotuberculata Inger 1954, Rana dubita Taylor
FIG. 2.—Live male (A) and female (B) Sanguirana mearnsi (formerly S.
albotuberculata [Inger 1954; Brown et al. 2000a, 2016]) from the
municipalities of (A) Burauen, Leyte Island, Leyte Province (deposited at
KU; RMB Field No. 21807; Photo by J. Fernandez), and (B) Gingoog City,
Mindanao Island, Misamis Oriental Province (KU 333014; Photo by RMB).
Note thickened dorsolateral dermal folds and rugose texture of skin (the
result of densely distributed keratinized asperities) in males (both character
states reduced in female) and the distribution of green pigment throughout
dorsal surfaces of males (limited to ventrolateral surfaces in females). A color
version of this ﬁgure is available online.
186 Herpetological Monographs 31, 2017
1920 (Type locality: Bunawan, eastern Mindanao) is also
hereby placed in synonymy with Sanguirana mearnsi
Taylor’s (1920) assignment of the name Rana mearnsi to
the West Visayan population (Negros Island) clearly was in
error, as noted correctly by Inger (1954). However, despite
the fact that he identiﬁed Taylor’s lapsus, Inger (1954) did
not formally act on the distinctiveness of the new species
from Negros, Masbate, and Panay islands. Given the
limited appreciation of among-faunal region variation at
that time (most Mindanao and Negros records were
referred to ‘‘Rana everetti everetti’’; Taylor 1922; Inger
1954), it is understandable that Inger (1954) conservatively
discounted the validity of R. mearnsi, placed it in synonymy
with R. e. everetti, and described the (same) species as R. e.
Furthermore, remarking on the paucity of available
specimens, Inger (1954:310) stated: ‘‘The Negros specimens
cannot be placed in any of the deﬁned subspecies with any
reasonable degree of assurance.’’ Acknowledging Inger’s
(1954) powers of observation and that his conservative
approach set the stage for this study, below we deﬁne the
unassigned population as a new species.
Continuously Varying Morphological Variation
Due to the similarity between the quantitative and
qualitative results for separately analyzed male and female
specimens, we report the details of the results for analyses
of males only. Although it took 10 principal components
(PCs) to account for .95% of the total variance, the ﬁrst
four PCs each had eigenvalues of more than 1.0 and
together accounted for 75% of the total variance (Table 1).
The ﬁrst principal component (PC1) loaded heavily on the
lengths of femur, tibia, tarsus, and feet, indicating that
differences in lower hindlimb morphology were responsible
for most of the variance (29.5%). The second (PC2; 19.5%)
loaded heavily on characters pertaining to head morphology
(head length, snout length, tympanic annulus diameter)
whereas PC3 and PC4 (26%) had signiﬁcant loadings for
the characters SVL, head width, forearm length, and
nuptial pad lengths. Ordination of the ﬁrst two components
showed taxon-based group structure evident in partial
separation between S. everetti versus S. igorata,S. tipanan,
FIG. 3.—Bivariate ordination of ﬁrst two components from a principal components analysis (PCA; A) and subsequent discriminant analysis of principal
components (DAPC; B) for 14 continuously varying morphometric variables (males only) selected by each variable’s within-species/OTU linear coefﬁcient
when regressed against snout–vent length (SVL). Character loadings (Table 1) indicate that distal limb dimensions contributed disproportionately to PC1
whereas dimensions related to head length contributed heavily to dispersion along PC2. See text for character deﬁnitions; pale polygon encompassing S. acai
points added to the PCA plot (A) for emphasis; inertia ellipses included in the DAPC plot (B) for emphasis. A color version of this ﬁgure is available online.
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
FIG. 4.—Adult male Sanguirana acai (paratype PNM 9807) and female (paratopotype KU 326383) in dorsal (A) and ventral (B) views. Scale bars ¼5 mm.
A color version of this ﬁgure is available online.
188 Herpetological Monographs 31, 2017
and S. mearnsi along the PC1 axis. The PC2 axis exhibited
separation between both S. mearnsi and S. luzonensis South
from both S. everetti and S. igorota (Fig.3A);additionally,
S. tipanan is distinct from S. igorota along this axis. The
new species clustered broadly in morphospace with S.
mearnsi,S. luzonensis North, and S. tipanan (Fig. 3A) along
both axes and, to a lesser extent with S. everetti and S.
luzonensis South. The DAPC analysis discriminated be-
tween groups, as expected, and supported S. mearnsi,S.
everetti,andS. igorata, and as distinct clusters, whereas the
new species and S. luzonensis North overlapped broadly
and the new species further overlapped minimally with S.
tipanan and S. luzonensis South (Fig. 3B).
Results of ANOVAs were highly signiﬁcant (P.0.0001)
for all 14 characters, and Tukey tests (or Tukey–Kramer
tests) detected differences among means of West Visayan ‘‘S.
everetti’’ and at least four (northern S. luzonensis) to as many
as all 14 (S. sanguinea) characters per species pair
The available multilocus estimate of phylogeny (Fig. 1B;
Brown et al. 2016) has demonstrated the phylogenetic
distinctiveness of the West Visayan islands (Negros,
Masbate, and Panay) population, which is not closely
related to S. everetti (the species with which it has long
been confused). Instead, this newly discovered lineage is
the sister lineage to a well-supported clade consisting of S.
igorota,S. tipanan, and two clades referred to S. luzonensis
(Fig. 1B; Brown et al. 2016). This strongly supported
estimate of genealogical afﬁnities bolsters the recognition of
the new species as distinct from all OTUs considered here
and leaves us with no doubt that that the West Visayan
islands ‘‘S. everetti’’ populations constitute a valid species,
new to science. For reference, mitochondrial uncorrected
genetic distances between the new species and all
congeners range from 6.4–12.1 (Table 2), which are
equivalent to or exceed those typically observed between
morphologically and acoustically well-differentiated anuran
lineages (e.g., Pulchrana moellendorfﬁ vs. P. mangyanum
FIG. 5.—Details of the palmar surfaces of the hand in Sanguirana acai (A,
male paratype PNM 9807; B, female paratopotype KU 326383) and plantar
surface of foot (C, D, same specimens). Scale bars ¼5 mm. A color version
of this ﬁgure is available online.
TABLE 1.—Character loadings for principal components (PC) analysis of 14 continuously varying morphometric characters, selected (from 19 total) on the
basis of each variable’s within-species/operational taxonomic unit (OTU) regression coefﬁcient (regressed against snout–vent length [SVL]). Heavily loading
characters in PC1 (lower limb dimensions) and PC2 (head shape), contributing disproportionately to group structure (see Fig. 1A), are bolded for emphasis.
Character PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9
SVL 0.172 0.088 0.434 0.058 0.378 0.3934 0.320 0.428 0.324
Head length 0.202 0.417 0.182 0.340 0.186 0.1077 0.053 0.024 0.113
Snout length 0.231 0.423 0.138 0.084 0.148 0.2394 0.257 0.078 0.270
Tympanic annulus diameter 0.126 0.475 0.224 0.204 0.068 0.0116 0.119 0.050 0.160
Head width 0.133 0.160 0.423 0.359 0.201 0.0735 0.100 0.451 0.44
Forearm length 0.028 0.358 0.392 0.008 0.261 0.2412 0.032 0.250 0.431
Femur length 0.419 0.065 0.099 0.327 0.090 0.0724 0.100 0.188 0.209
Tibia length 0.416 0.118 0.085 0.232 0.017 0.087 0.127 0.374 0.254
Tarsus length 0.380 0.078 0.066 0.015 0.000 0.7022 0.230 0.310 0.317
Foot length 0.422 0.172 0.127 0.064 0.084 0.055 0.072 0.048 0.136
Hand length 0.311 0.179 0.235 0.226 0.172 0.005 0.693 0.007 0.251
Nuptial pad length 0.156 0.068 0.406 0.126 0.705 0.089 0.319 0.189 0.224
Toe IV disc width 0.212 0.283 0.345 0.051 0.229 0.443 0.334 0.078 0.240
Finger III disc width 0.064 0.299 0.035 0.685 0.306 0.004 0.167 0.476 0.078
Standard deviation 2.02 1.65 1.61 1.07 0.83 0.71 0.68 0.67 0.64
Proportion 0.292 0.195 0.184 0.082 0.049 0.036 0.039 0.032 0.029
Cumulative 0.292 0.487 0.671 0.753 0.802 0.837 0.870 0.901 0.931
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
[Brown and Siler 2013]; Sanguirana igorota vs. S.
luzonensis [Brown et al. 2016]).
Justiﬁcation for the Recognition of a New Lineage-based
The new species clearly is distinct in multivariate space
from S. everetti,S. igorata,S. mearnsi, and southern
populations of S. luzonensis. With respect to these species/
OTUs, continuous variation of mensural body proportions
demonstrated discernable group structure (which lends
support to the recognition of the new taxon, emphasizing its
distinctiveness from most congeners). Separation was not
observed between the new species and northern S. luzonensis
populations or between the new species and S. tipanan (Fig.
3B). These allopatric northern Luzon populations are,
however, readily diagnosed from the new species on the basis
of ﬁxed color characters (see Diagnosis and Table 3).
The results of our previous phylogenetic analysis (dem-
onstrating the nonmonophyly of populations currently
referred to S. everetti and demonstrating the distinctiveness
of the West Visayan lineage from Luzon populations) require
the recognition of the new taxon. The fact that the
monophyletic West Visayan PAIC Sanguirana overlaps
broadly in morphospace with some Luzon taxa (northern
populations of S. luzonensis and S. tipanan) does not deter
us from recognizing it as a new species. This is because (1) it
is the monophyletic, strongly supported sister clade to a large
clade of three or four differentiated Luzon taxa (and not
closely related to S. everetti, the species with which it has
long been confused), and (2) it is isolated biogeographically
on the geologically separate West Visayan PAIC, which has
never been connected to the Luzon PAIC. Thus, even
without diagnostic continuously varying morphological traits
which distinguish it from all congeneric populations, we are
comfortable recognizing this allopatric, genetically distinct
evolutionary lineage as a taxon in which speciation has not
been accompanied by complete differentiation in continu-
ously varying morphological characters. However, in addi-
tion to the above, we have identiﬁed ﬁxed diagnostic
coloration characters (Table 3) that, together with phyloge-
netic and biogeographic evidence, support the recognition of
the West Visayan PAIC (Negros, Masbate, and Panay
islands) populations of ‘‘Sanguirana everetti’’ as a new
species, to be known as:
Sanguirana acai sp. nov.
Rana mearnsi Stejnegeri (1905): Taylor (1920: 251), in part.
Rana everetti Boulenger (1882): Sison et al. (1995: 48).
Rana cf everetti: Ferner et al. (2000: 12). [misidentiﬁcation].
Rana everetti everetti Boulenger (1882): Inger (1954: 310–
311), in part; Brown et al. (2000a: 85), in part.
Sanguirana everetti (Boulenger 1882): Fuiten et al. (2011:
99); Frost (2016). [misidentiﬁcation].
Holotype.—Adult male (PNM 9800, formerly KU
326381; Field Number RMB 3249), collected by RMB and
V. Yngente at 1745 hr on 14 April 2001, in the Philippines,
Negros Island, Negros Oriental Province, Municipality of
Valencia, Barangay Bongbong, below ‘‘ Camp Lookout,’’ in a
forested stream (‘‘Maite Creek’’) at 500 m elevation above
sea level on Mt. Talinis, Cuernos de Negros Mountain Range
(9.26678N, 123.20628E; Datum ¼WGS84).
Paratypes (paratopotypes).—Three adult males (TNHC
62794–96), adult male and female (KU 326382, 326383), all
with same collection data as holotype; two adult males
(USNM 228387 and CM 116128), same locality, collected by
C.A. Ross, 15 March 1981, and 10 August 1987, respectively.
Other paratypes.—Adult female (CAS-SU 16398),
collected by W.C. Brown, A.C. Alcala, and D. Empeso, 15
August 1954, Negros Island, Negros Oriental Province,
Municipality of Valencia, 4–5 km west of Valencia town, east
side Cuernos de Negros Mountain Range, Maite River
Gorge; adult female (CAS 131883), collected by Q. Alcala, 16
August 1963, same locality; ﬁve adult males (CAS 18144–48),
collected by D. Empeso, 28 April 1957, Municipality of
Dauin, 15 km north of Dauin Town, southwest side of
Cuernos de Negros Mountain Range; three adult males and
an adult female (TNHC 62797, 62798, KU 326382, and
326383) and two juveniles of undetermined sex (KU 326384,
326885), collected by RMB and V. Yngente, 14 April 2001,
Municipality of Valencia, Sitio Nasuji, Cuernos de Negros
Mountain Range, Mt. Talinis, 1150 m, PNOC/EDC
watershed area; two adult males (TNHC 62798, 62799),
collected by RMB and V. Yngente, 2 December 2001; one
adult male (USNM 228440), collected by C.A. Ross, 21
March 1980, and four adult females and two immature males
(CAS 137498–503), collected by L.C. Alcala and party, 19–
23 September 1972, Municipality of Sibulan, Barangay
Janya-janya, Sitio Balinsasayo, Cuernos de Negros Mountain
Range, Mt. Talinis 850–900 m above sea level, Lake
Balinsasayo; three adult males (CAS 138144, 147326,
147327), collected by Q. Alcala and party, 19–20 January
1964, Municipality of Palaypay, Barangay Pamplona; two
immature males (CAS 147328, 147329), and two adult males
(CAS 147330, 147331), collected by A. C. Alcala and party,
TABLE 2.—Uncorrected percent sequence divergence for mitochondrial data (12S–16S) among species of the genus Sanguirana. Intraspeciﬁc
mitochondrial sequence divergences along the diagonal are bolded for emphasis; note intraspeciﬁc divergence within S. acai (Negros vs. Panay populations).
S. acai S. mearnsi S. aurantipunctata S. everetti S. igorota S. luzonensis S. sanguinea S. tipanan
S. acai 0.0–6.3
S. mearnsi 6.8–9.9 0.1–1.8
S. aurantipunctata 9.0–10.9 6.7–9.0 4.5
S. everetti 8.0–9.1 4.0–5.9 8.4–9.3 0.1–0.6
S. igorota 6.4–7.8 6.4–8.7 9.0–10.3 7.3–8.2 0.9
S. luzonensis 6.0–7.3 5.3–8.3 8.6–9.7 7.0–7.7 4.1–5.0 0.6–3.5
S. sanguinea 10.8–12.1 10.3–12.1 12.2–13.1 10.6–11.5 11.3–12.0 10.8–11.2 5.8
S. tipanan 6.8–7.8 5.6–8.4 9.1–10.1 7.7–8.3 1.6–1.9 4.1–5.3 11.7–12.0 0.3
190 Herpetological Monographs 31, 2017
TABLE 3.—Distribution of selected diagnostic morphological characters in Sanguirana acai and all known congeners (þ¼present, ¼ absent, /þ¼variable). Geographical distribution and ranges of body
sizes (males, females: snout–vent length [SVL] in mm) are included for reference. Selected diagnostic mensural characters (expressed relative to body size, SVL; range, followed by ( ¯
X61 SD), are included
when comparison of ranges among the new species and one or more congeners (see text and Table 4, for details) contributed to recognition of the new species in the form of discrete character differences
Character S. acai S. everetti S. luzonensis S. aurantipunctata S. tipanan S. igorota S. mearnsi S. sanguinea
Range Negros, Masbate,
W. Mindanao Luzon PAIC Montane central
NE Luzon NW Luzon E. Mindanao, Samar,
Male SVL 45.8–57.6 61.7–79.5 43.4–67.1 47.3–53.7 46.0–53.5 49.6–58.3 58.3–68.6 36.3–42.8
Female 62.6–80.3 73.6–86.6 66.5–82.2 59.1–86.3 62.3–80.1 67.6–81.7 63.8–85.9 61.3–71.1
Flank coloration Sharp dark-light
Dark-light gradient Dark-light gradient Green-purple
Green with brown
Green with brown
Light to dark brown
Dorsal asperities þ, fine þ, fine þ, coarse
Infracloacal tubercles þþþþþ
Dorsal color Yellow, green, tan,
or light gray
Light green Brown, yellow,
green, tan, or light
with black flecks
or orange spots
Iridescent green or
Bright green with
Dark green with
Tibial bars þ/þþþþþ
Forearm bars þþþþþ
Dark lateral head
þCanthal stripe Canthal stripe Canthal stripe Canthal stripe Canthal stripe Canthal stripe
Snout Rounded Rounded Pointed Rounded Rounded Squarish Rounded Pointed
Ventrum tuberculate Limited to groin Groin Groin Throughout Groin Groin Groin –
Dorsolateral ridges Thin to indistinct Indistinct Indistinct Indistinct to
Moderate Thick, fleshy Moderate
Light on dark
Variable on dark Variable on dark Light on light Light on light Light on light Light on light Light on light
Snout/SVL 0.16–0.21 (0.19 6
0.16–0.18 (0.17 6
0.16–0.24 (0.20 6
0.15–0.17 (0.16 6
0.16–0.20 (0.18 6
0.14–0.16 (0.17 6
0.15–0.19 (0.17 6
0.17–0.19 (0.18 6
Head width/SVL 0.28–0.33 (0.31 6
0.31–0.33 (0.32 6
0.32–0.38 (0.34 6
0.33–0.38 (0.35 6
0.31–0.36 (0.34 6
0.30–0.36 (0.33 6
0.29–0.33 (0.31 6
0.31–0.33 (0.32 6
Forearm/SVL 0.22–0.30 (0.26 6
0.18–0.23 (0.20 6
0.18–0.31 (0.25 6
0.20–0.27 (0.21 6
0.19–0.26 (0.23 6
0.18–0.23 (0.21 6
0.18–.022 (0.21 6
0.19–0.22 (0.21 6
Tibia/SVL 0.58–0.68 (0.63 6
0.57–0.66 (0.62 6
0.53–0.70 (0.17 6
0.52–0.57 (0.54 6
0.52–0.62 (0.57 6
0.54–0.61 (0.57 6
0.52–0.69 (0.57 6
0.59–0.67 (0.63 6
Nuptial pad/SVL 0.10–0.15 (0.13 6
0.11–0.16 (0.14 6
0.13–0.19 (0.16 6
0.12–0.16 (0.14 6
0.11–0.17 (0.14 6
0.11–0.16 (0.15 6
0.13–0.16 (0.14 6
0.16–0.18 (0.17 6
Female:male SVL 1.3–1.4 (1.4 60.0) 1.2–1.5 (1.4 60.0) 1.3–1.4 (1.3 60.1) 1.3–1.4 (1.3 60.0) 1.3–1.5 (1.4 60.1) 1.1–1.1 (1.1 60.0) 1.1–1.2 (1.2 60.0) 1.8–2.1 (2.0 60.5)
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
21 December 1960, Pamplona town, east bank of Pinanlaya-
an River; adult female (CAS-SU 19541), collected by A.C.
Alcala and party, 27 December 1958, Municipality of Siaton,
Bantolinao, 4 km NW of Bondo Barrio; adult male (CAS
139275), collected by L.C. Alcala and party, 11 April 1962,
Negros Occidental Province, Municipality of Biak na bato, 6
km. N.W. Biak na Bato town, above Sition Tinago; adult and
immature male (CAS 185565, 185566), collected by L.C.
Alcala and party, 11 April 1962, Negros Occidental Province,
Municipality of Tuyom, Bagtik River; three adult males and
two adult females (CAS–SU 18134–38), collected by A.C.
Alcala and Q. Alcala, 12–21 April 1957, Municipality of
Tuyom, 17 km SW of Tuyom town, Bagtik River; adult
female (PNM 9801, formerly KU 323855), collected by
CDS, M. Yngente, V. Yngente, and J. Fernandez, 16 July
2009, Municipality of Silay City, Barangay Patag, Mt.
Bungol; two adult males and a juvenile of undetermined
sex (PNM 9802, 9803 [formerly KU 323860, 323862], and
KU 323918); six adult males and one adult female (KU
323861, 323864, 323866–70), same locality and collectors, 21
July 2009; three adult males (KU 323873–75), an adult male,
and two juvenile of undetermined sex (PNM 9804–06
[formerly KU 323863, 323865, 323871]), same collectors,
24 July 2009, same locality; three adult males (PNM 9807–
09, formerly KU 323872, 323876, and 323886), two adult
males and two adult females (KU 323856–59), and four adult
males (KU 323877–80), same collectors, 25 July 2009, same
locality; two adult females and three adult males (KU
323881–85), and two adult males, and a juvenile of
undetermined sex (KU 323887, 323888, 323918), collectors,
26 July 2009, same locality; two adult males (PNM 1372,
1373), collected by R.V. Sison, August 1991, Panay Island,
Aklan Province, Municipality of Libacao Nacolon, Barangay
Rosal, Sitio Belen; 22 males (PNM 3800–03, 3806–15, 3817–
24), collected by R.V. Sison, 27 February 1994, Antique
Province, Municipality of San Remigio, Barangay Aningalan,
Sitio Iganyao; two adult females and two adult males (KU
306863–66), collected by CDS, 13–15 March 2006, same
locality; adult female (PNM 3913), collected by R.V. Sison,
12 March 1994, Tipuluan Mountain Range; immature
female (PNM 8527), collected G. Operiano, 15 May 2004,
Municipality of Sebaste, Barangay Alegre; adult male (PNM
8550), collected by N. Paulino, 18 April 2004, Municipality
of Pandan, Sito Nanling.
Other referred specimens.—Immature male (CAS
124213), collected by L.C. Alcala and party, 6 May 1969,
Calagna-an Island, Iloilo Province, Municipality of Carles,
Barangay Barangcalan; three adult males (CAS 144267, CAS
144269, and USNM 305499), collected by L.C. Alcala and
party, 13–14 June 1976, Masbate Island, Masbate Province,
Municipality of Mobo, ‘‘ Mapuyo Barrio, Pulangkahoy’’ ; two
juvenile specimens of undetermined sex (FMNH 61530,
61531), collected by D.S. Rabor, 25 May 1949, on Negros
Diagnosis.—Sanguirana acai differs from all other
members of this Philippine endemic genus by the (1)
presence of dark pigmentation covering the majority of
lateral head surfaces (vs. absence or presence but limited to
a canthal stripe); (2) absence of dark color pattern on dorsum
and dorsolateral body surfaces (vs. presence); (3) presence of
an abrupt dark-above, light-below color stratiﬁcation (abrupt
transition) on the ﬂanks, the position of which is marked with
TABLE 4.—Results of Tukey post hoc tests (or Tukey–Kramer tests, for unequal sample sizes,) on individual morphometric characters, testing the hypothesis of no difference between means of Sanguirana
acai mensural characters versus those eight other OTUs (all considered full species except northern vs. southern populations of S. luzonensis; see text for details). Table entries include difference between
sample means (negative sign indicates comparisons in which congener means were large than those of Sanguirana acai) and P-values (in parentheses). Bolded text indicates signiﬁcant differences.
S. igorota S. mearnsi S. everetti S. tipanan S. luzonensis–South S. luzonensis–North S. aurantipunctata S. sanguinea
SVL 0.0491 (0.4245)0.1218 (,0.0001) 0.2355 (,0.0001) 0.1191 (,0.0001) 0.0290 (0.8366) 0.0542 (0.1512)0.0857 (0.0056)0.3320 (,0.0001)
Head length 0.0522 (,0.0001) 0.0080 (0.98124) 0.0285 (0.41816) 0.0079 (0.9707) 0.0641 (,0.0001) 0.0120 (0.8647) 0.1549 (,0.0001) 0.2945 (,0.0001)
Snout length 0.0891 (,0.0001) 0.0045 (0.9997)0.0084 (0.9991) 0.0037 (0.99987) 0.1186 (,0.0001) 0.0126 (0.9393) 0.1533 (,0.0001) 0.3194 (,0.0001)
Tympanum diam. 0.3009 (,0.0001) 0.1705 (,0.0001) 0.0285 (0.9998) 0.0776 (0.0008) 0.1450 (,0.0001) 0.0547 (0.0965)0.2762 (,0.0001) 0.2677 (,0.0001)
Head width 0.0674 (,0.0001) 0.0033 (0.9999) 0.0427 (0.0767) 0.0648 (,0.0001) 0.0496 (0.0005) 0.0499 (0.0003) 0.0492 (0.2131) 0.2929 (,0.0001)
Forearm length 0.0443 (0.2660) 0.0019 (0.9999) 0.1441 (,0.0001) 0.1097 (,0.0001) 0.1569 (,0.0001) 0.0249 (0.7621) 0.0715 (0.0340) 0.3430 (,0.0001)
Femur length 0.0159 (0.8951) 0.1034 (,0.0001) 0.0588 (0.0156) 0.0660 (,0.0001) 0.0405 (0.0126) 0.0400 (0.0114) 0.1422 (,0.0001) 0.3159 (,0.0001)
Tibia length 0.0794 (,0.0001) 0.0602 (,0.0001) 0.0515 (0.0395) 0.0826 (,0.0001) 0.0265 (0.2335) 0.0571 (,0.0001) 0.1975 (,0.0001) 0.3037 (,0.0001)
Tarsus length 0.0187 (0.9477) 0.0224 (0.8068) 0.0887 (0.0050) 0.0457 (0.0322) 0.0248 (0.7216) 0.0027 (0.9999) 0.1261 (0.0005) 0.2757 (,0.0001)
Foot length 0.0163 (0.9721) 0.0579 (0.0073) 0.1038 (0.0004) 0.1130 (,0.0001) 0.0223 (0.8080) 0.0131 (0.9806) 0.107(0.0069) 0.2845 (,0.0001)
Hand length 0.0425 (0.3602) 0.0144 (0.9842) 0.1411 (,0.0001) 0.0716 (0.00035) 0.0322 (0.5559) 0.0198 (0.9185) 0.0413 (0.5065) 0.5215 (,0.0001)
Nuptial pad length 0.0710 (0.0454) 0.0704 (0.0153) 0.0476 (0.7170) 0.0082 (0.9994) 0.0985 (,0.0001) 0.0651 (0.0272) 0.0533 (0.3422)0.1489 (0.0008)
Toe IV disc 0.1474 (0.0008) 0.2643 (,0.0001) 0.0728 (0.6940) 0.0082 (0.9999) 0.1130 (,0.0001) 0.0425 (0.8035) 0.0567 (0.5125)0.2476 (,0.0001)
Finger III disc 0.0860 (0.0476) 0.06636 (0.1283) 0.0413 (0.9275) 0.0039 (0.9999) 0.0716 (0.0003) 0.0448 (0.5473) 0.1939 (0.0004) 0.9838 (,0.0001)
192 Herpetological Monographs 31, 2017
a dark brown line or row of dark spots (vs. absence of abrupt
stratiﬁcation, transition gradual); (4) presence of transverse
dark bars on hind limbs but indistinct on forearms (vs.
absence or presence on both); (5) presence of uniquely dark
plantar surfaces of hand and foot, with bright white
subarticular and supernumerary tubercles (vs. more uni-
formly pigmented ventral hand and foot surfaces); and (6)
presence of boldly patterned, contrasting dark humeral
patches (vs. absence or indistinct).
Comparisons.—The critical comparisons for the diagno-
sis of the new species are to the distantly allopatric and
unrelated Sanguirana everetti, the taxon with which it has
long been confused taxonomically, and S. luzonensis, the
species to which it is most-closely related (Brown et al. 2016:
Fig. 1B), geographically most proximate, and phenotypically
most similar. From S. everetti the new species differs by its
much smaller, nonoverlapping body size (Tables 3, 4) and by
the absence of greatly enlarged infracloacal tubercles (vs.
presence in .90% of specimens); from S. everetti and S.
luzonensis by the presence of abruptly stratiﬁed ﬂank
coloration (vs. absence), presence of distinct white sub-
articular tubercles (Fig. 5A, B) on dark brown palmar and
plantar surfaces of the hand and foot in males (vs. tubercle
color similar to palmar and plantar surfaces of hands and
feet), and presence of bold humeral patches (vs. diffuse,
indistinct or absent; Fig. 4B). Relative forearm length nearly
distinguishes the new species from S. everetti and its
relatively narrower head width abuts, but does not overlap,
the range of variation observed in S. luzonensis (Tables 3, 4).
Sanguirana acai is distinguished from S. tipanan,S. igorota,
and S. mearnsi by the absence of dermal asperities on dorsal
and lateral body surfaces (vs. presence; Brown et al. 2000a:
Fig. 4), the presence of yellow, tan, or light gray dorsal
ground coloration (vs. iridescent green, with a brown
reticulum in S. tipanan [Brown et al. 2000a: Fig. 3C,D],
vibrant dark green with large dark brown osceli or purplish
spots in S. igorota [Brown et al. 2012a: Fig. 31] or metallic
bright green with bright yellow dorsolateral folds in S.
mearnsi [Diesmos et al. 2015: Fig. 39F]); further, from S.
igorota, the new species differs by its relatively shorter snout
and forearm (Tables 3, 4) and from S. mearnsi by its
relatively shorter forearm, by the absence of greatly enlarged
infracloacal tubercles, absence of raised, ﬂeshy dorsolateral
folds (vs. presence; Diesmos et al. 2015: Fig. 39H), and
presence of transverse tibial bars (vs. absence; Diesmos et al.
2015: Fig. 39F,H). The range of sexual size dimorphism
exhibited by the new species distinguishes it from both taxa
as well (Table 3). From S. aurantipunctata, the new species
is distinguished by having a pointed snout (vs. rounded;
Diesmos et al. 2015: Fig. 39G), a relatively narrower head
and shorter tibia (Table 3), glandular ventral texture limited
in distribution to the groin (vs. spanning entire ventrum),
yellow, green, tan, or light gray dorsal ground coloration (vs.
bright green-yellow with black ﬂecks [males] or bright
orange spots [females and some males]; Fuiten et al. 2011:
Fig. 2) and purple ﬂank coloration (vs. abruptly stratiﬁed
ﬂank coloration), by the presence of dark tibial bars (vs.
absent) and dark lateral head coloration (vs. bright green),
the absence of enlarged infracloacal tubercles (vs. presence),
and the absence of thickened postaxial dermal ﬂanges on
posterior surfaces of the hind limbs (vs. presence). Finally,
from S. sanguinea the new species differs by its larger,
nonoverlapping body size (Table 3), relatively shorter
forearm and nuptial pad (Table 3), having glandular (vs.
smooth) ventral texture around the groin, and by exhibiting
FIG. 6.—Sanguirana acai in life (from the Municipality of Valencia,
southern Negros Island): (A) adult male holotype (PNM 9800); (B) adult
female paratopotype (KU 326383). Photos by RMB.
FIG. 7.—Sanguirana acai, in life, photographed in the Municipality of
Sebaste, Antique Province, Panay Island (specimen not collected) in the
species, typical, stream-side vegetation perch microhabitat. Photo by MG. A
color version of this ﬁgure is available online.
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
less-pronounced sexual size dimorphism (female/male SVL
¼1.2–1.4 [S. acai] vs. 1.9–2.1 [S. sanguinea]).
Description of holotype.—Adult male in excellent state
of preservation (Fig. 4A,B). Snout pointed, but terminally
rounded in dorsal proﬁle and extending well beyond lower
jaw in lateral view; snout/head length ¼0.45; head width
narrower than body width, slightly wider than long; head
width/head length ¼0.76; head length/SVL ¼0.41; canthus
rostralis sharply angular, straight in dorsal aspect; loreal
region slightly concave; nares slightly protuberant laterally,
anterodorsal in position, visible from ventral aspect;
interorbital/internarial distance ¼0.87; interorbital dis-
tance/eye diameter ¼1.2; labial region thin, barely visible
in dorsal aspect; interorbital region ﬂat, wider than eye
diameter; rostrum ﬂat; eyes moderate in size, oriented
anterolaterally beyond jaw when viewed in ventral aspect,
protuberant on top of head; tympanum distinct, located
immediately behind eye; tympanum smaller than eye;
tympanic annulus/eye diameter ¼0.86; supratympanic ridge
slightly evident, continuous with barely evident dorsolateral
ridges; postrictal tubercles irregular, continuous, elongate,
arching ventrally, composed of enlarged ﬂeshy tubercles.
Dentigerous processes oriented transversely. Vomerine
teeth in row of four atop dentigerous process of each vomer;
dentigerous processes just posteromedial to choanae,
separate for a distance equal to width of one choana;
choanae moderate in size, suboval, widely separated, nearly
obscured by maxilla when viewed from ventral aspect;
premaxillary and maxillary teeth present; vocal slits absent;
tongue elongate (length twice that of width), free for two
thirds its length, posterior margin deeply notched.
Skin of dorsum smooth (Fig. 4A), asperities absent;
posterior two thirds of venter glandular; skin of cloacal
region coarsely glandular, especially adjacent to groin;
cloacal region lacking prominently enlarged infracloacal
tubercles; cloacal opening round, with transverse supra-
cloacal cutaneous ﬂap.
Upper arm slender; humeral glands absent; forearms
robust (Fig. 4A,B); forearm/hand length ¼0.70; forearm
length/SVL ¼0.24; ﬁngers in increasing order of length
II,III,V,IV (II much shorter than III); Fin2L/Fin4L ¼
0.47; interdigital webbing absent; lateral fringes present on
all digits of hand, most prominent on distal portions of
Fingers III–V; terminal phalanges widely dilated distally, 3–
53width of penultimate phalanges; disks with circum-
marginal grooves; ventral pads on Fingers III–V pointed,
protruding beyond distal edge of dorsal surface, visible from
dorsal aspect; penultimate phalanges with rounded supra-
articular cutaneous ﬂap.
Subarticular tubercles of hand large, raised, rounded,
protuberant (Fig. 5A); digit (Roman numerals) and tubercle
number (Arabic numbers): II (1), III (1), IV (2), V (2);
supernumerary tubercles present basally on each ﬁnger,
moderate in size, slightly raised, elongated on Fingers IV and
V, with medial constriction; thenar (inner palmar metacar-
pal) tubercle elongate, 0.43length of Finger II, separate
from medial and outer palmar tubercles; thenar tubercle
1.33length of large, subcircular, ﬂat medial palmar tubercle
and 2.13length of narrow, elongate outer metacarpal
tubercle (Fig. 4A); entire medial edge of thenar tubercle
covered by translucent, velvety nuptial pad; nuptial pad
continuing distally to just beyond articulation of penultimate
and ultimate phalanges; nuptial pad wrapping around
preaxial side of Finger II entirely and nearly in contact with
subarticular tubercle on its anterior edge; nuptial pad length/
Finger I length ¼0.98.
Hind limbs slender; tibia length/SVL ¼0.66; forearm
length/SVL ¼0.58; forearm/tibia length ¼0.87; tarsus/
forearm length ¼0.65; tarsus/foot length ¼0.64; foot/tibia
length ¼0.88; heels overlap when thigh segment of hind
limbs held at right angles to body; tibiotarsal articulation of
adpressed limb reaching beyond rostrum; toes long, in
increasing order of length I,II,IIIV,IV (III @V);
Toe4L/FL ¼0.79; toe disks smaller than those of ﬁngers;
Toe IV/Finger III disc width ¼0.56; interdigital webbing of
foot nearly complete (Fig. 4B), homogeneous, acrenulate;
modal webbing formula of toes (Savage and Heyer 1969,
–0V; webbing diminishing dis-
tally to form wide fringes along lateral edges of distal
phalanges on portions free of web; tarsal fold distinct,
continuous with postaxial dermal ﬂange on edge of Toe V;
subarticular tubercles of foot large, round or occasionally
subelliptical, nearly pointed; digit (Roman numerals) and
tubercle number (Arabic numbers): I (1), II (1), III (2), IV
(3), V (2); inner metatarsal tubercle oval, 33longer than
minute, round, outer metatarsal tubercle; supernumerary
tubercles absent from pes.
Measurements of holotype (mm).—SVL 52.1; head
length 21.3; head width 16.1; snout length 9.6; interorbital
distance 5.9; internarial distance 6.8; eye diameter 5.4;
tympanic annulus diameter 5; head width 16.1; forearm
length 12.4; femur length 30.2; tibia length 34.6; tarsus
length 19.6; foot length 30.5; hand length 17.6; Toe IV length
24.2; Finger I length 6.1; Finger III length 12.8; Toe IV disc
width 1.9; Finger III disc width 3.4; nuptial pad length 6.6.
Coloration of holotype in life.—(Based on ﬁeld notes
and photographs of RMB; see similarly patterned para-
topotype; Fig. 6A) Ground color of dorsal surfaces
homogenous light green; limbs slightly yellowish-green with
evenly distributed tiny, dark, grayish-purple spots and ﬂecks;
trunk with pale yellow pigment on faint dorsolateral folds (¼
faintly raised dermal ridges; Fig. 5A); dark transverse bars on
hind limbs (numbering four on femur, four on tibial segment
of limb); dorsal head color similar to body; pigment along
canthus rostralis, lateral head surface, pre- and postocular
regions, and tympanum solid dark brown; labial region bright
pale yellow, lightening to nearly white below eye, starkly
contrasting with dark brown lateral surfaces of head;
postrictal tubercles yellow.
Dorsolateral surfaces of body light green above, with
sharp transition lateral stratiﬁcation or transition to pale
yellow ventrolaterally; position of dark-above, light-below
ﬂank stratiﬁcation marked by ﬁne dark greenish-brown line
(Fig. 6A); lateral inguinal region heavily blotched with dark
gray markings on pale yellow background; tibio-tarsal
articulation bright white with ﬁne gray markings; dorsal
surfaces of hand and foot fade from lighter cream to white on
Finger I to yellowish-green on Finger II, then to dark gray
on Fingers III and IV; nuptial pad velvety gray; dorsal
humerus yellow between dark green transverse bands,
lightening to white by articulation with tibia; dorsal tibial
segment nearly white between dark green transverse bands;
dorsal surface of foot dark green, interdigital webbing dark
gray with faint darker patches of pigment.
194 Herpetological Monographs 31, 2017
Ventral surfaces lighter than dorsal surfaces; throat
homogeneous pale yellow; sternal region white with boldly
contrasting dark brown humeral patches; venter yellow
anteriorly, fading to cream with white glandular surfaces
Ventral surfaces of forearms white with starkly contrasting
dark brown ventrolateral coloration, darker distally at wrist;
palmar surface of hand dark brown, with grayish-purple
palmar and carpal tubercles and nuptial pad; ventral surfaces
of ﬁngers homogenous dark brown, with bright white
subarticular and supernumerary tubercles; ventral surfaces
of outer terminal ﬁnger discs light gray, ventral surfaces of
Fingers I and II discs pale yellow; ventral surfaces of femur,
tibia, and shank yellow with boldly contrasting dark brown
patches on posterior surfaces; tarsus purple; plantar surface
of foot purple with grayish-purple subarticular tubercles;
plantar surfaces of foot dark brown, with bright yellow to
cream subarticular tubercles, ventral toe discs white
proximally, dark gray distally; interdigital webbing of foot
dark brown, boldly patterned with distinct white patches
Coloration of holotype in preservative.—In preserva-
tive, the holotype’s color pattern has been retained, but
ventral colors have shifted to white or pale cream (yellow
lost), dark brown coloration somewhat lightened. Other than
loss of bright yellows and green (e.g., dorsal green coloration,
accent colors of the postrictal tubercles, and dorsolateral
ridges), difference between live and preserved coloration is
minimal (Fig. 4A,B).
Color variation.—Dorsal ground surfaces of body
varying shades of brown, from light brown (Negros Island
male KU 323885, 323887; female KU 323858; Panay Island
male KU 306863) to dark brown (Negros Island males KU
323868, 323873, 323883, 326382; Panay Island female KU
306864), immaculate or homogenous (most specimens) or
with distinct darker spots (Negros Island females KU
306649, 323882) or indistinct darker blotches (Panay Island
male KU 306863; Negros Island males KU 306437, 323675,
323866, 323869, 323880, 323885, 323887). Masbate Island
specimens (CAS 144267, 144269) are patterned more boldly
and exhibit stronger contrast between light and dark
pigmentation than do Negros and Panay specimens.
Most specimens have some transverse dark bars on tibial
and radio-ulnar segments of fore and hind limbs, respec-
tively. Five specimens lack dark bars on limbs altogether
(Negros Island males KU 306438, 323864, 323866, 323874;
Panay Island male KU 306863); specimens with dark dorsal
coloration have darkest transverse limb bars (Negros Island
males KU 323868, 323870, 323873, 323880, 323883, 326382;
Panay Island male KU 306865). Most remaining specimens
have faint transverse limb bars on all limbs, but some
specimens exhibit faint tibial bars and lack forearm bars
(Negros Island females KU 323858, 323859, 323881, 326383;
Negros Island males KU 323861, 323877, 323878–79; Panay
Island female KU 306864).
Ventral body surfaces range from light, immaculate cream
with dark pigment absent throughout (Negros Island males
KU 306437, 306438, 323859, 323864, 323869, 323875,
323884–85, 323887) to cream with distinct dark spots
scattered across all ventral surfaces and concentrated on
throat and pectoral region (Negros Island males KU 323873,
323877, 323883, female KU 323881). The remaining
specimens have scattered light brown and indistinct
speckling throughout ventral surfaces (Fig. 4B), some with
darker congregation of dark pigment on throat (Panay Island
males KU 306863, 306865–66; female KU 306864).
Lateral surfaces of heads grayish-blue, lacking canthal
stripes (most individuals) or with very faint canthal stripe
(KU 325898, 325905, 325912, 325916–17). Adult males
lacking transverse limb bars (most) or with thin, faint, light
gray bars (ﬁve or six) across forelimbs (KU 325913, 325916–
17, 325923, 325926, 325944, PNM 9735); hind limbs lacking
transverse bars (most) or with ﬁve or six thin, light gray bars
(KU 325913, 325926). Flank coloration more clearly
partitioned in females than in males, with sharper demar-
cation between dorsal grayish-blue and ventral grayish-pink.
Palmar surfaces of hand range from dark gray with
yellowish subarticular tubercles (Panay Island female KU
306864 and males KU 306863, 306865–66; Negros males KU
306437, 323866, 323868, 323873, 323878, 323880, 323883,
326382), to dark brown with bold white tubercles (Figs. 3A,
4A; males KU 323861, 323875, 323879, 323887), to very light
gray to yellowish-cream with little contrast between surface
of hand and subarticular tubercles (Fig. 4A, B; Negros Island
females KU 323857–59). Remaining (majority) specimens
have light gray palmar surface of the hand with distinguish-
able, brighter, yellowish-cream subarticular tubercles. Plan-
tar surface of the foot ranges from relatively homogeneous
dark gray-brown with yellowish-cream subarticular tubercles
(Panay Island female: KU 306864 and males KU 306863,
306865–66) or dark gray tubercles (Negros Island females:
KU 323858, 323867, 323881–82 and males KU 306437,
323866, 323868, 323870, 323873, 323877–79) to homoge-
neous light gray with slightly lighter subarticular tubercles
(Negros Island females KU 306649, 323856–57, 323859,
326383; males KU 306438, 323861, 323864, 323869,
323874–75, 323885, 323887).
Infracloacal rugosity slightly variable in size and shape
with the following exceptions: some possess minute tuber-
culation (KU 325896, 325903, 325912, 325919) whereas
others have enlarged and irregularly shaped glandular
patches (KU 325923, 325928, PNM 9733, 9736).
Distribution.—The new species is known from Negros,
Masbate, and Panay islands in the central Philippines (Fig.
1A). Other small islands of the West Visayan PAIC may also
harbor populations of Sanguirana acai, if appropriate habitat
can be located (e.g., Bantayan, Guimaras, Poro, San
Francisco), but we are reasonably certain that the new
species does not occur on Siquijor (several surveys in the last
10 yr have failed to detect its presence) and that it does not
occur—or no longer occurs—on the heavily deforested and
well-studied island of Cebu (Brown and Alcala 1970, 1986).
A population referred to S.‘‘ everetti’’ has been reported on
Bohol (Brown and Alcala 1970), but as of yet no genetic
tissue samples have been obtained and so its position in
phylogeny (Brown et al. 2016) cannot be ascertained. We
would expect, based on PAIC-structured Philippine bioge-
ography (Brown and Diesmos 2002, 2009), that the Bohol
population should be conspeciﬁc with the species docu-
mented on Leyte, Samar, and eastern Mindanao islands (S.
mearnsi), but this expectation remains untested. Sanguirana
acai has been documented from 375 m above sea level to
1350 m on the large mountains of southern Negros, northern
Negros, northwest Panay, and the western coastal mountains
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
of Panay (Taylor 1922; Inger 1954; Alcala 1962; Brown and
Alcala 1970; Ferner et al. 2000; Gaulke 2011).
Natural history.—Frogs of the genus Sanguirana are
stream breeders with indirect aquatic larval development
and reasonably well characterized larval biology (Taylor
1920, 1922; Inger 1954; Alcala 1962; Brown and Alcala
1982a,b; Brown et al. 2000a; Gaulke 2011). Individuals of
Sanguirana acai were found at night along forested
mountain streams or in disturbed, regenerating, or second-
growth forest, provided that it was adjacent to primary forest.
The new species perches on rocky stream banks, on
midstream boulders, and on rocks along lakeshores but is
most frequently encountered perched on branches and
leaves of streamside vegetation (Inger 1954; Ferner et al.
2000; Gaulke et al. 2008; Gaulke 2011). Brown and Alcala
(1955, 1961) described a variety of semiarboreal substrates
for this species, including branches a few meters high in
trees and away from water, but emphasized that ovulating
females primarily were located near water (lake shores and
pools of highly oxygenated streams). Eggs are not laid
together in masses but are scattered and adhere to rocks,
branches, pebbles, and other submerged debris (Alcala
1962). Gravid females carry between 800–1000 eggs (Alcala
1962; Brown and Alcala 1982b). Alcala (1962) provided a full
technical description of S. acai tadpoles including notes on
growth rates, morphological characteristics, diet, and behav-
ior. Gaulke (2011) described the live coloration of S. acai
tadpoles (bronze-green, with white scattered granules; larvae
have a maximum body length of nearly 70 mm) and
metamorphs (similar to that of adult). The new species
appears to have a relatively broad season of reproductive
activity; newly laid eggs and/or gravid females have been
collected from February to December although amplexus
has only been observed in April and May. Newly emerged
metamorphs have been collected in May, June, July, and
November (Alcala 1962; Gaulke 2011).
Species of Sanguirana lack vocal sacs (Inger 1954), but
vocalizations have been reported in breeding aggregations of
S. luzonensis (Brown et al. 2000b) and recently documented
in S. mearnsi (RMB, personal observations). To the best of
our knowledge, vocalizations of S. acai have not been
previously reported in the literature. Our recordings of the
new species include at least two distinct call types (see
Sympatric species of anurans that have been recorded
from at least parts of the new species range (Brown and
Alcala 1961, 1964; Gaulke 2011) include Kaloula pulchra
(introduced; Diesmos et al. 2015), K. picta (widespread,
endemic), K. conjuncta negrosensis (West Visayan PAIC
endemic), K. cf. kalingensis (West Visayan PAIC endemic
and potentially undescribed species; Blackburn et al. 2013),
Platymantis dorsalis (widespread, endemic), P. corrugatus
(widespread, endemic), P. negrosensis (West Visayan PAIC
endemic), P. hazelae (West Visayan PAIC endemic), P.
paengi (northwest Panay endemic), P. spelaeus (southern
Negros endemic), Limnonectes visayanus (West Visayan
PAIC and Romblon Island Group endemic), L. leytensis
(widespread endemic), Philautus surdus (widespread en-
demic), Kurixalus appendiculatus,R. pardalis (widespread
nonendemic natives; Brown and Alcala 1982a, 1994), and the
three introduced species Hoplobatrachus rugulosus,Rhinella
marina, and Hylarana erythraea (Diesmos et al. 2006, 2015).
Vocalizations.—The advertisement call of Sanguirana
acai has been recorded on two occasions. The ﬁrst segment
(9 April 2001; ML 224181) was recorded at ‘‘Camp
Lookout,’’ 500 m elevation (ambient temperature 22.98C;
cloacal temperature 248C), Barangay Bongbong, Municipal-
ity of Valencia (the type locality). The second segment (2
December 2001; ML 224348) was recorded at Lake
Balinsasayo, 865 m elevation (ambient temperature
20.18C), Barangay Janya-janya, Municipality of Sibulan. Both
sites are on the slopes of Mt. Talinis in the Cuernos de
Negros Mountain Range. In the ﬁrst instance, an adult male
(TNHC 62794; not vocalizing when ﬁrst observed), captured
at 2000 h and held in an inﬂated plastic bag inside a tent,
began calling at 0300 h the next morning, apparently
stimulated by the sound of light rain striking the tent (¼
Type 1, a dull, amplitude modulated ‘‘ rattle’’ call). Over a 6-
min period, TNHC 62794 called eight times and eventually
ceased as the shower abated. Twelve subsequent calls were
elicited artiﬁcially by RMB by simulating the approximate
frequency of the rain by wrinkling paper and shaking the
walls of the tent. The second unvouchered recording was
captured from a dugout canoe upon approaching the
lakeshore of Lake Balinsasayo (2000 h). In this instance,
two or three males were observed in close proximity to a few
larger females and surrounded by an estimated .15
additional males perched in nearby shrub-layer vegetation;
two distinct call types were captured. In this segment,
presumed advertisement calls (rattles) from two alternating
males are interspersed with numerous high frequency, brief,
tonal, frequency-modulated vocalizations (Type 2, chirping
‘‘peeps’’ and ‘‘ squeaks’’ ) from other males perched in close
proximity (RMB, personal observation).
The stereotyped presumed advertisement call vocalization
of S. acai is a moderately rapid, dull, amplitude-modulated
pulsed train, sounding to the human ear like a hollow
wooden rattle, initially shaken quickly then more slowly, with
a gradual decline in pulse repetition rate (Fig. 8). Over the
course of the ~0.5–2.5-s call, call amplitude climbs with
successive pulses to maximum (Fig. 8C) as they simulta-
neously decline in pulse repetition rate (i.e., increase to
maximum interpulse interval). Calling rate ([total number of
calls 1] / time from beginning of ﬁrst call to beginning of
last) in the unvouchered specimen at Lake Balinsasayo was
0.133 calls/s (in the presence of calling conspeciﬁcs), and
TNHC 62794 called at 0.028 calls/s in response to rain and
then at 0.038 calls/s in response to an artiﬁcial stimulus.
Mean calling duration ranged from 0.89 60.31 SD (0.57–
1.79; n¼8) in the vouchered specimen to 1.03 60.48 SD
(0.33–2.29; n¼20) s in TNHC 62794. Individual calls
contained 8–16 ( ¯
X¼8.2 63.7 SD) distinct pulses (Fig. 8D)
in the unvouchered specimen and 4–25 ( ¯
X¼10.5 65.4 SD)
pulses in TNHC 62794. Pulse repetition rate ([total number
of pulses 1] / time from beginning of ﬁrst pulse to
beginning of last) ranged from 0.06 to 0.09 ( ¯
SD) pulses/s in the unvouchered specimen and 0.061 to
0.122 ( ¯
X¼0.096 60.017 SD) pulses/s in TNHC 62794.
Within-call declines in pulse repetition rate are reﬂected in
increasing interpulse intervals, which were brief at the start
of each call (0.04–0.10, ¯
X¼0.07 60.02 SD in the
unvouchered recording; 0.03–0.12, ¯
X¼0.08 60.02 SD in
TNHC 62794), increased by a within-call average of 140% at
midcall (0.07–0.12, ¯
X¼0.10 60.02 SD in the unvouchered
196 Herpetological Monographs 31, 2017
recording; 0.07–0.14, ¯
X¼0.12 60.02 SD in TNHC 62794),
and increased further to an average of 290% of the initial
interpulse interval at the call’s terminus (0.09–0.19, ¯
60.03 SD in the unvouchered recording; 0.13–0.21, ¯
0.27 60.03 SD in TNHC 62794; Fig. 8C). Spectral
properties of the advertisement call are structured and
apparently invariant across multiple calls from a single
individual (Fig. 8A,B), but frequency differences are
apparent between the two recorded individuals. Throughout
the call, energy is apparent at multiple, distinct frequency
components (Fig. 8B) with the fundamental frequency
(lowest) either the dominant (possessing the highest energy
of any of the call’s frequency components; Fig. 8B) or
apparently subequal to the fourth frequency band in some
calls. The call of TNHC 62794 had between three (Fig.
8A,B) to seven detectable frequency components in some
calls with highest energy in the fundamental, dominant
frequency (relative power, in decibels [dB], included in
parentheses) of 0.9 kHz (78–79), 1.8 (73–74), 2.3 (72–73), 2.9
(76–78), 3.6 (64–65), 4.2 (59–62), and 5.0 (54–56) kHz. The
unvouchered Lake Balinsasayo male’s call had three to six
distinct frequency components, peaking at 0.9, 1.7, 2.6, 3.4,
4.1, and 5.9 kHz, respectively. Toward the end (the last 3–5
pulses) of 4/20 calls recorded for TNHC 62794, the majority
of the call’s energy shifted up into the fourth frequency
component, with energy levels that rose above the funda-
mental (80–82 dB).
The second call type (chirping peeps and squeaks) initially
was thought to represent female response calls until it was
discovered that they originated from the large group of
nearby males. In this single instance, RMB observed
alternating calling males on rocks, each facing nearby
females (~10–15 cm). In the background, interspersed
between and overlapping rattle calls, we recorded a rapid
sequence of chirps. Type 2 chirping calls overlapped
temporally (multiple males vocalizing at the same time,
temporally overlapping one another and Type 1 calls), unlike
the nature of the assumed Type 1 male advertisement call in
which males calling in close proximity alternate and do not
overlap temporally. These tonal chirping vocalizations (Fig.
9) took the form of brief (0.05–0.07) frequency arcs, rising
from 0.6–0.7 to 1.5–1.7 kHz (n¼14), with subsequent
declines back to 0.6–0.7 kHz, constant frequency tones (2.6–
3.2 kHz; n¼9) with durations of 0.09–1.1 s, followed by a
steep frequency sweep (terminating at 0.9–1.0 kHz) or
simple frequency sweeps from 2.9–3.1 to 1.0–1.1 kHz over
an interval of 0.04–0.06 s (n¼19). The concordance
FIG. 9.—Audiospectrograms of the complex acoustic repertoires of
Sanguirana mearnsi (A, B; from Municipality of Burauen, northern Leyte
Island; RMB Field No. 21807; deposited at KU) and Sanguirana acai (C; from
Lake Balinsasayo, Cuernos de Negros Mountain Range, southern Negros
Island; voucher not collected, ML 224348). In both species, the structured,
presumably advertisement rattle vocalizations (Type 1 calls) differ from chirping
frequency arcs and sweeps (Type 2 calls) of unknown function. A third
vocalization, quacks (B), has been recorded only in S. mearnsi;inthelower
panel, calls of orthopterans overlap vocalization of S. acai at 2.2 and 4.7 kHz.
FIG. 8.—Male advertisement call (Type 1, rattle call) of Sanguirana acai
(male paratype TNHC 62794; ML 224181) recorded from the type locality,
Barangay Bongbong, the Municipality of Valencia, southern Negros Island
(9 April 2001; body temperature 248C). An expanded sonogram (A:
frequency in kHz vs. time in ms) and waveform (relative amplitude vs.
time in ms) of two notes from midcall, and relative power spectrum (B: from
a Fast Fourier Transformation, relative amplitude vs. frequency in kHz) and
a full call as depicted in a 1.8-s oscillogram (C: relative amplitude vs. time in
s) and corresponding audiospectrogram (D: frequency in kHz vs. time in s)
of a typical 18-note call.
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
between observed Type 2 vocalizations in S. acai and similar
calls reported for S. luzonensis (Brown et al. 2000b) suggests
that calls reported previously for S. luzonensis were Type 2
vocalizations (also observed in large aggregations of males);
to date, Type 1 calls have not been observed or reported in S.
luzonensis. In contrast, both Type 1 (rattles) and Type 2
(chirps) have been reported in S. mearnsi (Sanguila et al.
2016), although in that study it was also assumed these
represented male advertisement calls and female response
vocalizations. Our recent ﬁeld work on Samar and Leyte
conﬁrms our revised interpretation, namely that males of
Sanguirana acai and S. mearnsi both produce multiple
classes of vocalizations which we term Type 1 and Type 2.
Additionally, to date, only documented (vouchered) Type 2
calls (chirps) have been conﬁrmed in males of S. luzonensis.
The advertisement calls of all other Sanguirana species
Finally, the true social context and ultimate function of
Sanguirana call variation remains poorly understood. Type 1
rattle calls have been recorded in S. acai and S. mearnsi in
solitary males (suggesting advertisement, mate attraction)
but also in instances of one or a few males, vocalizing in close
vicinity to females (suggesting courtship) and at times when
nearby, large aggregations of males were producing only
Type 2 calls (suggesting chorusing behavior, possibly longer-
distance mate attraction, or even agonistic interactions). The
interpretation of multiple call types with distinct functions
has been reported in other anuran communication studies
(Narins and Capranica 1978; Rand and Ryan 1981) and is
supported by one recent observation of apparent female
phonotactic approach, over a 5-m stretch of stream, in the
direction of a solitary, Type 2-calling male S. luzonensis (J.
Binaday and RMB, personal observations, January 2017,
Sorsogon Province, Luzon).
Etymology.—We are pleased to name this new species
for our mentor, collaborator, and friend, Dr. Angel C. Alcala
of the Silliman University (Dumaguete City, Negros Island),
in recognition of his numerous contributions to Philippine
herpetology. Angel Alcala (known by friends and colleagues
by a nickname, derived from his initials ACA, pronounced
‘‘Ah-Kah’’ ) is one of the Philippines’ premier biodiversity
and conservation scientists whose lifelong dedication to
conservation of the country’s forests and coral reefs stands as
an inspiration to generations of Filipinos. Alcala’s earlier
ﬁeldwork (conducted in collaboration with the late Walter C.
Brown; Alcala 2004) resulted in the world’s most signiﬁcant
collection (.30,000 specimens) of Philippine herpetological
diversity (deposited at CAS), which forms the foundation of
what is known globally of the taxonomy (Diesmos et al.
2015), distribution (Brown and Alcala 1970, 1986), repro-
duction and ecology (Brown and Alcala 1961, 1964, 1982a,b,
1986), and conservation status (Alcala et al. 2012; Diesmos et
al. 2014; IUCN 2015) of the country’s endemic amphibians
and reptiles (Diesmos et al. 2015). The speciﬁc epithet is a
patronym and a masculine noun in the genitive case.
Suggested common name, Alcala’s West Visayan Stream
The recognition of the Sanguirana mearnsi as the valid
name for the Northeast Mindanao Stream Frogs (Inger
1954; Brown and Alcala 1970; Sanguila et al. 2016), and the
recognition of the West Visayan PAIC populations as a new
species (Brown et al. 2000a; Fuiten et al. 2011; Gaulke
2011), represent taxonomic solutions that are long overdue
(Brown 2007; Diesmos and Brown 2011; Diesmos et al.
2014, 2015). It is not surprising that neither the northeast
Mindanao PAIC lineage (S. mearnsi) nor the West Visayan
lineage (S. acai) should be found to be distinct from the
nominal S. everetti of southwest Mindanao Island (Fig. 1;
Brown et al. 2016). With respect to the former, the sister
species pair S. mearnsi and S. everetti are parapatric,
separated by deep genetic divergence, are phenotypically
distinct, and show no evidence of reticulation or gene ﬂow
(Inger 1954; Brown et al. 2000a, 2016). With respect to the
latter, S. acai and S. everetti are distantly allopatric on
separate PAICs, are phenotypically distinct (Fig. 3), and are
distantly related (Brown et al. 2016). In contrast, as might be
expected, S. acai actually is phenotypically most similar (Fig.
3) to its closest relative, S. luzonensis. Previous studies have
suggested that the problematic and disjunct distribution of S.
everetti warranted scrutiny (Inger 1954; Ferner et al. 2000;
Fuiten et al. 2011; Gaulke 2011), and we ﬁnd it surprising
that this unresolved biogeographic anomaly (Brown and
Alcala 1970; Brown and Diesmos 2009; Brown et al. 2013a)
has not been addressed until now.
The eight recognized species of the genus Sanguirana
form a well supported clade (Bossuyt et al. 2006; Stuart
2008; Wiens et al. 2009; Holder et al. 2010; Brown et al.
2016), with most taxa distinguished from congeners by
diagnostic morphological character differences, morphomet-
ric and body size variation, degree of sexual size dimorphism,
allopatry on isolated (separated by deep marine channels)
island groups, and considerable genetic divergence (Table 2;
Brown et al. 2000a, 2016; Fuiten et al. 2011). As currently
understood, no other congeners occur in the West Visayan
PAIC and, therefore, none occur in sympatry with the
biogeographically isolated S. acai.
With the resolution of this taxonomic problem, all
available evidence (morphological diagnosability, genetic
distinctiveness, position in phylogeny, biogeography) points
to a logical PAIC-structured understanding of species
diversity in the genus Sanguirana of the central and southern
Philippines (Brown et al. 2000a,b, 2013a, 2016; Brown and
Diesmos 2002, 2009)—with a few lingering, minor excep-
tions. One remaining, unanswered question is the taxonomic
identiﬁcation of the central Bohol population of ‘‘S.
everetti.’’ Given that Leyte, Samar, and northeastern Mind-
anao populations of Sanguirana have all been identiﬁed
convincingly as S. mearnsi (Brown et al. 2000a, 2016), Bohol
amphibians are most-often allied with the Mindanao PAIC
(e.g., Brown and Alcala 1970; Brown and Siler 2013;
Gonzales et al. 2014), and that true S. everetti populations
(Taylor 1920; type locality ¼‘‘Zamboanga’’ [western Mind-
anao]) are now known only from southwestern Mindanao
(Inger 1954; Brown et al. 2000a, 2016), the allopatric Bohol
population of S. everetti should be re-examined. We would
not be surprised if this population was identiﬁed as S.
mearnsi, but it remains possible that it may represent an
additional, undescribed species.
Three unresolved questions still complicate our under-
standing of evolutionary relationships and Sanguirana
species diversity in the northern Philippines (Luzon PAIC).
198 Herpetological Monographs 31, 2017
First, S. luzonensis, as currently recognized, is widespread
across multiple islands within the Luzon PAIC, spanning
numerous marine channels (Fig. 1) and known fault zones
(Hall 2002; Yumul et al. 2003, 2009b), all of which have been
shown to be biogeographic barriers that deﬁne species
distributions in unrelated, codistributed groups (Brown and
Diesmos 2009; Welton et al. 2010; Brown and Siler 2013;
Brown et al. 2013a; Gonzales et al. 2014). To date,
systematists have not critically evaluated patterns of intra-
speciﬁc variation in S. luzonensis nor considered whether all
of the populations referred to S. luzonensis in this region are
in fact a single evolutionary lineage (species). One recent
study (Brown et al. 2016) has taken a ﬁrst step toward this
goal, ﬁnding extensive geographically structured genetic
variation in this species. However, because so much of
Luzon remains unsurveyed and no formal species delimita-
tion analyses were conducted, the population-level diversity
within S. luzonensis remains poorly understood (Fig. 1;
Brown et al. 2016).
Second, whether S. tipanan (Sierra Madre of Luzon;
Brown et al. 2000a,b) is a distinct species relative to S.
igorota (Central Cordillera of Luzon; Brown et al. 2016)
remains an open question. The phenotypic distinctiveness of
these two taxa is clear at the most-northern extent of their
ranges where they are separated by the wide, arid,
environmental barrier represented by the Cagayan Valley
(Taylor 1922; Brown et al. 2000a). However, much like
Brown and Siler’s (2013) recent ﬁndings from the Pulchrana
signata Complex (see Brown and Guttman 2002: Fig. 3),
variable and intermediate phenotypes have been document-
ed in the southern extent of their ranges where the
distributions of these two species abut in the Caraballo
Mountains of central Luzon (Fuiten et al. 2011; Brown et al.
2012a, 2013b). A recent phylogenetic analysis suggested that
S. igorota may be paraphyletic with respect to S. tipanan,an
arrangement that would require the placement of the latter
species in synonymy with the former, if veriﬁed with
additional geographic and gene sampling (Brown et al.
Finally, with small islands and isolated geological compo-
nents of large islands increasingly appreciated for their
tendency to support endemic species (Welton et al. 2010;
Sanguila et al. 2011, 2016; Brown et al. 2013a, 2015a), it
would not be surprising if additional species of Sanguirana
were discovered in the near future. Islands like Bantayan,
Basilan, Biliran, Bohol, Burias, Pacijan, Ponson, Poro, Ticao,
and the remaining islands of the Sulu Archipelago all deserve
amphibian biodiversity survey efforts if trained naturalists
can be provided access to the last remaining habitats on
these isolated landmasses. Likewise, the recent unexpected
discovery of a highly distinct evolutionary lineage of
Sanguirana in isolated mountains of central Luzon (S.
aurantipunctata; Fuiten et al. 2011) emphasizes the degree
to which this endemic and understudied Philippine genus is
prone to differentiation in montane habitats; all high-
elevation peaks of Luzon and Mindanao deserve particular
attention by ﬁeld biologists (Brown 2015).
Conservation efforts aimed at central Philippine amphib-
ians are plagued by near-complete removal of forests in the
West Visayan islands of Cebu (Brown and Alcala 1986;
Supsup et al. 2016), Guimaras, Negros (Brown and Alcala
1961, 1964; Alcala et al. 2004), Masbate, and Panay (Ferner
et al. 2000; Gaulke 2011), with wholesale conversion of
marginal habitats to agriculture (Brown and Alcala 1986) and
the archipelago-wide infection of amphibian populations by
chytrid fungus (Swei et al. 2011; Brown et al. 2012b;
Diesmos et al. 2012). We ﬁnd that the new species, with its
forested habitat severely fragmented and its patchy, but well
documented, distribution tied to clean water sources running
within, or at the margins of, intact vegetation cover (Ferner
et al. 2000; Gaulke 2011), qualiﬁes for classiﬁcation at a
formal, elevated level of conservation threat under IUCN
criteria: ‘‘Vulnerable’’ (VU, IUCN 2010: A2ac; B2ab[iii];
D2). Thus, the new species should be considered an
immediate conservation concern (Diesmos et al. 2014).
Given the absence of new data on the status or
distribution of Sanguirana everetti on central and southern
Mindanao Island (Diesmos and Brown 2011; Diesmos et al.
2014, 2015), the conservation status of true S. everetti
remains ‘‘Data Deﬁcient’’ (IUCN 2015). Studies of the
remaining populations of S. everetti (southwestern Mindanao
Island) and S. acai (West Visayan PAIC) are pressing
challenges for future ﬁeld surveys and conservation research
(Brown et al. 2012b). Both S. tipanan and S. igorota are
classiﬁed by IUCN (2015) at elevated conservation threat
levels, although new survey data suggest both species are
more widely distributed than previously thought and appear
tolerant to some level of disturbance (Brown et al. 2000a,b,
2012a, 2013b; Siler et al. 2011), suggesting that their status
needs to be reconsidered and revised (Diesmos et al. 2014).
The unexpected discovery of so many new amphibian species
on larger islands (Fuiten et al. 2011; Siler et al. 2011; Brown
2015; Brown et al. 2015b) emphasizes the need for an
accelerated pace of faunal inventories and ﬁeld-based
assessment of species boundaries, informed with basic
natural history data. These and other unexpected discoveries
of evolutionarily distinctive species of endemic Philippine
amphibians (e.g., Sanguila et al. 2011; Blackburn et al. 2013;
Brown 2015; Brown et al. 2015a) remind us that the only way
to solve persistent taxonomic and conservation status
questions of this kind is to encourage and support faunal
survey activities—necessarily including the collection of
properly preserved voucher specimens (Rocha et al. 2014),
advertisement calls, and genetic samples—in both the
unexplored and previously surveyed (yet still poorly under-
stood) islands of the Philippines (Brown et al. 2013a, 2016).
Acknowledgments.—The Biodiversity Management Bureau (BMB) of
the Philippine Department of Environment and Natural Resources (DENR)
facilitated research (Memorandums of Agreement KU2005, KU2009,
KU2015) and collection permits (GP 171, 185, 187, 201, 212, 221, 246,
258) necessary for this and related studies; and the University of Kansas’
Animal Use Committee approved and administered RMB’s IACUC
authorization (185-04). For the loans of specimens and photographs, or
assistance visiting collections we thank D. Blackburn (UF), D. Blackburn, J.
Vindum, and A. Leviton (CAS), A. Diesmos, R. Sison (PNM), J. Ferner
(CMNH), A. Resetar, R. Inger, and H. Voris (FMNH), K. de Queiroz, A.
Wynn, and R. Crombie (USNM), J. Hanken and J. Rosado (MCZ), T.
LaDuc and D. Cannatella (TNHC), and S. Rogers and J. Padial (CM).
Financial support for ﬁeldwork was provided by a National Science
Foundation (NSF) Doctoral Dissertation Improvement Grant (DEB
0073199) and a Biotic Surveys and Inventories grant (DEB 0743491) to
RMB; and by Fulbright and Fulbright-Hayes funding and an NSF Doctoral
Dissertation Improvement Grant (DEB 0804115) to CDS. Archival
deposition and digitization of RMB’s analog media collection was supported
by a grant from NSF’s Advancing Digitization of Biological Collections
Thematic Collections Network program (DEB 1304585) to RMB. During
the preparation of this manuscript AP was supported by funding from the
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
U.S. National Institute of Health, Research Initiative for Scientiﬁc
Enhancement (RISE) program. We thank V. Yngente, J. Fernandez, and
A. Diesmos for collaboration in ﬁeldwork, anonymous reviewers for critical
reviews of the manuscript, and our friend and mentor Angel C. Alcala for
many years of generous support and patient guidance over the course of our
fruitful international collaboration.
Alcala, A.C. 1962. Breeding behavior and early development of frogs of
Negros, Philippines Islands. Copeia 1962:679–726.
Alcala, A.C. 2004. A life of service to science: A tribute to Walter Creighton
Brown 1913–2002. Silliman Journal 45:258–262.
Alcala, E.L., A.C. Alcala, and C.N. Dolino. 2004. Amphibians and reptiles in
tropical rainforest fragments on Negros Island, the Philippines.
Environmental Conservation 31:254–261.
Alcala, A.C., A. Bucol, A.C. Diesmos, and R.M. Brown. 2012. Vulnerability
of Philippine amphibians to climate change. Philippine Journal of Science
Ariﬁn, U., D.T. Iskandar, D.P. Bickford, R.M. Brown, R. Meier, and S.N.
Kutti. 2011. Phylogenetic relationship within the genus Staurois
(Amphibia, Ranidae) based on 16S rRNA sequences. Zootaxa 2733:39–
52. DOI: http://dx.doi.org/10.11646/zootaxa.2733.4.1
Aurelio, M.A., R.E. Pe ˜
na, and K.J.L. Taguibao. 2013. Sculpting the
Philippine archipelago since the Cretaceous through rifting, oceanic
spreading, subduction, obduction, collision and strike–slip faulting:
Contribution to IGMA5000. Journal of Asian Earth Sciences 72:102–107.
Blackburn, D.C., D.P. Bickford, A.C. Diesmos, D.T. Iskandar, and R.M.
Brown. 2010. An ancient origin for the enigmatic ﬂat-headed frogs
(Bombinatoridae: Barbourula) from the islands of Southeast Asia. PLoS
ONE 5:e12090. DOI: http://dx.doi.org/10.1371/journal.pone.0012090
Blackburn, D.C, C.D. Siler, A.C. Diesmos, J.A. McGuire, D.C. Cannatella,
and R.M. Brown. 2013. An adaptive radiation of frogs in a Southeast
Asian island archipelago. Evolution 67:2631–2646.
Boettger, O. 1893. Drei neue Wasserfr ¨
osche (Rana) von den Philippinen.
Zoologischer Anzeiger 16:363–366.
Bossuyt, F., R.M. Brown, D.M. Hillis, D.C. Cannatella, and M.C.
Milinkovitch. 2006. Late Cretaceous diversiﬁcation resulted in conti-
nent-scale regionalism in the cosmopolitan frog family Ranidae.
Systematic Biology 55:579–594.
Boulenger, G.A. 1882. Catalogue of the Batrachia Salientia s. Ecaudata in
the Collection of the British Museum, 2nd ed. Taylor and Francis, UK.
Boulenger, G.A. 1894. On the herpetological fauna of Palawan and Balabac.
Annals and Magazine of Natural History (Series 6) 14:81–90.
Boulenger, G.A. 1896. Descriptions of new batrachians in the British
Museum. Annals and Magazine of Natural History (Series 6) 17:401–406.
Brown, R.M. 2007. Introduction to Robert F. Inger’s zystematics and
zoogeography of Philippine Amphibia. Pp. 1–17 in Systematics and
Zoogeography of Philippine Amphibia. Natural History Publications,
Brown, R.M. 2015. A new species of stream frog (genus Hylarana) from the
mountains of southern Mindanao Island, Philippines. Herpetologica
Brown, R.M., and A.C. Diesmos. 2002. Application of lineage-based species
concepts to oceanic island frog populations: The effects of differing
taxonomic philosophies on the estimation of Philippine biodiversity.
Silliman Journal 42:133–162.
Brown, R.M., and A.C. Diesmos. 2009. Philippines, biology. Pp. 723–732 in
Encyclopedia of Islands (R. Gillespie and D. Clague, eds.). University of
California Press, USA.
Brown, R.M., and J.C. Gonzalez. 2007. A new forest frog of the genus
Platymantis (Amphibia; Anura: Ranidae) from the Bicol Peninsula of
Luzon Island, Philippines. Copeia 2007:251–266.
Brown, R.M., and S.I. Guttman. 2002. Phylogenetic systematics of the Rana
signata complex of Philippine and Bornean stream frogs: Reconsideration
of Huxley’s modiﬁcation of Wallace’s Line at the Oriental–Australian
faunal zone interface. Biological Journal of the Linnean Society 76:393–
Brown, R.M., and C.D. Siler. 2013. Spotted stream frog diversiﬁcation at the
Australasian faunal zone interface, mainland versus island comparisons,
and a test of the Philippine ‘dual-umbilicus’ hypothesis. Journal of
Brown, R.M., and B.L. Stuart. 2012. Patterns of biodiversity discovery
through time: An historical analysis of amphibian species discoveries in
the Southeast Asian mainland and island archipelagos. Pp. 348–389 in
Biotic Evolution and Environmental Change in Southeast Asia (D.J
Gower, K.G. Johnson, J.E. Richardson, B.R. Rosen, L. R ¨
uber, and S.T.
Williams, eds.). Cambridge University Press, UK.
Brown, R.M., J.A. McGuire, and A.C. Diesmos. 2000a. Status of some
Philippine frogs related to Rana everetti (Anura: Ranidae), description of
a new species, and resurrection of Rana igorota Taylor 1922.
Brown, R.M., J.A. McGuire, J.W. Ferner, N. Icarangal, Jr., and R.S.
Kennedy. 2000b. Amphibians and reptiles of Luzon Island, II:
Preliminary report on the herpetofauna of Aurora Memorial National
Park, Philippines. Hamadryad 25:175–195.
Brown, R.M., A.C. Diesmos, and A.C. Alcala. 2008. Philippine amphibian
biodiversity is increasing in leaps and bounds. Pp. 82–83 in Threatened
Amphibians of the World (S.N. Stuart, M. Hoffmann, J.S. Chanson, N.A
Cox, R. Berridge, P. Ramani, and B.E. Young, eds.). IUCN – The World
Conservation Union, Switzerland; and Conservation International, USA.
Lynx Ediciones, Spain.
Brown, R.M., C.H. Oliveros, C.D. Siler, J.B. Fernandez, L.J. Welton, P.A.C.
Buenavente, M.L.D. Diesmos, and A.C. Diesmos. 2012a. Amphibians
and Reptiles of Luzon Island (Philippines), VII: Herpetofauna of Ilocos
Norte Province, Northern Cordillera Mountain Range. Check List 8:469–
Brown, R.M., A.C. Diesmos, M.B. Sanguila, C.D. Siler, M.L.D. Diesmos,
and A.C. Alcala. 2012b. Amphibian conservation in the Philippines.
Brown, R.M., C.D. Siler, C.H. Oliveros, . . ., A.C. Alcala. 2013a.
Evolutionary processes of diversiﬁcation in a model island archipelago.
Annual Review of Ecology, Evolution, and Systematics 44:411–435.
Brown, R.M., C.D. Siler, C.H. Oliveros, . . ., A.C. Diesmos. 2013b. The
amphibians and reptiles of Luzon Island, Philippines, VIII: The
herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre
Mountain Range. ZooKeys 266:1–120.
Brown, R.M., C.D. Siler, S. Richards, A.C. Diesmos, and D.C. Cannatella.
2015a. Multilocus phylogeny and a new classiﬁcation for Southeast Asian
and Melanesian forest frogs (family Ceratobatrachidae). Zoological
Journal of the Linnaean Society 174:130–168.
Brown, R.M., L.A. de Layola, A. Lorenzo, II, M.L.L. Diesmos, and A.C.
Diesmos. 2015b. A new species of limestone karst-inhabiting forest frog,
genus Platymantis (Amphibia: Anura: Ceratobatrachidae: subgenus
Lupacolus) from southern Luzon Island, Philippines. Zootaxa
4048:191–210. DOI: http://dx.doi.org/10.11646/zootaxa.4048.2.3
Brown, R.M., Y.-C. Su., B. Barger, C.D. Siler, M.B. Sanguila, A.C. Diesmos,
and D.C. Blackburn. 2016. Phylogeny of the island archipelago frog
genus Sanguirana: Another endemic Philippine radiation that diversiﬁed
‘Out-of-Palawan.’ Molecular Phylogeny and Evolution 91:531–536.
Brown, W.C., and A.C. Alcala. 1955. Observations on amphibians of the
Mount Halcon and Canlaon areas, Philippine Islands. Silliman Journal
Brown, W.C., and A.C. Alcala. 1961. Populations of amphibians and reptiles
in the submontane and montane forests of Cuernos de Negros, Philippine
Islands. Ecology 42:628–636.
Brown, W.C., and A.C. Alcala. 1964. Relationship of the herpetofaunas of
the non-dipterocarp communities to that of the dipterocarp forest on
southern Negros Island, Philippines. Senckenbergiana Biologie 45:591–
Brown, W.C., and A.C. Alcala. 1970. The zoogeography of the herpetofauna
of the Philippine Islands, a fringing archipelago. Proceedings of the
California Academy of Sciences 38:105–130.
Brown, W.C., and A.C. Alcala. 1982a. Reproductive biology of some species
of Philautus (Rhacophoridae) and other Philippine anurans. Kalikasan,
Philippine Journal of Biology 11:203–226.
Brown, W.C., and A.C. Alcala. 1982b. Modes of reproduction of Philippine
anurans. Pp. 416–428 in Advances in Herpetology and Evolutionary
Biology (A.G.J. Rodin and K. Miyata, eds.). Museum of Comparative
Brown, W.C., and A.C. Alcala. 1986. Comparison of the herpetofaunal
species richness on Negros and Cebu Islands, Philippines. Silliman
Brown, W.C., and A.C. Alcala. 1994. Philippine frogs of the family
Rhacophoridae. Proceedings of the California Academy of Sciences
Chan, K.O., D.C. Blackburn, R.W. Murphy, B.L. Stuart, D.A. Emmett, C.T.
Ho, and R.M. Brown. 2013. A new species of narrow-mouthed frog of the
genus Kaloula from eastern Indochina. Herpetologica 69:329–341.
Chan, K.O., and R.M. Brown. 2017. Did true frogs ‘‘ dispersify’’ ? Biology
200 Herpetological Monographs 31, 2017
Cochran, D.M. 1961. Type specimens of reptiles and amphibians in the U.S.
National Museum. Bulletin of the United States National Museum
de Queiroz, K. 1998. The general lineage concept of species, species criteria,
and the process of speciation. Pp. 57–75 in Endless Forms: Species and
Speciation (D.J. Howard and S.H. Berlocher, eds.). Oxford University
de Queiroz, K. 1999. The general lineage concept of species and the deﬁning
properties of the species category. Pp. 49–89 in Species: New
Interdisciplinary Essays (R.A. Wilson, ed.). Massachusetts Institute of
Technology Press, USA.
de Queiroz, K. 2005. A uniﬁed concept of species and its consequences for
the future of taxonomy. Proceedings of the California Academy of
de Queiroz, K. 2007. Species concepts and species delimitation. Systematic
Diesmos, A.C., and R.M. Brown. 2011. Diversity, biogeography, and
conservation of Philippine Amphibians. Pp. 26–49 in Biology and
Conservation of Tropical Asian Amphibians (I. Das, A. Haas, and A.A.
Tuen, eds.). Proceedings of the Conference ‘‘Biology of the Amphibians
in the Sunda Region, South-east Asia.’’ Institute of Biodiversity and
Environmental Conservation, Universiti Malaysia Sarawak, Malaysia.
Diesmos, A.C., M.L. Diesmos, and R.M. Brown. 2006. Status and
distribution of alien invasive frogs in the Philippines. Journal of
Environmental Science and Management, Philippines 9:41–53.
Diesmos, M.L.D., A.C. Diesmos, C.D. Siler, V.T. Vredenburg, and R.M.
Brown. 2012. Detecting the distribution of chytrid fungus in the
Philippines. FrogLog 104:48–49.
Diesmos, A.C., A.C. Alcala, C.D. Siler, and R.M. Brown. 2014. Status and
conservation of Philippine amphibians. Pp. 310–336 in Conservation
Biology of Amphibians of Asia. Status of Conservation and Decline of
Amphibians: Eastern Hemisphere (H. Heatwole and I. Das, eds.).
Natural History Publications, Malaysia.
Diesmos, A.C., J.L. Watters, N.A. Huron, . . ., C.D. Siler. 2015. Amphibians
of the Philippines, Part I: Checklist of the species. Proceedings of the
California Academy of Sciences 62:451–531.
Dubois, A. 1992. Notes sur la classiﬁcation des Ranidae (Amphibiens
anoures). Bulletin Mensuel de la Soci ´
eenne de Lyon 61:305–352.
Evans, B.J., R.M. Brown, J.A. McGuire, J. Supriatna, N. Andayani, A.C.
Diesmos, D.T. Iskandar, D.J. Melnick, and D.C. Cannatella. 2003.
Phylogenetics of fanged frogs: Testing biogeographical hypotheses at the
interface of the Asian and Australian faunal zones. Systematic Biology
Ferner, J.W., R.M. Brown, R.V. Sison, and R.S. Kennedy. 2000. The
amphibians and reptiles of Panay Island, Philippines. Asiatic Herpeto-
logical Research 9:1–37.
Frost, D.R. 2016. Amphibian Species of the World: An Online Reference,
Version 6.0. Available at http://research.amnh.org/herpetology/amphibia/
index.html. Archived by WebCite at http://www.webcitation.org/
Frost, D.R., and D.M. Hillis. 1990. Species in concept and practice:
Herpetological applications. Herpetologica 46:87–104.
Fuiten, A., A.C. Diesmos, L.J. Welton, A. Barley, B. Oberheide, E.L.B.
Rico, and R.M. Brown. 2011. New species of stream frog from the
mountains of Luzon Island, Philippines. Herpetologica 67:89–103.
Gaulke, M. 2011. The Herpetofauna of Panay Island, Philippines. Edition
Gaulke, M., I. Frank, and B. Tacud. 2008. Zur Farbvariabilit ¨
Brutbiologie einiger philippinischer Anuren. Sauria 30:11–21.
Gonzales, P., Y.-C. Su, C.D. Siler, A. Barley, M.B. Sanguila, A.C. Diesmos,
and R.M. Brown. 2014. Archipelago colonization by ecologically
dissimilar amphibians: Evaluating the expectation of common evolution-
ary history of geographical diffusion in co-distributed rainforest tree frogs
in islands of Southeast Asia. Molecular Phylogenetics and Evolution
Hall, R. 2002. Cenozoic geological and plate tectonic evolution of SE Asia
and the SW Paciﬁc: Computer-based reconstructions, model and
animations. Journal of Asian Earth Sciences 20:353–431.
Hayek, L.-A.C., W.R. Heyer, and C. Gascon. 2001. Frog morphometrics: A
cautionary tale. Alytes 18:153–177.
Holder, M.T., J.A. Sukumaran, and R.M. Brown. 2010. Bayesian approaches
to phylogenetic analysis. Pp. 1–39 in Bayesian Modeling in Bioinformatics
(D.K. Dey, S. Ghosh, and B.K. Mallick, eds.). Chapman & Hall/CRC,
Taylor & Francis Group, USA.
Kaiser, H.F. 1960. The application of electronic computers to factor analysis.
Educational and Psychological Measurement 20:141–151.
Inger, R.F. 1954. Systematics and zoogeography of Philippine Amphibia.
Inger, R.F. 1999. Distribution of amphibians in southern Asia and adjacent
islands. Pp. 445–482 in Patterns of Distribution of Amphibians (W.E.
Duellman, ed.). John Hopkins University Press, USA.
International Union for the Conservation of Nature (IUCN). 2010.
Guidelines for Using the IUCN Red List Categories and Criteria,
Version 8.1. Available at http://intranet.iucn.org/webﬁles/doc/SSC/
RedList/RedListGuidelines.pdf. Archived by WebCite at http://www.
webcitation.org/6YZVTeL9Z on 16 May 2015.
IUCN. 2015. IUCN Red List of Threatened Species, Version 2013.2.
Available at http://www.iucnredlist.org. Archived by WebCite at http://
www.webcitation.org/6YZVxdVsE on 16 May 2015.
Jombart, T. 2008. Adegenet: A R package for the multivariate analysis of
genetic markers. Bioinformatics 24:1403–1405.
Jombart, T., S. Devillard, and F. Balloux. 2010. Discriminant analysis of
principal components: A new method for the analysis of genetically
structured populations. BMC Genetics 11:94.
Lee, J.C. 1982. Accuracy and precision in anuran morphometrics: Artifacts
of preservation. Systematic Zoology 31:266–281.
Lee, J.C. 1990. Sources of extraneous variation in the study of meristic
characters: The effect of size and inter-observer variability. Systematic
Narins, P.M., and R.R. Capranica. 1978. Communicative signiﬁcance of two-
note calls of the tree frog Eleutherodactylus coqui. Journal of
Comparative Physiology 127:1–9.
Oliver, L., E. Prendini, F. Kraus, and C.J. Raxworthy. 2015. Systematics and
biogeography of the Hylarana frog (Anura: Ranidae) radiation across
tropical Australasia, Southeast Asia, and Africa. Molecular Phylogenetics
and Evolution 90:176–192.
R Core Team. 2015. R: A Language and Environment for Statistical
Computing, Version 3.1.2. Available at http://www.r-project.org/. R
Foundation for Statistical Computing, Austria.
Rand, A.S., and M.J. Ryan. 1981. The adaptive signiﬁcance of a complex vocal
repertoire in a Neotropical frog. Zeitschrift f¨ur Tierpsychologie 57:209–214.
Rocha, L.A., A. Aleixo, G. Allen, . . ., C.C. Witt. 2014. Specimen collection:
An essential tool. Science 344:814–815.
Sabaj, M.H. 2016. Standard Symbolic Codes for Institutional Resource
Collections in Herpetology and Ichthyology: An Online Reference,
Version 6.5. American Society of Ichthyologists and Herpetologists, USA.
Available at http://www.asih.org/. Archived by WebCite at http://www.
webcitation.org/6otYSPsdM on 9 March 2017.
Sanguila, M.B., C.D. Siler, A.C. Diesmos, O. Nu ˜
neza, and R.M. Brown.
2011. Phylogeography and conservation implications of geographic
structure of genetic variation and potential species boundaries in
Philippine slender toads. Molecular Phylogenetics and Evolution
Sanguila, M.B., K.A. Cobb, C.D. Siler, A.C. Diesmos A.C. Alcala, and R.M.
Brown. 2016. The amphibians and reptiles of Mindanao Island, southern
Philippines, II: The herpetofauna of northeast Mindanao and adjacent
islands. ZooKeys 624:1–132.
Savage, J.M., and W.R. Heyer. 1969. Variation and distribution of the tree-
frog genus Phyllomedusa in Costa Rica, Central America. Beitrage zur
Neotropischen Fauna 5:111–131.
Savage, J.M., and W.R. Heyer. 1997. Digital webbing formulae for anurans:
A reﬁnement. Herpetological Review 28:131.
Setiadi, M.I., J.A. McGuire, R.M. Brown, M. Zubairi, D.T. Iskandar, N.
Andayani, J. Supriatna, and B.J. Evans. 2011. Adaptive radiation and
ecological opportunity in Sulawesi and Philippine fanged frog (Limno-
nectes) communities. American Naturalist 178:221–240.
Siler, C.D., J.D. McVay, A.C. Diesmos, and R.M. Brown. 2009. A new species
of fanged frog, genus Limnonectes (Amphibia: Anura: Dicroglossidae) from
southeast Mindanao Island, Philippines. Herpetologica 65:105–114.
Siler, C.D., A.C. Diesmos, C.W. Linkem, M.L. Diesmos, and R.M. Brown.
2010. A new species of limestone-forest frog, genus Platymantis (Amphibia:
Anura: Ceratobatrachidae) from central Luzon Island, Philippines. Zootaxa
2482:49–63. DOI: http://dx.doi.org/10.11646/zootaxa.4048.2.3
Siler, C.D., L.J. Welton, J.M. Siler, J. Brown, A. Bucol, A.C. Diesmos, and
R.M. Brown. 2011. Amphibians and reptiles, Luzon Island, Aurora
Province and Aurora Memorial National Park, northern Philippines: New
island distribution records. Check List 7:182–195.
Siler, C.D., J.R. Oaks, L.K. Welton, C.W. Linkem, J. Swab, A.C. Diesmos,
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA
and R.M. Brown. 2012. Did geckos ride the Palawan raft to the
Philippines? Journal of Biogeography 39:1217–1234.
Simpson, G.G. 1961. Principles of Animal Taxonomy. Columbia University
Sison, R.V., P.C. Gonzales, and J.W. Ferner. 1995. New records from Panay,
Philippines. Herpetological Review 26:48–49.
Stejneger, L. 1905. Three new frogs and one new gecko from the Philippine
Islands. Proceedings of the United States National Museum 28:343–348.
Stuart, B.L. 2008. The phylogenetic problem of Huia (Amphibia: Ranidae).
Molecular Phylogenetics and Evolution 46:49–60.
Supsup, C., N.M. Puna, A.A. Asis, B.R. Redoblado, M.F.G. Panaguinit, F.M.
Guinto, E.B. Rico, A.C. Diesmos, R.M. Brown, and N.A. Mallari. 2016.
Amphibians and reptiles of Cebu, Philippines: The poorly understood
herpetofauna of an island with very little remaining natural habitat. Asian
Herpetological Research 7:151–179.
Swei, A., J.J.L. Rowley, D. R ¨
odder, . . ., V.T. Vredenburg. 2011. Is
chytridiomycosis an emerging disease in Asia? PLoS ONE 6:e23179.
Taylor, E.H. 1920. Philippine Amphibia. Philippine Journal of Science
Taylor, E.H. 1922 Additions to the herpetological fauna of the Philippine
Islands. II. Philippine Journal of Science 21:257–303.
Thorpe, R.S. 1975. Quantitative handling of characters useful in snake
systematics with particular reference to intraspeciﬁc variation in the
Ringed Snakes Natrix natrix (L.). Biological Journal of the Linnean
Thorpe, R.S. 1983a. A review of the numerical methods for recognizing and
analyzing racial differentiation. Pp. 404–423 in Numerical Taxonomy:
Proceedings of a NATO Advanced Studies Institute NATO ASI series,
Volume G1 (J. Felsenstein, ed.). Springer Verlag, Germany.
Thorpe, R.S. 1983b. A biometric study of the effects of growth on the
analysis of geographic variation: Tooth number in green geckos (Reptilia:
Phelsuma). Journal of Zoology 201:13–26.
Turan, C. 1999. A note on the examination of morphometric differentiation
among ﬁsh populations: The Truss System. Turkish Journal of Zoology
Voris, H.K. 2000. Maps of Pleistocene sea levels in Southeast Asia:
Shorelines, river systems, time durations. Journal of Biogeography
Welton, L.J., C.D. Siler, D. Bent, A.C. Diesmos, M.R. Duya, R. Dugay, E.L.
Rico, M. van Weerd, and R.M. Brown. 2010. A spectacular new
Philippine monitor lizard reveals a hidden biogeographic boundary and a
novel ﬂagship species for conservation. Biology Letters 6:654–658.
Wiens, J.J., J.S. Sukumaran, R.A. Pyron, and R.M. Brown. 2009.
Evolutionary and biogeographic origins of high tropical diversity in old
world frogs (Ranidae). Evolution 64:1558–5646.
Wiley, E.O. 1978. The evolutionary species concept reconsidered.
Systematic Zoology 21:17–26.
Yumul, G.P., Jr., C.B. Dimalanta, R.A. Tamayo, Jr., and R.C. Maury. 2003.
Collision, subduction and accretion events in the Philippines: A synthesis.
Island Arc 12:77–91.
Yumul, G.P., Jr., C.B. Dimalanta, E.J. Marquez, and K.L. Quea ˜
Onland signatures of the Palawan microcontinental block and Philippine
mobile belt collision and crustal growth process: A review. Journal of
Asian Earth Sciences 34:610–623.
Yumul, G., C.B. Dimalanta, K.L. Quea ˜
no, and E.J. Marquez. 2009b.
Philippines, geology. Pp. 732–738 in Encyclopedia of Islands (R.
Gillespie and D. Clague, eds.). University of California Press, USA.
Zamoros, L.R., M.G.A. Montes, K.L. Quea ˜
no, E.J. Marquez, C.B. Dimalanta,
J.A.S. Gabo, and G.P. Yumul, Jr. 2008. Buruanga Peninsula and Antique
Range: Two contrasting terranes in northwest Panay, Philippines featuring
an arc–continent collision zone. Island Arc 17:443–457.
ZooBank.org registration LSID: 19B2F7D9-DC20-4821-991B-
Published on 12 October 2017
All specimens are from the Philippines.
Sanguirana acai.—See holotype and paratypes sections.
Sanguirana aurantipunctata.—LUZON ISLAND, NUEVA VIZCAYA PROV-
INCE,Municipality of Quezon, Barangay Maddiangat, Sitio Parola
E; datum ¼WGS84): PNM 9727 (holotype),
PNM 9728–45, KU 325894–932, 325934–45, 329950–51, 308655, 308665,
308667, 308687, 308705, 308706, 308712, 308775, 308776 (Paratopotypes);
AURORA PROVINCE,Municipality of San Luis, Barangay Real, Sitio Minoli:
KU 322548, 322549 (paratypes); Municipality of Dingalan, Mt. Mingan:
MVD 066, 068, 069, and 074, DSB 3728 and 3745 (six uncataloged
specimens, deposited at PNM)
Sanguirana everetti.—MINDANAO, LANAO DEL SUR PROVINCE,Lake
Lanao, Camp Keithley: CAS-SU 2141; ZAMBOANGA: CAS 61872; SOUTH
COTOBATO PROVINCE,‘‘near Saub,’’ MCZ 14083–84; Municipality of Tupi,
Barangay Kablon, Masbang creek: PNM 469; Municipality of Tiboli,
Barangay Salacafe, Lake Parker: PNM 3002–07, 3009–12., 3018–19, 3059,
3073; Municipality of Tampakan, Barangay Tablu, Sitio Datal Mangisi: KU
327523, 327527, 327529; Sitio Tukuymal: KU 327525, 327526, 327528.
Sanguirana igorota.—LUZON ISLAND: KALINGA–APAYAO PROVINCE,
KALINGA SUBPROVINCE,Municipality of Balbalan, Barangay Balbalan: CAS
61484 (EHT F789; holotype of Rana igorota); CAS 61483, 61485–89, MCZ
14096–98 (paratypes of Rana igorota); NUEVA VIZCAYA PROVINCE,Munici-
pality of Quezon, Barangay Maddiangat, Mt. Palali: KU 308688, 308707–11,
325843–93; Benguet Province, Municipality of Kabayan, Barangay Apunan:
PNM 158, 162; IFUGAO PROVINCE,Municipality of Banaue, Barangay
Bayninan, PNM 741, 742; ILOCOS PRO VINCE,Municipality of Adams,
Barangay Adams, Mt. Pao: KU 329824-89.
Sanguirana luzonensis.—LUZON ISLAND, AURORA PROVINCE,Munici-
pality of Carmen, Aurora National Park: PNM 5742–5765; CMNH 5605–11;
5612–30; Municipality of Maria Aurora, Aurora Memorial National Park
‘‘tower site,’’ KU 322566–67; Barangay Villa Aurora, Sitio Dimani, Aurora
Memorial National Park: KU 322568–87; Barangay Villa Aurora, Aurora
Memorial National Park, Mt. Dayap, area known locally as ‘‘Siete’’:KU
322588–90; Municipality of Baler, Barangay Zabali, Aurora State College of
Technology (ASCOT): KU 322591–619; Municipality of San Luis, Barangay
Real, Sitio Minoli: KU 322620–28, 322520–39, 322540–47; Barangay
Lipimental: KU 322550–65, 322503–19; MOUNTAIN PROVINCE,Municipality
of Bontoc: MCZ 10556; LAGUNA PROVINCE,Municipality of Los Ba ˜nos, Mt.
Makiling: MCZ 23178–79, 14142–45; ZAMBALES PROVINCE:Municipality of
Masinloc, Barangay Coto: CMNH 4171–72, 4279–85; PNM 2371, 2378–84,
2496–98; TRAIL BETWEEN FAMY (LAGUNA PROV.) AND INFANTA (TAYABAS PROV.):
CAS 61819 (holotype of R. tafti); BANGUET PROVINCE,Municipality of
Baguio, Baguio City: CM 3271, 3273–78, 3280–81, 3283, MCZ 10482–84
(topotypes of R. guerreroi); QUEZON PROVINCE,Municipality of Polillo,
Barangay Pinaglubayan: KU 302380, 307649–51; Barangay Salipsip, Sition
Kapilijan: KU 307652–60; QUEZON PROVINCE: 303561–63; CATANDUANES,
Municipality of San Miguel, Sulong: KU 308067, 308090–98; Municipality of
Gigmoto, Barangay San Pedro: KU 308121, 308139, 308158–69; CAMARINES
DEL SUR PROVINCE:Municipality of Tabaco, Barangay Comon: KU 306495–
98, 306503–06; CAMARINES DEL NORTE PROVINCE,Municipality of Labo,
Barangay Tulay na Lupa: KU 306499–502, 306507–306509; ISABELA
PROVINCE,Municipality of Cabagan, Barangay Garita, Mitra Ranch: KU
307636; NUEVA VIZCAYA PROVIN CE,Municipality of Quezon,Barangay
Maddiangat, Mt. Palali: KU 308655, 308665, 308667, 308687, 308705–06,
308712, 308774–76, 308835–36, 325501–40; CAMARINES NORTE PROVINCE:
Municipality of Labo, Barangay Tulay Na Lupa: KU 313647-313681;
POLILLO ISLAND: POLILLO PROVINCE,Burdeos: CAS 62448 (holotype of
Sanguirana mearnsi.—MINDANAO ISLAND: DAVAO ORIENTAL PROV-
INCE,Municipality of Baganga,‘‘Baganga River, east coast mountain range,
300–1500 m above sea level’’: USNM 35258 (holotype of Rana mearnsi);
AGUSAN DEL NORTE PROVINCE:Tagibo and Daydayan rivers: S. side of Mt.
Hilong-hilong: CAS 13922–25, 137533–34; Municipality of Remedios T.
Romualdez, Eye Falls, intersection of Dayhopan and Agan Rivers, Mt.
Hilong-hilong: KU 332972–007; Municipality of Cabadbaran, Barangay Tag-
Ibo, Dalaydayan River: USNM 305594–97; MISAMIS ORIENTAL PROVINCE,
Municipality of Gingoog City, Barangay Lumotan, Sitio San Isidro, Mt.
Balatukan: KU 319777–82; Barangay Lawan, Sitio Kibuko, Mt. Lumot: KU
333014–67; DAVAO DEL NORTE PROVINCE,Municipality of New Bataan, Sitio
Liboton, Mt. Puting Bato (Malaya River drainage): CMNH 5603–04; DAVAO
CITY PROVINCE,Municipality of Paquibato, Barangay Malambuon, Mt.
Makaayat: PNM 2880–81; DAVAO DEL SUR PROVINCE,Mt. Apo: KU 327521;
SAMAR ISLAND: NORTHERN SAMAR PROVINCE:Municipality of San Isidro:
Barangay Matuquinao: CAS-SU 18160, 18167–69, 18172–73; EASTERN
SAMAR PROVINCE:Municipality of Taft, Barangay San Rafael: KU 338613–
34, 338648–61, 310697–98; WESTERN SAMAR PROVINCE:Municipality of San
Jose de Buan; Barangay Uno, Mt. Huraw: KU 338021, 338635–45; LEYTE
ISLAND: CABALIAN: MCZ A-23190 (holotype of Rana everetti albotubercu-
lata), A-23188–89, A-132410–14, A-132416–19 (topotypes Rana everetti
202 Herpetological Monographs 31, 2017
albotuberculata); BOHOL ISLAND, BOHOL PROVINCE,Municipality of
Cantub, Sierra Bullones: CAS 137028.
Sanguirana sanguinea.—PALAWAN ISLAND: CMNH 3700–01, 3733,
3737; PALAWAN PROVINCE,Puerto Princesa City, Barangay Irawan: KU
308987, 309016, 309019–21, 309023–24, 309026, 309027–31, 309033,
309037, 309094; Municipality of Rizal, Mt. Bintangor: KU 311312;
Municipality of Brooke’s Point, Barangay Mainit, Mainit Falls: KU 309570;
Barangay Samarinana, Mt. Mantalingajan, area known locally as ‘‘Pitang’’ :
KU 309577, 309578, 309587.
Sanguirana tipanan.—LUZON ISLAND, AURORA PROVINCE,Municipal-
ity of San Luis, Barangay Villa Aurora, Aurora National Park: PNM 5727
(holotype of Rana tipanan), CMNH 5579–86, 5588, 5590–99, PNM 5720–
26, 5728–36, 5738–41 (paratypes of Rana tipanan); Municipality of Maria
Aurora: Aurora Memorial National Park: KU 322755–66; Barangay Villa
Aurora, Aurora Memorial National Park, Mt. Dayap, area known locally as
‘‘Siete’’: KU 322767–94; Municipality of San Luis: Barangay Lipimental: KU
322795–805, 322808–58, 323013; Barangay Real, Sitio Minoli: 322806–07,
BROWN ET AL.—NEW PHILIPPINE SANGUIRANA