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A new frog species from rapidly dwindling cloud forest streams of Sri Lanka—Lankanectes pera (Anura, Nyctibatrachidae)

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Abstract and Figures

The monotypic genus Lankanectes, considered an evolutionary long branch with India’s Nyctibatrachus as its sister lineage, is represented by L. corrugatus, a species widely distributed within the wet zone of Sri Lanka up to 1500 m asl, where it in- habits a variety of lotic and lentic habitats. Here, following an integrative taxonomic approach using DNA-based phyloge- nies, morphology, morphometry, and ecological niche models, we describe a new species—Lankanectes pera sp. nov. The new species is distinguished from its sister species mainly by its tuberculated throat and absence of dark patches on venter, throat, manus and pes. The uncorrected genetic distances between the two Lankanectes species for a fragment of the non- coding mitochondrial 16S rRNA gene is 3.5–3.7%. The new species has a very restricted climatic distribution with a total predicted area of only 360 km2 (vs. 14,120 km2 for L. corrugatus). Unlike L. corrugatus, which prefers muddy substrates and marshy areas, the new species is observed inhabiting only pristine streams flowing through canopy covered montane forests in the highest reaches of the Knuckles Mountain range. The specialized new species will need immediate conservation atten- tion due to its restricted distribution (montane isolate), specialized habit of inhabiting clear mountain streams, and small pop- ulation size.
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Accepted by M. Vences: 9 Jul. 2018; published: 23 Aug. 2018
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4461 (4): 519
A new frog species from rapidly dwindling cloud forest streams of
Sri Lanka—Lankanectes pera (Anura, Nyctibatrachidae)
Department of Organismal Biology & Anatomy, University of Chicago, Chicago IL, USA
Postgraduate Institute of Science, University of Peradeniya, Sri Lanka
Department of Molecular Biology & Biotechnology, Faculty of Science, University of Peradeniya, Sri Lanka
No. 308/7 A, Warathanna, Halloluwa, Sri Lanka
Department of Zoology, Faculty of Applied Sciences, University of Sri Jayewardenepura, Sri Lanka
Postgraduate Institute of Archaeology, 407 Bullers Road, Colombo 07, Sri Lanka
Melbourne School of Population and Global Health, University of Melbourne, Melbourne, Australia
National Institute of Fundamental Studies, Kandy, Sri Lanka
Guangxi Key Laboratory for Forest Ecology and Conservation, College of Forestry, Guangxi University, Nanning, China
Corresponding author. E-mail:
The monotypic genus Lankanectes, considered an evolutionary long branch with India’s Nyctibatrachus as its sister lineage,
is represented by L. corrugatus, a species widely distributed within the wet zone of Sri Lanka up to 1500 m asl, where it in-
habits a variety of lotic and lentic habitats. Here, following an integrative taxonomic approach using DNA-based phyloge-
nies, morphology, morphometry, and ecological niche models, we describe a new species—Lankanectes pera sp. nov. The
new species is distinguished from its sister species mainly by its tuberculated throat and absence of dark patches on venter,
throat, manus and pes. The uncorrected genetic distances between the two Lankanectes species for a fragment of the non-
coding mitochondrial 16S rRNA gene is 3.5–3.7%. The new species has a very restricted climatic distribution with a total
predicted area of only 360 km
(vs. 14,120 km
for L. corrugatus). Unlike L. corrugatus, which prefers muddy substrates and
marshy areas, the new species is observed inhabiting only pristine streams flowing through canopy covered montane forests
in the highest reaches of the Knuckles Mountain range. The specialized new species will need immediate conservation atten-
tion due to its restricted distribution (montane isolate), specialized habit of inhabiting clear mountain streams, and small pop-
ulation size.
Key words: Ecological niche models, General lineage concept, Knuckles Mountains, Montane-isolate
Compared to mainland India, where endemic lineages at the level of higher taxa, such as Nasikabatrachus,
Micrixalus and Nyctixalus abound (Biju & Bossuyt 2003; Roelants et al. 2004; Van Bocxlaer et al. 2011), only a
few endemic anuran lineages dating to before the Cretaceous-Paleogene extinction event seem to have survived in
Sri Lanka. The only Sri Lankan amphibian lineage that made it through this mass extinction event is Lankanectes,
a monotypic endemic genus represented by L. corrugatus (Peters 1863). Its closest living relatives are considered
to be the representatives of the family Nyctibatrachidae, which diverged prior to the Cretaceous-Paleogene
extinction event, about 72–73.5 mya (Van Bocxlaer et al. 2011; Pyron & Wiens 2011, 2013).
Lankanectes is now accepted as being substantially different morphologically from Euphlyctis, Limnonectus,
Taylorana, Occidozyga, Phrynoglossus and Nyctibatrachus, where it was previously placed (Peters 1863;
Boulenger 1920; Deckert 1938; Dubois 1981; Dubois & Ohler 2001). Molecular studies have confirmed not only
the distinctiveness of Lankanectes, but also its ancientness among South Asian anuran lineages (van der Meijden et
Zootaxa 4461 (4) © 2018 Magnolia Press
al. 2004; Delorme et al. 2004; Roleants et al. 2004). This widely distributed species, with its conspicuously loud
call, is found up to an elevation of about 1500 m asl, in submontane habitats in the southern, western and central
parts of Sri Lanka, (Frost 2017; Manamendra-Arachchi & Pethiyagoda 2006).
During our field work in the streams of the cloud forests of the Knuckles mountain region, we discovered a
population of Lankanectes that appeared to be morphologically distinct from Lankanectes corrugatus. Given that
many of the species of Nyctibatrachus are montane endemics (Van Bocxlaer et al. 2011), the possibility of the
existence of an undescribed species of Sri Lankan Lankanectes was high. Using an integrative taxonomic
approach, in the sense of the General Lineage Concept (De Queiroz 1998), from a lineage that was thought to be on
an evolutionary long branch with a single species, we show this population to be clearly distinct from L. corrugatus
and describe it as a new species—Lankanectes pera sp. nov.
Materials and methods
Specimen collection, DNA barcoding, and Phylogenetic analyses. Field surveys for Lankanectes were
conducted in Sri Lanka in the years 2012– 2016 (Table 1). Guided by calling males, sampling was carried out
mostly at night. Animals (adults and tadpoles) were euthanized in tricaine methanesulphonate (MS-222). Thigh
muscle tissue (~20 mg) from adults and a small part of the tail-fin of two tadpoles were stored in absolute ethanol
at -20 °C for subsequent molecular studies in the Department of Molecular Biology & Biotechnology (DZ),
University of Peradeniya. Adult specimens used in morphological studies were fixed in 4% formalin and later
preserved in 70% ethanol. The type specimens of the new species are deposited in the Department of Molecular
Biology & Biotechnology, University of Peradeniya.
Collected samples (Table 1) were DNA barcoded for the 16S ribosomal RNA (16S rRNA) mitochondrial gene
fragment to ascertain species identity. DNA was extracted from ethanol-preserved tissue using a standard protocol
(Sambrook et al. 1989). Portions of the mitochondrial 16S rRNA gene were amplified by PCR and sequenced
directly using dye-termination cycle sequencing. Primer sets, 16Sar and 16Sbr (Palumbi 1996) were used, which
amplified a ca. 600 bp fragment of the 16S rRNA gene. PCR conditions for amplification were as follows:
denaturation at 95 °C for 40 s, annealing at 45 °C for 40 s and extension at 72 °C for 50 s, 35 cycles, with a final
extension of 72 °C for 5 min. Newly generated sequences were visualized and checked using 4peaks (v. 1.7.1).
Twenty-eight taxa (Table 1) representing closely related congeners of Lankanectes (according to recently
published phylogenies—Van Bocxlaer et al. 2011) were also included in the dataset. Additionally, Nasikabatrachus
sahyadrensis and Micrixalus phyllophilus representing closely related basal nyctibatrachids (Pyron & Wiens 2011)
were used as an outgroup (Table 1). The compiled 16S rRNA dataset was aligned using ClustalW as implemented
in MEGA v. 7.0 (Kumar et al. 2016). Uncorrected pairwise distances were calculated using PAUP* 4.0b10
(Swofford 2002; Table 2). Regions that were highly variable were manually removed from the dataset; the final
dataset consisted of 485 bps. The best-fitted model was chosen using jModeltest v. 2.1.4 (Posada & Crandall 1998).
Maximum likelihood (ML) analysis was performed to infer relationships among the lineages and clades using the
software GARLI (Zwickl 2006) on the CIPRES Science Gateway (Miller et al. 2010), using the best model
(TIM2+I+G) parameters. Clade support was assessed using posterior probability (PP) and Maximum Parsimony
(MP) bootstrapping values. Bayesian inference as implemented in MrBayes (v.3.1.2; Huelsenbeck & Ronquist
2001) was used to assess posterior probability (PP) values for each node with the parameters of the best-fitted
model estimated as obtained from the jModelTest. Four Metropolis-Coupled Markov Chain Monte Carlo
(MCMCMC) chains were run for ten million generations. Burn-in of 5 million generations was estimated using
Tracer v. 1.6 (Bouckaert et al. 2014). Bootstrapping was done in a MP framework using PAUP, where a full
heuristic search was done, with 1000 replicates.
Morphology and Morphometrics. The suite of characters and character states used by Manamendra-Arachchi
& Pethiyagoda (2006) was considered. Measurements were made to the nearest 0.1 mm using dial Vernier calipers.
The following morphometric variables were measured: distance between back of eyes (DBE); distance between front
of eyes (DFE); eye diameter (ED); eye-to-nostril distance (EN); eye-to-snout length (ES); femur length (FEL); length
of finger 1 (FLI); length of finger 2 (FLII); length of finger 3 (FLIII); length of finger 4 (FLIV); pes length (FOL);
head length (HL); head width (HW); length of inner metatarsal tubercle (IML); internarial distance (IN); interorbital
distance (IO); lower-arm length (LAL); posterior mandible-to eye distance (MBE); least distance from mandible to
Zootaxa 4461 (4) © 2018 Magnolia Press
anterior eye (MFE); least distance from mandible to nostril (MN); nostril-to-snout length (NS); palm length (PAL);
snout–vent length (SVL); tibia length (TBL); length of toe 1 (TLI); length of toe 2 (TLII); length of toe 3 (TLIII);
length of toe 4 (TLIV); and length of toe 5 (TLV); length of upper arm (UAW); and width of upper eyelid (UEW).
Illustration of the webbing pattern follows Manamendra-Arachchi & Pethiyagoda (2006). All these measurements
were used in the Principal Components Analysis (PCA). Abbreviations used in the study: DZ, Department of
Molecular Biology and Biochemistry, Peradeniya, Sri Lanka; FR, Forest reserve; GS, Gayani Senevirathne; HUN,
Hunnasgiriya (field collection numbers); KM-A, Kelum Manamendra-Arachchi; KNU, Knuckles (field collection
numbers); MM, Madhava Meegaskumbura; NW, Nayana Wijayathilaka; WHT, Wildlife Heritage Trust of Sri Lanka,
Colombo, Sri Lanka; ZMB, Zoological Museum of Berlin.
Principal Components Analysis of the character correlation matrix was used to reduce dimensionality of the
continuous morphological variables and to identify those variables that best discriminate among morphologically
similar forms. Various axis rotations were tested, and one was selected for optimal interpretability of variation
among the characters. SYSTAT (Version 11.00.01) was used for statistical analysis.
Haplotype Network. Population genetic structure was determined by constructing haplotype networks using
available sequences of the 16S rRNA fragment as implemented in PopArt ( Given the
close relationships between populations and the two sister taxa, ambiguously aligned regions were absent among
Lankanectes sequences, and the full dataset (560 bp) was used for this analysis.
Adult osteology. Osteological preparation and descriptions for Lankanectes corrugatus, which would serve as
a description for the genus (type species for the genus), is carried out here based on cleared and stained
postmetamorphic adults (N = 2; DZ 1397; DZ 1399). Neutral-buffered formalin preserved specimens were stained
following the procedure by Taylor and Van Dyke (1985). Initial dehydration was done in 100% ethanol, followed
by submersion in alcian blue for cartilage staining. Excessive musculature was digested using an infusion of borax
and trypsin, and the specimens were subsequently stained in alizarin red for bone visualization. Preparations were
photographed and scored for bones and cartilage within 2–3 days following the clearing and staining procedure.
Osteological terminologies follow Duellman and Trueb (1986) and Senevirathne et al. (2016).
Niche modeling. We collected distribution records for Lankanectes both from the published literature (e.g.
Manamendra-Arachchi & Pethiyagoda 2006) and our own field records. The program MaxEnt, version 3.3.3k (Philips
et al. 2004) was used to predict the geographic distribution of each putative species. For this study, we used 24
presence locations for L. corrugatus and five for L. pera sp. nov. (Appendix 1). We downloaded 19 environmental
variables and an altitude layer with a 30 arc-second (ca. 1 km
) spatial resolution, from WorldClim dataset
( All layers were clipped to our study region bounded by 5.908° to 9.842 °N and 79.516° to
81.891 °E (which includes all of Sri Lanka). Highly correlated variables (r ≥ 0.8 Pearson correlation coefficient) were
eliminated from the analysis. Altogether, seven and four variables were selected to generate the predictive models of
L. corrugatus and L. pera sp. nov., respectively (Table 3). We used: the automatic mode with jackknife validation;
random seed option for all sample points to train the model; and 25% of the records to test it. We ran 10 replicates
using bootstrap function, and the average model was selected. The logistic method was used to obtain the values of
habitat suitability, in which the probability values ranged from 0 to 1. Resulting values were then transferred to binary
presence and absence values using Lowest Presence Threshold (LPT). Model performance was evaluated using area
under the Receiving Operator Curve (AUC), in which the value ranges from 0 to 1 (Fielding & Bell 1997).
DNA barcoding and phylogenetic analyses. The final dataset contained 16S rRNA mitochondrial gene sequences
from 41 putative species. Eleven of these represent Sri Lankan Lankanectes (9 tissues from adults and 2 tadpoles),
while 28 represent Nyctibatrachus, the sister group of Lankanectes. The Maximum Likelihood (ML) tree (Fig. 1) is
rooted using Nasikabatrachus sahydrensis and Micrixalus phyllophilus. TIM2+I+G was selected as the best-fitted
model for our dataset. The parameters of the nucleotide substitution model for the most likely tree were as follows:
rate matrix: shape parameter for gamma distributed rate variation among sites (alpha)= 0.5260; Tree length =
1.990892; R(A-C) = 9.0960; R(A-G) = 45.7402; R(A-T) = 15.2825; R(C-G) = 0.5040; R(C-T) = 142.4271; R(G-T)
= 1.000000; Nucleotide frequencies: (A) = 0.3279; (C) = 0.2297; (G) = 0.2073; (T) = 0.2350; likelihood= -
2631.6028. The ML tree had the same topology as the Distance, Bayesian, and Maximum Parsimony trees.
Zootaxa 4461 (4) © 2018 Magnolia Press
TABLE 1. GenBank Accession numbers and Voucher numbers of the taxa used in the study.
Species name Accession number Voucher number Location
Lankanectes corrugatus MH697874 DZ1396 Peradeniya
Lankanectes corrugatus MH697875 DZ1397 Peradeniya
Lankanectes corrugatus MH697876 DZ1409 Panwila
Lankanectes corrugatus (tadpole) MH697877 GS2_32 Galle
Lankanectes pera sp. nov. (tadpole) MH697873 DZ1320 Knuckles
Lankanectes pera sp. nov. MH697872 DZ1307 Knuckles
Lankanectes pera sp. nov. MH697871 DZ1290 Knuckles
Lankanectes corrugatus DQ346971 X Sri Lanka
Lankanectes corrugatus AF215393 X Sri Lanka
Lankanectes corrugatus AY880445 X Sri Lanka
Lankanectes corrugatus DQ019603 X Galle
Nyctibatrachus vrijeuni JN644783 X India
Nyctibatrachus sp. B JN644784 X India
Nyctibatrachus aliciae JN644785 X India
Nyctibatrachus poocha JN644786 X India
Nyctibatrachus minor JN644787 X India
Nyctibatrachus vasanthi JN644788 X India
Nyctibatrachus indraneili JN644789 X India
Nyctibatrachus karnatakaensis JN644790 X India
Nyctibatrachus deccanensis JN644791 X India
Nyctibatrachus petraeus JN644792 X India
Nyctibatrachus beddomii JN644793 X India
Nyctibatrachus sp. A JN644794 X India
Nyctibatrachus acanthodermis JN644795 X India
Nyctibatrachus deveni JN644796 X India
Nyctibatrachus minimus JN644797 X India
Nyctibatrachus sylvaticus JN644798 X India
Nyctibatrachus sanctipalustris JN644799 X India
Nyctibatrachus gavi JN644800 X India
Nyctibatrachus major JN644801 X India
Nyctibatrachus shiradi JN644774 X India
Nyctibatrachus dattatreyaensis JN644775 X India
Nyctibatrachus anamallaiensis JN644776 X India
Nyctibatrachus jog JN644777 X India
Nyctibatrachus humayuni JN644778 X India
Nyctibatrachus danieli JN644779 X India
Nyctibatrachus kempholeyensis JN644780 X India
Nyctibatrachus pillaii JN644781 X India
Nyctibatrachus grandis JN644782 X India
Nasikabatrachus sahyadrensis AY364381 X India
Micrixalus phyllophilus KJ711349 X India
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TABLE 2. Uncorrected percentage pairwise divergences of 16S rRNA between the Lankanectes corrugatus and L. pera
sp. nov. species used in the study (refer Table 1 for the corresponding locations of the voucher numbers).
Maximum parsimony bootstrap values and PP values at the node of the Lankanectes lineage indicates that it is
highly supported (Fig. 1). It results in two highly supported subclades: a widely distributed Lankanectes corrugatus
and the new species from the Knuckles region. The relationships of the taxa of the Lankanectes clade were
identical to the relationships from the Maximum Parsimony, Distance and Bayesian analyses. The uncorrected
pairwise distance between L. corrugatus and L. pera sp. nov. is 3.5–3.7%. (Table 2).
Haplotype Network. The entire fragment (560 bp) was used to construct the haplotype networks for the 6
populations (Knuckles, Galle, Agra, Peradeniya, Panwila, Morningside), representing the two species of
Lankanectes (Fig. 1). All haplotypes of Lankanectes corrugatus are shared between populations with 1–4
mutational changes. Lankanectes pera sp. nov. differs from L. corrugatus by 16 mutational steps with no sharing
of haplotypes.
Adult osteology. Cranium (Fig. 5): Cranium of Lankanectes corrugatus has a height of 69% of its width.
Frontoparietals are paired, quadrangular, invest the roof of the neurocranium dorsally, and posterolateral margins
reach prootics fusing synostically. Parasphenoid, is azygous, ventral, and T-shaped; furcated cultriform process
extends almost up to the palatines, but do not fuse with them. Perpendicular to the anterior cultriform process, alae
extend laterally, overlaying prootics and exoccipitals, and end bluntly; medially, a depression is present where alae
and cultriform process congregate. Vomers have four processes (dentary, anterior, prechoanal, and postchoanal) and
are medially separated from one another; dentary processes of the vomers are edentate and reach the center region of
the palatines; anterior process with a blunt terminal end extends towards the premaxillary-maxillary junction;
prechonal and postchonal processes extend laterally from the anterior process, and the bifurcation between the two
occurs at the base of the dentary process. Palatines are paired and invest the posterior margins of the planum
antorbitale; lateral blunt ends reach up to medial face of the pars dentalis of maxillae, but do not articulate with
maxillae. Anterior ramus of the pterygoids reaches the posterior margin of the planum antorbitale ventrally and is
connected to the dentary process of the maxillae; posterior ramus, equal to the length of the anterior ramus, is directed
towards the quadrate and has a cartilaginous epiphysis; medial ramus, shortest among the three rami, reaches the
anterior margins of the otic capsule ventrally. Squamosals have three distinct rami; ventral ramus reaches up to the
epiphysis of ventral ramus of the pterygoid; zygomatic ramus has a pointed terminus; otic ramus is blunt-ended.
Maxillary arcade: Premaxillae are separated from one another, but overlap with maxillae; alary process of the
premaxilla is about half of the length of pars dentalis; pars palatina is bifurcated and extend from the posteroventral
margin of dentate pars dentalis of premaxilla. Maxillae are composed of pars dentalis, pars facialis and pars
palatina; pars dentalis bear 30/38 (upper/lower) blunt teeth; pars palatina of maxillae do not articulate with the pars
palatina of premaxillae; posteriomost ends of maxillae articulate with the qudratojugals, which completes the
maxillary arcade posteriorly; pars facialis is well ossified and forms the lateral walls of the nasal capsules.
Qudratojugals are paired and complete the arcade laterally; they end with wide bases, which are connected to the
ventral ramus of the squamosal.
DQ346971 -
DZ1396 0 -
DZ1397 0 0 -
DZ1409 0 0 0
GS2 32 0.4 0.4 0.4 0.4
AF215393 0.2 0.2 0.2 0.2 0.2
AY880445 0.4 0.4 0.4 0.4 0.4 0.2
DQ019603 0.0 0 0 0 0.4 0.2 0.4
DZ1320 3.7 3.7 3.7 3.7 3.7 3.5 3.7 3.7
DZ1307 3.7 3.7 3.7 3.7 3.7 3.5 3.7 3.7 0
DZ1290 3.7 3.7 3.7 3.7 3.7 3.5 3.7 3.7 0 0 0
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FIGURE 1. (A) 16S rRNA Maximum-likelihood phylogram for 41 taxa with node support (posterior probabilities above nodes,
with values >95 indicated by a *; MP bootstrap values below nodes, with values >50) showing two clearly-defined clades
including L. corrugatus and L. pera (B) The uncorrected pairwise genetic distances range for 16S rRNA 3.5–3.7%; Haplotype
network analysis using the full dataset with 485 bp, shows all haplotypes shared across L. corrugatus; L. pera sp. nov. is
different by more than 16 mutational steps from the L. corrugatus populations. (C) The predicted distributions for L.
corrugatus is 360 km
and L. pera sp. nov. is 14120 km
on the northern hills of Sri Lanka. (D) Component loadings of the first
two principal component axes (PC1 vs PC2) shows clear separation of the males, but with slight overlap of females; L. pera sp.
nov. indicated in red, L. corrugatus in blue (filled circles denote females and open circles denote males).
Mandible: Cranial articulation of the manibular arch occurs via mineralized pars aricularis located at the
posterolateral ends. Lateral margins of the mentomeckelians are attached to dentaries, elongated Meckel’s
cartilages and angulosplenials. Dentaries invest lateral and anterodorsal margins of the Meckel’s cartilage.
Hyoid skeleton: Thin, mineralized hyoid plate houses U-shaped hyoglossal sinus, anterolateral processes,
posterolateral (slightly longer than anterolateral processes), ossified posteromedial processes (longest) and a pair of
curved hyales that allow attachment of hyoid skeleton to the cranium.
Postcranial skeleton: Eight presacral vertebrae, sacrum and urostyle form the axial skeleton. Transverse
processes of vertebrae III and IV are posterolaterally directed and are same as the length as of the sacral
diapophyses and end in cartilaginous epiphyses distally (Fig. 5). Rest of the transverse processes (II, V, VI, VII,
VIII) are laterally oriented and are shorter in length. Hypochord is fussed with the coccyx and form the urostyle
extending posteriorly, from the sacrum. Pectoral girdle has a firmisternal construct. Paired suprascapulae are flat,
thin blade-like, with its proximal ends being cartilaginous. Anterior margins of suprascapulae are ossified to give
Zootaxa 4461 (4) © 2018 Magnolia Press
rise to the paired cleithra. Ossified scapulae are connected together at the junction of glenoid fossa, via the
articulation with clavicles and coracoids. Paired clavicles occupy the whole procoracoid cartilages except for a thin
posterior margin. Epicoracoids connect each half of the pectoral girdle together. Epicoracoidal bridge is ossified.
Sternum and omosternum are well developed, with expanded, ossified proximal ends. Proximal end of the
omosternum is fused with the epicoracoidal bridge and the most distal cartilaginous end assumes a half-arched
shape. Inverted v-shaped, proximal end of the sternum is attached to the cartilaginous epicoracoid cartilage and the
wider distal part has irregular shaped margins, both anteriorly and posteriorly. Manus is composed of radiale,
ulnare, carpals (2, 3-4-5), element Y, prepollex and prepollical element I. Metacarpals decrease in length as
follows, 4> 3> 5> 2. Phalangeal digit formula is 3-3-4-4. Pes is composed of fused tarsals 2-3 (ossified), prehallical
elements, metatarsals and phalange digits. Prehallux is completely ossified and base of the prehallical element 1 is
ossified. Length of the metatarsals increase as; IV> V> III> II> I. Phalangeal formula is 3-3-4-5-4. Long ilia with
ilial crests articulate with epiphyses of sacral diapophyses. Ischia and pubis are fused with each other.
TABLE 3. Bioclimatic variables used to predict the distribution models of the two species.
Note. Highlighted variables selected through multi-collinearity test, values indicate the percentage contribution of the
variable to build the model.
Niche modeling. The Maxent models for the two species provided satisfactory results, with higher AUC
values; Lankanectes corrugatus = 0.959 and L. pera = 0.999. LPT values used for the two models of L. corrugatus
and L. pera were 0.145 (20.1%) and 0.23 (28.7%), respectively. Precipitation of Driest Month (Bio 14) contributed
most to the distribution model of L. corrugatus, whereas Altitude and Precipitation of Coldest Quarter (Bio19)
contributed most to that of L. pera sp. nov. (Table 3). Even though L. pera is restricted to the Knuckles Mountain
range, the predictive models show that the new species has suitable climatic conditions also in the northern high
region of the central mountains (L. pera sp. nov. is restricted to the Knuckles mountain range). Lankanectes
corrugatus is distributed throughout the wet-zone of the south-western quadrant of the island, extending also up the
lower (< 1500m asl) slopes of the Central mountains (Fig. 1).
Code Bioclimatic variables L. corrugatus L. pera sp. nov.
BIO1 Annual Mean Temperature x x
BIO2 Mean Diurnal Range 1.7 x
BIO3 Isothermality 1.8 x
BIO4 Temperature Seasonality x x
BIO5 Max Temperature of Warmest Month x x
BIO6 Min Temperature of Coldest Month x x
BIO7 Temperature Annual Range x x
BIO8 Mean Temperature of Wettest Quarter x x
BIO9 Mean Temperature of Driest Quarter x x
BIO10 Mean Temperature of Warmest Quarter x x
BIO11 Mean Temperature of Coldest Quarter x x
BIO12 Annual Precipitation x x
BIO13 Precipitation of Wettest Month x x
BIO14 Precipitation of Driest Month 76.8 x
BIO15 Precipitation Seasonality 2.1 x
BIO16 Precipitation of Wettest Quarter x x
BIO17 Precipitation of Driest Quarter x 0.9
BIO18 Precipitation of Warmest Quarter 7.7 0.1
BIO19 Precipitation of Coldest Quarter 0.5 42.3
Alt Altitude 9.3 56.7
Total number of variables used 7 4
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Lankanectes pera, sp. nov.
(Figs. 2,3,4, Appendix 2)
Type Material. Holotype: mature male, 66.0 mm SVL, DZ1858 (KNU01), Knuckles Peak, alt. 1580 m, 7.4646 °N
80.7409 °E. Collected by MM, KM-A 10
August, 2012.
Paratypes: mature female, 51.0 mm SVL, DZ1859 (KNU02), Knuckles Peak, alt. 1580 m (7˚4646’ N
80˚7409’E) collected by MM, KM-A 10
August, 2012; mature female, 42.4 mm SVL DZ1860 (HUN01),
Dothalugala (Hunnasgiriya Peak), alt. 1420 m (7˚3206’N 80˚8568’E), MM, KM-A, NW 12
December, 2012;
mature male, 68.7 mm SVL, DZ1307, Riverston Knuckles, alt 1330 m (7˚5233’N, 80˚7333’E), collected by MM,
NW, GS 15
August, 2013; mature female, 55.8 mm SVL, DZ1290, Riverston Knuckles, alt. 1260 m (7˚5180’N,
80˚7375’E), collected by MM, NW, GS 15
August, 2013; mature female, 47.3 mm SVL, DZ1302, Riverston
Knuckles, 1260 m (7˚5180’N, 80˚7375’E), collected by MM, NW, GS 15
August, 2013.
Diagnosis. Lankanectes pera sp. nov. can be distinguished from L. corrugatus by the following characters:
ventrally greyish (vs white with dark brown patches in L. corrugatus); white tubercles on throat (vs smooth throat
in L. corrugatus); edge of the upper lip uniform grey (vs white border with dark brown patches in L. corrugatus);
inner edge of toes grey (vs inner edge of I, II, III and IV toes white in L. corrugatus); inner edge of foot grey (vs
white in L. corrugatus); flank grey (vs. flank with dark brown and white patches in L. corrugatus).
FIGURE 2. Lankanectes pera sp. nov. in life, SVL 22.40 mm: (A) Dorsolateral view; (B) Dorsal view; (C) Habitat—clear
water stream habitat under montane forest canopy cover.
Zootaxa 4461 (4) © 2018 Magnolia Press
FIGURE 3. Illustrations of (A) Lankanectes corrugatus, ZMB4897, mature male, SVL 43.70 mm and (B) L. pera sp. nov.,
DZ1858, mature male, SVL 66.0 mm, anal region (top); manus (middle) and maxilla (lower).
Description (based on the holotype, DZ 1858; Figs. 2,3,4). Body stout. Head flat dorsally. Snout rounded
when viewed dorsally and laterally. Canthal edges indistinct. Loreal region convex. Edges of upper lip with distinct
tubercles; interorbital and internasal spaces convex. Nostrils oval; close to each other (19.3% of the width of the
skull); placed dorsally on snout; edges fleshy. Tympanum absent; pineal ocellus absent. Vomerine teeth present; the
vomerine teeth are tusk-like (more prominent in males), with an angle of 60º relative to body axis; less close to
choanae than to each other. Tongue large; emarginated; not bearing a lingual papilla; two tubercles on posterior
base of tongue. Two fang-like processes on the mandible. Internal vocal slits present, close to gape. Supratympanic
fold absent. Parotid glands absent. Head wide. Cephalic ridges absent. Cephalic knob on head. Skin on head not co-
ossified. Dorsal surface of head and body covered in numerous prominent dermal folds (corrugations) and
glandular, white-tipped warts. Corrugations present also on ventral surface of head and throat. Manus robust.
Zootaxa 4461 (4) © 2018 Magnolia Press
FIGURE 4. Lankanectes pera sp. nov., SVL 68.69 mm, DZ1307 (A–D), in preservation. (A) ventral view; (B) right ventral
foot; (C) ventral view of the head; (D) lateral view. Lankanectes corrugatus, SVL 55.28 mm, DZ1396 (E–H), in preservation.
(A) ventral view; (B) right ventral foot; (C) ventral view of the head; (D) lateral view.
Zootaxa 4461 (4) © 2018 Magnolia Press
Forearms short, strong; fingers thin. Tips of fingers rounded, enlarged; discs absent; finger-tips not wide compared
to finger width; no dermal fringe on inner or outer sides of fingers; no webbing on fingers; subarticular tubercles on
fingers prominent, oval, single; prepollex distinct. Two palmar tubercles, oval, distinct, convex. Supernumerary
tubercles on palm very small. Nuptial pad absent. Pes robust. Thigh and shank stout. Toes thin. Tips of toes
rounded, enlarged, discs absent; tips of toes not wide compared to toe width. Toes fully webbed (see Figs 2B &
3B). Dermal folds present on inner edge of toe I and outer edge of toe V. Subarticular tubercles on toes prominent,
rounded or oval, single. Inner metatarsal tubercle long, prominent, oval. Tarsal fold present. Outer metatarsal
tubercle absent. Supernumerary tubercles on foot absent. Tarsal tubercle absent. Snout between eyes and side of
head with folds and fine tubercles. Anterior and posterior part of back, and upper and lower flank with dermal
folds. Dorsolateral fold absent on body. Corrugations and glandular warts present on dorsal surface of legs, but are
less prominent; ventral surface of legs smooth. Lateral-line system present. Dorsal parts of forelimb and thigh with
corrugations. Dorsal part of shank and tarsus with corrugations and tiny tubercles. Chest, belly and ventral part of
thigh smooth. A cluster of macroglands (femoral glands) on inner surface of thigh. Possess vocal sacs and nuptial
Sexual dimorphism. Head of females narrower than males (see Appendix 4 for measurements); cephalic knob
and vocal slits absent.
FIGURE 5. Cleared and stained adult specimen of Lankanectes corrugatus (DZ1397, SVL 45.02 mm). (A) Cranium,
dorsal view. (B) Vertebral column, with the pelvic girdle attached to the sacral diapophysis via the cartilaginous epiphysis of
ilia, dorsal view. (C) Close-up of vertebrae (D) Hyobranchial skeleton, ventral view. (E) Pectoral girdle and forelimbs, ventral
view. Abbreviations: CR, coracoid; CT, cleithrum; CV, clavicle; EX, exoccipital; FP, frontoparietal; HP, hyoid plate; HY, hyale;
IL, ilium; IS, ischium; MX, maxillae; OS, omosternum; PM, premaxilla; PP, posteromedial process; PR, prezygapophysis; PZ,
postzygapophysis; PT, pterygoid; SC, suprascapula; SQ, squamosal; SL, scapula; ST, sternum; PV, vomer.
Zootaxa 4461 (4) © 2018 Magnolia Press
Coloration (in alcohol; Fig. 4)—Dorsally dark brown with unequal dark patches, edges of corrugations lighter
in color, some pale spots on dorsum. A pale-yellow bar with dark edges on inter-orbital area. Flank, inguinal zone,
loreal region and sides of back of head light brown, edges of corrugations pale. Throat, margin of throat and vocal
sacs pale brown with lighter spots. Chest, belly, ventral sides of thighs and webbing light brown.
Color in life: Dorsally chocolate brown with unequal dark-brown patches. Ridges of the numerous prominent
corrugations lighter in color, with interspersed light-brown spots. A light-brown bar edged with dark brown/black
colors in the interorbital area. Flank, inguinal zone, loreal region and sides of back of head light brown. Throat,
margin of throat and vocal sacs white with pale brown patches. Chest, belly ventral sides white. Ventral sides of
thighs light brown, with white patches. Underside of webbing light brown. Disks and tubercles of pes and manus
Measurements of Holotype (DZ1858 in mm). DBE, 17.2; DFE, 9.6; ED, 7.5; EN, 4.3; ES, 9.1; FEL, 29.7; FL
I, 5.8; FL II, 6.0; FL III, 7.9; FL IV, 6.9; FOL, 42.0; HL, 27.3; HW, 25.7; IML, 3.3; IN, 3.6; IO, 5.6; LAL, 13.2;
MBE, 12.6; MFE, 20.4; MN, 24.1; NS, 6.9; PAL, 15.8; SVL, 66.0; TBL, 28.4; TL I, 7.3; TL II, 9.0; TL III, 12.7;
TL IV, 15.9; TL V, 12; UAW, 9.8; UEW, 3.0.
Etymology. The specific epithet pera is applied as a noun in apposition. It is a reference to the University of
Peradeniya, Sri Lanka, affectionately referred to as “Pera” by its alumni.
Morphometrics. Unrotated principal components analysis separates the males of the two species on PC1, but
slight overlap is seen for females (Fig. 1D). Of the total variance, 92 % is explained by PC1, which is a size axis
(although the highest factor loading was for SVL, all other variables too, had high positive values); Lankanectes
pera sp. nov. is larger in size than L. corrugatus (see Appendix 4 for all material studied and measurements). Only
2.6% of the total variance is explained by PC2, which reflects mostly in FLI (length of first finger) and NS (nostril
to snout distance). This axis, however, is uninformative as the two species show nearly complete overlap (Fig. 1D,
Appendix 4).
Distribution: Lankanectes pera sp. nov. is restricted to streams flowing through the montane forests on
highest peaks of the Knuckles Mountain range—1100 m asl, in Dothalugala and Bamabarella and Riverston
Ecological notes and natural history. This species has so far only been observed inhabiting pristine streams
flowing through closed-canopy montane forests. These streams are characterized by clear, shallow and slow-
flowing water, and sand and rock-strewn substrates. Males are found under rocks or rock-crevices in flowing water.
Occasionally males call haltingly during daytime, but several males frequently vocalize in chorus at night,
especially after light rain. Tadpoles of these frogs are large (total length of Gosner stage 35 tadpoles range between
42.00–45.14 mm, N = 4), and occur in deeper regions (0.5 m) where decaying vegetation gathers.
All characters that typify the genus Lankanectes (Dubois and Ohler 2001) are present in L. pera, except for the
femoral glands, which were thought to be absent. Here, we have shown femoral glands to be present both in L.
corrugatus (Manamendra-Arachchci and Pethiyagoda 2006) and L. pera.
Lankanectes pera differs from L. corrugatus in all criteria (external morphology, genetics and climatic niche)
on which they were evaluated. They differ from each other by 3.5−3.7% uncorrected genetic distances for 16S
rRNA, which is consistent with the range of species-level genetic distances commonly observed in amphibian sister
taxa (Vences et al. 2005). Lankanectes pera differs from L. corrugatus in at least 16 mutational steps, with no
sharing of haplotypes, indicating the reciprocal monophyly of the two clades; only between one and four
mutational steps are observed within populations of L. corrugatus. Though we have data for only one mt-DNA
gene fragment, 16S rRNA is considered a conservative mitochondrial gene, and hence the patterns that are observed
here are expected to hold also for other mt-DNA fragments.
In morphology, there are several, consistent, but somewhat subtle differences between the two species—
tubercle distribution and several color and pattern related features (Figs. 2,3,4). However, in morphometry, in PCI,
which is explained mostly by size, only separates the males, with a slight overlap of females, with L. pera being
larger than L. corrugatus.
In contrast to L. corrugatus, which occurs commonly in muddy substrates, including marshes and rice paddies,
Zootaxa 4461 (4) © 2018 Magnolia Press
where they burrow into soft mud and leaf litter, L. pera has so far only been found under more pristine conditions—
sand and rock strewn clear-water mountain streams flowing under canopy cover, where they hide in rock crevices.
This niche specialization partitioning between the two species can be an important factor that prevents the
specialized L. pera from spreading more widely.
The large tadpole (maximum total length of ca. 45 mm) that was collected from the habitat of L. pera was
DNA barcoded and confirmed to be of this species (Fig. 1). There seems to be resource partitioning between the
adults and tadpole (stage 35) habitats—tadpoles are found in pockets of deeper pools with detritus, while adults
prefer rocky and sandy regions. The tadpole of L. corrugatus has been described (Ukuwela & Bandara 2009), and
in external morphology, the tadpole of L. pera is similar to that of L. corrugatus.
Ecological niche models suggest Lankanectes pera to be a montane isolate. Its predicted distribution is limited
(360 km
) due to its adaptations to high-altitude bioclimatic conditions. Though suitable climatic conditions are
predicted by the niche model also to be present in the northern region of the central mountains, this area is
climatically and ecologically isolated from the presently known range of the species. Due to its small area of
occupancy and extent of occurrence (sensu IUCN Redlist Criteria 2001), together with the small population sizes
observed during this study, L. pera can be evaluated as a Critically Endangered species. In contrast, the present area
occupied by and predicted for L. corrugatus is much larger (14,120 km
). Due to its wide distribution and large
population size, L. corrugatus is considered as a Least Concern species.
Hence, Lankanectes pera is in need of immediate conservation attention due to its specialized habitat
requirements and climatic conditions, which is predicted to deteriorate under the current predicted global climatic
warming models. The Knuckles range is already highlighted as a mountain refuge for as many as eight micro-
endemic frog species that are already considered to be critically endangered or endangered (Meegaskumbura &
Manamendra-Arachchi 2005, Manamendra-Arachchi & Pethiyagoda 2006, MOE 2012, Senevirathne &
Meegaskumbura 2015). Given that the habitat requirements of L. pera is different from that of the highly
threatened micro-endemics highlighted so far, the conservation strategy for amphibians of the Knuckles mountains
must consider this new knowledge, i.e. the conservation of streams of the mountains.
Given that Lankanectes being endemic to Sri Lanka, occupying a position on a phylogenetic long branch as two
distinct species, in the absence of a common name to highlight these frogs, we propose calling them Corrugated
Frogs, which describes the numerous and prominent transverse skin folds on of both L. corrugatus and L. pera.
We thank the following individuals and organizations: an anonymous reviewer and Miguel Vences for suggesting
improvements to the manuscript; Department of Wildlife Conservation and Forest Department of Sri Lanka for
research permits and logistical support at their respective reserves; National Research Council of Sri Lanka (NRC
11-124) for graduate student support; members of the EES lab for field support. GS acknowledges the Rufford
Small Grant (No. 192721) and NW, the Nagao Natural Environment Foundation for funding part of this work.
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Presence locations used to make predicted distribution maps for the two species, Lankanectes corrugatus and L. pera
Species District Locality Altitude Coordinates
(m) Latitude (
N) Longitude (
L. corrugatus Nuwara-Eliya Bogawanthalawa 1300 6.7977 80.6783
Nuwara-Eliya Bogawanthalawa 1300 6.7952 80.6865
Nuwara-Eliya Moray Estate 1430 6.7961 80.5238
Ratnapura Deniyaya 470 6.3380 80.5516
Ratnapura Kudawa 560 6.4201 80.4196
Ratnapura Dodammuluwa 410 6.4220 80.4498
Ratnapura Morningside Forest Reserve 1050 6.4068 80.6133
Ratnapura Suriyakanda 1100 6.4426 80.6199
Ratnapura Batadombalena 350 6.7747 80.3946
Ratnapura Ehaliyagoda 140 6.8501 80.1965
Ratnapura Mahawalathenna 540 6.5889 80.7494
Ratnapura Samanala Nature Reserve 1040 6.7986 80.4690
Kandy Doluwa 550 7.1832 80.6109
Kandy Peradeniya 520 7.2527 80.5997
Kegalle Deraniyagala 150 6.9368 80.3416
Kegalle Avissawella 70 6.9620 80.2485
Galle Kottawa 80 6.0985 80.3155
Galle Hiniduma 50 6.2515 80.3397
Kaluthara Athwelthota 80 6.5336 80.2927
Matara Diyaduwa 300 6.3544 80.4971
Galle Hiniduma 60 6.3096 80.3238
Matale Rattota 390 7.5183 80.6858
Matale Kumbiyangoda 500 7.4542 80.5925
Colombo Avissawella 30 6.9333 80.1876
L. pera sp. nov. Matale Knuckles F.R. 1330 7.5233 80.7333
Matale Knuckles F.R. 1260 7.5180 80.7375
Matale Knuckles F.R. 1580 7.4646 80.7409
Kandy Meemure 1100 7.4104 80.8240
Kandy Hunnasgiriya 1420 7.3209 80.8568
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APPENDIX 2. Ventral view of Lankanectes pera (A, DZ1858) and L. corrugatus (B, DZ1399). The scale bar represents
20 mm.
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APPENDIX 3. Component loadings and the variances explained by the loadings for the Lankanectes Principal
Components Analysis.
Variables Component Loading 1 Component Loading 2 Component Loading 3
SVL 0.996 0.048 -0.008
TL3 0.994 0.039 -0.007
PAL 0.994 -0.059 -0.055
FOL 0.993 0.077 -0.001
TL4 0.992 -0.012 0.062
TBL 0.992 0.067 -0.029
TL2 0.992 0.004 0.034
ES 0.991 -0.025 0.029
MN 0.990 -0.009 0.023
HW 0.989 0.065 -0.054
TL5 0.989 0.030 0.058
HL 0.988 0.010 0.015
FL3 0.987 -0.122 -0.055
MFE 0.987 -0.026 0.014
TL1 0.985 -0.090 -0.047
DFE 0.979 -0.010 -0.000
DBE 0.974 0.105 -0.140
FL2 0.973 -0.007 -0.138
FL4 0.969 -0.100 -0.171
MBE 0.964 0.044 0.043
LAL 0.954 0.120 0.091
ED 0.952 0.043 -0.189
IO 0.943 -0.145 0.084
EN 0.926 -0.109 0.252
FLI 0.924 -0.334 0.099
FEL 0.922 0.127 -0.140
IN 0.919 -0.284 0.041
NS 0.913 -0.324 0.027
IML 0.865 0.289 0.360
UAW 0.849 0.155 -0.210
UEW 0.808 0.519 0.046
Variance explained by components 28.51 0.810 0.414
% of total variance explained by components 91.96 2.60 1.34
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APPENDIX 4. Variables used in morphometric analysis; measurements in mm.
……continued on the next page
Species Voucher Number
L. corrugatus WHT3007 11 6.7 4.6 3.4 6.6 19.1 3.7 3.9 5.2 3.6 27.7 18.9 17.1 3.5 2.5
L. corrugatus WHT3008 11.3 7.1 4.9 3.4 6.9 17.6 3.8 3.5 5 4.2 27.2 18.2 17.5 3.3 3
L. corrugatus WHT3009 8.1 4.8 3.8 2.7 5.1 10.6 2.3 2.7 3.7 2.8 18.8 12.8 11.7 2 2.2
L. corrugatus WHT3018 7.6 4.7 3.5 2.4 4.6 13.2 2.8 2.8 3.9 3.2 18.2 11.8 11.8 1.8 2.2
L. corrugatus WHT3019 9.6 5.7 3.9 2.4 5.1 14.7 2.8 2.9 4 3.3 19.3 13.6 13.2 2 2.3
L. corrugatus WHT3013 10.3 5.8 4.6 2.9 5.7 16 2.8 3.3 4.4 3.6 22.9 15 15.2 2.1 2.5
L. corrugatus WHT3014 10.5 6.1 4.9 2.7 6 18.3 3.3 3.6 4.9 4 25.1 16.3 15.2 2.3 2.7
L. corrugatus WHT3015 11.1 5.8 4.7 2.7 5.7 16.1 3.1 3.2 4.6 4 22.6 16.5 15.1 2 2.6
L. corrugatus WHT3010 8.4 5 3.8 2.4 4.7 12.7 2.4 2.4 3.7 3 17.5 11.5 11.4 1.8 2.2
L. corrugatus WHT3011 7.7 5.1 3.5 2.4 4.5 11.9 2.3 2.4 3.5 2.5 17.9 11.2 11.4 1.8 2.1
L. corrugatus WHT2641 9.2 5.2 4.3 2.7 4.8 13.6 3 2.7 4 3.3 22 14.2 13.5 2.5 2.3
L. corrugatus ZMB4897 11.7 6.5 4.5 3.3 6.3 21.9 3.3 3.4 4.7 3.8 28.7 16.6 16.8 3.1 2.7
L. corrugatus ZMB62772 8.9 5.4 3.8 2.6 4.9 14.7 2.7 2.7 3.8 3.2 20.8 12.7 12.2 2.6 2.2
L. corrugatus ZMB62771 10.6 5.7 5.1 2.8 5.4 18 2.8 3.3 4.5 3.9 24 14.9 14 2.2 2.6
L. pera DZ1858 17.23 9.6 7.5 4.28 9.09 29.7 5.8 6 7.9 6.9 42.03 27.35 25.71 3.32 3.6
L. pera DZ1859 13.99 7.72 5.38 3.34 7.55 24.61 4.45 4.5 6.1 5.2 30.78 20.29 19.61 3.18 3.1
L. pera DZ1860 11.14 6.76 5.23 3.01 6.23 18.81 3.9 4 5.4 4.7 26.12 16.9 16.65 2.4 2.8
L. pera DZ1307 17.25 8.71 6.72 3.81 9.3 31.47 5.74 5.19 7.59 6.62 41.33 29.87 28.01 4.1 3.74
L. pera DZ1290 16.02 8.1 6.5 3.1 7.53 28 3.07 5 5.92 5.32 35.7 22.48 22.86 3.62 2.5
L. pera DZ1302 13.35 6.62 4.83 2.92 6.6 29.39 3.88 3.9 5.12 4.48 29.38 17.3 19.08 2.98 2.79
Zootaxa 4461 (4) © 2018 Magnolia Press
APPENDIX 4. (Continued)
L. corrugatus
WHT3007 3.3 9.8 9.4 13.5 16.6 3.8
44.9 18.5 4.3 5.9 8.6 11.2 8.6 7.7 2.8 Morningside FR
L. corrugatus
WHT3008 3.4 8.4 8.2 12.1 15 3.5
43.7 19.5 4.5 5.9 8.4 10.9 8.2 7.2 2.8 Morningside FR
L. corrugatus
WHT3009 2.1 6.1 6.3 9.1 11.4 2.6
28.6 13.1 3.2 4.1 6 7.8 5.9 7.2 1.9 Morningside FR
L. corrugatus
WHT3018 2.3 6.1 5.8 8.6 10.4 2.3
29.1 12.3 2.9 3.6 5.5 7.4 5.4 6.7 1.6 Agra Elbadda, Agarapathana
L. corrugatus
WHT3019 2.3 6.8 6.1 9.1 10.7 2.8
31 13.3 3.1 3.9 6 7.5 5.3 5.8 1.9 Agra Elbadda, Agarapathana
L. corrugatus
WHT3013 2.9 7.2 7.2 10.8 12.9 3.1
36.3 16.1 3.6 4.8 7 9 6.9 9 2.5 Agra Elbadda, Agarapathana
L. corrugatus
WHT3014 2.6 6.9 8.4 12 14.5 3.3
38.1 17.4 4.4 5.2 7.4 9.8 7.1 5.6 2.5 Agra Elbadda, Agarapathana
L. corrugatus
WHT3015 2.2 7 8.5 12 14.1 3.4
38.4 16.7 3.6 4.7 6.8 8.8 6.3 6.9 2.3 Agra Elbadda, Agarapathana
L. corrugatus
WHT3010 2.1 5.3 4.8 7.8 9.8 2.4
27.2 12.3 3.1 4 5.8 7.7 5.8 5.5 2.1 Morningside FR
L. corrugatus
WHT3011 2.5 5.7 5.6 8.2 11.5 2.2
27.5 12.3 2.7 3.7 5.2 7.1 5 5.6 1.8 Morningside FR
L. corrugatus
WHT2641 2.7 6.5 6.5 10.4 12.8 2.3
33.4 15.3 3.3 4.7 6.7 9.2 6.4 5.6 2.2 Kottawa FR galle
L. corrugatus
ZMB 4897 3 9.6 8.9 12.7 15.3 3.4 43.8 19.1 3.9 5.4 8.3 10.9 7.7 7.4 2.7 Syntype, Ramboda
L. corrugatus
ZMB 62772 2 6.7 6.8 9.3 11.1 2.6
32.3 13.6 3.3 4.5 6 8.2 6.3 5.3 2.4 Syntype, Ramboda
L. corrugatus
ZMB 62771 2.3 8 7.2 11.1 13.3 2.8 37.8 15.8 3.6 4.5 7 9.1 7.1 6.4 2.9 Syntype, Ramboda
L. pera DZ1858 5.59
66 28.44
7.3 9 12.7 15.9 12 9.82 3.05 Knuckles
L. pera DZ1859 4.13
8.7 10.39
51 21.49
5.4 7 10.2 12.8 9.4 7.73 2.55 Knuckles
L. pera DZ1860 3.07
7.78 8.79 12.42
4.6 5.5 8.5 10.8 7.9 7.61 2.23 Hunnasgiriya
L. pera DZ1307 6.41
8.71 12.57 15.71 11.5 10.7 3.63 Riverston
L. pera DZ1290 3.6 10.91
7.28 10.92 12.77 9.79 9.79 3.93 Riverston
L. pera DZ1302 3.15
8.1 7.5 12.08
3.3 47.3 20.98
5.93 8.72 11.15 8.03 8.48 2.28 Riverston
Zootaxa 4461 (4) © 2018 Magnolia Press
Other material studied: mature female, 34.5 mm SVL, WHT1299B, Yogama; mature male, 36.8 mm SVL, WHT1299A; mature
female, 38.7 mm SVL, WHT816, Kalatuwawa, Labugama; mature female, 37.1 mm SVL, WHT945, Koskulana, Panapola;
mature male, 44.4 mm SVL, WHT912, Ambalamahena, Athwelthota; mature female, 59 mm SVL, WHT5132, Agra, Elbedda;
mature male, 56.6 mm SVL, Agra, Elbedda; mature male, 56.6 mm SVL, Agra, Elbedda; mature male, 30.8 mm SVL,
WHT875C, Kanneliya forest reserve; mature female, 33.3 mm SVL, WHT882, Devon ford estate; mature female, 44.6 mmm,
Kanneliya FR, mature female, 25.7 mm SVL, WHT875B, Kanneliya FR; mature male, 26.4 mm SVL, WHT2474, Morningside
FR; mature female, 22.1 mm SVL, WHT810, Namunukula, gonkale; mature male, 22.5 mm SVL, WHT3017, Agra, Elbedda;
mature female, 50.0 mm SVL, WHT5130, Agra, Elbedda; mature male, 57.7 mm SVL, WHT 5129, Agra, Elbedda; mature
male, 45.3 mm SVL, WHT5128, Agra, Elbedda; mature female, 34.4 mm SVL.
... No information is available on the larval biology or reproductive modes of Astrobatrachus. Astrobatrachus differs from Lankanectes in lacking dorsal skin ridges, lacking a lateral line system as an adult, having a distinct tympanum and canthus rostralis, lacking odontoid fangs, having widely spaced nasals, having a long ridge of vomerine teeth, having separate anterior and posterior portions of the vomer with the latter fused to the neopalatine, lacking a forked omosternum, lacking a prominent dorsal crest on the ilium, and lacking both a tarsal fold and webbed pedal phalanges Senevirathne et al., 2018;Figs. 5 and 6). ...
... Leaf-litter dwelling and habitat distinguishes A. kurichiyana from many species of Nyctibatrachus that are torrential frogs and prefer to live in water or next to perennial streams (Biju et al., 2011). While its terrestrial habits are somewhat similar to some small-bodied Nyctibatrachus species (see Garg et al., 2017), the new lineage differs strongly from the two Lankanectes species which are aquatic (Senevirathne et al., 2018). ...
... The genus Nyctibatrachus contains predominantly torrent-dwelling species (Biju et al., 2011;Garg et al., 2017). The Sri Lankan Lankanectes is almost entirely aquatic, but resides in slower-moving streams (Delorme et al., 2004;Senevirathne et al., 2018). The only known species of Astrobatrachus comprises entirely terrestrial forest-dwellers that prefer leaf-litter habitats. ...
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The Western Ghats (WG) is an escarpment on the west coast of Peninsular India, housing one of the richest assemblages of frogs in the world, with three endemic families. Here, we report the discovery of a new ancient lineage from a high-elevation massif in the Wayanad Plateau of the southern WG. Phylogenetic analysis reveals that the lineage belongs to Natatanura and clusters with Nyctibatrachidae, a family endemic to the WG/Sri Lanka biodiversity hotspot. Based on geographic distribution, unique morphological traits, deep genetic divergence, and phylogenetic position that distinguishes the lineage from the two nyctibatrachid subfamilies Nyctibatrachinae Blommers-Schlösser, 1993 and Lankanectinae Dubois & Ohler, 2001, we erect a new subfamily Astrobatrachinae subfam. nov. (endemic to the WG, Peninsular India), and describe a new genus Astrobatrachus gen. nov. and species, Astrobatrachus kurichiyana sp. nov. The discovery of this species adds to the list of deeply divergent and monotypic or depauperate lineages with narrow geographic ranges in the southern massifs of the WG. The southern regions of the WG have long been considered geographic and climatic refugia, and this new relict lineage underscores their evolutionary significance. The small range of this species exclusively outside protected areas highlights the significance of reserve forest tracts in the WG in housing evolutionary novelty. This reinforces the need for intensive sampling to uncover new lineages and advance our understanding of the historical biogeography of this ancient landmass.
... The high-risk zone (Figure 1) comprises three out of the five amphibian zones of Sri Lanka (Batuwita et al., 2019; MoMD&E, 2019): the Central Highlands, the Knuckles Massif, and the Rakwana Hills. The importance of these regions as local amphibian hotspots is illustrated by the many new species described from them in the recent past (Manamendra-Arachchi & Pethiyagoda, 2005; Meegaskumbura & Manamendra-Arachchi, 2005; Meegaskumbura & Manamendra-Arachchi, 2011; Wickramasinghe et al., 2013b;Senevirathne et al., 2018), as well as the rediscovery of species(Wickramasinghe et al., 2013a). Thus, the spread of Bd in these areas could have devastating consequences for the amphibians of Sri Lanka.The present scientific consensus is that Bd is Korean in origin, and mainland Asian frog populations have a low prevalence and load of Bd(O'Hanlon et al., 2018;Sreedharan & Vasudevan, 2021). ...
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Chytridiomycosis, caused by Batrachochytrium dendrobatidis ( Bd ), constitutes a major threat to many amphibian species worldwide. Predicting the species and regions of highest geographical risk is critical for the early detection and mitigation of chytrid emergence. In this study, using a niche modelling approach, the most conducive habitat for Bd within Sri Lanka (a high‐risk zone) was modelled. The distribution of 69 amphibian species was then modelled and their overlap with the high‐risk zone (area Bd ) was calculated. Using area Bd and a biotic index (BI), created using ecological traits of each species, a risk index (RI) was calculated. Using this RI, a high‐risk species index (HRSI) was developed to identify the species most at risk. The results indicate that the high elevations of Sri Lanka (>600 m a.s.l.) are highly conducive for Bd . The HRSI includes 35 species, with Minervarya greenii being the species most at risk. All species in the HRSI are globally Critically Endangered ( n = 14) or Endangered ( n = 21). We propose active conservation measures such as the routine monitoring of HRSI species and other proactive measures to identify and prevent the spread of Bd . We believe our findings would promote the establishment of pre‐emptive mitigation measures both within Sri Lanka and elsewhere, to counter the threat of chytridiomycosis and to conserve amphibian species.
... Sri Lanka is an amphibian hotspot with a high level of endemism (Ellepola et al. 2021;Meegaskumbura et al. 2002Meegaskumbura et al. , 2009Meegaskumbura et al. , 2019Senevirathne et al. 2018;Wijayathilaka et al. 2016). Being the most diverse and speciose amphibian genera of the island, Pseudophilautus shows a high degree of adaptation to the diverse microhabitats and climatic conditions. ...
Pseudophilautus conniffae from Lowland wet zone of Sri Lanka, was described as a new species in 2019. The validity of the new species was questionable and was often challenged as it shares strong morphological resemblance with P. limbus. Moreover, the phylogenetic placement of P. conniffae was unknown as no molecular data was available until now. Here, we generated 16S DNA sequences and re-examined the external morphological characters to assess its taxonomic distinctiveness. Pseudophilautus conniffae was recovered as being close to P. limbus with strong posterior probability and bootstrap support. The uncorrected pairwise genetic distance between P. conniffae and P. limbus was negligible, being less than 0.3% for the 16S gene fragment. Further two molecular species delimitation methods, ABGD and mPTP suggested that P. conniffae and P. limbus are a single operational taxonomic unit. The Principal Component Analysis of the morphometric characters also resulted in overlapping clusters. These results suggest that the newly described P. conniffae is not a valid species and we conclude that P. conniffae as a junior synonym of P. limbus.
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We used citizen science and inexpensive methodology to assess the distribution of the jungle cat Felis chaus , a relatively common species in Sri Lanka but the least studied of the four wild cat species occurring in the country. We obtained three types of records of the jungle cat: geo-referenced photographs of the species from the public; sightings obtained from print and social media, and provided via an online sighting form; and sightings by field biologists. We combined the 112 unique records obtained in this way with the 21 records from the 2012 National Red List distribution map of the species, and used MaxEnt to predict habitat suitability for the species. The new sightings were primarily in drier regions, expanding the known extent of occurrence for this species in Sri Lanka. Of the new sightings, 7.1% were road kills. Distance to nearest riverine forest, annual precipitation and distance to the nearest reservoir were the most important variables explaining habitat suitability. These findings validate our hypotheses that the species is more widespread than demonstrated previously and also ranges in human-dominated landscapes outside protected areas. Our study provides a model for how ecological and behavioural information for common species can be obtained inexpensively and incorporated into species distribution models. Studies of species such as the jungle cat, which are neither threatened nor charismatic, will help ensure that we keep common species common.
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Background Sri Lanka is a continental island separated from India by the Palk Strait, a shallow-shelf sea, which was emergent during periods of lowered sea level. Its biodiversity is concentrated in its perhumid south-western ‘wet zone’. The island’s freshwater fishes are dominated by the Cyprinidae, characterized by small diversifications of species derived from dispersals from India. These include five diminutive, endemic species of Pethia ( P. bandula , P. cumingii , P. melanomaculata , P. nigrofasciata , P. reval ), whose evolutionary history remains poorly understood. Here, based on comprehensive geographic sampling, we explore the phylogeny, phylogeography and morphological diversity of the genus in Sri Lanka. Results The phylogenetic analyses, based on mitochondrial and nuclear loci, recover Sri Lankan Pethia as polyphyletic. The reciprocal monophyly of P. bandula and P. nigrofasciata , and P. cumingii and P. reval , is not supported. Pethia nigrofasciata , P. cumingii , and P. reval show strong phylogeographic structure in the wet zone, compared with P. melanomaculata , which ranges across the dry and intermediate zones. Translocated populations of P. nigrofasciata and P. reval in the Central Hills likely originate from multiple sources. Morphological analyses reveal populations of P. nigrofasciata proximal to P. bandula , a narrow-range endemic, to have a mix of characters between the two species. Similarly, populations of P. cumingii in the Kalu basin possess orange fins, a state between the red-finned P. reval from Kelani to Deduru and yellow-finned P. cumingii from Bentara to Gin basins. Conclusions Polyphyly in Sri Lankan Pethia suggests two or three colonizations from mainland India. Strong phylogeographic structure in P. nigrofasciata , P. cumingii and P. reval , compared with P. melanomaculata , supports a model wherein the topographically complex wet zone harbors greater genetic diversity than the topographically uniform dry-zone. Mixed morphological characters between P. bandula and P. nigrofasciata , and P. cumingii and P. reval , and their unresolved phylogenies, may suggest recent speciation scenarios with incomplete lineage sorting, or hybridization.
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Purpose : The nomenclature of the wild strawberries inhabited in Sri Lanka is ambiguous. In Sri Lanka, this species is still named Duchesnea indica which needs a revision. Wild strawberries grow well in natural habitats of upcountry in Sri Lanka. Since the commercial strawberry cultivations gain a popularity in upcountry, the studies on wild strawberry is essential for crop improvement and management. Research Method : In the present study, we conducted extensive field sampling followed by a phylogenetic analysis with the DNA barcoding markers ITS and, trnL-F by using a representative sample of wild strawberry plants in Sri Lanka. The distribution of the species was identified using maximum entropy modeling approaches. Findings : Sri Lankan wild strawberry got placed at subtribe: Potentilla, and clade: Reptans and show a shallow divergence with the species Potentilla indica reported. Thus, we reposition the genus of wild strawberries in Sri Lanka from Duchesnea to Potentilla and hereafter name it as P. indica. The niche model analysis predicted a highly restricted distribution of Sri Lankan wild strawberry in Nuwara-Eliya district over an area of 166.36 km 2 in the altitude range of 1546-2524 m in a small climatic envelop highlighting the need for urgent conservation measures. Research Limitations : The pop-set for available in literature of P. indica is limited for comparison. Extensive studies based on DNA sequencing is needed for further validation. Originality / Value : Taxonomy, narrow distribution, need of conservation, and phylogenetic distance to Fragaria chiloensis, a progenitor species of cultivated strawberry, are defined for Sri Lankan wild strawberries.
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The amphibian fauna of Sri Lanka comprises 120 species, including 107 (~90.0%) endemic species. They belong to five families: Bufonidae, Dicroglossidae, Ichthyophiidae, Microhylidae, and Rhacophoridae. Based on distribution, we recognized five zoogeographic zones for them, Central Hills, Dry Zone, Knuckles Range, Lowland Wet Zone, and Rakwana Hills. Fifty three species were reported from the Central Hills (48 endemics [90.6%] and 42 [79.2%] threatened species). 47 species were recorded from the Lowland Wet Zone, including 36 (76.6%) endemics and 28 (59.6%) threatened species. The Knuckles Range had 25 species, of which, 19 (76.0%) were endemics and 15 (60.0%) are threatened species. 19 species were reported from Dry Zone including seven endemics (36.8%) and four threatened species (21.1%). Out of 29 species, which inhabited in the Rakwana Hills, 26 were endemics (~89.7%) including 24 (82.8%) threatened species. Species diversity along the elevational gradient was also observed with the highest species richness in the mid-elevational localities. Family Ichthyophiidae can be considered as the least studied family. Recent rediscoveries and studies have helped to reduce the number of extinct species from 21 to 18. It is speculated that some of the other extinct species have to be rediscovered or probably were misidentified as other species. About 90% of Sri Lankan amphibians occur in the regions with the highest human populations where there are established agricultural lands. Loss of habitats, competition due to anthropogenic species and invasive species, pollution (cause for malformations, parasites, and other diseases), and climate change appear to be major threats.
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We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line. The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput and scripted analysis. Both versions are available from free of charge.
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The Indian Purple frog, Nasikabatrachus sahyadrensis, occupies a basal phylogenetic position among neobatrachian anurans and has a very unusual life history. Tadpoles have a large ventral oral sucker, which they use to cling to rocks in torrents, whereas metamorphs possess adaptations for life underground. The developmental changes that underlie these shifts in habits and habitats, and especially the internal remodeling of the cranial and postcranial skeleton, are unknown. Using a nearly complete metamorphic series from free-living larva to metamorph, we describe the postembryonic skeletal ontogeny of this ancient and unique monotypic lineage. The torrent-dwelling larva possesses a dorsoventrally flattened body and a head with tiny dorsal eyes, robust lower and upper jaw cartilages, well-developed trabecular horns, and a definable gap between the trabecular horns and the tip of the snout. Unlike tadpoles of many other frogs, those of Nasikabatrachus retain larval mouthparts into late metamorphic stages. This unusual feature enables the larvae to maintain their clinging habit until near the end of metamorphosis. The subsequent ontogenetic shift from clinging to digging is correlated with rapid morphological changes and behavioral modifications. Metamorphs are equipped with a shortened tibiafibula and ossified prehallical elements, which likely facilitate initial digging using the hind limbs. Subsequently, the frogs may shift to headfirst burrowing by using the wedge-shaped skull, anteriorly positioned pectoral girdle, well-developed humeral crests and spatula-shaped forelimbs. The transition from an aquatic life in torrents to a terrestrial life underground entails dramatic changes in skeletal morphology and function that represent an extreme in metamorphic remodeling. Our analysis enhances the scope for detailed comparative studies across anurans, a group renowned for the diversity of its life history strategies.
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Eight new species of Sri Lankan frogs of the genus Philautus are described (P. mooreorum, P. poppiae, P. hoffmanni, P. mittermeieri, P. frankenbergi, P. hallidayi, P. steineri and P. stuarti). The species are diagnosed on the basis of mitochondrial DNA sequence, morphological features and, in two cases, bioacoustics data. Six of the eight species are confined to high elevation cloud forest isolates on the three main mountain ranges of central Sri Lanka. Because of their limited geographic distribution and small extent of remaining habitat, these species are classified as Endangered under the IUCN Red List criteria. These descriptions bring the total number of Sri Lankan Philautus to 61 species, 44 of which are extant.
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Nannophrys Günther, 1868, a group of flat-bodied frogs, is an endemic Sri Lankan genus bearing three extant and one extinct species, adapted to live among narrow and horizontal rock crevices adjacent to clear water streams. One of these species, Nannoprhys marmorata Kirtisinghe, 1946 is mostly restricted to the rock strewn streams of the Knuckles region (200-1200 m asl). Here, we re-describe the osteology of Nannophrys marmorata highlighting apomorphies and adaptations for life between narrow spaces. Previous studies on skeletal morphology of the genus Nannophrys include Gunther (1869), Boulenger (1882, 1890), Noble (1931), Kirtisinghe (1946), Clarke (1983) and Scott (2005). Basic descriptions of the skeleton of N. marmorata have been done (Kirtisinghe 1946; Clarke 1983), on which we build and elaborate. We describe the osteology using three adult specimens (SVL= 35.2-36.5 mm) of N. marmorata, stained differentially for bone and cartilage following the procedure by Taylor and Van Dyke (1985); we follow the osteological terminology of Trueb (1973), Duellman and Trueb (1986), and Pugener and Maglia (1997, 2009).
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We present a new open source, extensible and flexible software platform for Bayesian evolutionary analysis called BEAST 2. This software platform is a re-design of the popular BEAST 1 platform to correct structural deficiencies that became evident as the BEAST 1 software evolved. Key among those deficiencies was the lack of post-deployment extensibility. BEAST 2 now has a fully developed package management system that allows third party developers to write additional functionality that can be directly installed to the BEAST 2 analysis platform via a package manager without requiring a new software release of the platform. This package architecture is showcased with a number of recently published new models encompassing birth-death-sampling tree priors, phylodynamics and model averaging for substitution models and site partitioning. A second major improvement is the ability to read/write the entire state of the MCMC chain to/from disk allowing it to be easily shared between multiple instances of the BEAST software. This facilitates checkpointing and better support for multi-processor and high-end computing extensions. Finally, the functionality in new packages can be easily added to the user interface (BEAUti 2) by a simple XML template-based mechanism because BEAST 2 has been re-designed to provide greater integration between the analysis engine and the user interface so that, for example BEAST and BEAUti use exactly the same XML file format.
A list of the nominal genera and subgenera of Ranoidea (Amphibia Anura) of the world is given, with identification of their type-species. The nomenclatural consequences of the rediscovery of overlooked names or type-species designations are discussed. The following genus-group names are resurrected: Limnonectes Fitzinger, 1843 as a subgeneric name for Rana kuhlii Dumeril & Bibron, 1841 and its allies; Occidozyga Kuhl & Van Hasselt, 1822a in the place of Ooeidozyga Kuhl & Van Hasselt, 1822b; Nyctixalus Boulenger, 1882 in the place of Edwardtayloria Marx, 1975; and Tachycnemis Fitzinger, 1843 in the place of Megalixalus Gunther, 1869. The familial and subfamilial classification and nomenclature of the Ranoidea and the generic and subgeneric classification and nomenclature of the Rani-nae are also discussed. Five families are recognized in the superfamily Ranoidea: the Ranidae (Raninae, Phrynobatrachinae and Mantellinae), the Rhacophoridae (Rhacophorinae and Philautinae, the latter subfamily being created here), the Arthroleptidae (Astylosterninae and Arthroleptinae), the Hyperoliidae (Leptopelinae, Kassininae and Hyperoliinae) and the Hemi-sidae.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.