Journal of
Threatened Taxa | www.threatenedtaxa.org | 26 January 2019 | 11(1): 13120–13131
Description of a new species of
Pseudophilautus (Amphibia: Rhacophoridae) from southern Sri Lanka
Sudesh Batuwita 1, Madura De Silva 2 &
Sampath Udugampala 3
1 The Society for the Biodiversity
Conservation Sri Lanka, 63/4, Adikaramwatta, Yaggahapitiya, Gunnepana, Kandy
20270, Sri Lanka.
1 Present address: e: 130, Al Rustaq,
Rustaq, Welaiyh, Sultanate of Oman.
2,3 Wildlife Conservation Society-Galle, Biodiversity
Education and Research Center, Hiyare Reservoir, Hiyare 80000, Sri Lanka.
1 sudesh.batuwita@gmail.com (corresponding
author), 2 dsmanusha@gmail.com, 3 s.udugampal@gmail.com
Abstract: We describe a new Pseudophilautus
species, P. conniffae sp. nov. from southern Sri Lanka.
It was previously confused with Pseudophilautus
rus (Manamendra-Arachchi & Pethiyagoda). The new
species differs from the latter by the combination of the following
characters: fourth toe webbing to penultimate subarticular tubercle on inner
and outer sides (vs. fourth toe webbing in between penultimate and
anetpenultimate subarticular tubercles on inner and outer sides), presence of conical median lingual
process (vs. absent), and black patches on the posterior flank, anterior and
posterior edges of the thigh (vs. black patches on the anterior surface of the thigh). Pseudophilautus conniffae sp. nov. may be sympatric with P.
limbus (Manamendra-Arachchi & Pethiyagoda), which shares certain
characters with the new species. The new
species is, however, distinguished from P. limbus by the following characters: having supernumerary tubercles on manus (vs. lacking), absence
of frontoparietal ridges (vs. presence), fourth toe webbing to penultimate
subarticular tubercle on both sides (vs. fourth toe webbing between penultimate
and anetpenultimate subarticular tubercle on both sides), third toe webbing to
distal subarticular tubercle on both sides (vs. distal subarticular tubercle on
outer side and below penultimate subarticular tubercle on inner side), and having the dorsum light brown with dark
brown patches (vs. black and yellow variegated pattern on dorsum). The new species may be
restricted to the southwestern wet zone of Sri Lanka. It is compared with all known Pseudophilautus
species and also provided with a field key to identify it from those
species that are sympatric with it or inhabit the southwestern wet zone.
Keywords: Dediyagala, lowland rainforest, Pseudophilautus
rus, Pseudophilautus limbus.
doi: https://doi.org/10.11609/jott.3903.11.1.13120-13131 | ZooBank: urn:lsid:zoobank.org:pub:9C5058E7-5356-4F3E-A85B-EF2BFBD837E9
Editor: S.K. Dutta, Retired Professor of Zoology,
Bhubaneswar, India. Date of publication: 26
January 2019 (online & print)
Manuscript details: #3903 |
Received 16 November 2017 | Final received 18 November 2018 | Finally accepted
07 January 2019
Citation: Batuwita, S., M.D. Silva & S. Udugampala (2019). Description of a new species of Pseudophilautus
(Amphibia: Rhacophoridae) from southern Sri Lanka. Journal
of Threatened Taxa 11(1):13120–13131; https://doi.org/10.11609/jott.3903.11.1.13120-13131
Copyright: Batuwita et al. 2019. Creative Commons
Attribution 4.0 International License. JoTT allows unrestricted use,
reproduction, and distribution of this article in any medium by adequate credit
to the author(s) and the source of publication.
Funding: Co-funding by the Biodiversity Secretariat of the
Ministry of Environment & Renewable Energy of Sri Lanka (BDS) and the
Nations Trust Bank PLC Sri Lanka.
Competing interests: The authors declare no competing interests.
Author
Details: Sudesh
Batuwita is the President of the Society for the
Biodiversity Conservation of Sri Lanka. His interests are the taxonomy, ecology
and conservation of fauna of Sri Lanka. Madura
de Silva is the President of the Wildlife Conservation Society - Galle
Sri Lanka (WCSG). He is engaging on the taxonomy, conservation and management
of threatened fauna of Sri Lanka. Sampath
Udugampala is a Research Officer at the WCSG. He is working on the
taxonomy of vertebrates of Sri Lanka and engaging in a wild animal rescue
project.
Author
Contribution: SB, MDS &
SU - designed the research, did the field work and
wrote the manuscript.
Acknowledgements: We are grateful to the Department of Forest and the Department of
Wildlife Conservation (DWC) Sri Lanka for giving us permits (ref. WL/3/2/1/18)
to conduct this research. We are also grateful to the Biodiversity Secretariat
of the Ministry of Environment and Natural Resources (BDS) and the Nations
Trust Bank PLC Sri Lanka for providing financial support. Thanks to N.
Wickramasinghe, M. Goonathilake, and C. Munasinghe for giving us permission to
examine the WHT type material in their care. We are grateful to S. Seneviratne
(University of Colombo) for providing facilities and for his support to record
frog calls in the field. We are also grateful to Dharma Sri Kandamby for his
continuous guidance throughout our research. Thanks to S. Akmeemana, S.
Gunasingha, S. Darshana, L. Siriwardene, S. Deshaprema, and V. Pushpamal (all
of WCSG) for the fieldwork. Finally, we wish to extend our appreciation to
anonymous reviewers for constructive comments that help to improve the paper.
INTRODUCTION
Sri Lanka possesses highly diverse endemic
fauna and flora and is a reservoir of a unique evolutionary history (Myers et
al. 2000; Bossuyt et al. 2004). It is
also included in the revised hotspots of the world along with the Western Ghats
of India (Mittermeier et al. 1998). It
was estimated that India’s Western Ghats along with Sri Lanka lost more than
70% of its original habitat due to the rapid growth of human population (Myers
et al. 2000). In Sri Lanka, the most
sensitive and unique amphibian biodiversity is found to be
confined to the southwestern wet zone stretching up to the southwestern
side of the central massif covering the Peak Wilderness and the Horton Plains
(Manamendra-Arachchi & Pethiyagoda 2005; Meegaskumbura &
Manamendra-Arachchi 2005, 2011; Meegaskumbura et al. 2007, 2009, 2012;
Wickramasinghe et al. 2013, 2015).
South Asian Shrub Frog genus Pseudophilautus
is restricted to India and Sri Lanka (Biju & Bossuyt 2009;
Manamendra-Arachchi & Pethiyagoda 2006; Dinesh et al. 2017). Seventy-nine valid species are known from
this genus of which 76 occur in Sri Lanka (Manamendra-Arachchi &
Pethiyagoda 2005; Meegaskumbura & Manamendra-Arachchi 2005, 2011;
Meegaskumbura et al. 2007, 2009, 2012; Wickramasinghe et al. 2013a,b,c, 2015; Biju & Bossuyt 2009; Dinesh et al. 2017), all
of which are endemic to the island and all but one is restricted to the wet
zone (annual rainfall > 2,000mm) (Meegaskumbura et al. 2012). Interestingly, about 60% of the Sri Lankan
members of the genus were described in the recent past (Manamendra-Arachchi
& Pethiyagoda 2005; Meegaskumbura & Manamendra-Arachchi 2005 , 2011; Meegaskumbura et al. 2007, 2009; Wickramasinghe
et al. 2013a, 2015). Major forest reserves includes in the southwestern wet zone are Haycock,
Sinharaja, and Kanneliya-Dediyagala-Nakiyadeniya forest reserves (KDN
complex). Highest altitude is around
< ~600m in this region. These forests
are evergreen with no significant seasonal change. A total of 44 species of amphibians were
recorded from the lowland wet zone in the southwestern Sri Lanka, of which 34
(77.3%) are endemic to Sri Lanka (Manamendra-Arachchi & Pethiyagoda 2006;
Meegaskumbura & Manamendra-Arachchi 2005, 2011; Meegaskumbura et al. 2009;
Wickramasinghe et al. 2012). Sixteen
species of Pseudophilautus were hitherto recorded from the lowland rain
forests of the island’s southwestern wet zone (Manamendra-Arachchi &
Pethiyagoda 2005; Meegaskumbura & Manamendra-Arachchi 2005, 2011;
Meegaskumbura et al. 2009).
The new species described herein was first
observed during a biodiversity survey of the Dediyag--ala Forest Reserve by the
Wildlife Conservation Society-Galle (WCSG).
MATERIALS AND METHODS
Materials
used for this study are deposited in the collection of WCSG and will be deposited in the collection of the National Museum
Colombo (NMSL). Measurements were taken with KWB dial
vernier calipers to the nearest 0.1mm under a stereoscope. The methodology of measurements and anatomical
nomenclature follow Manamendra-Arachchi & Pethiyagoda (2005), except for
the lingual papillae. For lingual
papillae, we follow the terminology introduced by Grant et al. (1997), the
median lingual process. Images were
taken using Canon IXUS 50 and Nikon D700 digital cameras. Specimens were fixed in 10% formalin
bath and
were subsequently stored in 70% ethanol. Comparisons were made with the materials
preserved in the collections of the NMSL, the Wildlife Heritage Trust of Sri
Lanka (WHT, now in NMSL), Zoologisches Museum Berlin (ZMB), and also with published
descriptions of Bossuyt & Dubois (2001), Manamendra-Arachchi & Pethiyagoda (2005), Meegaskumbura &
Manamendra-Arachchi (2005), Biju & Bossuyt (2009), and Meegaskumbura et
al. (2009).
Bioacoustics data were recorded in the
field using a Marantz PMD660 recorder with a Sennheiser MKH20 microphone
mounted on to a parabola. Ambient
relative humidity and temperature were noted during the recordings. Calls were analyzed using Raven Lite
software, Raven Pro: Interactive Sound Analysis Software, Version 1.0 (The
Cornell Lab of Ornithology, Ithaca, NY).
The calls of the new species (n= 5) were recorded at 76% relative
humidity and 27.50C air temperature (during 2000–2300 hr), while the
calls of P. rus (n= 4) were recorded at 80% relative humidity and
260C (during 1800–2300 hr).
For call character terminology, we followed Cocroft & Ryan (1995).
RESULTS
Pseudophilautus conniffae sp. nov.
urn:lsid:zoobank.org:act:15697184-70F3-4E3D-A8DB-58FC94ACCD88
Conniff’s Shrub Frog (Images 1, 2A–E; Figs. 1A, 2)
Material examined: Holotype: NMSL WCSG 0005, male, 20.v.2012, 23.5mm SVL, Dediyagala Forest
Reserve, Matara District, Southern Province, Sri Lanka, 6.166⁰N, 80.433⁰E, 80m, coll. S.
Batuwita & S. Udugampala. Paratypes (all from type locality): NMSL WCSG 0001,
male, 25.6mm SVL, 14.v.2012, coll. S. Batuwita & S. Udugampala; NMSL WCSG
0003 & NMSL WCSG 0004, 29.3mm SVL & 24.1mm SVL, female & male,
10.vi.2012, coll. S. Batuwita, S. Udugampala & V. Pushpamal; NMSL WCSG
0002, female, 30.7mm SVL, 11.vi.2012, coll. S. Batuwita, M. de Silva & S.
Darshana; NMSL WCSG 0006, male, 22.9mm SVL, 11.vi.2012, coll. S. Batuwita &
S. Udugampala.
Diagnosis: Mature male holotype 23.5mm in SVL.
Tympanum discernible. Dorsal surface of body glandular, with prominent warts. Supratympanic fold
prominent. Canthal edges more or
less straight. Skin on head co-ossified
with cranium. Nuptial
pads absent. Median
lingual process present, small and conical. Tarsal tubercle present. Supernumerary tubercles
present on manus and pes. Toes
medially webbed. Chest
and belly heavily granular.
Large, dark brown more or less square-shaped blotch on middorsum between
forelimbs. Anterior
and posterior surfaces of thigh and posterior flank region with black and white
patches (Image 1D).
Description of holotype: (see Table 1 for mensural data). Snout rounded in lateral aspect (Image 1C), obtuse in dorsal aspect (Image 1A), angle of snout in dorsal
view ~900. Canthal edges more
or less straight (Image 1A), canthus rostralis
moderate (Image 1A). Loreal region concave
(Image 1A). Interorbital region flat. Snout, interorbital region, sides of head, and dorsum with
isolated glandular warts. Internarial
region slightly convex, nostril closer to snout tip
than eye (Image 1C). Eyes prominent, large,
laterally oriented (Image 1C). Pineal ocellus absent.
Tympanum distinct, oval, oblique (Image 1C); supratympanic fold
prominent (Image 1C). Skin on head co-ossified
with cranium. Tongue ovoid, distal end deeply divided; a median
lingual process present, conical and small.
Vomerine ridge present, bearing a few small teeth, angled at about 450
relative to body axis, shorter than
the distance between ridges.
Skin of ventral surface of throat with isolated granules; chest, belly, and
underside of thigh heavily granular (Image 1B);
flank with isolated, indistinct granules.
Dorsolateral fold absent. Cloacal opening directed posteriorly at the ventral level of thighs, unornamentated (Image1A).
Forelimbs moderately slender (Image 1A), their dorsal
and ventral surfaces smooth. Relative length of fingers:
III>IV>II>I (Image 2A); tips of fingers rounded, lateral dermal fringe
absent; circum-marginal groove
present. Subarticular tubercles rounded
to ovoid, distributed on fingers as follows: one on fingers І, ІІ, and ІV, two
on finger III (Images 2B); inner palmar tubercle ovoid (Image 2B); outer almost
granular, minute; supernumerary palmar tubercles absent. Nuptial pads absent (Image 2A).
Hind limbs slender (Image 1A,B); tarsal fold absent; tarsal tubercle present. Dorsal areas of thigh,
shank, and pes with a few scattered granules. Toes long, slender, their relative length:
IV>III~V>II>I (Image 2C); toes medially webbed (Image 2C), their tips
rounded; inner metatarsal tubercle elongate, no external tubercle (Image 2B). Subarticular tubercles rounded, small,
distributed on toes as follows: one on toes I and II, two on toes III and V,
three on toe IV (Image 2B,C).
Colour in life: (See Images 2D,E). Dorsum light brown with dark brown
patches. Middle
of tympanum dark brown, border lighter.
Upper lip dark brown with pale and ashy brown patches. Loreal region dark brown. Anterior mid dorsal region
of the body with a large, dark brown, square-shaped patch. Upper flank light brown to chestnut; lower
flank spotted in females. Dorsal and lateral regions of forelimb and dorsal region of thigh,
shank, and pes brown with dark brown crossbars. Throat, chest, belly, and
underside of thigh light yellow with small dusky spots, throat light yellow in
males. Webbing dark brown.
Colour in
preservative: (based on holotype,
NMSL WCSG 0005, see Image 2A,B). Dorsum greyish with dark
grey-brown patches. Anterior mid-region of the body with a large, dusky, square-shaped
patch. The region below
supratympanic fold dusky brown than that above, tympanum dusky brown than the
area surrounding it, lower margin of tympanum whitish. Loreal region greyish. Anterior flank greyish;
posterior flank, anterior, and posterior thigh marbled in black and white. Dorsal and lateral sides of forelimb, dorsal
side of thigh, dorsal side of shank, and dorsal side of pes brown with dark
brown crossbars. Throat
pale white with dark brown pigments; margins of lower jaw, anterior chest,
belly, and underside of thigh pale white with dark brown pigments. Webbing black with small grayish patches.
Vocalization: Pseudophilautus conniffae sp. nov. is able to emit two different
call types, of which one consists of a long series of unpulsed click-like notes
repeated at regular intervals. The other
call type consists of a shorter series of very short whistling notes repeated
in fast succession (Fig. 1A). The latter
call appears to be the advertisement call of P. conniffae sp. nov. The
advertisement call of the species may be distinguished from that of P. rus
by the greater call length (0.693–1.052 s vs. 0.408–0.427 s in P. rus),
higher frequency (4593–5111 Hz vs. 4222–4444 Hz), greater pulse length
(0.161–0.189 s vs. 0.046–0.047 s), and greater number of pulses per call (6–7
vs. 2–3) (Fig. 1A vs 1B).
Variation: Single specimen had less prominent warts on dorsum (NMSL WCSG 0003):
snout, interorbital region, sides of head, and dorsum had isolated
glandules. The angle of snout in dorsal
view was approximately 800 in females and ~900 in males.
Etymology: The species name is an eponym honoring Karen Iynn Conniff for her
generous support for the establishment of the WCSG and for her efforts to
document the Odonata of Sri Lanka.
Distribution: Pseudophilautus conniffae sp. nov. was recorded from Galle
and Matara districts (~80–300 m elevation; Fig. 2). It is confined to the rainforest areas in
both districts. Pseudophilautus
conniffae sp. nov. appears
to be a habitat specialist as it was often observed in bamboo Davidsea
attenuata vegetation (on dry bamboo leafs).
It was observed calling from leaves and bamboo branches less than 1m
above the ground.
Conservation status: Extent of occurrence: about 200km2, Dediyagala,
Kottawa-Kombala, and Hiyare forest reserves.
Outcome: Near Threatened.
Remarks
Pseudophilautus conniffae sp. nov. keys out as P. limbus according to the key of
Manamendra-Arachchi & Pethiyagoda (2005).
It may be distinguished from P. limbus, however, in having less prominent throat granules (vs. heavily granular in P.
limbus; see Manamendra-Arachchi & Pethiyagoda 2005: Fig. 95c vs. Image 1B), having
supernumerary tubercles on manus (vs. lacking), having the dorsum
light brown with dark brown patches (vs. black and yellow variegated pattern on
dorsum; see Manamendra-Arachchi & Pethiyagoda 2005: Fig. 93 vs. Image 2D,E),
lacking frontoparietal ridges (vs. having), fourth toe webbing to penultimate
subarticular tubercle on both sides (vs. below the penultimate subarticular
tubercle on both sides), and third toe webbing to distal subarticular tubercle
on both sides (vs. distal subarticular tubercle on outer side and below
penultimate subarticualr tubercle on inner side; see
Manamendra-Arachchi & Pethiyagoda 2005: Fig. 95f vs. Image 2B,C).
We note in passing that although Manamendra-Arachchi
& Pethiyagoda (2005) stated that nuptial pads were absent in males of P.
silvaticus, we observed these to be present in all male specimens of
its type series. While examining the type
specimens of the species described recently by Wickramasinghe et al. (2013), we
observed a few discrepancies: the authors mentioned that the tympanum of P.
newtonjayawardanei is wanting, but we
observed the presence of tympanum in the holotype and its upper one-third is
covered by supratympanic fold. Wickramasinghe et al. (2013) stated that P.
dayawansai lacks vomerine teeth, whereas we observed the same in the
holotype.
Comparisons: The new species is immediately distinguished from the following
species that are sympatric in the type locality
(only opposing suites of characters are
mentioned):
P. abundus (Manamendra-Arachchi & Pethiyagoda, 2005): throat heavily granular and
presence of a distinct suborbital white marking; P. auratus (Manamendra-Arachchi
& Pethiyagoda, 2005): median lingual process absent and vomerine teeth
absent; P. cavirostris (Günther, 1869): presence of tuberculated
fringe on posterior margin of lower arm and tarsus, and cloaca ornamented with
spine-like tubercles; P. folicola (Manamendra-Arachchi &
Pethiyagoda, 2005): a dark lateral stripe from nostrils to base of upper arm and two
fronto-parietal ridges present; P. hoipolloi (Manamendra-Arachchi &
Pethiyagoda, 2005): vomerine teeth absent and head skin not co-ossified with
cranium; P. limbus (Manamendra-Arachchi
& Pethiyagoda, 2005):
supernumarary tubercles absent on manus and throat heavily granular; P. mittermeieri (Meegaskumbura
& Manamendra-Arachchi, 2005): presence of tuberculated fringe on posterior
margin of lower arm and tarsus and
absence of vomerine teeth; P. nemus (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine teeth absent and median lingual process
absent; P. reticulatus (Günther, 1864):
presence of calcar and head skin not co-ossified with cranium; P. schneideri Meegaskumbura &
Manamendra-Arachchi, 2011:
absence
of median lingual process, head skin not co-ossified with cranium and presence
of nuptial pads; P. singu (Meegaskumbura, Manamendra-Arachchi & Pethiyagoda, 2009): vomerine teeth absent and head skin not
co-ossified with cranium
P.
sordidus (Manamendra-Arachchi
& Pethiyagoda, 2005): throat and chest smooth; P. stictomerus (Günther,
1876): median lingual process absent, and nuptial pads present in males; P. tanu (Meegaskumbura,
Manamendra-Arachchi & Pethiyagoda, 2009): absence of vomerine teeth, no
median lingual process and head skin not co-ossified with cranium.
The new
species is also distinguished from the following non-sympatric species (only opposing
suites of characters are mentioned): P. alto (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine
teeth absent and calcar present; P.
asankai (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine
teeth absent and tympanum not discernible; P. bambaradeniyai Wickramasinghe et
al., 2013: absence of supernumerary tubercles on manus and pes, absence of vomerine teeth, presence of nuptial pads and
absence of median lingual process;
P. caeruleus (Manamendra-Arachchi & Pethiyagoda, 2005): supratympanic fold not prominent and
absence of vomerine teeth; P. cuspis (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine
teeth absent and calcar present; P. dayawansai Wickramasinghe et al., 2013: presence of nuptial pads and absence of median lingual process; P. decoris (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine
teeth absent and calcar present; P. dilmah Wickramasinghe et al., 2015:
calcar present and vomerine teeth absent; P. femoralis (Günther, 1864):
vomerine teeth absent and supernumerary tubercles absent on pes; P. fergusonianus (Ahl, 1927): presence of nuptial pads and
absence of a median lingual process;
P. frankenbergi (Meegaskumbura & Manamendra-Arachchi, 2005):
supernumerary tubercles absent on pes and vomerine teeth absent; P. fulvus (Manamendra-Arachchi
& Pethiyagoda, 2005): calcar
present and nuptial pads present in males; P. hallidayi (Meegaskumbura
& Manamendra-Arachchi, 2005): vomerine teeth absent and nuptial pads
present in males; P. hankeni Meegaskumbura & Manamendra-Arachchi, 2011: presence
of dermal fringe on tarsal and absence of vomerine teeth; P. hoffmanni (Meegaskumbura &
Manamendra-Arachchi, 2005): skin on head not co-ossified with cranium and
vomerine teeth absent; P. hypomelas (Günther, 1876):
supernumerary tubercles absent on pes and vomerine teeth absent; P. jagathgunawardanai Wickramasinghe et
al., 2013: absence of vomerine teeth, presence of nuptial pads and
absence of median lingual process; P. karunarathnai Wickramasinghe et al.,
2013: absence of supernumerary tubercles on manus and pes, presence of nuptial pads and
absence of median lingual process;
P. lunatus (Manamendra-Arachchi & Pethiyagoda, 2005): calcar present and skin on head not
co-ossified with cranium; P. macropus (Günther, 1869):
skin on head not co-ossified with cranuim and toes fully webbed; P.
microtympanum (Günther, 1859):
skin on head not co-ossified with cranium and nuptial pads present in males; P.
mooreorum (Meegaskumbura & Manamendra-Arachchi, 2005): absence of
vomerine teeth and absence of supratympanic fold; P. newtonjayawardanei Wickramasinghe et al., 2013: nuptial pads present, absence of both vomerine
teeth and median lingual process; P.
ocularis (Manamendra-Arachchi & Pethiyagoda, 2005): throat heavily granular and skin on head
not co-ossified with cranium. P. papillosus (Manamendra-Arachchi & Pethiyagoda, 2005): calcar and nuptial pads present; P.
pleurotaenia (Boulenger,
1904): vomerine teeth absent and toes fully webbed; P. poppiae (Meegaskumbura
& Manamendra-Arachchi, 2005): vomerine teeth and supernumarary tubercles
absent on pes; P. popularis (Manamendra-Arachchi & Pethiyagoda, 2005): nuptial pads present in males and vomerine
teeth absent; P. procax (Manamendra-Arachchi & Pethiyagoda, 2005): supernumarary tubercles absent on pes and
a suborbital pale blotch present; P. puranappu Wickramasinghe et
al., 2013: presence of nuptial pads and
supernumarary tubercles absent on pes;
P. regius (Manamendra-Arachchi & Pethiyagoda, 2005): throat heavily granular and nuptial pads
present in males; P. rus (Manamendra-Arachchi & Pethiyagoda, 2005): absence of conical median
lingual process and anterior surface of thigh with black patches; P. samarakoon Wickramasinghe et al, 2013: nuptial pads present, absence of both
median lingual process and vomerine teeth; P. sarasinorum (Müller, 1887): skin on head not co-ossified
with cranium and vomerine teeth absent; P. schmarda (Kelaart, 1854):
prominent lateral dermal finge on lower arm and tarsus and vomerine teeth
absent; P. semiruber (Annandale, 1913): vomerine teeth absent and
supratympanic fold feebly developed; P. silus (Manamendra-Arachchi
& Pethiyagoda, 2005): throat
heavily granular and skin on head not co-ossified with cranium; P. silvaticus (Manamendra-Arachchi &
Pethiyagoda, 2005): skin on head not co-ossified with cranium and presence of a
V-shaped tubercular pattern starting behind eyes extending towards midline of dorsum; P. simba (Manamendra-Arachchi
& Pethiyagoda, 2005): skin on head
not co-ossified with cranium and no vomerine teeth; P. sirilwijesundarai
Wickramasinghe et al., 2013: absence of vomerine teeth, presence of nuptial pads and
absence of median lingual process;
P. steineri (Meegaskumbura & Manamendra-Arachchi, 2005): skin on
head not co-ossified with cranium and nuptial pads present in males; P.
stellatus (Kelaart, 1853): supratympanic fold indistinct and spotted dorsal
colouraton; P. stuarti (Meegaskumbura & Manamendra-Arachchi, 2005): vomerine
teeth absent and skin on head not co-ossified with cranium; P. viridis (Manamendra-Arachchi
& Pethiyagoda, 2005): vomerine
teeth absent and skin on head not co-ossified with cranium; P. zorro (Manamendra-Arachchi
& Pethiyagoda, 2005): calcar present
and vomerine teeth absent.
Pseudophilautus
conniffae sp. nov.
differs from following putative extinct
species (only opposing suite
of characters are mentioned): P.
adspersus (Günther, 1872): vomerine teeth absent, median lingual process
absent and throat heavily granular; P. dimbullae (Shreve, 1940): median lingual process absent and skin on head not co-ossified with cranium; P. eximius (Shreve, 1940): throat smooth and vomerine teeth
absent; P. extirpo (Manamendra-Arachchi & Pethiyagoda, 2005): throat heavily granular and
vomerine teeth absent; P. halyi (Boulenger, 1904): vomerine
teeth absent, median lingual process absent and skin on head not co-ossified with cranium; P. leucorhinus (Lichtenstein, Weinland
& Von Martens, 1856): vomerine teeth absent and median lingual process
absent; P. maia (Meegaskumbura,
Manamendra-Arachchi, Schneider & Pethiyagoda, 2007): absence of both tarsal
tubercle and median lingual process; P. malcolmsmithi (Ahl, 1927): median
lingual papilla absent; P. nasutus (Günther, 1869): calcar
present, median lingual process absent and vomerine teeth absent; P. oxyrhynchus (Günther, 1872): calcar present and vomerine teeth absent; P. pardus (Meegaskumbura, Manamendra-Arachchi, Schneider & Pethiyagoda,
2007): absence of both vomerine teeth and median lingual process; P. rugatus (Ahl, 1927): median lingual process absent, vomerine teeth absent
and throat heavily granular; P. temporalis (Günther, 1864): median lingual process absent and skin on head not
co-ossified with cranium; P. nanus (Günther, 1869): median lingual
process absent, nuptial pads present and throat heavily granular; P. variabilis (Günther, 1859): median lingual process absent, vomerine teeth
absent and throat heavily granular; P. zal (Manamendra-Arachchi & Pethiyagoda, 2005): vomerine teeth absent
and median lingual process absent; P. zimmeri (Ahl, 1927): nuptial pads present in males, median lingual
process absent and W-shaped tubercle pattern on head.
The new species can
also compared with three Indian species of Pseudophilautus (Dinesh et
al. 2017) (only opposing characters are mentioned): P. amboli (Biju
& Bossuyt, 2009): presence of nuptial pads in males and absence of median
lingual process; P. kani (Biju & Bossuyt, 2009): absence of nuptial
pads in males and median lingual process; P. wynadensis (Jerdon, 1853):
absence of both vomerine teeth and median lingual process.
Table 1. Mensural data of Pseudophilautus
conniffae sp. nov. (in
mm)
|
|
Holotype |
Paratypes |
||||
|
|
NMSL WCSG 0005 |
NMSL WCSG 0001 |
NMSL WCSG 0003 |
NMSL WCSG 0004 |
NMSL WCSG 0002 |
NMSL WCSG 0006 |
|
Sex |
Male |
Male |
Female |
Male |
Female |
Male |
|
DBE |
8.8 |
8.5 |
9.7 |
9.3 |
10.2 |
8.1 |
|
DFE |
4.7 |
4.9 |
5.6 |
4.9 |
5.5 |
4.4 |
|
DL |
1.2 |
1.2 |
1.7 |
1.3 |
1.4 |
1.4 |
|
DW |
0.9 |
1.0 |
1.5 |
1.2 |
1.4 |
1.1 |
|
ED |
3.4 |
3.4 |
4.1 |
3.6 |
5.2 |
3.7 |
|
EN |
2.5 |
2.5 |
2.7 |
2.6 |
3.0 |
2.7 |
|
ES |
4.2 |
4.7 |
5.3 |
4.4 |
5.5 |
4.4 |
|
FEL |
12.7 |
14.3 |
16.3 |
12.7 |
17.3 |
12.1 |
|
FL |
1.7 |
2.0 |
2.7 |
2.5 |
2.3 |
1.7 |
|
FOL |
16.9 |
17.9 |
22.3 |
17.5 |
22.1 |
16.7 |
|
HL |
11.0 |
11.4 |
13.2 |
11.6 |
14.2 |
11.2 |
|
HW |
10.6 |
11.0 |
13.1 |
11.2 |
12.9 |
10.3 |
|
IML |
1.0 |
1.0 |
1.4 |
1.1 |
1.3 |
1.1 |
|
IN |
2.4 |
2.2 |
2.5 |
2.5 |
2.6 |
2.5 |
|
IO |
2.7 |
3.1 |
3.2 |
2.7 |
3.0 |
2.7 |
|
LAL |
5.5 |
6.3 |
7.6 |
6.2 |
7.8 |
6.2 |
|
MBE |
4.8 |
4.2 |
5.6 |
4.7 |
4.5 |
4.3 |
|
MFE |
7.3 |
7.3 |
8.5 |
7.1 |
8.9 |
6.8 |
|
MN |
9.6 |
9.5 |
11 |
9.8 |
12.0 |
9.6 |
|
NS |
1.7 |
1.2 |
1.6 |
1.5 |
2.1 |
1.7 |
|
PAL |
6.3 |
6.8 |
8.4 |
7.0 |
8.4 |
5.6 |
|
SVL |
23.5 |
25.6 |
29.3 |
24.1 |
30.7 |
22.9 |
|
TAD |
1.1 |
1.7 |
1.4 |
1.2 |
1.7 |
0.8 |
|
TBL |
13.5 |
14.4 |
17.3 |
14.1 |
17.5 |
12.7 |
|
TL |
2.0 |
2.0 |
2.8 |
2.3 |
2.4 |
1.9 |
|
TND |
6.8 |
7.4 |
8.8 |
7.3 |
8.8 |
7.4 |
|
TPD |
4.7 |
4.7 |
5.5 |
4.9 |
5.3 |
4.6 |
|
TYE |
2.2 |
2.3 |
2.8 |
2.0 |
2.5 |
1.8 |
|
UAW |
4.6 |
4.6 |
6.5 |
4.7 |
6.5 |
5.2 |
|
UEW |
3.2 |
2.7 |
3.6 |
2.7 |
3.5 |
2.7 |
Table 2. Mensural data of Pseudophilautus
rus and P. silvaticus (in mm)
|
|
Pseudophilautus rus (paratypes) |
P. silvaticus (type series) |
||||||||
|
|
WHT 5438 |
WHT 5434 |
WHT 5436 |
WHT 5440 |
WHT 5432 |
WHT 3295, holotype |
WHT 3279 |
WHT 3310 |
WHT 3378 |
WHT 3462 |
|
Sex |
Male |
Male |
Male |
Male |
Male |
Male |
Male |
Male |
Male |
Male |
|
DBE |
6.4 |
7.3 |
7.5 |
7.4 |
7.5 |
9.3 |
8.8 |
10.2 |
9.2 |
7.6 |
|
DFE |
4.5 |
3.8 |
4.4 |
4.1 |
3.8 |
4.6 |
4.9 |
5.6 |
4.8 |
4.1 |
|
DL |
0.9 |
1.0 |
1.0 |
1.0 |
0.8 |
0.9 |
1.5 |
1.2 |
1.5 |
1.4 |
|
DW |
0.9 |
1.0 |
1.1 |
1.1 |
1.0 |
0.7 |
1.4 |
1.2 |
1.3 |
1.0 |
|
ED |
3.6 |
3.6 |
3.5 |
3.4 |
3.5 |
4.5 |
4.1 |
3.8 |
4.4 |
3.8 |
|
EN |
2.1 |
2.2 |
2.7 |
2.4 |
2.6 |
2.5 |
3.0 |
3.0 |
2.9 |
2.3 |
|
ES |
3.8 |
3.6 |
3.9 |
3.9 |
4.0 |
4.0 |
4.6 |
5.2 |
5.0 |
4.1 |
|
FEL |
12.5 |
12.9 |
12.9 |
12.7 |
13.0 |
16.1 |
17.0 |
16.0 |
17.3 |
14.1 |
|
FL |
2.4 |
2.4 |
3.1 |
3.0 |
2.3 |
2.3 |
1.8 |
2.6 |
2.7 |
1.9 |
|
FOL |
17.3 |
17.2 |
18.2 |
16.2 |
17.5 |
20.6 |
21.0 |
22.9 |
21.5 |
17.8 |
|
HL |
9.7 |
9.4 |
9.9 |
9.4 |
9.9 |
11.9 |
12.1 |
13.0 |
12.7 |
10.3 |
|
HW |
8.8 |
8.6 |
8.9 |
8.5 |
8.4 |
11.0 |
11.3 |
12.7 |
12.0 |
9.3 |
|
IML |
1.0 |
0.8 |
1.1 |
1.0 |
1.0 |
0.9 |
1.2 |
1.2 |
1.2 |
0.9 |
|
IN |
1.8 |
1.6 |
1.7 |
1.6 |
1.6 |
2.3 |
2.2 |
1.9 |
2.4 |
1.5 |
|
IO |
2.3 |
2.2 |
2.2 |
2.3 |
2.2 |
2.9 |
3.0 |
2.9 |
3.3 |
2.5 |
|
LAL |
4.8 |
5.0 |
5.3 |
4.8 |
4.6 |
6.8 |
6.7 |
7.0 |
7.5 |
6.4 |
|
LPH |
– |
– |
– |
– |
– |
0.5 |
0.2 |
0.3 |
– |
0.1 |
|
LPW |
– |
– |
– |
– |
– |
0.5 |
0.4 |
0.3 |
– |
0.4 |
|
MBE |
2.9 |
2.8 |
3.0 |
3.3 |
3.0 |
4.1 |
4.7 |
4.4 |
4.1 |
3.5 |
|
MFE |
6.0 |
6.0 |
6.3 |
6.1 |
6.5 |
7.6 |
8.1 |
7.9 |
8.2 |
6.9 |
|
MN |
8.1 |
8.1 |
8.5 |
7.8 |
8.2 |
9.8 |
10.3 |
11 |
10.7 |
8.6 |
|
NS |
0.7 |
1.0 |
0.9 |
1.0 |
1.1 |
1.1 |
1.4 |
1.2 |
1.7 |
1.0 |
|
PAL |
6.6 |
6.3 |
6.7 |
7.0 |
7.1 |
8.1 |
7.7 |
8.5 |
7.3 |
6.8 |
|
SVL |
21.8 |
20.5 |
21.8 |
22.8 |
22.8 |
27.4 |
27.1 |
31.4 |
28.3 |
24.8 |
|
TAD |
0.2 |
0.5 |
0.4 |
0.4 |
0.5 |
0.3 |
0.6 |
0.6 |
0.6 |
0.4 |
|
TBL |
13.3 |
12.3 |
13.3 |
12.7 |
13.1 |
16.7 |
17.7 |
17.4 |
18.0 |
14.3 |
|
TL |
1.5 |
1.6 |
2.0 |
1.8 |
2.2 |
2.4 |
1.8 |
2.6 |
2.0 |
1.7 |
|
TND |
5.8 |
5.5 |
6.3 |
6.2 |
6.5 |
7.3 |
7.4 |
7.6 |
7.6 |
6.6 |
|
TPD |
3.5 |
3.6 |
5.7 |
3.8 |
4.0 |
4.9 |
4.6 |
4.8 |
4.8 |
4.0 |
|
TYE |
1.4 |
1.4 |
1.5 |
1.6 |
1.6 |
1.8 |
2.1 |
1.9 |
2.2 |
1.8 |
|
UAW |
4.1 |
3.5 |
4.5 |
4.1 |
4.8 |
5.8 |
6.5 |
6.2 |
6.1 |
4.4 |
|
UEW |
1.9 |
2.1 |
2.4 |
2.6 |
2.3 |
2.1 |
2.9 |
2.3 |
2.8 |
2.5 |
DISCUSSION
Pseudophilautus comprises a highly diverse
amphibian species assemblage in Sri Lanka (Manamendra-Arachchi &
Pethiyagoda 2006). Currently, about 79
species are recognized from India and Sri Lanka (Manamendra-Arachchi &
Pethiyagoda 2005; Meegaskumbura & Manamendra-Arachchi 2005, 2011;
Meegaskumbura et al. 2007, 2009, 2012; Wickramasinghe et al. 2013a,b,c, 2015; Dinesh et al. 2017). The discovery of P. conniffae sp. nov. increases the assemblage of frog fauna of the lowland wet
zone of Sri Lanka to 17.
Among its Sri Lankan congeners, P. conniffae sp. nov. superficially resembles P. rus, P. fergusonianus, and P.
silvaticus. Pseudophilautus conniffae sp. nov. is distinguished
clearly from P. rus based on morphometric data (Table 1 vs. Table 2). Pseudophilautus
conniffae sp. nov. is
also distinguished from P. silvaticus in having a relatively acute snout
(~800–900 in dorsal view vs. 950–1000
in P. silvaticus), presence (vs. absence) of skin on head
co-ossified with cranium, and absence (vs. presence; Image 2F) of a V-shaped
tubercular pattern on dorsum.
Based on their
morphological and mensural data (Table 1 vs. Table 2), Pseudophilautus
conniffae sp. nov.
is easily distinguished from P. rus. To support our conclusion, however, we here
analyzed the call structure of both species.
The two species are easily distinguished by their call
length, frequency, pulse length, and
the pulse per call (Fig.
1A
vs. 1B). In addition, the type series of
Pseudophilautus rus were collected from Kandy (7.283⁰N & 80.583⁰E, ~500m) in the Kandy District, whereas the type
locality of P. conniffae sp. nov. is
Dediyagala (6.166⁰N & 80.433⁰E, 80m) in Matara District.
The two localities are at a ~200km straight-line distance and also in
different zoogeographic zones (Central
Hills and lowland wet zone, respectively). We confirmed that P. conniffae sp.
nov. is restricted to
lowland wet zone rain
forests
(in Galle and Matara districts), while P. rus is confined to forests and
forest buffer zones of the Central Hills
around Kandy District (see Fig. 2) and to
Morningside in the Rakwana Hills
(Meegaskumbura et al.
2012). We, however, did not observe P. rus from
Sinharaja, Millawa, Diyadawa, and the surrounding forests. Thus, P. rus and P. conniffae sp.
nov. are two discrete
species based on the geography as well.
In addition, P. rus is a habitat generalist, common in
anthropogenic habitats (e.g., buffer zones of Gannoruwa and Hantana mountains and home gardens), whereas P.
conniffae sp. nov. is a habitat specialist, distributed in lowland wet zone
forests such as Kottwa-Kombala, Kanneliya, and Dediyagala.
Grant et al. (1997)
stated no intraspecific variation in the median lingual process in frogs and
also mentioned that both sexes and all sizes of frogs had this character. Interestingly, we observed a paratype of P.
silvaticus (WHT 3378, an adult male) wanting this character (see also
Manamendra-Arachchi & Pethiyagoda 2005; e.g., P. folicola and P.
viridis). Thus, the individual
variation of such characters, including the median lingual process, in Sri
Lankan Pseudophilautus needs to be studied in the future.
Based on its
present distribution data, Pseudophilautus conniffiae sp. nov. can be a Near Threatened
species. The species has a restricted
distribution and is highly
vulnerable due to habitat fragmentation (Manamendra-Arachchi & Pethiyagoda
2005). Hence, conservation assessment of the species needs to be initiated in
the future.
For
images/figures – click here
Key to the species of Pseudophilautus that are or may be sympatric with the new species
1 A. Presence of a distinct suborbital white
marking/s
..........................................................................................................................
2
.. B.
Absence of suborbital marking/s ................................................................................................................................................
3
2 A. Absence
of
vomerine teeth/ ridge ................................................................................................................................
P. hoipolloi
B.
Presence of vomerine teeth/ ridge ................................................................................................................................................
4
3 A. Presence of tuberculated fringe on
posterior margin of lower arm and tarsus ..............................................................................
5
B.
Absence of tuberculated fringe on posterior
margin of lower arm and tarsus
................................................................................
6
4 . A. Dorsum
light brown color with scattered dark brown blotches
................................................................................
P. sordidus
B. Dorsum
dark green color with scattered dusky green blotches ......................................................................................
P. abundus
5 A. Presence of
vomerine teeth
.........................................................................................................................................
P. cavirostris
B.
Absence of vomerine teeth .......................................................................................................................................
P. mittermeieri
6 . A. Absence
of
vomerine teeth .........................................................................................................................................................
7
B.
Presence of vomerine teeth ..........................................................................................................................................................
8
7 . A. Longitudinal
dermal ridges on dorsum ...........................................................................................................................
P. nemus
B.
No longitudinal dermal ridges on
dorsum
.....................................................................................................................................
9
8 A. Presence
of calcar or tarsal tubercle ............................................................................................................................................
10
B.
Absence of calcar or tarsal tubercle ...........................................................................................................................................
11
9 A. Prominent
tubercle on upper eyelid ..................................................................................................................................
P. singu
. B.
No prominent tubercle on upper eyelid
....................................................................................................................................
12
10 A. Presence
of calcar ......................................................................................................................................................
P. reticulatus
B. Presence
of tarsal tubercle
............................................................................................................................................................
13
11 A.
Skin on head not co-ossified with cranium .................................................................................................................................
14
B. Skin on head co-ossified with cranium ........................................................................................................................................
15
12 A.
Skin on head co-ossified with cranium ...................................................................................................................................
P. tanu
B. Skin on head not co-ossified with cranium ..................................................................................................................................
16
13 A. Supernumarary tubercles absent on manus, two
frontoparietal ridges present; fourth toe webbing to below the penultimate subarticular
tubercle on both sides; third toe webbing to distal subarticular tubercle on
outer side and below penultimate
subarticualr
tubercle on inner side ......................................................................................................................................
P. limbus
B. Supernumerary tubercles present on manus, no frontoparietal
ridges; fourth toe webbing to penultimate subarticular tubercle
on both sides;
third toe webbing to distal subarticular tubercle on both sides ...................................................
P. conniffae
sp. nov.
14 A. Nuptial
pads absent in males; dorsum light brown color with scattered dark brown
blotches or spots .......................... P. sordidus
B.
Nuptial pads present in males; no
spots or blotches on dorsum .............................................................................
P. stictomerus
15 . A. Nuptial
pads present in males
..................................................................................................................................
P. schneideri
... B.
Nuptial pads absent in males .......................................................................................................................................
P. folicola
16 A. Median
lingual process present ...................................................................................................................................
P. hoipolloi
B. Median
lingual process absent ........................................................................................................................................
P. auratus
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Appendix
1. Comparative material examined
Pseudophilautus abundus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype, WHT 3006; Labugama Forest Reserve, Labugama.
Pseudophilautus auratus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype, WHT 2356; Morningside Forest Reserve (near Rakwana).
Pseudophilautus cavirostris (Günther, 1869): topotypes, WHT 1294, Haycock (Hiniduma, Galle); WHT
2318; Kosmulla near Neluwa, 450m.
Pseudophilautus cuspis (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 1177(e); paratypes: WHT 1177(a), 1177(b); Koskulana (near
Panapola).
Pseudophilautus decoris (Manamendra-Arachchi & Pethiyagoda,
2005): paratypes: WHT 3194, WHT 3257, WHT 3258; Morningside Forest Reserve
(near Rakwana).
Pseudophilautus fergusonianus (Ahl, 1927): topotypes, WHT 731, WHT 2233; Kumaradola,
Monaragala; WHT 3178, WHT 3179; Puwakpitiya, Knuckles Range.
Pseudophilautus folicola (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 2645; paratypes: WHT 2646, 2647; Kottawa (Galle).
Pseudophilautus hoipolloi (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 2673; paratypes, WHT 2674, 2675, Haycock (Hiniduma).
Pseudophilautus limbus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype, WHT 2700; Haycock.
Pseudophilautus lunatus (Manamendra-Arachchi &
Pethiyagoda, 2005): holotype, WHT 3283; Handapan Ella Plains.
Pseudophilautus microtympanum (Günther, 1859): topotypes, WHT 2710, WHT 3446; Horton Plains
National Park.
Pseudophilautus mittermeieri (Meegaskumbura & Manamendra-Arachchi,
2005): holotype: WHT 3522; paratype: WHT 3526; Kottawa, Galle.
Pseudophilautus nemus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 1319; Haycock (Hiniduma).
Pseudophilautus ocularis (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 2360; Morningside Forest Reserve (near Rakwana).
Pseudophilautus papillosus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype: WHT 3284; Handapan Ella Plains (near Rakwana).
Pseudophilautus procax (Manamendra-Arachchi & Pethiyagoda,
2005): paratypes: WHT 2786, 2787, WHT 3277, 3278; Morningside Forest Reserve
(near Rakwana).
Pseudophilautus reticulatus (Günther, 1864): topotypes, WHT
2288, 2289; Dediyagala Forest reserve, Akuressa; WHT 3366, Haycock (Hiniduma).
Pseudophilautus rus (Manamendra-Arachchi & Pethiyagoda,
2005): paratypes, WHT 5432, WHT 5434, WHT5436, WHT 5438, WHT 5440; Pilimatalawa
(near Peradeniya).
Pseudophilautus silvaticus (Manamendra-Arachchi & Pethiyagoda,
2005): holotype, WHT 3295; paratypes: WHT 3378, 3379, 3462; Handapan Ella
Plains (near Suriyakanda); WHT 3310, Sinharaja World Heritage Site (near
Kudawa).
Pseudophilautus
stictomerus (Günther, 1876): topotypes: WHT 1173, WHT
2402; Kottawa (Galle).
Pseudophilautus zimmeri (Ahl, 1927): holotype: ZMB 6111; Point de
Galle.