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Limnonectes kadarsani (Amphibia: Anura: Ranidae), a new frog from the Nusa Tenggara Islands

Authors:
  • Indonesian Academy of Sciences

Abstract

Limnonectes kadarsani, a new large species described here is known from Lombok, Sumbawa, Flores, and Adonara in the Nusa Tenggara Islands. This species is distinctive in having robust but slight elongated body in males and subadult specimens, skin covered with few elongated tubercles and a pair of short dorsolateral folds. Adult males have wider head, a distinct enlargement of the exoccipital crest and wider interorbital space and the eye-tympanum distance. Limnonectes kadarsani is believed to be closely related to L. macrodon, and to a lesser extent to L. modestus.
LIMNONECTES KADARSANI (AMPHIBIA: ANURA: RANIDAE),
A NEW FROG FROM THE NUSA TENGGARA ISLANDS
ABSTRACT. - Limnonectes kadarsani, a new large species described here is known
from Lombok, Sumbawa, Flores, and Adonara in the Nusa Tenggara Islands. This species
is distinctive in having robust but slight elongated body in males and subadult specimens,
skin covered with few elongated tubercles and a pair of short dorsolateral folds. Adult males
have wider head, a distinct enlargement of the exoccipital crest and wider interorbital space
and the eye-tympanum distance. Limnonectes kadarsani is believed to be closely related to
L. macrodon, and to a lesser extent to L. modestus.
When reviewing the amphibian collection at Museum Zoologicum Bogoriense (MZB)
in 1976, one of us (DTI) examined a number of specimens from Lombok, Sumbawa, Flores
and Adonara, identified as Limnonectes (=Rana) macrodon or L. modestus. They are distinct
in several ways from L. macrodon, especially by the distinctly enlarged exoccipital crest in
adult males. This new form is a large species up to 120 mm in both sexes. At that time, only
few species from other regions had been examined. After participating in a number of
expeditions to Sumatra, Java, Ambon and Sulawesi, we have now seen an adequate number
of species and are now able to understand that the L. macrodon populations from Nusa
Tenggara islands represent a new species.
Djoko T. Iskandar Laboratory of Biosystematics, Jurusan Biology, Faculty of Mathematics and
Natural Sciences, Institute of Technology, Bandung 10, Jalan Ganesha, Bandung 40132, Indonesia.
Boeadi, Mumpuni Sancoyo Museum Zoologicum Bogoriense, Research and Development Centre
for Biology, The Indonesian Institute of Sciences, 18, Jalan Jr. H. Juanda, Bogor 16002, Indonesia.
For correspondence, please refer to the first author.
Limnonectes kadarsani, new species
(Fig. 1)
Rana macrodon, Barbour, 1912 (part); Boulenger, 1920 (part); van Kampen, 1923 (part); Mertens,
1927 (part); Darevskyi, 1964.
Rana modesta, Barbour, 1912 (part); van Kampen, 1923 (part); Dunn, 1928.
Material examined. - Holotype - Adult male (MZB Amph. 2940 (ex.2654», Jaran Pasang, Lombok,
colI. D. Hardjono, 14 Mar. 1984.
Paratypes - LOMBOK: - male and female (MZB Amph. 2941-2942 (ex. 2652», - 2 males and
female (MZB Amph. 294~2945 (ex. 2651» - 2 males and female (MZB Amph. 2946-2948 (ex. 2653»
- male (MZB Amph. 2654), Jaran Pasang, colI. D. Hardjono, 8-14 Mar. 1984 - 2 males and female
(MZB Amph. 2949-2951 (ex. 2645», from West Lombok, no exact locality, colI. Rachmatun,
May. 1982. SUMBA WA: female (MZB Amph. 120), Batu Dulang, Sumbawa, colI. R. Mertens, Sunda
Exp. Rensch, 6 May. 1927 - females (MZB Amph, 2935-2936 (ex. 193) - males (MZB Amph. 2959,
2960 (ex. 215», (partly cleaned), Semangkat Atas, colI. R. Mertens, Sunda Exp. Rensch, 11 May. 1927
- males and females (MZB Amph. 2952-2956 (ex. 2507», Sariberu, nearSimpasai market, Monta
county, Bima, colI. M. Siluba, 7 Feb. 1984 - male and female (MZB Amph. 2957-2958 (ex. 2805»,
along stream creek near Air Merah, Batu Hijau, colI. K. Martin, Feb.1994. FLORES: 2 males and
female, (MZB Amph. 009a-c (3 of5 ex.), Wolowaru, colI. R. Mertens, Sunda-Exp. Rensch, 16 Jul.1927
- female (MZB Amph. 214), Bajawa, colI. R. Mertens, Sunda Exp. Rensch, 6 Jul.1927 - 2 females
and male (MZB Amph. 2937-2939 (ex.398», Rana Mese, colI. Fr. J.M. Vianney, 10 Apr.1958.
ADONARA: 2 males (MZB Amph. 1025-1026), Hinga, colI. Fr. J.M. Vianney, 10 May. 1959. - male
(MZB Amph. 1024), Wure, colI. Fr. J.M. Vianney, 3 May.1959.
The description is based on 35 adult specimens, from Lombok (13 ex.), Sumbawa (12 ex.), Flores
(9 ex.) and Adonara (3 ex.). In addition, we also examined 28 specimens of young and subadults from
the same localities, all from the MZB collection, but not considered as a part of the type series.
Diagnosis. - Body is usually much longer than broad, except in gravid females. The legs
are moderate in length and the foot is fully webbed to the toe disk. The colour is brownish
to grayish black. A short, fme dorsolateral fold is located at the anterior part of dorsum. A
distinct
"I \"
or "w" shaped scapular marking is present between the shoulders. Skin is
usually smooth in adult specimens, but young specimens have two rows of enlarged tubercles
in the mid dorsal of the sacral region.
Description. - A large frog, body much longer than broad, adults up to about 120 mID.
Head much wider than long in adult males, in females and young males it is nearly as wide
as long (see Fig. 1); distinctly enlarged odontoid processes at end of mandible, larger in
adult males; snout rounded, feebly projecting; nostril closer to tip of snout than to eye; canthus
rounded, loreal region oblique, slightly concave or straight; eyes pointing obliquely upwards,
interorbital narrower than eye diameter; vomerine teeth in two oblique series just behind the
level of choanae; tympanum distinct, smaller than eye-nostril distance. Limbs stout, fingers
short, length of fingers 3> 1>4>2; a distinct fringe of skin along both sides of second and
third fingers, not movable; subarticular tubercles well developed. Tips of finger distinctly
enlarged, devoid of circum-marginal groove, dorsal groove present. Hindlimbs short; heels
slightly overlapping when legs flexed at right angle to the body; tibia about one-half of
SVL; toes fully webbed up to swollen tips; toe disks distinctly enlarged; a narrow tarsal fold
and a flap along the outer toes and metatarsal. Elongated inner metatarsal tubercle followed
with a non-movable skin fold, compressed; no outer metatarsal tubercle.
Skin slightly rugose,with tubercles on the sides of body, a short dorsolateral fold on the
anterior portion of the back,a "/ \"or "w" shaped scapular markings. A few short folds,
spinules or small tubercles,especially on the posterior part of the back and on the legs.
Young specimens with two rows of tubercles on the dorsum between the dorsolateral
(paravertebral) region, especially on the sacral region. These tubercles always disappeared
in adults, leaving only few elongated folds. Upper eyelids with tubercles, supra-tympanic
fold thick.Lower parts of body and legs smooth.
Fig.1. Holotype of Limnonectes kadarsani.A. dorsal aspect;B.ventral aspect; C.Lateral view of the
head to show the exoccipital crest enlargement.
THE RAFFLES BULLETIN
OF
ZOOLOGY
199644(1)
Skin slightly rugose, with tubercles on the sides
of
body, a short dorsolateral fold on the
anterior portion
of
the back, a
"/
\" or "w" shaped scapular markings. A few short folds,
spinules or small tubercles, especially on the posterior part
of
the back and on the legs.
Young specimens with two rows
of
tubercles on the dorsum between the dorsolateral
(paravertebral) region, especially on the sacral region. These tubercles always disappeared
in adults, leaving only few elongated folds. Upper eyelids with tubercles, supra-tympanic
fold thick. Lower parts
of
body and legs smooth.
Eggs moderate (2.2 mm in female
of
91
mm), with darker animal pole.
B
Fig.
1.
Holotype
of
Limnonectes kadarsani.
A.
dorsal aspect;
B.
ventral aspect;
C.
Lateral view of the
head to show the exoccipital crest enlargement.
23
Colouration in alcohol. - Uniform grayish black, indistinct bar between the eyes,
supratympanic fold widely black coloured. Dorsum more or less uniformly reddish brown
to grayish brown, lips with vertical bars; throat and breast dusted with gray. Skin is covered
with light tipped tubercles.
Secondary sex characters. - Males are slight larger than females, vocal sacs absent.
Females with eggs measure 53-107 mm; males with secondary growth of exoccipital crest
measure 54-120 mm. The head is wider in males than females. The head is distinctly enlarged
Table 1. Morphometric data and ratios for Limnonectes kadarsani.
CHARACTERS MALES· FEMALES RATIOS MALES FEMALES
(IN=18) (N=14) (N=18) (N=14)
SVL 94.06 ± 17.93 86.47 ± 14.75 HW/HL 1.10 ± 0.05 1.14 ± 0.05
54.0 - 120.0 53.5 - 107.0 1.03 - 1.19 1.05 - 1.20
HW 39.66 ± 8.00 33.36 ± 6.08 IN
I
10
1.06 ± 0.19 1.15±0.15
23.7 - 55.2 20.0 - 42.0 0.78 - 1.21 0.86 - 1.30
HL 37.94 ± 7.69 29.40 ± 5.39 IO
lEY
0.63 ± 0.11 0.56 ± 0.05
21.6 - 46.5 18.3 - 36.0 0.40 - 0.83 0.44 - 0.68
FE 44.96 ± 8.27 42.72 ± 7.28 TY
lEY
0.58 ± 0.04 0.53 ± 0.07
28.6 - 57.8 27.1 - 51.6 0.51 - 0.65 0.47 - 0.75
TI 47.49 ± 7.99 45.04 ± 7.41 EY
I
SL 1.03 ± 0.10 1.06 ± 0.06
29.7 - 60.2 29.2 - 55. 0.86 - 1.18 0.95 - 1.10
FL 68.57 ± 13.72 66.92 ± 12.47 TI
I
SVL 0.51 ± 0.Q3 0.53 ± 0.02
42.6 - 89.5 40.4 - 84.9 0.46 - 0.55 0.49 - 0.56
SL 11.54 ± 1.81 10.69 ± 2.04 HW
ISVL
0.43 ± 0.03 0.39 ± 0.02
8.0 - 15.0 6.6 - 13.7 0.37 - 0.48 0.36 - 0.41
TY 6.92 1.29 6.23 ± 1.78 HL/SVL 0.39 ± 0.02 0.34 ± 0.02
4.5 - 9.0 3.4 - 7.5 0.34 - 0.40 0.30-0.36
EY 12.01 ± 2.28 11.29 ± 2.24 FL
I
SVL 0.74 ± 0.05 0.78 ± 0.02
7.1 - 14.6 7.2 - 13.9 0.65 - 0.81 0.73 - 0.81
10
7.64 ± 2.09 6.33 ± 1.25 TY
I
SVL 0.07 ± 0.01 0.07 ± 0.01
4.1 - 11.3 3.7 - 8.0 0.06 - 0.08 0.05 - 0.12
EN 8.24 ± 1.71 7.81 ± 1.33 IN
I
SVL 0.08 ± 0.01 0.08 ± 0.01
5.4 - 10.6 5.1 - 9.7 0.08 - 0.10 0.07 - 0.09
IN 7.81 ± 1.51 7.18 ± 1.18 EN
I
SVL 0.08 ± 0.01 0.09 ± 0.01
5.0-11.0 5.1 - 9.2 0.07 - 0.11 0.08 - 0.12
ET 7.84 ± 2.66 4.74 ± 1.22 ET
I
SVL 0.08 ± 0.02 0.05 ± 0.01
3.4 - 14.5 2.5 - 6.7 0.06 - 0.13 0.04 - 0.07
The first line is mean ± SD; second line is the range. All morphometrical values are in millimeters.
Abbreviations of variables are as follows: SVL = snout-vent length; HW = head width; HL = head
length; FE = femoral length; TI = tibial length; FL = foot length; SL = snout length;
10
= interorbital
space; IN = intemarial distance; EN = eye - nostril distance; EY = eye diameter; ET = eye - tympanum
distance; TY = tympanum diameter.
24
~SVIo
~HW
"- HL
~IO
~ET
~SVL
~HW
-.- HL
-<>-10
~ET
Fig. 2. Some morphometrical differences between males (A) and females (B) of Limnonectes kadarsani
in relation to snout-vent length. See Table I for abbreviations of variables.
in adult males, resulting from secondary growth in the posterior portion of the head. The
distance between eye and tympanum in extra large males could attain 15 mm, but in large
females only between 4 to 6 mm. The measurements of this character are biased by the
secondary growth when the specimens reach adulthood as shown in Table 1. The anterior
part of the post-occipital is slightly bulging in adult males so that the interorbital becomes
wider and the exoccipital crest is distinctly enlarged and could be seen as a raised, visible
rounded knob at the posterior part of the head. The bodies of gravid females are stocky and
are larger than the head; in the adult males, the head is wider than the body. Some other
characters that are significantly different between male and female specimens and are not
clear in Table 1, are presented in Fig. 2.
Etymology. - It is our pleasure to name this new species in honor of Dr. Sampurno
Kadarsan, the former Director of Museum Zoologicum Bogoriense (MZB, Bogor) during
the periods of 1960-1962; 1964-1968 and 1971-1977. For his achievements, he received
about eight medals of honour from the Indonesian government. Dr. Kadarsan retired in
September 1994, after serving MZB for 40 years (1955-1994). His contributions towards
the progress of Museum Zoologicum Bogoriense up until present are immeasurable and
incomparable to any Indonesian biologist.
Barbour (1912), van Kampen (1923) and Dunn (1928) reported the occurence of
Limnonectes modestus from Lombok Island, presumably based on young specimens. Their
records was concured by Mertens (1930) and considered that the record was based on young
specimens of L. macrodon. The specimens examined by Dunn (1928) according to Mertens
(1929) were in fact L. dammermani. After examining all of the Sulawesi an and Moluccan
species and comparing them to the Nusa Tenggara populations, we understand why at least
a part of the Lombok population was assigned to L. modestus. Although maturity is reached
at about 55 mm in both species, L. modestus is a smaller species (max 80 mm) compared to
L. kadarsani. There is no distinct secondary sex characters in the overall morphology of L.
modestus, except for the presence of paired vocal sacs. The differences of SVL and head
width between sexes or in young L. modestus are not significant. Unless dissected, it is
impossible to distinguish young males from young females of both species. In L. modestus,
both sexes have enlarged odontoid processes, but this structure is only present in males of
L. kadarsani. Males of {-.modestus have only slight larger head and more prominent odontoid
processes than femald: The odontoid processes of males of L. modestus directed upwards
but in adult females directed backwards at about 45°; the odontoid bones are exposed in both
sexes. Beside the enlargement of post-occipital knob and the enlarged odontoid processes,
adult males of L. kadarsani are much larger with distinct enlarging and widening of head.
Young specimens of L. kadarsani usually have two rows of tubercles as in L. modestus, but
these tubercles tend to disappear in adults, leaving only a few small elongated tubercles or
spinules. The presence of the tubercles in young specimens seemed to be the main reason
why some authors attributed the Lombok's population to L. modestus (see Barbour, 1912;
van Kampen, 1923; Dunn, 1928). They probably examined some subadult specimens, and
could not find the diagnostic characters of L. kadarsani.
The presence of two rows of tubercles at the mid-dorsal region between dorsolateral of
the sacral regions is not only known in Limnonectes modestus, but equally present in several
smaller forms related to L. microdiscus from Sulawesi as well as in L. dammermani. Another
species, L. dammermani is the only other Limnonectes in the Nusa Tenggara Islands. The
less toe webbings and small size will distinguish L. dammermani from the fully webbed and
the much larger sized L. kadarsani.
Limnonectes macrodon, to which L. kadarsani was attributed (Barbour, 1912; Boulenger,
1920; van Kampen, 1923; Mertens, 1927, 1930; Darevskyi, 1964), is very similar in the
overall physiognomy of the body and legs, width of the head and skin texture of the adults.
If young specimens of both forms are compared side by side, the prominent skin tubercles
in young L. kadarsani will distinguish these two forms immediately. Adults of L. macrodon
could be distinguished immediately from L. kadarsani based on the absence of enlargements
of exoccipital crest and the much larger size (see note below).
All other large South-East Asian species, Limnonectes ingeri, L. blythi, L. ibanorum, L.
malesianus, L. grunniens, L.macrocephalus and L. magnus were all easily distinguished by
the absence of visible exoccipital crest, hence different from L. kadarsani in this regard. In
contrast, most young specimens or smaller species are practically indistinguishable from the
similar sized L.kadarsani. The presence of vocal sacs in males of L. acanthi, L.
microtympanum, L. heinrichi, L. modestus or L. visayanus will distinguish these species
from L. kadarsani, but female specimens still remain a problem. The presence of small and
sharp odontoid processes in females of L. modestus, L. heinrichi and L. microtympanum will
be very useful to distinguish them from females and young males of L. kadarsani.
According to Dubois's arrangements (1986; 1992), this species should be placed within
or close to members of the subgenus Bourretia based on the enlargements of the exoccipital
crest and postoccipital bones. As shown above, this new species is very closely related to
L.
modestus or L. macrodon. Two other species, L. blythi and L. malesianus have also slight
enlargement of the exoccipital crest in very big males although not distinctly seen, bulging
from the surface of the skin, and the interorbital space is not widened. Keeping these species
together within the subgenus Bourretia will only obscure the relationships between East
Indonesian Limnonectes and such arrangement is not supported by geographical evidences.
Note. - The types of Limnonectes macrodon is a composite. To avoid confusion, Kiew
(1978) chose a lectotype to solve the problem. All specimens from Java at our disposition
consist of L. blythi like species, which will be discussed elsewhere (Inger
&
Iskandar, in
prep). Here we present additional data result from our examination of the type series. In fact
the type series is also 'as complicated as its name. Aside the lectotype and some of the
paralectotypes, ttk rest of the type series (not examined by Kiew) consists of one specimen
from Ambon, one other from Sulawesi, another single individual from the Philippines and
two others from Borneo. The Ambonese specimen belongs to L. grunniens, the Sulawesian
specimen belongs to an undescribed spe.cies (Iskandar, 1996), and the Philippine specimen
probably to L. visayanus. Those from Borneo, presumed to be young specimens are in fact
adult specimens of L. paramacrodon.
I thank R.F. Inger, for important comments on drafts of the manuscript. A. Dubois and
A.M. Ohler kindly helped me during my visit to MNHN (Paris). The work is partly supported
by Basic Science Grant No. 601P4M/DPPMIL-3311/MS/1993 from the Indonesian Directorate
General of Higher Education, Ministry of Education originated from World Bank IBRD-
Loan No. 2944 and 331l-IND.
Barbour, T., 1912. A contribution to the zoogeography of the East Indian islands. Mem. Mus. Compo
Zool. Harvard, XLIV (I): 1-168, 18 tabl, 8 pis.
Boulenger, G.A., 1920. A monograph of the South-Asian, Papuan, Melanesian and Australian frogs
of the Genus Rana. Rec. Indian Mus., 20:1-223.
Dubois, A., 1992. Notes sur la classification des Ranidae (Amphibiens Anoures). Bull. mens. Soc.
linn. Lyon, 1992,
61
(10): 305-352.
Dunn, E.R., 1927. Results of the Douglas Burden expedition to the Island of Komodo. IV. Frogs from
the East Indies., Amer. Mus. Novit., 315: 9 pp.
Iskandar, D.T., 1996. Limnonectes (Amphibia, Anura, Ranidae), a new large frog from
Central Sulawesi. Alytes (submitted).
Kiew, B.H., 1978. The nomenclature and identity of the javanese frog Rana macrodon Dumeril and
Bibron. Malayan Nat.,
31
(4): 219-229.
Mertens, R., 1927. Neue Amphibien und Reptilien aus dem Indo-Australischen Archipel.
Senckenbergiana, 9 (6): 234-242.
Mertens, R., 1929. Herpetologische Mitteilungen.
xxv.
Zur Kenntnis der Rana microdisca Boettger
und ihrer Rassen. Zool. Anz., 86: 66-68.
Mertens, R., 1930. Die Amphibien und Reptilien der inseln Bali, Lombok, Sumbawa und Flores. Abh.
Senck. Naturf. Ges., 42: 115-344. 9 pIs.
Van Kampen, P.N., 1907. Amphibien des indischen Archipels. in M. Weber, Zool. Erg. Niederlandisch
Ost-Indien, 4 (2): 383-418, pI. 16.
Van Kampen, P.N., 1923. The Amphibians of the Indo-Australian Archipelago. Amsterdam, E.J. Brill.
304 pp.
Received 27 Apr 1995
Accepted 25 Oct 1995
... TWA Kerandangan juga memiliki spesies amfibi endemik yaitu Limnonectes kadarsani [13]. Spesies ini hanya terdistribusi di Nusa Tenggara [14]. Berdasarkan IUCN Red List [15], trend populasi dari Limnonectes kadarsani adalah decreasing. ...
... Pengumpulan sampel dimulai dari pukul 20.00 -24.00. sampel dalam penelitian ini adalah amfibi dewasa yang ditentukan berdasarkan Snout to venth length (SVL) mengacu pada [14,20]. Secara khusus, sampel untuk spesies Limnonectes kadarsani mencakup individu dewasa dan juvenile. ...
... Keterangan: Kr = kelimpahan relatif (%) ni = jumlah individu spesies i N = jumlah individu seluruh spesies H' = indeks keanekaragaman spesies pi = ni/N E = indeks kemerataan spesies S = kekayaan spesies D = indeks dominansi spesies Piramida dibuat berdasarkan jumlah amfibi juvenile dan dewasa. Amfibi juvenile dan dewasa ditentukan berdasarkan Snout to Vent length (SVL) tiap spesies mengacu pada [14,20,21,22]. Jumlah individu pada fase juvenile menjadi dasar/pondasi piramida, dan fase dewasa sebagai puncaknya. ...
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... The Lesser Sunda Islands contain 18 species of anurans including two species of fanged frogs of the genus Limnonectes (Family Dicroglossidae), which are common throughout Southeast Asia (Inger, 1999). Both species are restricted to the Sunda Arc Islands (Figure 1), with L. dammermani (Mertens, 1929) reported from Lombok (with unconfirmed reports from Sumbawa and Flores) and Limnonectes kadarsani reported from Lombok, Sumbawa, Flores, Adonara and Lembata (Iskandar, Boeadi, & Sancoyo, 1996;Iskandar & Mumpuni, 2004a,b). Research on Limnonectes from other parts of Southeast Asia, such as the Greater Sunda Islands, Sulawesi and the Philippines have shown that these frogs are not only able to colonize oceanic islands but that they also have frequently diverged ecologically and morphologically, radiating into new species that can be sympatrically distributed (Evans et al., 2003;Iskandar, Evans, & McGuire, 2014;McLeod, 2010;Setiadi et al., 2011). ...
... kadarsani is on Lombok (Iskandar et al., 1996), L. kadarsani species B-E will require new names should these lineages be formally described (which we will not pursue here). That L. kadarsani is composed of multiple independently evolving lineages is consistent with the idea that species diversity in the Lesser Sundas is substantially underestimated, and that the western Lesser Sundas is an area of endemism representing a distinct Wallacean biogeographical unit (Michaux, 2010). ...
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Aim: The Lesser Sunda Islands are situated between the Sunda and Sahul Shelves, with a linear arrangement that has functioned as a two-way filter for taxa dispersing between the Asian and Australo-Papuan biogeographical realms. Distributional patterns of many terrestrial vertebrates suggest a stepping-stone model of island colonization. Here we investigate the timing and sequence of island colonization in Asian-origin fanged frogs from the volcanic Sunda Arc islands with the goal of testing the stepping-stone model of island colonization. Location: The Indonesian islands of Java, Lombok, Sumbawa, Flores and Lembata. Taxon: Limnonectes dammermani and L. kadarsani (Family: Dicroglossidae) Methods: Mitochondrial DNA was sequenced from 153 frogs to identify major lineages and to select samples for an exon-capture experiment. We designed probes to capture sequence data from 974 exonic loci (1,235,981 bp) from 48 frogs including the outgroup species, L. microdiscus. The resulting data were analysed using phylogenetic, population genetic and biogeographical model testing methods. Results: The mtDNA phylogeny finds L. kadarsani paraphyletic with respect to L. dammermani, with a pectinate topology consistent with the stepping-stone model. Phylogenomic analyses of 974 exons recovered the two species as monophyletic sister taxa that diverged ~7.6 Ma with no detectable contemporary gene flow, suggesting introgression of the L. dammermani mitochondrion into L. kadarsani on Lombok resulting from an isolated ancient hybridization event ~4 Ma. Within L. kadarsani, the Lombok lineage diverged first while the Sumbawa and Lembata lineages are nested within a Flores assemblage composed of two parapatrically distributed lineages meeting in central Flores. Biogeographical model comparison found strict stepping-stone dispersal to be less likely than models involving leapfrog dispersal events. Main conclusions: These results suggest that the currently accepted stepping-stone model of island colonization might not best explain the current patterns of diversity in the archipelago. The high degree of genetic structure, large divergence times, and absent or low levels of migration between lineages suggests that L. kadarsani represents five distinct species.
... Manfaat dari amfibi yang besar tersebut tidak berbanding lurus dengan upaya konservasi terhadap potensi keanekaragaman hayati tersebut (Iskandar and Erdelen, 2006). Fakta ini dapat dibuktikan dengan tidak adanya spesies dari kelas amfibi yang berstatus dilindungi UU (Kusrini, 2003;Ariza dkk., 2014), padahal terdapat spesies di Pulau Lombok yang terancam karena mengalami trend penurunan populasi (Iskandar et al., 1996;Iskandar and Mumpuni, 2004). Banyak dari aspek ekologi amfibi di Indonesia belum diketahui (Iskandar and Erdelen, 2006) karena studi tentang kelompok ini masih jarang (Kusrini, 2003). ...
... Fenomena semakin meluasnya daerah urban memberikan luasan habitat yang lebih besar terhadap Bufo melanostictus yang berasosiasi kuat dengan aktivitas manusia manusia (Iskandar, 1998 (Mistar, 2003). Spesies ini bahkan merupakan satu-satunya spesies yang dapat hidup di air payau (Gordon and Tucker, 1965 (Handayani, 2011;Kadir, 2011) sampai dengan hutan dataran tinggi di Gunung Rinjani (Iskandar, 1996). Sedangkan Oreophryne monticola yang memiliki ukuran populasi paling rendah hanya dapat ditemukan p-ISSN: 1411-9587 e-ISSN:2549-7863 di Gunung Rinjani (Iskandar, 1998) dan di Pulau Bali (Kurniati dan Hamidy, 2014 ) Kepadatan spesies dari kelas amfibi di Pulau Lombok yang memiliki kisaran dari 0.01 sampai dengan 0.30 individu/are lebih kecil dibandingkan dengan yang ditemukan di Iberian Peninsula (Schall and Pianka, 1977). ...
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Abstrak. Aspek ekologi dari amfibi dan trend populasi dari Limnonectes dammmermani yang belum banyak diketahui, disebabkan oleh kurangnya studi tentang hewan tersebut di habitatnya. Update data ukuran populasi spesies lainnya juga dibutuhkan untuk kegiatan konservasi amfibi di Pulau Lombok. Penelitian ini bertujuan untuk menganalisis densitas 12 spesies dari kelas amfibi yang ditemukan di Pulau Lombok. Survey lapangan dilakukan di 10 lokasi sampling yang tersebar di Pulau Lombok menggunakan metode Visual Encounter Survey (VES) selama bulan Maret – Juli 2016. Hasil penelitian menunjukkan bahwa Bufo melanostictus adalah spesies yang memiliki kepadatan paling tinggi (0.30 individu/are), kemudian Fejervarya cancrivora dan Limnonectes kadarsani (0.24 individu/are), sedangkan yang paling rendah adalah Oreophryne monticola (0.01 individu/are). Kata kunci: densitas, amfibi, Pulau Lombok Abstract. The ecological aspect of the amphibians and population trends of Limnonectes dammmermani are not widely known, due to the lack of studies of these animals in their habitats. Update data on other species population size is also required for amphibious conservation activities on Lombok Island. This study aims to analyze the density of 12 species of amphibian classes found on the island of Lombok. The field survey was conducted in 10 sampling sites spread on Lombok Island using Visual Encounter Survey (VES) during March - July 2016. The results showed that Bufo melanostictus was the highest density species (0.30 indi./are), then Fejervarya cancrivora and Limnonectes kadarsani (0.24 indi./are), while the lowest is Oreophryne monticola (0.01 indi./ are). Key word: density, amphibians, Lombok Island
... Manfaat dari amfibi yang besar tersebut tidak berbanding lurus dengan upaya konservasi terhadap potensi keanekaragaman hayati tersebut (Iskandar and Erdelen, 2006). Fakta ini dapat dibuktikan dengan tidak adanya spesies dari kelas amfibi yang berstatus dilindungi UU (Kusrini, 2003;Ariza dkk., 2014), padahal terdapat spesies di Pulau Lombok yang terancam karena mengalami trend penurunan populasi (Iskandar et al., 1996;Iskandar and Mumpuni, 2004). Banyak dari aspek ekologi amfibi di Indonesia belum diketahui (Iskandar and Erdelen, 2006) karena studi tentang kelompok ini masih jarang (Kusrini, 2003). ...
... Fenomena semakin meluasnya daerah urban memberikan luasan habitat yang lebih besar terhadap Bufo melanostictus yang berasosiasi kuat dengan aktivitas manusia manusia (Iskandar, 1998 (Mistar, 2003). Spesies ini bahkan merupakan satu-satunya spesies yang dapat hidup di air payau (Gordon and Tucker, 1965 (Handayani, 2011;Kadir, 2011) sampai dengan hutan dataran tinggi di Gunung Rinjani (Iskandar, 1996). Sedangkan Oreophryne monticola yang memiliki ukuran populasi paling rendah hanya dapat ditemukan p-ISSN: 1411-9587 e-ISSN:2549-7863 di Gunung Rinjani (Iskandar, 1998) dan di Pulau Bali (Kurniati dan Hamidy, 2014 ) Kepadatan spesies dari kelas amfibi di Pulau Lombok yang memiliki kisaran dari 0.01 sampai dengan 0.30 individu/are lebih kecil dibandingkan dengan yang ditemukan di Iberian Peninsula (Schall and Pianka, 1977). ...
Article
Abstrak. Aspek ekologi dari amfibi dan trend populasi dari Limnonectes dammmermani yang belum banyak diketahui, disebabkan oleh kurangnya studi tentang hewan tersebut di habitatnya. Update data ukuran populasi spesies lainnya juga dibutuhkan untuk kegiatan konservasi amfibi di Pulau Lombok. Penelitian ini bertujuan untuk menganalisis densitas 12 spesies dari kelas amfibi yang ditemukan di Pulau Lombok. Survey lapangan dilakukan di 10 lokasi sampling yang tersebar di Pulau Lombok menggunakan metode Visual Encounter Survey (VES) selama bulan Maret – Juli 2016. Hasil penelitian menunjukkan bahwa Bufo melanostictus adalah spesies yang memiliki kepadatan paling tinggi (0.30 individu/are), kemudian Fejervarya cancrivora dan Limnonectes kadarsani (0.24 individu/are), sedangkan yang paling rendah adalah Oreophryne monticola (0.01 individu/are). Kata kunci: densitas, amfibi, Pulau Lombok Abstract. The ecological aspect of the amphibians and population trends of Limnonectes dammmermani are not widely known, due to the lack of studies of these animals in their habitats. Update data on other species population size is also required for amphibious conservation activities on Lombok Island. This study aims to analyze the density of 12 species of amphibian classes found on the island of Lombok. The field survey was conducted in 10 sampling sites spread on Lombok Island using Visual Encounter Survey (VES) during March - July 2016. The results showed that Bufo melanostictus was the highest density species (0.30 indi./are), then Fejervarya cancrivora and Limnonectes kadarsani (0.24 indi./are), while the lowest is Oreophryne monticola (0.01 indi./ are). Key word: density, amphibians, Lombok Island
... Upon the discovery of this new species, the distribution of B. cynodon should be removed from the snake species lists of the Nusa Tenggara Islands. The distribution of B. hoeseli is similar to Coelognathus subradiatus, Cylindrophis opistorhodus (Boulenger, 1897), Limnonectes kadarsani (Iskandar et al.,1996), and L. dammermani (Mertens, 1929), which are restricted to the Nusa Tenggara Islands (Mertens, 1929;How and Kitchener, 1997;Iskandar et al., 1996;Colijn, 2000, 2001). ...
... Upon the discovery of this new species, the distribution of B. cynodon should be removed from the snake species lists of the Nusa Tenggara Islands. The distribution of B. hoeseli is similar to Coelognathus subradiatus, Cylindrophis opistorhodus (Boulenger, 1897), Limnonectes kadarsani (Iskandar et al.,1996), and L. dammermani (Mertens, 1929), which are restricted to the Nusa Tenggara Islands (Mertens, 1929;How and Kitchener, 1997;Iskandar et al., 1996;Colijn, 2000, 2001). ...
Article
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We describe a new cat snake species of the genus Boiga from the Nusa Tenggara Islands, Indonesia. The new species is superficially similar to Boiga cynodon, as it was identified previously. It differs from the latter species by the following combination of characteristics: only reaching half of the size of B. cynodon (up to 1250 mm SVL), higher number of dorsal scales; lower ventral and subcaudal counts and having only a very fine postorbital stripe.
... Faivovich Appendix S3. Sources for codification of character 1. Limnonectes_kadarsani * Iskandar et al. (1996) Limnonectes_kohchangae * , Limnonectes_kuhlii * , ; ; Emerson & Barrigan (1993); Limnonectes_leporinus * Limnonectes_leytensis * ; , Emerson & Barrigan (1993) Limnonectes_limborgi * Limnonectes_macrocephalus * Limnonectes_macrodon * ; Limnonectes_magnus * , Limnonectes_microdiscus * Limnonectes_microtympanum * Limnonectes_modestus * ; Limnonectes_namiyei * , Emerson & Barrigan (1993) Limnonectes_palavanensis * Smith (1927); , Laliostoma_labrosum * , Glaw & Vences (2007) Mantella_aurantiaca * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantella_baroni * Glaw & Vences (2007) Mantella_bernhardi * Glaw & Vences (2007) Mantella_betsileo * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantella_cowanii * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantella_crocea * Glaw & Vences (2007) Mantella_ebenaui * Glaw & Vences (2007) Mantella_expectata * Glaw & Vences (2007) Mantella_haraldmeieri * Glaw & Vences (2007) Mantella_laevigata * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantella_madagascariensis * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantella_manery * Glaw & Vences (2007) Mantella_milotympanum * Glaw & Vences (2007) Mantella_nigricans * Glaw & Vences (2007) Mantella_pulchra * Glaw & Vences (2007) Mantella_viridis * Glaw & Vences (2007) Mantidactylus_aerumnalis * Glaw & Vences (2007) Mantidactylus_albofrenatus * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantidactylus_ambreensis * Glaw & Vences (2007) Mantidactylus_argenteus * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantidactylus_bellyi * Glaw & Vences (2007) Mantidactylus_betsileanus * Glaw & Vences (2007) Mantidactylus_biporus * Mantidactylus_brevipalmatus * Glaw & Vences (2007) Mantidactylus_charlotteae * Glaw & Vences (2007) Mantidactylus_cowanii * Glaw & Vences (2007) Mantidactylus_curtus * Blommers-Schlösser & Blanc (1991), Glaw & Vences (2007) Mantidactylus_delormei * Glaw & Vences (2007) Mantidactylus_femoralis * Amolops_chunganensis * , Yang (1991), Amolops_compotrix * Amolops_cremnobatus * Amolops_cucae * Amolops_daiyunensis * , Amolops_daorum * Amolops_granulosus * Yang (1991) Amolops_hainanensis * Yang (1991), Amolops_hongkongensis * Yang (1991) Amolops_indoburmanensis * Amolops_iriodes * Amolops_jinjiangensis * Yang (1991) Amolops_kangtingensis Amolops_manztorum , Yang (1991) Amolops_larutensis * Yang (1991) Amolops_liangshanensis * Amolops_lifanensis * Yang (1991) Amolops_loloensis * Yang (1991) Amolops_mantzorum * , Yang (1991) Amolops_marmoratus * Boulenger (1882) , Babina_subaspera * Clinotarsus_alticola * , Clinotarsus_curtipes * Boulenger (1882), , , Huia_cavitympanum * , , Yang (1991), Huia_masonii * Yang ( ...
Article
Cascades and fast-flowing streams impose severe restrictions on acoustic communication, with loud broadband background noise hampering signal detection and recognition. In this context, diverse behavioural features, such as ultrasound production and visual displays, have arisen in the evolutionary history of torrent-dwelling amphibians. The importance of the vocal sac in multimodal communication is being increasingly recognized, and recently a new vocal sac visual display has been discovered: unilateral inflation of paired vocal sacs. In the diurnal stream-breeding Hylodidae from the Atlantic forest, where it was first described, this behaviour is likely to be enabled by a unique anatomical configuration of the vocal sacs. To assess whether other taxa share this exceptional structure, we surveyed torrent-dwelling species with paired vocal sacs across the anuran tree of life and examined the vocal sac anatomy of exemplar species across 18 families. We found striking anatomical convergence among hylodids and species of the distantly related basal ranid genera Staurois, Huia, Meristogenys and Amolops. Ancestral character state reconstruction identified three new synapomorphies for Ranidae. Furthermore, we surveyed the vocal sac configuration of other anuran species that perform visual displays and report observations on what appears to be unilateral inflation of paired vocal sacs, in Staurois guttatus-an extremely rare behaviour in anurans.
... Information for the rest of the archipelago is comparatively thinner, despite the efforts of various researchers over the course of the last one hundred years. Recent publications, de Lang's (2011) synthesis of the snakes and the results of the Western Australian Museum/Museum Zoologicum Bogoriense expeditions conducted during 1987-1993(e.g., How et al. 1996a, 1996b1998;How and Kitchener 1997), provide the most modern overview of the herpetofauna, and supported by Merten's works during early to mid-1900s (e.g., Mertens 1927a1927b;1928;1957), other species-specific or taxonomic snippets (e.g., Das 1993;Iskandar et al.1996;Wüster 1996), and the baseline data of seminal publications, such as de Rooij (1915;1917), Boulenger (1897) and van Kampen (1923), this forms the body of our herpeto-zoogeographical knowledge for the Lesser Sundas. ...
... Information for the rest of the archipelago is comparatively thinner, despite the efforts of various researchers over the course of the last one hundred years. Recent publications, de Lang's (2011) synthesis of the snakes and the results of the Western Australian Museum/Museum Zoologicum Bogoriense expeditions conducted during 1987-1993(e.g., How et al. 1996a, 1996b1998;How and Kitchener 1997), provide the most modern overview of the herpetofauna, and supported by Merten's works during early to mid-1900s (e.g., Mertens 1927a1927b;1928;1957), other species-specific or taxonomic snippets (e.g., Das 1993;Iskandar et al.1996;Wüster 1996), and the baseline data of seminal publications, such as de Rooij (1915;1917), Boulenger (1897) and van Kampen (1923), this forms the body of our herpeto-zoogeographical knowledge for the Lesser Sundas. ...
... Information for the rest of the archipelago is comparatively thinner, despite the efforts of various researchers over the course of the last one hundred years. Recent publications, de Lang's (2011) synthesis of the snakes and the results of the Western Australian Museum/Museum Zoologicum Bogoriense expeditions conducted during 1987-1993(e.g., How et al. 1996a, 1996b1998;How and Kitchener 1997), provide the most modern overview of the herpetofauna, and supported by Merten's works during early to mid-1900s (e.g., Mertens 1927a1927b;1928;1957), other species-specific or taxonomic snippets (e.g., Das 1993;Iskandar et al.1996;Wüster 1996), and the baseline data of seminal publications, such as de Rooij (1915;1917), Boulenger (1897) and van Kampen (1923), this forms the body of our herpeto-zoogeographical knowledge for the Lesser Sundas. ...
... Within this clade there is full support (both analyses) for sister rela- tionships between L. gyldenstolpei and L. dabanus, and between L. plicatellus and L. macrognathus. The caruncule-bearing Limnonectes are sister to the clade containing Limnonectes microdiscus (Boettger, 1892), Limnonectes kadarsani Iskandar, Boeadi & Sancoyo, 1996, andLimnonectes laticeps (Boulenger, 1882). Results of the BA present these two clades as recip- rocally monophyletic, but in the ML analysis, the clade containing L. microdiscus and L. kadarsani is sister to L. laticeps, which is sister to the caruncular Limnonectes. ...
Article
The males of four species of the Asian frog genus Limnonectes [Limnonectes dabanus (Smith, 1922a), Limnonectes gyldenstolpei (Andersson, 1916), Limnonectes macrognathus (Boulenger, 1917), and Limnonectes plicatellus (Stoliczka, 1873)] exhibit remarkable ornamentation in the form of a swollen, or cap-like, structure (caruncle) on the top of their heads. These caruncles vary in their appearance among species, and neither their function nor their actual systematic value is known. We compared their anatomy via dissections, morphometrics, radiography, and histology, and analysed the available mitochondrial DNA sequences as well as new data to place these species within the context of a larger phylogenetic hypothesis for Limnonectes. Despite the externally different morphology, the underlying histological structure is virtually identical. Beneath skin that is densely packed with mucous glands lies a pad of connective tissue overlaying the parietal bone. The actual function of the caruncle, however, remains enigmatic. In addition to the presence of the caruncle, independent evidence from osteological characters and molecular data support the monophyly of a clade comprising of L. dabanus, L. gyldenstolpei, L. macrognathus, and L. plicatellus. The caruncles are therefore interpreted as a robust autapomorphy for this clade, and suggest that the subgenus Elachyglossa should be restricted to the four species in question. © 2014 The Linnean Society of London
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Une nouvelle classification provisoire de la superfamille des Ranoidea, et plus particulierement de la famille des Ranidae et du genre Rana, est proposee. Le genre Rana est ici subdivise, principalement sur la base de criteres phenetiques, en trente-trois groupes, auxquels est ici provisoirement attribue le rang de sous-genres. Le but principal de cette classification provisoire, qui doit etre comprise comme un outil de travail et non pas comme une proposition definitive, est de servir de guide pour de futures etudes sur la phylogenie de cette famille tres vaste et a repartition quasi cosmopolite : cette classification aidera a selectionner des especes representatives de chaque groupe phenetique pour l'etude des etats de caracteres, et a choisir les hors-groupes appropries pour l'etablissement de la polarite des morphoclines de caracteres. Ce n'est que lorsque la phylogenie de la famille dans son ensemble sera resolue qu'il sera possible d'elaborer une classification plus stable, qui s'averera peut-etre tres differente de celle qui est presentee ci-dessous.
Results of the Douglas Burden expedition to the Island of Komodo. IV. Frogs from the East Indies
  • E R Dunn
Dunn, E.R., 1927. Results of the Douglas Burden expedition to the Island of Komodo. IV. Frogs from the East Indies., Amer. Mus. Novit., 315: 9 pp.
Herpetologische Mitteilungen. xxv. Zur Kenntnis der Rana microdisca Boettger und ihrer Rassen
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Mertens, R., 1929. Herpetologische Mitteilungen. xxv. Zur Kenntnis der Rana microdisca Boettger und ihrer Rassen. Zool. Anz., 86: 66-68.
The Amphibians of the Indo-Australian Archipelago
  • P N Van Kampen
  • E J Amsterdam
  • Brill
Van Kampen, P.N., 1923. The Amphibians of the Indo-Australian Archipelago. Amsterdam, E.J. Brill. 304 pp. Received 27 Apr 1995 Accepted 25 Oct 1995
Neue Amphibien und Reptilien aus dem Indo-Australischen Archipel
  • R Mertens
Mertens, R., 1927. Neue Amphibien und Reptilien aus dem Indo-Australischen Archipel.
Amphibien des indischen Archipels
  • P N Van Kampen
Van Kampen, P.N., 1907. Amphibien des indischen Archipels. in M. Weber, Zool. Erg. Niederlandisch Ost-Indien, 4 (2): 383-418, pI. 16.
The nomenclature and identity of the javanese frog Rana macrodon Dumeril and Bibron
  • B H Kiew
Kiew, B.H., 1978. The nomenclature and identity of the javanese frog Rana macrodon Dumeril and Bibron. Malayan Nat., 31 (4): 219-229.