TAXONOMIC REVISION OF BRACHYMELES SAMARENSIS

 
 
Herpetological Monographs, 25, 2011, 76–112
E 2011 by The Herpetologists’ League, Inc.

      PHYLOGENY-BASED SPECIES DELIMITATION IN PHILIPPINE
       SLENDER SKINKS (REPTILIA: SQUAMATA: SCINCIDAE) II:
      TAXONOMIC REVISION OF BRACHYMELES SAMARENSIS AND
              DESCRIPTION OF FIVE NEW SPECIES
          CAMERON D. SILER1,3, ALLISON M. FUITEN1, ROBIN M. JONES1, ANGEL C. ALCALA2,
                                                           1
                                        AND RAFE M. BROWN
  1
   Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, 1345 Jayhawk
                                     Boulevard, Lawrence, KS 66045-7593, USA
        2
          SUAKCREM Marine Laboratory, Silliman University, Bantayan, Dumaguete City, Philippines 6200

          ABSTRACT: With robust new datasets from morphology and DNA sequences, we review the limbed,
       nonpentadactyl species of the Brachymeles samarensis complex (now known to include B. cebuensis, B.
       minimus, and B. lukbani), and describe five new species in this highly limb-reduced, endemic Philippine
       clade of scincid lizards. For more than four decades, B. samarensis has been recognized as a single
       ‘‘widespread’’ species. This perception of the species’ peculiar geographic range has persisted as a result of
       weak sampling and similar gross morphology (body sizes, scale pigmentation) among populations. However,
       previous authors have noted morphological variation between different island populations, and our new data
       build on these observations and extend them to delimit new proposed species boundaries. Our data indicate
       that the ‘‘widespread’’ species B. samarensis is actually a complex of six distinct lineages, some of which are
       not each others’ closest relatives, and each of which is genetically unique. The taxa we define possess
       allopatric geographic ranges and differ from their congeners by numerous diagnostic characters of external
       morphology, and therefore should be recognized as full species in accordance with any lineage-based species
       concept. Species diversity in the genus has doubled in the last 3 yr, with these six taxa increasing the total
       known number of species of Brachymeles to 30.
         Key words: Biodiversity; Endemism; Faunal region; Fossorial lizards; Limb reduction; Species
       delimitation; Taxonomy




   FEW GENERA of scincid lizards are known to                     B. tridactylus, and B. wrighti), and five are
possess species representing a full spectrum of                   entirely limbless (B. apus, B. minimus, B.
body forms, from fully limbed, pentadactyl                        miriamae, B. lukbani, and B. vermis).
species to limbless species (see Siler et al.,                       Among the nonpentadactyl species, numer-
2011b; Siler and Brown, 2011). In the genus                       ous studies have documented a wide range of
Brachymeles, all but two of the 26 recognized                     limb- and digit-reduced states, from minute
species are endemic to the Philippines, with                      limbs with 0–3 digits (B. bonitae, B. cebuensis,
the exceptions being a single species (B. apus)                   B. muntingkamay, B. samarensis, B. tridacty-
from northern Borneo and another (B.                              lus), to moderately developed limbs with
miriamae) from Thailand (Brown and Alcala,                        four to five digits on the hands and feet (B.
1980; Hikida, 1982; Siler, 2010; Siler et al.,                    elerae, B. pathfinderi, B. wright; Duméril and
2009, 2010a,b, 2011a,b,c; Siler and Brown,                        Bibron, 1839; Brown, 1956; Brown and Rabor,
2010, 2011). Thirteen species are pentadactyl                     1967; Brown and Alcala, 1980; Taylor, 1917,
(B. bicolor, B. boholensis, B. boulengeri, B.                     1918, 1925; Siler, 2010; Siler et al., 2009,
gracilis, B. kadwa, B. makusog, B. mind-                          2010a, 2011a,b,c; Siler and Brown, 2011). All
orensis, B. orientalis, B. schadenbergi, B.                       species are semifossorial and typically found
talinis, B. taylori, B. tungaoi, and B. vindumi),                 in dry, rotting material inside or underneath
eight are nonpentadactyl with incompletely                        decaying logs or in loose soil, forest floor
developed limbs and reduced numbers of                            detritus, and leaf litter.
digits (B. bonitae, B. cebuensis, B. elerae, B.                      Although the genus was named well over
muntingkamay, B. pathfinderi, B. samarensis,                      150 yr ago (Duméril and Bibron, 1839), the
                                                                  rate of Brachymeles species descriptions
  3
      CORRESPONDENCE: e-mail, camsiler@ku.edu                     reached an apparent asymptotic maximum in

                                                             76
2011]                           HERPETOLOGICAL MONOGRAPHS                                        77


1980 (Brown and Alcala, 1980). The one             diversity in the genus for 30 yr, until Taylor
exception is B. minimus, a legless species         published a series of herpetofaunal descriptions
described in 1995 (Brown and Alcala, 1995).        in the early 1900s (Taylor, 1917, 1918, 1922,
For more than a century, limited numbers of        1925). It would be 50 yr before Brown (1956)
specimens in museum collections, combined          described B. samarensis from a single juvenile
with the similar body plans and external           specimen (FMNH 44472) collected in Guiuan,
morphological features among species of            Samar Island, Philippines in 1945. At the time of
Brachymeles, limited assessments of species-       description, Brown (1956) hypothesized the
level diversity (Brown, 1956; Brown and            species was most closely related to B. elerae
Alcala, 1980; Brown and Rabor, 1967; Taylor,       due to similarities in the number of paraverte-
1917). Recent studies have revealed the            bral scale rows. This single juvenile would
species-level diversity within Brachymeles to      remain the only vouchered specimen of this
be drastically underestimated, and have iden-      unique bidactyl species from Samar Island (Siler
tified numerous nonmonophyletic species            and Brown, 2010).
complexes within the Philippines (Siler and           By the time Brown and Rabor (1967)
Brown, 2010, 2011; Siler et al., 2011a).           revised the genus Brachymeles, samples of
Additionally, several rare, mid-to-high eleva-     specimens morphologically similar to B. sa-
tion species long represented by only a few        marensis had been collected from the islands
specimens (e.g., B. bicolor, B. elerae, B.         of Luzon and Leyte. Additionally, Brown and
wrighti, B. pathfinderi) have recently been        Rabor (1967) reported on a second specimen
rediscovered (Siler, 2010) and redescribed as      from Samar Island; however, no information
valid taxa (Siler et al., 2011a,c). Together,      on where the specimen was deposited, or its
these studies coupled with increased sampling      museum catalog number, was provided. Al-
throughout the Philippines and a new, robust       though Brown and Rabor (1967) treated B.
molecular dataset, allow us to begin evaluating    samarensis as a single widespread species,
variation across the isolated populations of       they referred to the species as a ‘‘complex,’’
widespread species in the Philippines.             suggesting that they suspected it contained
   In a recent study, Siler and Brown (2010)       multiple species. They also noted several
revised two polytypic species (B. boulengeri       distinct morphological differences between
and B. schadenbergi) and one widespread            island populations, including differences in
species (B. talinis), and inferred the presence    fore- and hind-limb digit number and head
of 10 genetically and morphologically distinct     scale patterns.
allopatric evolutionary lineages (species). Sev-      Additional island populations of B. samar-
eral other species are still recognized as         ensis were subsequently sampled by the time
having widespread distributions that span          Brown and Alcala (1980) revised the genus,
historical faunal demarcations in the Philip-      including populations from the Lapinig Group
pines (Brown and Diesmos, 2002, 2009;              islands off the northeast coast of Bohol Island
Brown and Guttman, 2002; Heaney, 1985),            (Fig. 1). Ross and Gonzales (1992) would later
including B. samarensis and B. bonitae             report on observations of B. samarensis from
(Brown, 1956; Brown and Rabor, 1967; Brown         Catanduañes Island off the northeast coast of
and Alcala, 1980). One of these species (B.        the Bicol Peninsula (Fig. 1) and in 2001, RMB
samarensis) is the focus of this study.            (unpublished data) recorded B. samarensis on
                                                   the southern tip of the Bicol Peninsula in the
              TAXONOMIC HISTORY                    foothills of Mt. Bulusan.
  The genus Brachymeles was first described by        To date, Brachymeles samarensis remains a
Dumeril and Bibron (1839) for the small, limb-     widespread species spanning islands of the
reduced species B. bonitae. Three additional       Luzon and Mindanao Pleistocene Aggregate
species (Senira bicolor Gray, 1845; Eumeces        Island Complexes (PAICs; Brown and Dies-
(Riopa) gracilis Fischer, 1885; E. (R.) schaden-   mos, 2002; Brown and Guttman, 2002;
bergi Fischer, 1885) were transferred to the       Fig. 1). Widespread distributions such as this
genus by Boettger (1886) and Boulenger (1887).     have been the focus of many recent studies
These four species represented the known           (Brown et al., 2000; Siler and Brown, 2010;
78                                      HERPETOLOGICAL MONOGRAPHS                                          [No. 25


                                                               distinct evolutionary lineages (species). In this
                                                               paper we provide a phylogenetic analysis of all
                                                               of these taxa, fully describe each species, clarify
                                                               taxonomic boundaries, and provide the first
                                                               illustrations of all included species. We also
                                                               provide information on each species’ natural
                                                               history, ecology, and geographic distribution.

                                                                           MATERIALS AND METHODS
                                                                    Field Work, Sample Collection, and
                                                                             Specimen Preservation
                                                                  Fieldwork was conducted on Catanduañes,
                                                               Lapinig Grande, Leyte, Luzon, and Samar
                                                               islands, all in the Philippines (Fig. 1) between
                                                               2001 and 2009. Specimens were collected
                                                               between 900 and 1600 h, euthanized in
                                                               aqueous chloretone, dissected for tissue sam-
                                                               ples (liver preserved in 95% ethanol or flash
                                                               frozen in liquid nitrogen), fixed in 10%
                                                               formalin, and eventually (,2 mo) transferred
                                                               to 70% ethanol. Newly sequenced specimens
                                                               are deposited in US and Philippine museum
                                                               collections (see Acknowledgments and Ap-
                                                               pendix I, Additional Specimens Examined);
                                                               voucher information corresponding to data
   FIG. 1.—Map of the Philippine islands, with island
labels provided for islands with representative samples
                                                               from GenBank sequences is included in
used for this study. The five recognized major Pleistocene     Table 1. Museum abbreviations for specimens
Aggregate Island Complexes (PAICs), major island               examined follow Leviton et al. (1985).
groups, and additional deep-water islands are labeled for
reference. Islands of the Romblon Island Group are              Taxon Sampling and Outgroup Selection for
designated by the first letter of the island name (T, Tablas                Phylogenetic Analyses
Island; R, Romblon Island; S, Sibuyan Island). Current
islands in the Philippines are shown in medium grey; light        Because our primary goal was to estimate
gray areas enclosed in black 120-m bathymetric contours        phylogenetic relationships among the various
indicate the hypothesized maximum extent of land during        populations of Brachymeles samarensis we
the mid- to late Pleistocene.                                  sequenced only a few exemplars (2–4) per
                                                               sampled population. We included samples of
Siler et al., 2010a,b; Welton et al., 2009,                    Lygosoma bowringi as an outgroup based on
2010a,b), which have revealed that few                         relationships presented in recent phylogenetic
endemic Philippine reptiles actually possess                   studies of the genus Brachymeles (Siler et al.,
broad distributions spanning these regional                    2011b; Siler and Brown, 2011). Additionally, we
faunistic boundaries (reviewed by Brown and                    included samples of B. apus, B. bonitae, B.
Diesmos, 2009).                                                cebuensis, B. lukbani, and B. minimus to explore
   The goal of the present study is to revise the              the sister group relationships within the B.
taxonomy of the B. samarensis complex such                     samarensis complex (Siler et al., 2011b; Siler
that individual units (species) represent inde-                and Brown, 2011). A total of 28 ingroup samples
pendently evolving, cohesive lineage segments                  were used in the phylogenetic analyses.
(sensu de Queiroz, 1998, 1999; Frost and
Hillis, 1990; Simpson, 1961; Wiley 1978).                               DNA Extraction, Purification,
Comprehensive examination of all recently                                     and Amplification
collected specimens from throughout the                           We extracted total genomic DNA from
known range of B. samarensis results in the                    tissues (Table 1) using the modified guanidine
reorganization of the species complex into six                 thiocyanate extraction method of Esselstyn et al.
2011]

TABLE 1.—Summary of specimens corresponding to genetic samples included in the study, general locality, and GenBank accession number. SP 5 Sabah Parks Reference
Collection, KU 5 University of Kansas Natural History Museum, LSUHC 5 La Sierra University Herpetological Collections, TNHC 5 Texas Natural History Collections of
                                                   the Texas Memorial Museum of the University of Texas at Austin.

                                                                                                                                GenBank accession numbers

               Species          Voucher                                   Locality                                    ND1       ND2           a-enolase      PTGER4

Lygosoma bowringii           LSUHC 6970   West Malaysia                                                            JN981975   JN981989     JN981962         JN982001
Lygosoma bowringii           LSUHC 6998   West Malaysia                                                            HQ907328   HQ907430     HQ906969            —
Brachymeles apus             SP 06915     Malaysia, Borneo, Sabah, Mt. Kinabalu National Park                      HQ907331   HQ907433     HQ906972         HQ907533
Brachymeles bonitae          KU 307747    Philippines, Polillo Island, Municipality of Quezon                      JN981976   JN981990     JN981963         JN982002
Brachymeles bonitae          KU 326080    Philippines, Polillo Island, Municipality of Quezon                      JN981977   JN981991     JN981964         JN982003
Brachymeles minimus          KU 308131    Philippines, Catanduanes Island, Municipality of Gigmoto                 HQ907404   HQ907508     HQ907039         HQ907608
Brachymeles minimus          KU 308129    Philippines, Catanduanes Island, Municipality of Gigmoto                 HQ907405   HQ907509     HQ907040         HQ907609
Brachymele lukbani           KU 313602    Philippines, Luzon Island, Municipality of Labo                          HQ907407   HQ907511     HQ907042         HQ907611
Brachymele lukbani           KU 313596    Philippines, Luzon Island, Municipality of Labo                          HQ907408   HQ907512     HQ907043         HQ907612
Brachymeles cebuensis        KU 320419    Philippines, Cebu Island, Municipality of Carcar                         HQ907410   HQ907514     HQ907045         HQ907614
Brachymeles cebuensis        KU 320421    Philippines, Cebu Island, Municipality of Carcar                         HQ907411   HQ907515     HQ907046         HQ907615
Brachymeles libayani2        KU 320430    Philippines, Lapinig Grande Island, Municipality of Carlos P.   Garcia   JN981978      —         JN981965            —
Brachymeles libayani2        KU 320458    Philippines, Lapinig Grande Island, Municipality of Carlos P.   Garcia   JN981979   JN981992        —             JN982004
Brachymeles libayani1        KU 320466    Philippines, Lapinig Grande Island, Municipality of Carlos P.   Garcia   JN981980   JN981993     JN981966         JN982005
Brachymeles libayani2        KU 320445    Philippines, Lapinig Grande Island, Municipality of Carlos P.   Garcia   JN981981      —         JN981967            —
Brachymeles bicolandia1      KU 324003    Philippines, Luzon Island, Municipality of Tobaco                        JN981982   JN981994     JN981968         JN982006
Brachymeles bicolandia2      KU 324016    Philippines, Luzon Island, Municipality of Tobaco                        JN981983   JN981995     JN981969         JN982007
Brachymeles bicolandia2      KU 324010    Philippines, Luzon Island, Municipality of Tobaco                        JN981984   JN981996     JN981970         JN982008
Brachymeles bicolandia2      KU 324004    Philippines, Luzon Island, Municipality of Tobaco                        JN981985   JN981997     JN981971         JN982009
Brachymeles brevidactylus3   TNHC 62469   Philippines, Luzon Island, Municipality of Sorsogon                      JN981986   JN981998     JN981972         JN982010
                                                                                                                                                                       HERPETOLOGICAL MONOGRAPHS




Brachymeles brevidactylus1   KU 324017    Philippines, Luzon Island, Municipality of Sorsogon                      JN981987   JN981999     JN981973         JN982011
Brachymeles cobos2           KU 324025    Philippines, Catanduanes Island, Municipality of San Miguel              JN981988   JN982000     JN981974         JN982012
Brachymeles cobos2           KU 324020    Philippines, Catanduanes Island, Municipality of San Miguel              HQ907349   HQ907450     HQ906989         HQ907550
Brachymeles cobos2           KU 324022    Philippines, Catanduanes Island, Municipality of San Miguel              HQ907350   HQ907451     HQ906990         HQ907551
Brachymeles samarensis       KU 310850    Philippines, Samar Island, Municipality of Taft                          HQ907416   HQ907520     HQ907051         HQ907620
Brachymeles samarensis       KU 310851    Philippines, Samar Island, Municipality of Taft                          HQ907417   HQ907521     HQ907052         HQ907621
Brachymeles samarensis       KU 310849    Philippines, Samar Island, Municipality of Taft                          HQ907413   HQ907517     HQ907048         HQ907617
Brachymeles paeforum2        KU 311225    Philippines, Leyte Island, Municipality of Baybay                        HQ907414   HQ907518     HQ907049         HQ907618
Brachymeles paeforum2        KU 311226    Philippines, Leyte Island, Municipality of Baybay                        HQ907418   HQ907522     HQ907053         HQ907622
Brachymeles paeforum2        KU 311227    Philippines, Leyte Island, Municipality of Baybay                        HQ907419   HQ907523     HQ907054         HQ907623
 1
     Holotype.
 2
     Paratopotype.
 3
     Paratype.
                                                                                                                                                                       79
80                                            HERPETOLOGICAL MONOGRAPHS                                             [No. 25

TABLE 2.—Models of evolution selected by Akaike Information Criterion (AIC) and applied for partitioned, Bayesian
                                            phylogenetic analyses.1

            Partition                           AIC Model                  Model applied          Number of characters

ND1, 1st codon position                      GTR + I + G                    GTR   +   G                  322
ND1, 2nd codon position                      GTR + I + G                    GTR   +   G                  322
ND1, 3rd codon position                      GTR + I + G                    GTR   +   G                  322
ND2, 1st codon position                      TVM + I + G                    GTR   +   G                  287
ND2, 2nd codon position                      GTR + I + G                    GTR   +   G                  287
ND2, 3rd codon position                      TVM + I + G                    GTR   +   G                  287
a-enolase                                    TVMef + G                      GTR   +   G                  261
PTGER4                                       HKY + I + G                    HKY   +   G                  490
 1
     The model GTR + G was used for partitioned RAxMLHPC analyses.


(2008). The mitochondrial NADH dehydroge-                            random addition-sequence replicates and tree
nase subunit 1 (ND1), NADH dehydrogenase                             bisection and reconnection (TBR) branch
subunit 2 (ND2), and the protein-coding nuclear                      swapping. To assess heuristic support, non-
loci, a-enolase and PTGER4, were completely                          parametric bootstrapping was conducted us-
sequenced for nearly all samples using the                           ing 1000 replicates, each with 100 random
primers and protocols provided in Siler et al.                       addition-sequence replicates and TBR branch
(2011b) and Siler and Brown (2011). We                               swapping.
purified polymerase chain reaction templates                            Partitioned maximum likelihood (ML) anal-
with 1 mL of a 20% solution of ExoSAP-IT                             yses were conducted in RAxMLHPC v7.04
(US78201, Amersham Biosciences, Piscataway,                          (Stamatakis, 2006). The alignment was parti-
NJ). Cycle-sequencing reactions were complet-                        tioned into eight regions consisting of the
ed with the same primers and ABI Prism                               codon positions of ND1 and ND2, and the
BigDye Terminator chemistry (Ver. 3.1; Applied                       two nuclear loci, a-enolase and PTGER4,
Biosystems, Foster City, CA). Cycle-sequenc-                         following the methods of Siler et al. (2011b)
ing products were purified with Sephadex Me-                         and Siler and Brown (2011). Analyses that
dium (NC9406038, Amersham Biosciences)                               partition protein-coding genes by codon posi-
in Centri-Sep 96 spin plates (CS-961, Prince-                        tion have been shown to improve resulting
ton Separations, Princeton, NJ). We analyzed                         inferences (Brandley et al., 2005). The partitions
purified products using an ABI Prism 3130xl                          were run under the same generalized time-
Genetic Analyzer (Applied Biosystems), and                           reversible model (GTR + C) with 100 replicate
gene sequences were assembled with Sequen-                           best-tree inferences. Each inference was per-
cher 4.8 (Gene Codes Corp., Ann Arbor, MI).                          formed with a random starting tree and relied
                                                                     on the rapid hill-climbing algorithm (Stamatakis,
     Alignment and Phylogenetic Analysis                             2006). Clade support was assessed with 1000
   Initial alignments of the gene regions were                       bootstrap pseudoreplicates. We considered
produced in Muscle v3.7 (Edgar, 2004) and                            branches receiving $70% bootstrap support to
manual adjustments were made in MacClade                             be well supported (Wilcox et al., 2002).
4.08 (Maddison and Maddison, 2005). No                                  The Akaike Information Criterion, as im-
instances of insertions or deletions, or ambig-                      plemented in jModeltest v0.1.1 (Guindon and
uously aligned regions, were observed in the                         Gascuel, 2003; Posada, 2008), was used to
data, and all data were used for analyses. The                       select the best model of nucleotide substitu-
final alignment consisted of 2570 nucleotide                         tion for each partition (Table 2). The best-
positions.                                                           fit model for each of the eight partitions
   Phylogenetic analyses were conducted us-                          (Table 2) was used for Bayesian analyses
ing parsimony and likelihood optimality crite-                       performed in MrBayes 3.1 (Ronquist and
ria, as well as Bayesian methods. Parsimony                          Huelsenbeck, 2003). The same partitioning
(MP) analyses were conducted in PAUP* 4.0                            strategy used for ML analyses was used for
(Swofford, 2002) with all characters weighted                        Bayesian inferences. Searches over tree space
equally. Most-parsimonious trees were esti-                          were conducted with four runs, each with four
mated using heuristic searches with 1000                             chains, and were run for 2 3 107 generations.
2011]                          HERPETOLOGICAL MONOGRAPHS                                         81


Trees were sampled every 1000 generations,        1978). We consider as distinct lineages those
with 4000 samples discarded as burn-in; this      populations that are morphologically and
left 16,001 post–burn-in trees from each run      genetically distinct, especially if allopatric.
included in the posterior distribution of         Lineage-based species concepts have been
topologies. Visual inspection for chain statio-   employed in the recognition of Philippine
narity and high effective sample size (ESS)       biodiversity (Brown et al., 2000, 2008b, 2009;
values (equal to or greater than 800) was         Brown and Diesmos, 2002; Brown and Gutt-
conducted within the program Tracer v1.4          man, 2002; Gaulke et al., 2007; Siler and
(Rambaut and Drummond, 2007). Additional-         Brown, 2010; Welton et al., 2009, 2010a,b)
ly, correlations of split frequencies and cumu-   due to the highly partitioned nature of the
lative split frequencies were examined using      archipelago (Brown and Diesmos, 2009), and
the program AWTY (Nylander et al., 2008). We      because the geological history of the islands
considered topologies with posterior probabil-    has been so well documented (Hall, 2002;
ities $0.95 to be well supported (Leaché and     Voris, 2000; Yumul et al., 2009). In this study
Reeder, 2002; Wilcox et al., 2002).               we use an estimate of phylogenetic relation-
                                                  ships as a guide for delimiting species but
              Morphological Data                  restrict our diagnoses of new species to those
   We examined fluid-preserved specimens          populations that are clearly identified by
(Appendix I) for variation in qualitative and     diagnostic differences in nonoverlapping mor-
mensural characters. Sex was determined by        phological character states.
gonadal inspection, and measurements were
taken to the nearest 0.1 mm with digital
                                                                      RESULTS
calipers by C. D. Siler. X-rays were taken with
a company cabinet X-ray on Kodak MIN-R                               Phylogeny
2000 film exposed at 5 mA and 30 V for 1 min         Of 2570 mitochondrial and nuclear charac-
15 s.                                             ters, 848 were parsimony-informative. The
   Meristic and mensural characters were          maximum parsimony analysis inferred 10 most
chosen based on Siler et al. (2009, 2010a,b):     parsimonious trees (tree length 5 2084;
snout–vent length (SVL), axilla–groin distance    topology not shown; bootstrap support sum-
(AGD), total length, midbody width (MBW),         marized in Fig. 2). The resulting 100 inferenc-
midbody height (MBH), tail length (TL), tail      es from the partitioned RAxML ML analysis
width (TW), tail height (TH), head length         show an average likelihood score 2lnL
(HL), head width (HW), head height, snout–        12011.371112, with a single inference having
forearm length (SnFa), eye diameter (ED),         the highest likelihood score of 2lnL
eye–narial distance (END), snout length           12011.367644. Trees recovered from ML,
(SNL), internarial distance (IND), forelimb       MP, and Bayesian analyses are topologically
length (FLL), hind limb length (HLL), mid-        identical. No inferences support the monophy-
body scale-row count, paravertebral scale-row     ly of Brachymeles samarensis. All analyses
count, axilla–groin scale-row count, Finger-III   recover two reciprocally monophyletic clades
lamellae count, Toe-IV lamellae count, supra-     within the B. samarensis complex, each char-
labial count, infralabial count, supraciliary     acterized by the presence of multiple highly
count, and supraocular count. Additionally,       divergent, genetically distinct lineages (Fig. 2).
we counted the number of presacral vertebrae         The Leyte Island and Lapinig Group
from X-ray images of specimens. In the            Islands populations were recovered as a clade,
description, ranges are followed by mean 6        sister to B. cebuensis from Cebu Island
SD in parentheses.                                (Fig. 2). Topotypic B. samarensis from Samar
                                                  Island were recovered as sister to a clade of
               Species Concept                    two limbless species of Brachymeles (B.
  We follow the general lineage concept of        minimus and B. lukbani) and the Luzon and
species (de Queiroz, 1998, 1999) as a logical     Catanduañes island populations of B. samar-
extension of the evolutionary species concept     ensis (Fig. 2). Within Luzon Island, two se-
(Frost and Hillis, 1990; Simpson, 1961; Wiley,    parate lineages were recovered from the Bicol
82                                   HERPETOLOGICAL MONOGRAPHS                                            [No. 25




  FIG. 2.—Maximum likelihood (ML) estimate of combined mitochondrial and nuclear data for samples of Brachymeles
used for this study (preferred ML tree, 2lnL 12011.367644; ND1, ND2, a-enolase, PTGER4). Nodes are shown with
numerical values corresponding to maximum parsimony bootstrap proportions, maximum likelihood bootstrap
proportions, and Bayesian posterior probabilities, respectively. Terminals are labeled with taxonomic names, fore-
and hind limb digit states, and number of presacral vertebrae.

Peninsula, with no support for their mono-                both limbless species, nested within a didactyl
phyly (Fig. 2).                                           clade of species formerly part of B. samarensis
   In addition to the finding of paraphyly of B.          (topotypic B. samarensis, B. sp. nov. [Catan-
samarensis complex members, several other                 duañes Island], B. sp. nov. [Southern Bicol
recognized species were recovered as part of              Peninsula, Luzon Island], B. sp. nov. (Central
this clade. Brachymeles cebuensis, one of only            Bicol Peninsula, Luzon Island]; Fig. 2). All
two recognized species to have unequal                    three previously recognized species (B. ce-
numbers of fingers and toes (Brown and                    buensis, B. lukbani, B. minimus) have geo-
Alcala, 1980), was supported as part of a clade           graphical distributions that overlap, or are in
of species (B. cebuensis + B. sp. nov. [Leyte             close proximity to, the known ranges of other
Island] + B. sp. nov. [Lapinig Group Islands])            species in the B. samarensis complex (Fig. 3).
with three fingers and two or three toes                     All analyses result in the strong support
(Fig. 2). Sister to this three-finger clade, all          of six genetically distinct lineages within
analyses recovered B. minimus and B. lukbani,             the Brachymeles samarensis species complex
2011]                                 HERPETOLOGICAL MONOGRAPHS                                                   83




   FIG. 3.—(Left) Map of the Philippine islands showing previously recognized distribution of Brachymeles samarensis
(indicated by black shaded islands), and recognized distributions of other members of the B. samarensis complex
(indicated by dark gray shapes). (Right) Hypothesized distributions of B. bicolandia, B. brevidactylus, B. cobos, B.
libayani, B. paeforum, and B. samarensis in the eastern-central Philippines. The sampling localities are indicated by
black shapes, and the hypothesized geographic range of each species is indicated by shaded islands and dashed lines,
with shapes and shades of islands corresponding to the map’s key.

(Fig. 2). Uncorrected pairwise sequence di-                 Mitochondrial sequence divergences among
vergences are low within the lineages defined               the other three lineages within the B. samar-
here as species and high between these                      ensis species complex (B. samarensis [Samar
lineages (Table 3). Percentage of divergences               Island], B. sp. nov. [Lapinig Group islands], B.
for the mitochondrial and for the nuclear data,             sp. nov. (Leyte Island]) are greater than 9.2%
respectively, show that the monophyletic                    (Table 3; Fig. 2). Intraspecific sequence diver-
lineages defined by our phylogenetic analyses               gences are low (0.0–1.9%) in comparison to
(B. samarensis, B. sp. nov. [Leyte Island], B.              divergences among monophyletic lineages.
sp. nov. [Lapinig Group Islands], B. sp. nov.               Additionally, moderate levels of sequence diver-
[Catanduañes Island], B. sp. nov. [Southern                gence are observed for nuclear sequence data
Bicol Peninsula, Luzon Island], B. sp. nov.                 (Table 3).
(Central Bicol Peninsula, Luzon Island]) are
distinguished from congeners by levels of                                    Morphology
genetic divergence similar to, or greater than,               Variation in morphological characters
those between previously defined species—                   (Tables 4–6) mirrors the results observed in
viz., B. bonitae, B. cebuensis, B. minimus, and             phylogenetic analyses, and supports the rec-
B. lukbani (Table 3; Fig. 2). The two most                  ognition of six B. samarensis group lineages.
closely related lineages, B. sp. nov. (Catan-               Characters differing among these six lineages
duañes Island) and B. sp. nov. (Southern Bicol             include digit number, presacral vertebrae
Peninsula, Luzon Island), are separated by                  number, degree of digit development, head
4.1–4.7% mitochondrial sequence divergence.                 and body scale counts and patterns, and
84                                                                                                                                                                                                       HERPETOLOGICAL MONOGRAPHS                             [No. 25


     paeforum, B. libayani, B. bicolandia, B. cobos, B. brevidactylus, B. bonitae, B. cebuensis, B. lukbani, and B. minimus (Fig. 2). Percentages on the diagonal represent                                           pigmentation patterns (Tables 4–6; species
     TABLE 3.—Uncorrected pairwise sequence divergence (%) for mitochondrial data (below diagonal) and nuclear data (above diagonal), for Brachymeles samarensis, B.

                                                                                                                                                                                                                      accounts below), all of which are typical
                                                                                                                                                                              B. minimus




                                                                                                                                                                                                 3.2–4.4
                                                                                                                                                                                                 3.2–3.6

                                                                                                                                                                                                 3.4–3.5


                                                                                                                                                                                                 0.7–0.8
                                                                                                                                                                                                   0.0
                                                                                                                                                                                                   3.4
                                                                                                                                                                                                   3.3


                                                                                                                                                                                                   3.5

                                                                                                                                                                                                   3.2
                                                                                                                                                                                                   3.2
                                                                                                                                                                                                                      morphological diagnostic characters employed
                                                                                                                                                                                                                      historically by taxonomists working with this
                                                                                                                                                                                                                      genus (review: Brown and Alcala, 1980). We
                                                                                                                                                                                                                      observed no intraspecific mensural or meristic
                                                                                                                                                                              B. lukbani

                                                                                                                                                                                                 2.7–3.1

                                                                                                                                                                                                 3.0–3.6
                                                                                                                                                                                                 2.6–3.2
                                                                                                                                                                                                 2.8–3.1
                                                                                                                                                                                                 2.7–3.1
                                                                                                                                                                                                 2.6–3.0
                                                                                                                                                                                                 2.8–3.0

                                                                                                                                                                                                 4.9–5.0
                                                                                                                                                                                                                      differences between the sexes of any of the six



                                                                                                                                                                                                   0.1
                                                                                                                                                                                                   3.0


                                                                                                                                                                                                                      species.
                                                                                                                                                                                                                         Superficially, the six lineages within the B.
                                                                                                                                                                                                                      samarensis complex appear morphologically
                                                                                                                                                                                                                      similar in overall body size and general shape;
                                                                                                                                                                              B. cebuensis




                                                                                                                                                                                                 1.4–2.8
                                                                                                                                                                                                 1.1–1.6

                                                                                                                                                                                                 1.4–1.6



                                                                                                                                                                                                                      however, upon closer inspection, three dis-
                                                                                                                                                                                                   0.0
                                                                                                                                                                                                 13.4
                                                                                                                                                                                                 13.7
                                                                                                                                                                                                   1.2
                                                                                                                                                                                                   1.3


                                                                                                                                                                                                   1.5

                                                                                                                                                                                                   2.2



                                                                                                                                                                                                                      tinct body forms are observed. Among the six
                                                                                                                                                                                                                      lineages, two are tridactyl (B. sp. nov. [Leyte
                                                                                                                                                                                                                      Island] and B. sp. nov. [Lapinig Group
                                                    intraspecific genetic diversity for mitochondrial data (bolded for emphasis).




                                                                                                                                                                                                 16.5–16.7
                                                                                                                                                                                                 15.0–15.1
                                                                                                                                                                                                 15.0–15.1




                                                                                                                                                                                                                      Islands]), three are bidactyl (B. samarensis,
                                                                                                                                                                              B. bonitae




                                                                                                                                                                                                  2.2–4.8
                                                                                                                                                                                                  2.3–2.7

                                                                                                                                                                                                  2.6–2.8
                                                                                                                                                                                                    1.4
                                                                                                                                                                                                    2.4
                                                                                                                                                                                                    2.6


                                                                                                                                                                                                    2.7




                                                                                                                                                                                                                      B. sp. nov. [Catanduañes Island] and B. sp.
                                                                                                                                                                                                                      nov. [Central Bicol Peninsula, Luzon Island]),
                                                                                                                                                                                                                      and one is bidactyl, but with small, highly
                                                                                                                                                                                                                      reduced, and nearly imperceptible claws (B.
                                                                                                                                                                              B. brevidactylus




                                                                                                                                                                                                 15.7–16.3
                                                                                                                                                                                                 14.8–14.9
                                                                                                                                                                                                 11.8–11.9
                                                                                                                                                                                                  1.6–2.0
                                                                                                                                                                                                  1.2–3.3
                                                                                                                                                                                                  0.4–1.1
                                                                                                                                                                                                  0.7–1.2




                                                                                                                                                                                                                      sp. nov. [Southern Bicol Peninsula, Luzon
                                                                                                                                                                                                    1.9



                                                                                                                                                                                                  11.6
                                                                                                                                                                                                    1.5




                                                                                                                                                                                                                      Island]). Additionally, numerous nonoverlap-
                                                                                                                                                                                                                      ping differences were detected in meristic,
                                                                                                                                                                                                                      mensural, osteological, and color pattern
                                                                                                                                                                                                                      characters for each complex member, readily
                                                                                                                                                                                                 15.7–16.2


                                                                                                                                                                                                 11.0–11.2
                                                                                                                                                                                                  0.0–0.2
                                                                                                                                                                                                  1.6–3.6
                                                                                                                                                                                                  0.5–0.9

                                                                                                                                                                                                  9.3–9.5
                                                                                                                                                                              B. cobos




                                                                                                                                                                                                                      defining six distinct lineages within the com-
                                                                                                                                                                                                    1.6
                                                                                                                                                                                                    2.0




                                                                                                                                                                                                   14.7
                                                                                                                                                                                                   11.2




                                                                                                                                                                                                                      plex (Tables 4–6). In summary, each lineage
                                                                                                                                                                                                                      (most of which are allopatric) possesses unique
                                                                                                                                                                                                                      and nonoverlapping suites of diagnostic char-
                                                                                                                                                                              B. bicolandia




                                                                                                                                                                                                 15.8–16.5
                                                                                                                                                                                                 14.6–14.8
                                                                                                                                                                                                 11.5–11.9
                                                                                                                                                                                                 11.6–12.0




                                                                                                                                                                                                                      acter states of morphology, perfectly cor-
                                                                                                                                                                                                  0.4–0.7
                                                                                                                                                                                                  1.1–1.5
                                                                                                                                                                                                  1.8–2.2
                                                                                                                                                                                                  1.4–2.8

                                                                                                                                                                                                  4.1–4.7
                                                                                                                                                                                                  9.4–9.6




                                                                                                                                                                                                                      responding to the six clades defined in the
                                                                                                                                                                                                                      phylogenetic analyses of DNA sequence data.
                                                                                                                                                                                                                                  Taxonomic Conclusions
                                                                                                                                                                                                 14.9–15.8
                                                                                                                                                                                                 14.5–14.8
                                                                                                                                                                                                 14.7–15.4
                                                                                                                                                                                                 15.5–16.5
                                                                                                                                                                                                 12.0–12.4
                                                                                                                                                                                                 14.8–15.3
                                                                                                                                                                                                 13.8–14.3
                                                                                                                                                                              B. libayani




                                                                                                                                                                                                  0.0–0.1
                                                                                                                                                                                                  1.8–2.8
                                                                                                                                                                                                  0.2–2.8




                                                                                                                                                                                                                         Our estimate of phylogeny (Fig. 2); biogeo-
                                                                                                                                                                                                                      graphically separate ranges of island or region
                                                                                                                                                                                                                      endemic species; diagnostic, nonoverlapping
                                                                                                                                                                                                                      morphological character states; and genetic
                                                                                                                                                                                                                      distances between the taxa (Table 3) indicate
                                                                                                                                                                              B. paeforum




                                                                                                                                                                                                  9.2–10.5
                                                                                                                                                                                                 14.1–14.5
                                                                                                                                                                                                 14.4–14.6
                                                                                                                                                                                                 14.6–14.9
                                                                                                                                                                                                 15.0–15.6
                                                                                                                                                                                                 11.6–11.7
                                                                                                                                                                                                 14.1–14.4
                                                                                                                                                                                                 14.0–14.2
                                                                                                                                                                                                  0.0–0.7




                                                                                                                                                                                                                      the distinctiveness of a new species from
                                                                                                                                                                                                    2.2




                                                                                                                                                                                                                      Catanduañes Island, two new species from the
                                                                                                                                                                                                                      Bicol Peninsula of Luzon Island (one from the
                                                                                                                                                                                                                      central Bicol region and one from the extreme
                                                                                                                                                                              B. samarensis




                                                                                                                                                                                                 14.6–15.2
                                                                                                                                                                                                 14.5–14.7
                                                                                                                                                                                                 14.4–14.5
                                                                                                                                                                                                 14.9–15.0
                                                                                                                                                                                                 16.9–17.3




                                                                                                                                                                                                                      southern tip of the peninsula), a new species
                                                                                                                                                                                                    0.0
                                                                                                                                                                                                  14.3




                                                                                                                                                                                                  16.2
                                                                                                                                                                                                  14.1
                                                                                                                                                                                                  14.1




                                                                                                                                                                                                                      from the Lapinig Group Islands, and a new
                                                                                                                                                                                                                      species from Leyte Island (Table 3; Fig. 2).
                                                                                                                                                                                                                      Each of the six species of the B. samarensis
                                                                                                                                                                                                 brevidactylus




                                                                                                                                                                                                                      complex is morphologically distinct from each
                                                                                                                                                                                                 samarensis


                                                                                                                                                                                                 bicolandia
                                                                                                                                                                                                 paeforum




                                                                                                                                                                                                 cebuensis




                                                                                                                                                                                                                      other and all other known species in the
                                                                                                                                                                                                 minimus
                                                                                                                                                                                                 libayani




                                                                                                                                                                                                 lukbani
                                                                                                                                                                                                 bonitae




                                                                                                                                                                                                                      genus, and the 11 species of Brachymeles
                                                                                                                                                                                                 cobos




                                                                                                                                                                                                                      included in phylogenetic analyses also are
                                                                                                                                                                                                                      genetically distinct. All monophyletic lineages,
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
                                                                                                                                                                                                 B.
2011]                               HERPETOLOGICAL MONOGRAPHS                                           85


with the exception of the two occurring on               (Tables 4 and 5); from B. brevidactylus by
the Bicol Peninsula of Luzon Island, are                 having fewer presacral vertebrae and fewer
endemic to single islands within two isolated            paravertebral scale rows (Tables 4 and 5); and
PAICs, thereby providing additional support              from B. bonitae by having longer relative hind
for the distinctiveness of each clade’s evo-             limb lengths, fewer presacral vertebrae, fewer
lutionary history and lineage integrity. Ac-             paravertebral scale rows, six supraciliaries, five
cordingly, we recognize B. samarensis as a               supraoculars, and the presence of contact
species that occurs only on Samar Island in              between frontoparietals (Tables 4 and 5).
the eastern Visayan (central) Philippine islands            Brachymeles samarensis can be distin-
(e.g., Mindanao PAIC; Fig. 3), and hereby                guished from all limbless species of Brachy-
recognize the five additional lineages within            meles (B. apus, B. lukbani, B. minimus, B.
the B. samarensis species complex as new                 miriamae, B. vermis) by having limbs; and
species.                                                 from all pentadactyl species of Brachymeles
                                                         (B. boholensis, B. boulengeri, B. bicolor, B.
               TAXONOMIC ACCOUNTS                        gracilis, B. kadwa, B. makusog, B. mind-
    Brachymeles samarensis Brown 1956: 6                 orensis, B. orientalis, B. schadenbergi, B.
                      Figs. 3–4                          talinis, B. taylori, B. tungaoi, B. vindumi) by
   Brachymeles samarensis, Brown, 1956,                  having nonpentadactyl limbs, shorter adult
Type locality: Guinuan, Samar Island, Philip-            forelimb lengths (less than 2.6 mm vs. greater
pines (holotype: FMNH 44472); Brown and                  than 5.9 mm), shorter adult hind limb lengths
Alcala, 1970, 1980; Brown and Rabor, 1967.               (less than 3.1 mm vs. greater than 10.3 mm),
   Diagnosis.—Brachymeles samarensis can                 and a narrower body (less than 6.4 mm vs.
be distinguished from congeners by the                   greater than 7.9 mm), and by the absence of a
following combination of characters: (1) body            postnasal scale and auricular opening (vs.
size small (SVL 57.9–66.1 mm), (2) limbs                 presence).
bidactyl, (3) limb length short, (4) supralabials           Description (based on holotype description
six, (5) infralabials six, (6) supraciliaries six, (7)   and six referred specimens).—Details of the
supraoculars five, (8) midbody scale rows 19–            head scalation of an adult female are shown in
22, (9) axilla–groin scale rows 66–69, (10)              Fig. 5. Measurements and character states of
paravertebral scale rows 86–88, (11) pineal              the holotype are provided below in square
eye spot present, (12) prefrontals not contact-          brackets. Body small, slender; SVL 57.9 mm
ing on midline, (13) frontoparietals contact,            for males, maximum SVL 66.1 mm for
(14) mental/first infralabial fusion absent, (15)        females, [43.5 mm, juvenile] (Tables 4 and
postnasals absent, (16) enlarged chin shields            5); head weakly differentiated from neck,
in three pairs, (17) nuchal scales differentiat-         nearly as wide as body, HW 7.3–9.2% (8.3 6
ed, (18) fourth supralabial below eye midline,           0.7) SVL, 91.4–117.8% (102.7 6 10.8) HL;
(19) auricular opening absent, (20) presacral            HL 36.6–42.5% (38.8 6 2.1) SnFa; SnFa
vertebrae 45, and (21) uniform body color                18.8–23.5% (20.9 6 1.6) SVL; snout short,
(Tables 4 and 5).                                        bluntly rounded in dorsal and lateral profile,
   Comparisons.—Characters distinguishing                SNL 50.9–55.3% (53.3 6 1.8) HL; ear
Brachymeles samarensis from all nonpenta-                completely hidden by scales; eyes small, ED
dactyl, limbed species of Brachymeles are                1.3–1.6% (1.4 6 0.1) SVL, 17.0–18.7% (17.6
summarized in Tables 4 and 5. Brachymeles                6 0.6) HL, 42.6–48.0% (45.8 6 2.1) END,
samarensis most closely resembles B. bicolan-            pupil subcircular; body slightly depressed,
dia, B. cobos, B. brevidactylus, and popula-             nearly uniform in thickness, MBW 109.1–
tions of B. bonitae, the only other bidactyl             150.6% (130.4 6 14.9) MBH; scales smooth,
species. However, B. samarensis differs from             glossy, imbricate; longitudinal scale rows at
these four taxa by having midbody scale rows             midbody 19–22 [22]; paravertebral scale rows
as few as 19 and axilla–groin scale rows as few          86–88 [86]; axilla–groin scale rows 66–69;
as 66 (Table 5). Brachymeles samarensis                  limbs short, poorly developed, with digits
further differs from B. bicolandia by having             reduced to two claws on both forelimbs and
fewer presacral vertebrae and six infralabials           hind limbs, finger and toe lamellae absent;
86                                         HERPETOLOGICAL MONOGRAPHS                                                                      [No. 25

TABLE 4.—Summary of meristic and mensural characters in all known limbed, nonpentadactyl species of Brachymeles.
Sample size, body length, and total length (TotL) among males and females, and general geographical distribution
(Pleistocene Aggregate Island Complexes [PAIC], sensu Brown and Diesmos, 2002) are included for reference. SVL 5
snout–vent length, TotL 5 total length, TL 5 tail length, FLL 5 forelimb length, HLL 5 hind limb length, ToeIVlam
5 toe-IV lamellae count. (SVL, TotL, MBW, FLL, and HLL given as range over mean 6 SD; all body proportions given
                                          as percentage over mean 6 SD).

                B. samarensis (1   B. paeforum (3 males, B. libayani (10 males, B. bicolandia (6 males,       B. cobos          B. brevidactylus (1
                male, 5 females)         9 females)            25 females)            10 females)            (9 females)         male, 2 females)

                                                              Lapinig Group           Central Bicol                              Southern Bicol
     Range       Samar Island           Leyte PAIC               Islands               Peninsula          Catanduañes Island      Peninsula

SVL (female)    62.4–66.1              47.2–61.4             52.8–66.1              46.4–67.4               54.0–64.4              54.0, 60.0
                (63.4 6 1.5)          (56.5 6 4.2)           (58.6 6 3.3)           (59.0 6 6.7)            (58.7 6 3.5)
SVL (male)          57.9               62.4–66.1             52.7–57.4              56.4–66.1                   —                  56.8
                                      (63.4 6 1.5)           (56.0 6 1.6)           (61.7 6 3.5)
TotL (female)   97.7–112.9             99.5–108.5            91.4–111.2             94.1–112.7             102.2–109.4             92.3, 95.2
                (107.3 6 8.3)         (102.6 6 5.1)          (102.2 6 6.4)          (102.1 6 8.8)           (106.2 6 2.7)
TotL (male)         93.0              106.7–114.6            92.1–103.4             99.6–107.9                  —                 102.0
                                      (110.6 6 5.6)          (99.4 6 4.5)           (104.1 6 4.2)
TL/SVL           57–81                   69–79                 63–84                  58–93                   76–96                59–80
                (68 6 12)             (75 6 4)               (77 6 6)               (75 6 12)               (88 6 8)             (70 6 11)
FLL             1.1–2.6                 1.3–1.7               1.1–1.8                1.1–1.9                 1.4–2.1              1.1–1.5
                (1.8 6 0.5)           (1.5 6 0.1)            (1.3 6 0.2)            (1.4 6 0.3)             (1.7 6 0.2)          (1.5 6 0.2)
FLL/SVL           2–4                     2–4                   2–3                    2–4                     3                    2–3
                (3 6 1)               (3 6 0)                (2 6 0)                (2 6 0)                 (3 6 0)              (3 6 0)
HLL             2.5–3.1                 2.3–3.0               2.0–2.7                1.9–3.1                 2.5–3.6              2.1– 2.7
                (2.9 6 0.2)           (2.6 6 0.3)            (2.4 6 0.2)            (2.6 6 0.3)             (3.0 6 0.3)          (2.5 6 0.3)
HLL/SVL           4–5                     4–5                   3–5                    3–5                     4–6                  4–5
                (5 6 0)               (4 6 0)                (4 6 0)                (4 6 1)                 (5 6 1)              (4 6 0)
ToeIVlam              0                   0                     0                      0                       0                    0


FLL 2.4–5.7% (3.9 6 1.3) AGD, 1.8–3.9%                                  anterior loreal about as long as and slightly
(2.9 6 0.9) SVL; HLL 5.3–7.2% (6.2 6 0.7)                               higher than posterior loreal; preocular one;
AGD, 4.0–5.0% (4.6 6 0.4) SVL [6.9]; tail not                           presubocular one; supraciliaries six, the ante-
as wide as body, gradually tapered towards                              riormost contacting prefrontal and separating
end, TW 70.2–82.6% (76.7 6 5.0) MBW, TL                                 posterior loreal from first supraocular, poste-
56.5–80.6% (68.4 6 11.6) SVL.                                           riormost extending to posterior edge of fifth
   Rostral projecting onto dorsal snout to level                        supraocular; subocular scale row single, com-
in line with middle of nasal, broader than                              plete, in contact with supralabials; lower
high, in contact with frontonasal; frontonasal                          eyelid with one row of scales; supralabials
wider than long; nostril ovoid, in center of                            six, first twice the anteroposterior length of
single trapezoidal nasal, longer axis directed                          others, fourth below eye midline; infralabials
anteroventrally and posterodorsally; suprana-                           six (Fig. 4).
sals present, large, broadly separated; postna-                            Mental wider than long, in contact with first
sals absent; prefrontals moderately separated;                          infralabials; postmental single, enlarged, its
frontal octagonal-shaped, its anterior margin                           width equal to width of mental; followed by
in moderate contact with frontonasal, in con-                           three pairs of enlarged chin shields, first pair
tact with first two anterior supraoculars, 33                           in broad medial contact, second pair wider
wider than anterior supraocular; supraoculars                           than first, broadly separated by single medial
five; frontoparietals moderate, in broad medi-                          scale, third pair separated by three medial
al contact, each frontoparietal in contact with                         scales (Fig. 4).
supraoculars 2–4; interparietal moderate,                                  Scales on limbs smaller than body scales;
its length roughly equal to midline length of                           scales on dorsal surfaces of digits wrapping
frontoparietal, longer than wide, diamond-                              around lateral edges of digits; lamellae absent;
shaped, wider anteriorly; parietals broader                             palmar surfaces of hands and plantar surfaces
than frontoparietals, in broad contact behind                           of feet with several small, irregular scales,
interparietal; nuchals enlarged; loreals two,                           each with irregular raised anterior edges;
2011]                                          HERPETOLOGICAL MONOGRAPHS                                                                     87


                                                          TABLE 4.—Extended




  B. muntingkamay   B. tridactylus (9 males,   B. bonitae (6 males,    B. cebuensis   B. elerae (2 males,     B. pathfinderi      B. wrighti (1
     (12 females)         11 females)               7 females)         (8 females)         1 female)        (14 m, 23 females)   male, 1 female)

                                                Mindoro & Luzon                                                 Mindanao
    Luzon Island         Visayan PAIC               PAICs              Cebu Island       Luzon Island            Island          Luzon Island

    61.8–81.3           45.5–59.1                49.7–59.8             51.5–67.9        68.2, 71.9            55.8–68.3           113.0 1
    (73.6 6 5.9)       (52.1 6 5.0)             (56.4 6 3.9)                                                  (62.0 6 3.4)
       N/A              55.7–78.3                65.1–80.0             (61.8 6 5.3)        71.5               54.5–65.1             25.8
                       (68.5 6 7.4)             (73.5 6 6.4)                N/A                               (59.4 6 3.8)
   107.4–136.0         102.6–154.1               93.4–150.4           104.3–128.0      109.9, 131.9          111.1–133.2          205.6
    (124.0 6 8.6)      (132.6 6 14.0)           (126.7 6 19.9)        (119.0 6 8.5)                           (119.7 6 8.2)
       N/A             105.3–133.67             102.6–144.5                                  N/A             101.4–107.0          216.4
                       (115.9 6 15.4)           (121.3 6 15.6)              N/A                               (104.2 6 4.0)
       50–79              69–112                    35–93                78–115            61–84                69–95               72, 82
    (65 6 10)          (92 6 12)                (69 6 18)             (92 6 13)          (72 6 16)            (84 6 10)
      2.4–3.0            1.5–2.5                   1.0–1.5              1.1–1.8           3.3–3.5              4.4–6.9             7.5, 7.5
    (2.7 6 0.2)        (2.0 6 0.3)              (1.3 6 0.1)           (1.5 6 0.3)        (3.4 6 0.1)          (5.8 6 0.5)
        3–4                2–3                       1–2                  2–3               5–5                  8–11                6, 7
     (4 6 0)           (3 6 0)                  (2 6 0)               (2 6 0)            (5 6 0)              (10 6 1)
      5.3–6.0            2.6–3.6                   1.3–2.0              2.3–3.0           4.3–5.4              8.4–12.9          10.9, 13.9
     (5.7 6 0.2)       (3.1 6 0.3)              (1.6 6 0.2)           (2.7 6 0.3)        (5.0 6 0.6)          (10.8 6 1.0)
        7–9                3–6                       2–3                  3–5               6–8                 15–21               10, 11
     (8 6 1)           (5 6 1)                  (2 6 0)               (4 6 0)            (7 6 1)              (18 6 1)
         0                 0                         0                    0                 3                    5–8                 4, 5



fingers equal in size; toes unequal in size,                           minor differences, including a dark brown
middle digit greatest in length, first and third                       body color and dark brown to black streaks of
digits equal in length.                                                pigmentation.
   Coloration in preservative.—Body ground                                Variation.—Morphometric variation of the
color medium brown, each dorsal scale light                            series is summarized in Table 6. We observed a
brown posteriorly, with a dark auburn streak                           single instance of digit variation, where one
on the anterior two-thirds to half of the scale.                       specimen (KU 310849) has no fingers and two
Dark streaks on each scale consist of four to                          toes. All specimens have two loreals with the
seven thin longitudinal lines with smudges                             exception of a single specimen (KU 310852),
between lines. Streaks on scales present                               which has a single loreal on the right side of the
around entire body, more distinct on venter,                           body resulting from the fusion of the two scales
where posterior ends of scales are cream,                              in this position. Additionally, the first and
giving greater contrast. Posterior edge of all                         second pairs of enlarged chin shields are equal
body scales transparent. Forelimb and hind                             in width among all specimens with the exception
limb scales same color as body scales.                                 of a single specimen (KU 310850), in which the
Precloacal scale coloration matches surround-                          width of the second pair of enlarged chin shields
ing ventral scale coloration. Head scales                              is greater than the width of the first pair.
mottled light and dark brown, matching dorsal                             Distribution.—Brachymeles samarensis is
background coloration. Supraocular, rostral,                           known only from Samar Island (Fig. 3).
nasal, supranasal, and supralabials scales gray-                          Ecology and natural history.—Brachymeles
cream. Mental, infralabial, postmental, and                            samarensis occurs in primary- and secondary-
chin shields scales cream with slight brown                            growth forest habitats. In contrast to the other
mottling, lighter than bordering ventral scales.                       members of the B. samarensis complex, this
   Coloration in life.—Coloration in life close-                       species appears to be a forest obligate, and
ly matches the coloration in preservative with                         was only observed within rotting logs in
88                                                  HERPETOLOGICAL MONOGRAPHS                                                                    [No. 25

TABLE 5.—Summary of qualitative diagnostic characters (present, absent) in all known limbed, nonpentadactyl species of
Brachymeles. The pairs of enlarged scales posterior to the postmental scale are abbreviated as chin shield pairs with
reference to the first, second, and third pairs (when present). In cases of scale count variation within species, numbers of
individuals showing specific counts are given in parentheses. PSV 5 presacral vertebrae, MBSR 5 midbody scale-row
count, AGSR 5 axilla–groin scale-row count, PVSR 5 paravertebral scale-row count, SL 5 supralabial count, IFL 5
                           infralabial count, SC 5 supraciliary count, SO 5 supraocular count.

                               B. samarensis (1 male,     B. paeforum         B. libayani (10      B. bicolandia (6      B. cobos       B. brevidactylus
                                     5 females)       (3 males, 9 females)   males, 25 females)   males, 10 females)    (9 females)   (1 male, 2 females)

Number of digits                       2/2                   3/3                   3/3                   2/2               2/2               2/2
   (fore/hind)
PSV                                   45                    47                    47                  46–49               45               47–48
MBSR                                19–22                 21–22                 22–23                 20–22              21–22              20
AGSR                                66–69                 71–74                 72–75                 68–73              68–72             73–77
PVSR                                86–88                 93–96                 90–92                 85–90              85–89             90–94
SL                                  6 (6)                 6 (12)                6 (35)                6 (16)             6 (9)             6 (3)
IFL                                 6 (6)               5 (4) 6 (8)             5 (35)             5 (11) 6 (5)          6 (9)             6 (3)
SC                                  6 (6)                 6 (12)                6 (35)                6 (16)             6 (9)             6 (3)
SO                                  5 (6)                 5 (12)                5 (35)                5 (16)             5 (9)             5 (3)
Pineal eyespot                    Present               Present              Present               Present              Present           Present
Prefrontal contact                Absent                Absent               Point contact           Absent             Absent            Absent
                                                                                or Absent
Frontoparietal           Present                        Present              Present                Present or          Present           Present
   contact                                                                                             Absent
First chin shield        Present                        Present or           Present or             Present or          Present           Absent
   pair contact                                            Absent                Absent                Absent
Thirdrd chin shield pair Present                        Present              Present                 Present            Present          Present
Chin shield pair size    3,1#2                         3,1#2                  3,1,2                3,1,2               3,1#2            3,1,2
Chin shield pair         1(0); 2(1);                   1(0/1); 2(1);         1(0/1); 2(1);         1(0/1); 2(1);       1(0); 2(1);      1(0); 2(1);
   separation1                3(3)                          3(3)                  3(3)                  3(3)                3(3)             3(3)
First/second loreal      Present or                     Present or           Present or             Present or          Absent           Absent
   fusion                   Absent                         Absent                Absent                Absent
Mental/first IFL fusion Absent                          Present or           Present                Present or          Absent            Absent
                                                           Absent                                      Absent
Differentiated nuchals            Present               Present              Present                Present             Present           Present
Continuous subocular              Present               Present              Present                Present             Present           Present
   scale row
Auricular opening                 Absent                Absent               Absent                 Absent              Absent            Absent
Dorsolateral stripes              Absent                Absent               Absent                 Absent              Absent            Absent
Longitudinal rows                 Absent                Absent               Absent                 Absent              Absent            Absent
   of dark spots
     1
     Parentheses show the number of small ventral scale rows separating each enlarged pair of chin shields.
     2
     Due to head damage in the nuchal region for both known specimens of B. wrighti, the presence of differentiated nuchals remains tentative.




secondary-growth forest. Three species of                                              Other sympatric lizard species observed on
Brachymeles have been confirmed to occur                                            Samar Island include the following: (Agamidae)
on Samar Island: B. gracilis hilong, B. orien-                                      Bronchocela cristatella, Draco bimacula-
talis, and B. samarensis (Brown and Alcala,                                         tus, D. ornatus, D. reticulates, Gonocephalus
1980; Siler and Brown, 2010).                                                       semperi, Hydrosaurus pustulatus; (Gekkoni-
   We have evaluated this species against the                                       dae) Cyrtodactylus annulatus, C. sumoroi,
International Union for Conservation of                                             Gehyra mutilata, Gekko gecko, Gek. mind-
Nature (IUCN) criteria for classification, and                                      orensis, Hemidactylus frenatus, H. platyurus,
find that it does not qualify for Critically                                        Hemiphyllodactylus typus, Lepidodactylus aureo-
Endangered, Endangered, Vulnerable, or                                              lineatus, L. planicaudus, Pseudogekko compressi-
Near Threatened status. Brachymeles samar-                                          corpus; (Scincidae) Emoia atrocostata, Eutropis
ensis has been documented to have a broad                                           multicarinata, Eu. multifasciata, Lamprolepis
geographic distribution across southern Samar                                       smaragdina, Lipinia pulchella, Li. quadrivittata,
Island. We therefore classify this species as                                       Sphenomorphus acutus, S. cumingi, S. fasciatus,
having Least Concern status (IUCN, 2011).                                           S. jagori, S. cf. mindanensis, S. steerei, S.
2011]                                        HERPETOLOGICAL MONOGRAPHS                                                                      89


                                                         TABLE 5.—Extended




  B. muntingkamay       B. tridactylus        B. bonitae         B. cebuensis   B. elerae (2 males,     B. pathfinderi     B. wrighti (1 male,
     (12 females)   (9 males, 11 females) (6 males, 7 females)   (8 females)        1 females)      (14 males, 23 females)     1 female)

        3/3                 3/3               0–2/0–2                3/2               4/4                  5/4                   4/4

      42, 44               47                  47–57                 45               43                    34                   —
      22–24              22–24                 21–23               22–24            22–24                 23–25                28, 28
      65–70              70–79                 73–90               65–69            63–67                 44–48                85, 85
      85–90              88–98                90–109               84–88            84–87                 64–67               106, 108
      6 (12)          6 (12) 7 (8)          6 (12) 7 (1)            6 (8)            6 (3)                6 (37)             6 (1) 7 (1)
      6 (12)          6 (12) 7 (8)       5 (1) 6 (10) 7 (2)      6 (5) 7 (3)         6 (3)                6 (37)                7 (2)
   6 (10) 7 (2)          5 (20)             5 (12) 6 (1)            6 (8)         5 (2) 6 (1)         5 (17) 6 (19)               6
   5 (11) 6 (1)          4 (20)                4 (13)               5 (8)         4 (2) 5 (1)             5 (37)                  5
 Absent              Present                Present              Present        Absent                 Present               Present
 Present             Absent                 Absent               Present or     Present                Absent                Present or
                                                                    Absent                                                      Absent
 Absent              Absent                  Absent              Present        Present                 Present or           Present
                                                                                                           Absent
 Absent              Present or              Absent              Present        Absent                  Absent               Absent
                          Absent
 Present             Present                 Present             Present        Present                 Absent               Present
   3,1,2              3,1,2                  3,2,1               153,2            1,3,2                   1,2                  2,1
   1(1); 2(1);        1(0/1); 2(1);          1(1); 2(1);         1(0); 2(1);      1(1); 2(1);           1(1); 2(1)           1(1); 2(3)
        3(3)               3(3)                   3(3)                3(3)             3(3)
 Absent              Absent                  Absent              Absent         Absent                  Absent               Absent
 Absent              Absent                  Present or          Absent         Absent                  Absent               Absent
                                                Absent
 Absent              Present                 Present             Present        Absent                  Absent               Present2

 Absent              Absent                  Present             Present        Present                 Present              Present

 Absent             Absent                   Absent              Absent         Absent                  Present              Absent
 Absent             Absent                   Absent              Absent         Absent                  Present              Absent
 Present, around    Present, vague           Absent              Absent         Present, around         Present, 6           Present
       body           to indistinct                                                body



variegatus, Tropidophorus misaminus; (Varani-                          collected between 29 October and 8 Novem-
dae) Varanus cumingi samarensis.                                       ber 2007.
                                                                          Other paratypes.—One adult male (CAS-
          Brachymeles paeforum sp. nov.
                                                                       SU 26120), four adult females (CAS-SU
                  Figs. 3, 6, 7
                                                                       26110, 26112, 26121–22), and two juveniles
   Holotype.—PNM 9746 (CDS Field                                       (CAS-SU 26115, 26123) collected between 1
No. 3418, formerly KU 311228), adult female,                           May and 4 June 1964, in Barrio Tambis,
collected under rotting logs in secondary-growth                       Municipality of Burauen, Leyte Province, Leyte
forest (1000 to 1230 h) on 8 November 2007 in                          Island, Philippines (11u009370N, 124u529190E;
the Sitio San Vicente Tree Nursery, Barangay                           WGS-84), by D. S. Rabor; one adult male
Pilim, Baybay City, Leyte Province, Leyte                              (CAS-SU 26771), two adult females (CAS-SU
Island, Philippines (10u439350N, 124u499050E;                          26770, 26772), and one juvenile (CAS-SU
WGS-84), by CDS and J. Fernandez.                                      26773), collected between 10 June and 17 July
   Paratopotypes.—One adult male (KU                                   1964, in the Municipality of Mahaplag, Leyte
311225), one adult female (KU 311229), and                             Province, Leyte Island, Philippines (10u359420N,
three juveniles (KU 311224, PNM 9747–48)                               124u599130E; WGS-84), by D. S. Rabor.
TABLE 6.—Summary of univariate morphological variation among mensural characters in series of Brachymeles samarensis, B. paeforum, B. libayani, B. bicolandia, B. cobos,                                      90
and B. brevidactylus. SVL 5 snout–vent length, AGD 5 axilla–groin distance, TotL 5 total length, MBW 5 midbody width, MBH 5 midbody height, TL 5 tail length, TW
5 tail width, TH 5 tail height, HL 5 head length, HW 5 head width, HH 5 head height, SnFa 5 snout–forearm length, ED 5 eye diameter, END 5 eye–narial distance,
                                  SNL 5 snout length, IND 5 internarial distance, FLL 5 forelimb length, HLL 5 hind limb length.

                                                                                                                                                                                        B. brevidactylus (1
                 B. samarensis (1 male, 5 females)   B. paeforum (3 males, 9 females)   B. libayani (10 males, 25 females) B. bicolandia (6 males, 10 females)   B. cobos (9 females)    male, 2 females)

SVL (male)               57.9               59.7–64.1 (61.8 6 2.2)   52.7–57.4 (56.0 6 1.6)  56.4–66.1 (61.7 6 3.5)             —                                                          56.8
SVL (female)      62.4–66.1 (63.4 6 1.5)    47.2–61.4 (56.5 6 4.2)   52.8–66.1 (58.6 6 3.3)  46.4–67.4 (59.0 6 6.7)    54.0–64.4 (58.7 6 3.5)                                              54.0,   60.0
AGD (male)               40.3               44.1–48.2 (46.0 6 2.1)   39.5–43.6 (42.1 6 1.2)  42.9–52.0 (47.4 6 3.4)             —                                                          43.8
AGD (female)      45.3–50.3 (47.6 6 1.8)    35.6–46.5 (43.2 6 3.1)   39.4–50.6 (44.7 6 2.8)  32.9–52.3 (44.8 6 5.8)    41.0–49.1 (44.6 6 2.3)                                              45.5,   45.8
TotL (male)              93.0             106.7–114.6 (110.6 6 5.6) 92.1–103.4 (99.4 6 4.5) 99.6–107.9 (104.1 6 4.2)            —                                                         102.0
TotL (female)    97.7–112.9 (107.3 6 8.3) 99.5–108.5 (102.7 6 5.1) 91.4–111.2 (102.2 6 6.4) 94.1–112.7 (102.1 6 8.8) 102.2–109.4 (106.2 6 2.7)                                             92.3,   95.2
MBW (male)                 5.7                5.0–5.3 (5.1 6 0.2)      4.0–4.7 (4.4 6 0.3)      4.2–5.1 (4.5 6 0.3)             —                                                           4.0
MBW (female)        5.2–6.4 (5.7 6 0.5)       3.7–5.0 (4.2 6 0.4)      3.4–5.9 (4.5 6 0.6)      3.8–4.8 (4.4 6 0.3)      4.2–5.4 (4.8 6 0.4)                                                3.2,   4.8
MBH (male)                 4.0                2.9–4.5 (3.8 6 0.8)      3.2–4.7 (3.6 6 0.5)      3.0–5.1 (3.6 6 0.8)             —                                                           3.2
MBH (female)        4.3–4.8 (4.5 6 0.2)       3.0–4.7 (3.9 6 0.5)      2.9–5.3 (3.7 6 0.8)      2.9–5.0 (3.7 6 0.8)      3.4–4.4 (3.8 6 0.4)                                                2.5,   4.3
TL (male)                35.1               45.0–50.5 (47.8 6 3.9)   39.3–47.1 (43.6 6 3.4)  38.7–51.5 (43.2 6 7.2)             —                                                          45.2
TL (female)       35.3–50.4 (44.5 6 8.1)    41.0–47.1 (43.9 6 3.1)   38.3–50.1 (44.2 6 4.1)  32.3–52.1 (42.5 6 7.4)    45.6–53.5 (49.5 6 3.1)                                              38.3,   35.2
TW (male)                  4.1                3.8–4.1 (4.0 6 0.2)      3.1–3.9 (3.4 6 0.2)      3.4–4.3 (3.7 6 0.3)             —                                                           3.7
TW (female)         4.1–4.6 (4.4 6 0.2)       3.3–3.8 (3.4 6 0.2)      2.8–4.6 (3.5 6 0.5)      2.8–4.2 (3.5 6 0.4)      3.6–4.5 (4.0 6 0.3)                                                2.4,   3.6
TH (male)                  3.5                3.2–3.4 (3.3 6 0.1)      2.8–3.5 (3.1 6 0.2)      2.6–3.6 (3.0 6 0.4)             —                                                           2.6
TH (female)         3.5–3.9 (3.8 6 0.2)       2.9–3.4 (3.1 6 0.2)      2.5–4.5 (3.1 6 0.5)      2.3–3.4 (2.8 6 0.4)      2.7–3.9 (3.1 6 0.4)                                                2.3,   3.2
HL (male)                  5.3                4.5–4.8 (4.6 6 0.2)      3.7–4.7 (4.1 6 0.4)      3.6–4.2 (3.8 6 0.2)             —                                                           3.9
HL (female)         4.8–5.2 (5.0 6 0.2)       3.5–4.4 (4.1 6 0.3)      3.5–4.5 (4.0 6 0.3)      3.5–5.3 (4.1 6 0.6)      3.8–5.5 (4.3 6 0.6)                                                3.1,   4.2
HW (male)                  5.1                 4.6–4.7 (4.6 6 0.1)     3.9–4.6 (4.2 6 0.2)      3.9–4.9 (4.2 6 0.4)             —                                                           4.0
HW (female)         4.6–5.8 (5.2 6 0.5)       3.8–4.6 (4.1 6 0.2)      3.6–5.2 (4.2 6 0.4)      3.7–5.4 (4.2 6 0.5)      4.0–5.3 (4.4 6 0.4)                                                3.5,   4.2
HH (male)                  3.5                 3.0–3.8 (3.4 6 0.4)     2.7–3.1 (2.9 6 0.2)      2.8–3.6 (3.0 6 0.3)             —                                                           2.9
HH (female)         3.4–4.2 (3.9 6 0.3)       2.9–3.4 (3.1 6 0.2)      2.5–3.9 (3.0 6 0.4)      2.5–3.9 (3.0 6 0.5)      2.9–3.9 (3.3 6 0.3)                                                2.5,   3.1
                                                                                                                                                                                                              HERPETOLOGICAL MONOGRAPHS




SnFa (male)              13.6               11.6–12.8 (12.0 6 0.6)   10.9–12.2 (11.6 6 0.4)  11.3–12.3 (11.8 6 0.3)             —                                                          11.9
SnFa (female)     11.7–14.1 (12.9 6 0.9)    10.1–11.9 (11.2 6 0.6)   10.9–12.5 (11.6 6 0.5)  10.6–13.0 (11.7 6 0.8)    11.3–13.2 (11.8 6 0.6)                                              11.5,   12.1
ED (male)                  0.9                1.0–1.1 (1.0 6 0.1)      1.0–1.1 (1.0 6 0.0)      0.8–0.9 (0.9 6 0.0)             —                                                           0.9
ED (female)         0.8–1.0 (0.9 6 0.1)       0.8–1.1 (1.0 6 0.1)      0.9–1.1 (1.0 6 0.1)      0.8–0.9 (0.9 6 0.1)      0.9–1.1 (1.0 6 0.1)                                                1.0,   1.0
END (male)                 1.9                 1.1–1.7 (1.5 6 0.3)     1.5–1.9 (1.7 6 0.1)      1.6–1.9 (1.7 6 0.1)             —                                                           1.6
END (female)        1.8–2.2 (1.9 6 0.1)       1.6–1.8 (1.7 6 0.1)      1.5–1.9 (1.7 6 0.1)      1.4–1.9 (1.7 6 0.2)      1.6–1.9 (1.8 6 0.1)                                                1.7,   1.9
SNL (male)                 2.7                1.8–2.5 (2.2 6 0.4)      2.1–2.5 (2.4 6 0.1)      2.2–2.5 (2.4 6 0.1)             —                                                           2.3
SNL (female)        2.6–2.9 (2.7 6 0.1)       2.2–2.6 (2.3 6 0.1)      2.2–2.6 (2.4 6 0.1)      2.1–2.7 (2.4 6 0.2)      2.3–2.6 (2.4 6 0.1)                                                2.4,   2.6
IND (male)                 1.4                1.2–1.4 (1.3 6 0.1)      1.2–1.4 (1.3 6 0.1)      1.1–1.5 (1.3 6 0.1)             —                                                           1.1
IND (female)        1.1–1.4 (1.3 6 0.1)       1.2–1.4 (1.3 6 0.1)      1.1–1.5 (1.3 6 0.1)      1.0–1.4 (1.3 6 0.1)      1.1–1.5 (1.3 6 0.1)                                                1.1,   1.3
FLL (male)                 2.3                 1.4–1.7 (1.6 6 0.1)     1.1–1.5 (1.3 6 0.1)      1.1–1.6 (1.3 6 0.2)             —                                                           1.4
FLL (female)        1.1–2.6 (1.7 6 0.5)       1.2–1.7 (1.4 6 0.1)      1.1–1.8 (1.3 6 0.2)      1.1–1.9 (1.5 6 0.3)      1.4–2.1 (1.7 6 0.2)                                                1.1,   1.5
HLL (male)                 2.9                2.3–3.0 (2.6 6 0.4)      2.1–2.4 (2.3 6 0.1)      2.4–2.8 (2.6 6 0.2)             —                                                           2.6
HLL (female)        2.5–3.1 (2.8 6 0.2)       2.3–2.9 (2.6 6 0.2)      2.0–2.7 (2.4 6 0.2)      1.9–3.1 (2.7 6 0.4)      2.5–3.6 (3.0 6 0.3)                                                2.1,   2.7
                                                                                                                                                                                                              [No. 25
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