Zootaxa, A new species of Litoria (Amphibia: Anura: Hylidae)

Zootaxa 2277: 1–13 (2009)
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ISSN 1175-5326 (print edition)
Article
Copyright © 2009 · Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
A new species of Litoria (Amphibia: Anura: Hylidae) from the foothills of the
Foja Mountains, Papua Province, Indonesia
STEPHEN J. RICHARDS1,5, PAUL M. OLIVER1,2, KELIOPAS KREY3 & BURHAN TJATURADI4
1
Terrestrial Vertebrates, South Australian Museum, North Terrace, Adelaide, S.A. 5000, Australia, and Conservation International,
Atherton, Qld 4883, Australia
2
Australian Centre for Evolutionary Biology and Biodiversity, Adelaide University
3
Department of Biology, Unversity of Manokwari, Papua, Indonesia
4
Komp Ariau Dunlop, Sentani, Papua, Indonesia.
5
Corresponding author. E-mail: [email protected]
Abstract
Litoria gasconi sp. nov. is described from low, forest-covered ridges on the southern edge of the Foja Mountains, Papua
Province, Indonesia. It is most similar to Litoria multiplica (Tyler, 1964) but can be differentiated from that species and
all other described Litoria by a unique combination of characters including moderate size (males 39.3–41.6 SVL), green
dorsum with yellow spots in life, relatively large eyes (EYE/SVL 0.12–0.15), dermal ridges below the vent and on the
posterior edge of both fore and hindlimbs, complete absence of blue thigh and lateral colouration, and its unique
advertisement call consisting of a single soft, distinctly pulsed chirp. New data on the morphology and ecology of the
superficially similar and poorly known species Litoria multiplica are also presented. Recent surveys in the Foja
Mountains have revealed a diverse frog fauna with numerous unrecognised or poorly known taxa; these ranges are likely
to be a previously unrecognised and largely unexplored centre of tropical vertebrate endemism.
Key words: Litoria, Indonesia, New Guinea, new species, Papua
Introduction
The Foja Mountains are a series of steep rainforest-covered peaks in northern Papua Province, Indonesian
New Guinea. Isolated from New Guinea’s central cordillera by the extensive lowland forests and swamps of
the Mamberamo Basin, the rugged, inaccessible and uninhabited slopes of the Fojas remain one of the most
poorly explored tropical regions on earth (Diamond 1982). In November 2005 a Rapid Assessment Program
biological survey by Conservation International and the Indonesian Institute of Sciences (LIPI) in this isolated
range revealed a wealth of undescribed taxa including the first new bird species discovered in New Guinea for
nearly 70 years (Beehler et al. 2007). Prominent amongst the new taxa were a large number of frog species
new to science.
One of the most distinctive new frogs was a moderately sized, predominantly green treefrog of the genus
Litoria that was found at two sites on the lower slopes of the range. The most similar described species is
Litoria multiplica (Tyler, 1964), a poorly known taxon from moderate altitudes on the slopes of the central
cordillera in Papua New Guinea. In order to adequately compare the new Litoria from the Fojas with L.
multiplica we gathered new data on the latter species based on pre-existing museum specimens, and on new
collections made by the senior author. Presented herein is a formal taxonomic description of the new species
from the Foja Mountains and new data on the morphology and ecology of Litoria multiplica.
Accepted by S. Castroviejo: 7 Oct. 2009; published: 30 Oct. 2009
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Materials and methods
Frogs were collected at night through audio-location of calling males by B. Tjaturadi and S. J. Richards.
Specimens were photographed before being fixed in 10% formalin and stored in 70% ethanol. For some
specimens liver samples were taken and stored in 70% ethanol. Collections were made in the foothills of the
Foja Mountain Range near Marina Valen Village in July 2004 and around Kwerba Village in November 2005.
Preserved animals are lodged in the Museum Zoologicum Bogoriense (MZB) and tissues are stored at the
MZB and the South Australian Museum (SAMA).
Measurements (to the nearest 0.1mm) were taken with dial calipers and a stereomicroscope fitted with an
ocular micrometer, and follow Richards et al. (2006). Measurements taken are SVL (snout-vent length), TL
(tibia length), HW (head width at tympanum), HL (head length from tip of snout to posterior edge of
tympanum), EYE (horizontal eye diameter), TYM (horizontal tympanum diameter), IN (inter-narial distance),
EN (distance between anterior edge of eye and posterior edge of naris), 3FD (widest transverse diameter of 3rd
finger disc), 3FP (widest transverse diameter of penultimate phalanx), 4TD (widest transverse diameter of 4th
toe disc) and 4TP (widest transverse diameter penultimate phalanx).
Most advertisement calls were recorded with a Sony TCM-5000 tape recorder and a Sennheiser ME66
microphone; several L. multiplica calls were recorded with a Sony Pro-Walkman WMD-6C tape recorder and
Sony ECM-Z200 microphone. Thirty one calls of the new species and 35 calls of L. multiplica were analyzed
using the AVISOFT SAS-Lab Pro sound analysis program. Air temperatures adjacent to calling males were
measured with a Miller & Weber quick-reading thermometer and all coordinates presented were taken using
the GPS datum WGS 84.
Comparative material from the following institutions was examined; the South Australian Museum,
Australia (SAMA), the Natural Sciences Resource Centre of the University of Papua New Guinea, Port
Moresby (UPNG), the Natural History Museum, London (BM), Museum Zoologicum Bogoriense, Cibinong,
Indonesia (MZB), Naturalis (formerly Rijksmuseum van Natuurlijke), Leiden (RMNH), and the Queensland
Museum, Brisbane (QM). Field numbers (FN) for field collections by SJ Richards (SJR) are also presented for
cross-reference.
Systematics
Litoria gasconi new species
(Figs. 1A–D, 2B, 3)
Holotype. MZB Amph. 15839 (FN SJR 6018), adult male, calling at time of collection (16 July 2004), tissue
preserved in 70% ethanol, camp near Marina Valen Village (02o22.230'S, 138o12.753'E), 500 m a.s.l., southern
foothills of the Foja Mountains, Papua Province, Indonesia, collected by Stephen Richards and Burhan
Tjaturadi.
Paratopotype. Male, calling when collected, MZB Amph. 15840 (FN SJR 6019), with same date, locality
and collectors as holotype.
Paratype. Male, calling when collected, MZB Amph. 12036 (FN SJR 9805), 17 November 2005,
'Kwerba Camp' (2o38.467' S, 138o24.916'E), 200 m a.s.l., near Kwerba Village, southern foothills of the Foja
Mountains, Papua Province, Indonesia, collected by Stephen Richards and Burhan Tjaturadi.
Diagnosis. Litoria gasconi sp. nov. can be distinguished from all other Litoria by the combination of
moderate size (males 39.3–41.6 mm), predominately green dorsal colouration interspersed with numerous
yellow spots (light blue in preservative), white bilobed dermal ridge below the vent, and further ridges
extending from heel to fifth toe and on the forearms, large eyes (EYE/SVL 0.12–0.15), thighs, groin and
axillary spots orange in life without any trace of blue, absence of black vermiculations on thigh and venter,
absence of a canthal stripe between the eye and nares, and advertisement call consisting of a single soft,
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distinctly pulsed chirp containing 4–6 pulses and lasting 0.02–0.03 s.
Description of holotype. Adult male with vocal slits and calling when collected (with measurements
given in Table 1). Body moderately slender, limbs moderately short (TL/SVL 0.56), head wider than body in
dorsal profile, distinct from neck (HW/SVL 0.36). Snout rounded in dorsal profile, slightly rounded but nearly
truncate in lateral profile, upper jaw protruding marginally over lower jaw. Canthus rostralis slightly curved,
not sharply defined, loreal region slightly concave, nares oriented anterio-laterally, closer to tip of snout than
to eyes, labial region flared. Eyes large (EYE/SVL 0.15), prominent, clearly protruding in lateral and dorsal
views, pupil horizontal. Tympanum clearly visible, small (TYM/SVL 0.06), approximately half diameter of
eye (TYM/EYE 0.44), annulus distinct and bordered dorsally by supratympanic fold that runs from posterior
corner of eye and terminates above axilla. Choanae small and circular, close to lateral edge of palate;
vomerine teeth in two moderate-sized clumps medial to choanae; tongue fleshy and ovoid. Dorsal skin finely
granular; ventral skin smooth on throat, coarsely granular on abdomen; a fleshy bilobed dermal ridge below
vent, and dermal ridges along outside edge of tarsus and forearm.
TABLE 1. Measurements (in mm) for the type series of Litoria gasconi sp. nov.
MZB 15839 (SJR 6018)
MZB 15840 (SJR 6019)
MZB 12036 (SJR 9805)
Holotype
Paratopotype
Paratype
M
M
M
SVL
39.3
40.7
41.6
TL
21.9
22.6
23.4
HW
14.2
13.9
15.2
HL
13.8
13.2
12.7
EYE
5.7
5.7
4.7
TYM
2.5
2.4
2.4
IN
3.9
4.1
4.2
EN
2.5
3.1
3.7
3FD
2.4
2.3
2.3
3FP
1.7
1.7
1.6
4TD
2.0
2.0
1.9
4TP
1.6
1.6
1.4
Fingers moderately long in comparison to other Papuan Litoria (Tyler 1968); with relative lengths
III>IV>II>I; fleshy opaque webbing between all digits: in a narrow strip between I and II, and reaching to
penultimate phalanx on outer edges of II and III, to third phalanx on inner edge of III, and to disc on IV.
Terminal discs on all fingers prominent (3FP/3FD 0.71), with circum-marginal grooves. Nuptial pads without
pigmentation and very indistinct; single indistinct unpigmented bifid subarticular tubercles present on
penultimate phalanx of all fingers, and a series of additional subarticular tubercles present on IV; single
indistinct ovoid inner metacarpal tubercle present at base of I.
Toes long in comparison to other Papuan Litoria (Tyler 1968); relative lengths IV>III>V>II>I; with
extensive fleshy opaque webbing. Web extends to discs on outer edge of II, III and inner edge of V, to
penultimate phalanx on both sides of IV and inner edge of II and III, and to penultimate tubercle on I; dermal
flanges extending to disc on III and IV. Discs prominent (4TP/4TD 0.80) with circum-marginal grooves;
single rounded subarticular tubercles present on all digits, small ovoid metatarsal tubercle at base of toe I,
series of distinct rounded ventral tubercles of varying sizes present on proximal third of femur.
In preservative dorsal ground colour of head, body and legs slate blue with extensive randomly scattered
sky-blue spots; dorsal surfaces of fingers, and lower portions of forelimbs and toes largely unpigmented,
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though sometimes with small patches of blue pigment, especially on toes. Lateral surfaces slate blue heavily
flecked with bluish white, lateral surfaces of head distinctly lighter than dorsal surfaces of head, edge of upper
jaw with a continuous white margin from rictus to rictus, orbital bordered by a thin white ring. Ventral
surfaces of body and forelimbs off-white, hind limbs slightly darker with poorly defined brown stripe along
posterior edge and on to toe V; prominent dermal flange below vent grades from slate blue dorsally to white
ventrally, to bordered at ventral edge by a poorly defined brown stripe.
FIGURE 1. Dorsal (A) and ventral (B) views of holotype MZB Amph. 15839 of Litoria gasconi sp. nov. (scale = 10
mm), (C) lateral view of head of holotype MZB Amph. 15839 (scale = 5 mm), (D) paratype MZB amph 12036 (SJR
9805) in life showing bilobed dermal fold below vent and orange thigh and axillary colourations.
Variation. All types are very similar in overall proportions to the holotype; summary meristic data is
given in Table 1 and key ratios are given in Table 2. Paratopotype MZB Amph. 15840 (SJR 6019) is missing
most of the second finger on the left hand. Colouration in preservative is highly consistent, the dorsal surfaces
are always slate blue with extensive sky blue spotting, ventral and hidden surfaces are off-white except for
thin brown stripes below the venter and along the posterior ventral edge of the tarsus, and all specimens also
possess a distinctive white dermal flange below the vent, and a further white lateral dermal flange extending
from the heel to the fifth toe. All types are males that were calling when collected, but lack obvious pigmented
nuptial pads and further collecting may reveal this to be an additional diagnostic character for the species.
Appearance in life. Typical colouration in life is shown in Figure 2. The following description is based
on colour photographs of all three type specimens. Colour pattern is highly consistent: dorsum mottled bright
leaf green, densely flecked with small indistinct tiny yellowish dots, and much larger, scattered and
indistinctly edged yellow spots of varying size; lateral surfaces grading from heavily mottled with green and
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yellow dorsolaterally, to mottled green and white ventrolaterally; venter white. Small amount of brown
blotching present in subocular region of paratopotype MZB Amph. 15840 (SJR 6019). Exposed dorsal and
lateral surfaces of arms and legs mottled green and yellow, with scattered large yellow spots as on dorsum;
hidden parts of groin, thighs and axillary region bright orange. Toes and fingers predominately white with
some scattered green patches on dorsal surface of outer digits. Crenulated dermal flanges on arm and tarsus
white; bilobed dermal ridge and scattered tubercles below vent and thighs white. Iris white with fine brown
vermiculations, pupil horizontal.
TABLE 2. Average and range (in parentheses) of key measurements (in mm) and proportions for Litoria gasconi sp.
nov. and Litoria multplica.
L. gasconi sp. nov.
L. multiplica (male)
L. multiplica (female)
n=3
n = 22
n=8
SVL
40.5 (39.3–41.6)
36.4 (31.1–38.9)
41.9 (37.65–47.1)
TL
22.6 (21.9–23.4)
20.3 (16.4–21.9)
24.0 (20.4–27.4)
HL
13.2 (12.7–13.8)
11.3 (9.5–12.8)
12.3 (10.2–14.3)
HW
14.5 (13.9–15.2)
12.1 (9.8–14.0)
13.5 (11.7–15.9)
EYE
5.5 (5.0–5.7)
3.9 (3.3–4.4)
3.9 (3.4–4.2)
HW/SVL
0.36 (0.34–0.37)
0.33 (0.30–0.37)
0.32 (0.29–0.34)
HL/SVL
0.33 (0.31–0.35)
0.31 (0.29–0.33)
0.29 (0.27–0.32)
TL/SVL
0.56 (0.56–0.56)
0.56 (0.53–0.58)
0.57 (0.54–0.62)
EYE/SVL
0.13 (0.12–0.15)
0.11 (0.09–0.13)
0.09 (0.08–0.10)
Advertisement call. Thirty-one calls produced by paratype MZB Amph. 15840 (SJR 6019) were
recorded and analysed. The holotype and paratype MZB Amph. 12036 (SJR 9805) produced calls that were
indistinguishable to the ear, but recordings were of insufficient quality for detailed analysis. L. gasconi sp.
nov. produced two distinct call types, herein referred to as ‘short’ calls and ‘long’ calls. Twenty-seven of the
31 calls (87.1%) are ‘short calls’, and 4 are ‘long’ calls. ‘Short’ calls were produced on average every 9.6 s (n
= 27, range = 4.3–23.7 s, SD = 3.72) and they were all structurally similar, consisting of a single sharp chirp
lasting just 0.019–0.030 s (n = 27, mean = 0.024 s, SD = 0.003) and having a dominant frequency of 1520–
1890 Hz (n = 27, mean = 1750 Hz, SD = 67). Pulsed structure within these extremely short calls was not
evident in the field, but analyses revealed that each call contains 4–6 pulses (n = 27, mean = 5.3, SD = 0.6)
produced at a rate of 153.3–216.2/s (n = 27, mean = 200.5/s, SD = 12.9).
‘Long’ calls were produced less frequently than ‘short’ calls and were usually uttered in response to the
call of a nearby male, suggesting a territorial rather than a mate attracting function. Length of ‘long’ calls was
extremely variable (0.046–0.146 s, SD = 0.05, n = 4) but 3 of these calls had pulse rates within the range of
short calls (i.e. 187–193/s) indicating that the frogs are simply adding pulses to their calls at the same rate to
produce ‘territorial’ calls. A consequence of their longer duration is that in the field these calls sounded
distinctly (albeit finely) pulsed to the ear. One ‘long’ call produced by the holotype had a much slower pulse
rate than all other calls (120/s) and the highest number of pulses (18). Similar calls were heard in the field,
again predominantly in response to calls of nearby males, and were often produced in couplets or triplets
producing a soft ‘bleating’ effect. Typical ‘short’ and ‘long’ calls are illustrated in Figure 3.
Comparisons. The combination of medium size (39.3–41.6 mm), green dorsal colouration with large
yellow spots, and white dermal ridges on the forelimbs, hindlimbs and below the vent will readily distinguish
Litoria gasconi sp. nov. from all Litoria except L. multiplica. Litoria gasconi differs from Litoria multiplica
in lacking extensive dark markings on the groin, venter and lateral surfaces, in having bright orange groin,
thigh and axillary colouration (as opposed to bright blue, or orange mixed with bright blue, see Figure 2), in
being on average slightly larger, and in having proportionally larger eyes (see summary of meristic and
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FIGURE 2. Paratopotype of Litoria gasconi sp. nov. MZB Amph. 15840 (SJR 6019) (top) photographed in life at night,
note distinct dermal folds on posterior edge of forearm, and Litoria multiplica (bottom) Tualapa, Southern Highlands
Province, Papua New Guinea, in life, note black ventral patches and blue groin and thighs. All photographs by S. J.
Richards.
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FIGURE 3. Wave form (top) and spectrogram (bottom) of A) advertisement call of Litoria gasconi sp. nov. (MZB
Amph. 15840), B) ‘territorial’ call of Litoria gasconi sp. nov. (MZB Amph. 15840), both recorded at an air temperature
of 23 oC, and C) two components of the biphasic 'Type 1' advertisement call of Litoria multiplica (SAMA R64644)
recorded at an air temperature of 20.5 oC.
FIGURE 4. Primary foothill rainforest around Marina Valen Village, Papua Province Indonesia, type locality of Litoria
gasconi sp. nov.
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mensural comparisons in Table 2). The advertisement call of L. gasconi sp. nov. is also distinctly different,
consisting of a single short but distinctly pulsed note (see ‘advertisement call’ above), while that of L.
multiplica consists of a very loud 2-note call with a pulse rate that is much slower than that of L. gasconi sp.
nov. (30–47/s vs 153–216/s). A brief description of the call of L. multiplica is provided below.
There are a number of other predominately green, medium-sized Litoria found in New Guinea that could
be superficially confused with L. gasconi sp. nov. Litoria wapogaensis and L. christianbergmanni share with
L. gasconi sp. nov. distinct white dermal ridges below the vent and extending along the arms and legs.
However these species are much smaller (26.9 to 32.9 mm), and the former can be further distinguished by its
purplish brown thighs and the latter by its dark brown thighs and golden iris (see Günther 2008). Litoria
elkeae and other members of the L. gracilenta and L. aruensis group (sensu Menzies and Tyler 2004) are of
similar size, although generally slightly smaller (Menzies and Tyler 2004) and can be spotted, but are readily
distinguished by possession of a pale canthal stripe and absence of a prominent white dermal ridge around the
vent and on the fore and hindlimbs. Litoria singadanae can be distinguished by it’s transparent tympanic
membrane (Richards 2005), Litoria verae by its more extensively developed crenulated fold on the outer edge
of the limbs, greenish brown and finely tubercular (vs smooth) dorsum (Günther 2004), and Litoria rubrops
by its red eye and lack of a dermal fold below the vent (Kraus and Allison 2004). Frogs of the Litoria
graminea group (Litoria dux, Litoria graminea, Litoria huntorum and Litoria sauroni) are predominately
green but are easily distinguished by their much larger size (male SVL > 55 mm), and lack of both dorsal
spotting and a bilobed dermal ridge below the vent (Richards et al. 2006, Richards and Oliver 2006). Litoria
mystax, L. albolabris, L. umarensis and New Guinean members of the Litoria bicolor group (L. bibonius, L.
contrastens, L. chloristona, L. eurynastes, L. lodesdema, and L. viranula), are predominantly green but are
much smaller than the new species (male SVL < 35 mm) and again lack a bilobed dermal ridge below the vent
(Tyler 1968, Gunther 2004, Menzies et al. 2008).
Distribution and Natural History. Known only from two sites in the vicinity of Marina Valen and
Kwerba Villages, in the southern foothills of the Foja Mountain Range, Papua Province, Indonesia (Figure 4).
All specimens were collected in relatively undisturbed hill forest on steep ridges between 200 and 500 m a.s.l.
(Figure 5). Litoria gasconi sp. nov. was not found at higher elevations despite intensive searches between
1100 m and 1700 m over a 2-week period, and further searches at similar altitudes over a three week period in
2008. Males were detected by their very soft calls, uttered from large leaves on low shrubs or tree branches
between 1 and 6 m above the ground. They were found only along two streams that formed a series of
shallow, disconnected pools and seeps. This species was never seen or heard in the vicinity of clear, flowing
streams, or around temporary or permanent forest pools. Two other species of Litoria, L. humboldtorum and L
c.f. genimaculata, were detected in microsympatry along the same intermittent streams.
Etymology. The new species is named for Claude Gascon of Conservation International, in gratitude for
his support of our amphibian research in New Guinea, and in recognition of his long involvement in the
conservation of amphibians globally.
New data for Litoria multiplica (Tyler 1964)
Litoria gasconi sp. nov. is most similar morphologically to Litoria multiplica, a poorly known species from
lower montane forests along the southern side of the central cordillera in Papua New Guinea. In the process of
comparing these two forms we examined a large series of L. multiplica, including new specimens obtained by
the senior author during recent surveys in Papua New Guinea. Here we present new data for this taxon based
on recently collected and photographed specimens from the Crater Mountain Wildlife Management area (near
the type locality) and Tualapa in Southern Highlands Province, and on re-examination of older material held
in the South Australian Museum (see Appendix 1 for specimen and locality details).
Most specimens examined conform to the descriptions provided by Tyler (1964) and Tyler (1968). All
possess a prominent white bilobed dermal ridge around the vent, and white dermal ridges around the knees
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RICHARDS ET AL.
and along the outer edge of the tarsus. In combination with moderate size (males 31.1–38.9 mm) these dermal
ridges diagnose this species from almost all other Melanesian Litoria. The posterior-ventral edge of the femur
is also moderately to heavily tuberculate. All preserved specimens have extensive dark bluish-purple
blotching on the lateral surfaces, but the amount of pigmentation on the venter varies from almost plain to
extensively spotted or mottled with dark blueish-purple. All specimens show at least some degree of bluishgreen spotting on the dorsum in preservative. Of the sixteen males examined, none had pigmented nuptial
pads, including a number that were calling when collected. Close examination indicated that these specimens
did have unpigmented granular rugosities at the base of the inner finger. Unpigmented nuptial pads have not
previously been noted for this taxon and they provide an additional character to distinguish L. multiplica from
most other green Litoria. Key meristic and comparative data for Litoria multiplica and Litoria gasconi sp.
nov. are presented in Table 2.
FIGURE 5. Map of New Guinea showing the known distribution of Litoria gasconi sp. nov. and Litoria multiplica.
Based on photographs in life of two specimens from Tualapa (SJR 10631–32) (Figure 2), the dorsum is
uniformly green with scattered small pale yellowish-green spots; venter off-white, with or without extensive
dark purplish-black blotching; dark purplish black lateral spots or mottling present on all specimens,
sometimes also extending along hidden surface of legs. All specimens have prominent blue patches on
posterior lateral half of torso, on the inner and outer thighs, and sometimes extending to the distal edge of
tibia. The lower arms, axillary spots, inner webbing and toes (to about digit IV) and sometimes hidden
surfaces of lower legs are flushed with orange; fingers and outer toes are unpigmented. Prominent white
dermal flanges are clearly apparent below the vent and on the heels. The iris is off white with scattered very
fine brown vermiculations.
An unusually dark specimen from Magidobo (SAMA R R34360) in Southern Highlands Province is
currently referred to Litoria multiplica and resembles this species in having a bilobed dermal flap and clear
evidence of dark purplish colouration on thigh and lateral surfaces. However it lacks any obvious sign of
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dorsal blue spots, has a distinctive white marking along the posterior-ventral edge of the eye, and shows
comparatively little evidence of a dermal flange on the tarsus. Further specimens from this area need to be
examined, to assess whether this specimen is conspecific with Litoria multiplica.
We analysed 35 calls of three Litoria multiplica individuals from Maimafu Village, Crater Mountain
Wildlife Management Area, Eastern Highlands Province (6o30.082’ S, 145o01.977’ E, 1540 m a.s.l.) in 1997
and from Tualapa, Southern Highlands Province (05o18.245’ S, 142o30.704’ E, 1438 m a.s.l.) in 2008. Calls
from the three animals were consistent so they are combined in the description below. Like L. gasconi sp. nov.
L. multiplica produces two distinct call types. Type 1 is a very loud and ‘biphasic’ call consisting of 1) a series
of 6–14 (mean = 7.75, SD = 2.05, n = 16) harsh pulses lasting 0.16–0.28 s (mean = 0.19 s, SD = 0.03, n = 16)
and produced at a rate of 30.52–47.61/s (mean = 36.30 s, SD = 5.06, n = 16), followed by 2) a single sharp,
unpulsed ‘click’ lasting 0.012–0.023 s (mean = 0.017 s, SD = 0.003, n = 16). The click is separated from the
preceding pulses by an interval of 0.20–0.29 s (mean = 0.22 s, SD = 0.02, n = 16), which is approximately the
length of the entire first phase. The final pulse in the first ‘phase’ of the biphasic calls is frequently subdivided
into two ‘sub-pulses’, or nearly entirely divided into two immediately adjacent pulses. A typical biphasic call
is illustrated in Figure 3. Type 2 calls are a series of pulses without a terminal ‘click’. Unlike Type 1 calls,
which have an extremely consistent call structure among specimens and sites (see above), Type 2 calls are of
extremely variable length, pulse number and pulse rate. Individual calls contain 4–49 pulses and last 0.04–
0.63 s (mean = 0.21 s, SD = 0.14, n = 19). Pulse rate is also extremely variable, ranging from 28–128/s (mean
= 92.43 s, SD = 29.7, n = 19). However only one of 19 Type 2 calls (5.3%) had a pulse rate slower than Type
1 calls, and 10 (53%) had pulse rates above 100/s (vs 30.5–47.6). Our impression was that males produced
‘Type 2’ calls in response to calls of nearby males and that, like males of L. gasconi sp. nov., they were adding
pulses to calls in response to nearby males. However unlike L. gasconi sp. nov. which added pulses at about
the same rate so that call length of territorial calls increased but pulse rate stayed approximately the same (see
above), L. multiplica frequently increased the number of pulses per call by as many as 3 times without
changing call length. As a result pulse rates for many Type 2 calls (n = 9 or 47%) were more than 100/s
compared to 35–47/s for Type 1 calls of similar length.
To date almost nothing has been published about the ecology of Litoria multiplica. Field observations of
specimens from Maimafu Village and at Tualapa, revealed that males aggregate and form breeding choruses at
moderately high densities (up to 1 frog every 10 m) along clear, slow-flowing streams in rainforest where they
call from low vegetation (~ 1–3 m high) overhanging the stream bed. Preferred streams are less than 5 m wide
and typically have numerous deep pools separated by shallow riffles. Egg deposition sites remain unknown.
No eggs were found attached to overhanging vegetation along streams with large L. multiplica populations
despite intensive searches at two localities, but the suggestion by Menzies (2006) that this species may lay
arboreal eggs like members of the L. iris group cannot be discounted. Known elevations for L. multiplica
range between ~1000 and 1540 m a.s.l., and it is apparently widespread in lower montane forests of Papua
New Guinea along the central cordillera, between Eastern Highlands Province in the east and the Papua
border in the west (Figure 4,Tyler 1968, Menzies 2006).
Discussion
Based on their numerous shared characteristics, including similar size, dorsal colouration and pattern, distinct
bilobed dermal ridge around the vent, and possibly unpigmented nuptial pads, we suggest that Litoria gasconi
sp. nov. and Litoria multiplica are closely related. Unpublished molecular data also supports this assertion
(Price and Donnellan pers com). However the relationship of these two taxa to other New Guinea hylids is
uncertain. Menzies and Tyler (2004) placed Litoria multiplica into the Litoria chloris species group
containing L. chloris, L. graminea and L. xanthomera. Although these taxa do show some superficial
morphological similarities, there are also substantial morphological differences between them, and
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RICHARDS ET AL.
meaningful discussion of their systematic relationships will have to await a more taxon-intensive multilocus
phylogeny for Australopapuan hylids.
The Foja Mountains are isolated from other upland areas of New Guinea, and recent surveys suggest that
they hold a number of endemic taxa (Beehler et al. 2007, Richards, Helgen and Oliver pers obs.). Litoria
gasconi sp. nov. is currently only known from the lowest slopes of the Fojas, and the vast surrounding
lowlands of the Mamberamo basin have been relatively poorly surveyed. However the few surveys conducted
at other sites in the Mamberamo lowlands failed to find this species (e.g. Richards et al. 2002). Known sites
for L. gasconi sp. nov. around Kwerba and Marina Valen are also on steep ridges with shallow seepages, and
the species was never observed in the numerous streams or pools sampled in flat lowland habitats, further
indicating the species may not occur in the lowland swampy habitats typical of most of the Mamberamo Basin
lowlands.
The Foja Mountains are one of a series of isolated ranges that rise out of the extensive lowlands of
northern New Guinea (e.g. Cyclops, Bewanis, Torricellis). Due largely to geographic proximity these ranges
have often been discussed as a single biogeographic unit (Stattersfield et al. 1998, Helgen 2005) and several
taxa show distributions consistent with this biogeographic association (e.g. Dendrolagus pulcherrimus
(Helgen and Richards pers obs. 2005) and Xenorhina arboricola (Allison and Kraus 2000, Richards and
Tjaturadi pers. obs.). However, based on the number of putative endemic taxa, Beehler et al. (2007) argued
that the Fojas might be relatively highly differentiated from other north coast ranges. The large size of the
Fojas, their long isolation, and their intermediate geographic position among north-coast ranges (the Arfak
Mountains are further west still) suggests they may harbour geographic isolates and endemic taxa with a range
of geographic origins. At this stage L. gasconi appears to be endemic to this range however while the
surrounding ranges are better explored than the Fojas, they continue to reveal novel lineages (Flannery and
Groves 1998, Helgen 2005), and the possibility that L. gasconi sp. nov. inhabits at least some of them cannot
be discounted. New Guinea remains a frontier for both biological survey and phylogenetic analysis and
further discussion of patterns of endemism within yhe Fojas should await the results of work currently in
progress.
Acknowledgments
We are extremely grateful to the Indonesian Institute of Sciences (LIPI) for permission to undertake research
in Papua Province, Indonesia and for providing export permits. We are also grateful to Sancoyo Lanang for his
assistance, and to Ibu Mumpuni and Hellen Kurniati from the Museum Zoologicum Bogoriense for help
approving export permits and registering specimens. This research was part of Conservation International's
Rapid Assessment Program biodiversity surveys (RAP), and we greatly appreciate their assistance. We are
particularly grateful to Bruce Beehler, Neville Kemp, Dr Yatna Supriatna, and Dr Yance de Fretes of
Conservation International for their support and assistance during the survey. The communities of Marina
Valen and Kwerba generously hosted our field studies in 2004 and 2005 respectively, for which we are most
grateful. Bruce Beehler, Kris Helgen, Henk van Mastrigt and Wayne Takeuchi provided lively discussions
around the camp mud. During preparation of this manuscript SJR was supported by funding from
Conservation International. Additional support was provided by National Geographic, the Mark Mitchell
Foundation, the Winifred Violet Scott Trust and the South Australian Museum Board. Luke Price and Steve
Donnellan provided access to unpublished molecular data. The curators of the following museums provided
access to specimens in their care: Barry Clarke (BMNH, London), Marinus Hoogmoed and Pim Arntzen
(RMNH, Leiden), and Giuliano Doria (MSNG, Genoa). Mark Hutchinson and Carolyn Kovach provided
access to specimens, and numerous other courtesies, at the South Australian Museum and Lisa Capon kindly
produced Figure 2.
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Zootaxa 2277 © 2009 Magnolia Press ·
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(Anura: Hylidae) in the Papuan Region. Australian Journal of Zoology, 56, 257–280.
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Appendix 1. Specimens examined.
Litoria albolabris (Wandolleck, 1911). Papua New Guinea. Sandaun Province, Aitape, SAMA R4947 (syntype).
Litoria auae Menzies and Tyler, 2004. Papua New Guinea: Gulf Province, Purari River, near McDowell Is, UP2490
(holotype), SAMA R57262–3 (paratypes).
Litoria chloronota (Boulenger, 1911). Indonesia. Papua Barat, Arfak Mountains, BM1947.2.31.20 (syntype).
Litoria contrastens (Tyler, 1968). Papua New Guinea. Eastern Highlands Province, Barabuna, SAMA R5845 (holotype),
SAMA R6450 (paratype); Western Highlands Province, Noreikova, SAMA R5847 (5 specimens).
Litoria elkeae Günther and Richards, 2000. Indonesia. Papua Province, Siewa, MZB Amp3866–9, QMJ70490–2 (all
paratypes).
Litoria havina Menzies, 1993. Papua New Guinea. Western Province, Ok Kam, UP7281 (holotype); Western Province,
Ok Ma, UP 8406–7 (paratypes); Western Province, Ok Kam, SAMA R38596–7; Southern Highlands Province,
Agogo, SAMA R60173–7.
Litoria iris (Tyler, 1963). Papua New Guinea. Chimbu Province, Bamna, BM 1961.1226 (holotype); Southern Highlands
Province, Tari, UP3115–35; Eastern Highlands Province, Ubaigubi, UP8289–90; Enga Province, Porgera, UP 7148–
67; Sandaun Province, Telefomin, SAMA R5423, 5874.
Litoria kumae Menzies and Tyler, 2004. Papua New Guinea. Southern Highlands Province, Tari, UP3108 (holotype),
SAMA R52760–61 (paratypes).
Litoria leucova (Tyler, 1968). Papua New Guinea. Sandaun Province, Mt Stolle, SAMA R44091–2, UP8604–6.
Litoria longicrus (Boulenger, 1911). Indonesia. Papua Province, Wendessi, BM 1947.2.22.60–61 (syntypes).
Litoria majikthise Johnston and Richards, 1994. Papua New Guinea. Western Province, Tabubil, SAMA R44093
(holotype), UP6734, 7305–9, 8501–8, 8602–3, SAMA R 44094–44101 (paratypes).
Litoria mucro Menzies, 1993. Papua New Guinea. East Sepik Province, Near Rauit Village, UP2741–3, 2745–56
(paratypes).
L. multiplica (Tyler, 1964). Papua New Guinea. Eastern Highlands Province, Kassam, Kratke Mountains, SAMA
R04946 (paratype); Crater Mountain Wildlife Management Area, SAMA R64647-656; Sandaun Province,
Telefomin area, SAMA R05422, SAMA R05277, SAMA R11835–9; Imigebip, Hindenburg Ranges, SAMA
R11150; Western Highlands Province, Amulua, Jimi Valley, SAMA R08923; Chimbu Province, Bomai, SAMA
R06303, R11833; Dege, SAMA R06169, R11832; Elmagale, SAMA R06171; Southern Highlands Province, Ialibu,
SAMA R09148, R11834; Tualapa, SAMA R64644-6; Tumia, 25 mls from Mendi, SAMA R06828; Magidobo,
SAMA R34360.
Litoria mystax (van Kampen, 1906). Indonesia. Papua Province, Moaif, RMNH 4632 (holotype).
Litoria nigropunctata (Meyer, 1875). Papua New Guinea. Morobe Province, Lae, SAMA R09296; Madang Province,
Binek near Madang, SAMA R11794. Indonesia. Papua Province, Gebeh Island, Hyla bernsteini RMNH 4421
(syntype); Yapen Island, Hyla ouwensi MCZ 2434 (holotype).
Litoria ollauro Menzies, 1993. Papua New Guinea. Milne Bay Province, Agaun, UP4644 (holotype), UP4634–43, 4645–
51, 5070–89 (paratypes).
L. richardsi Dennis and Cunningham, 2006. Papua New Guinea. Western Province, Tabubil, SAMA R60283 (holotype).
Indonesia. Papua Province, Tiri River, Mamberamo Drainage, MZB Amph. 11823 (paratype).
L. singadanae Richards, 2005. Papua New Guinea. Morobe Province, Ridge above Surim Camp, eastern Finisterre
Mountains, SAMA R60172 (holotype), UP 9968, SAMA R60171 (paratypes).
L. wapogensis Richards and Iskandar, 2001. Indonesia. Papua Province, Wapoga River Alpha exploration camp, MZB
Amp. 3873 (holotype), MZB Amp. 3874–76, SAMA R54595–98 (paratypes).
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