Description of a striking new Mantophryne species (Amphibia, Anura, Microhylidae) from Woodlark Island, Papua New Guinea

Abstract

We describe a striking new species of the microhylid frog genus Mantophryne from Woodlark Island in Milne Bay Province, Papua New Guinea. It is most similar to M. lateralis but is distinguished from that species by its more slender body, longer shanks, larger discs on the toes, and unique advertisement call. Most known specimens had, in life, a striking golden tan mid-dorsum bordered by broad blackish dorsolateral bands. The new species is currently known only from the rainforests of Woodlark Island, where males call from elevated perches up to 4 m above the ground from climbing Freycinetia plants, from crevices and hollows in elevated limestone outcrops, and from tree buttresses and on top of fallen logs on the forest floor. It is the most arboreal member of this predominantly terrestrial genus discovered to date.

Full text

museum für naturkunde
Description of a striking new Mantophryne species (Amphibia,
Anura, Microhylidae) from Woodlark Island, Papua New Guinea
Rainer Günther
1
, Stephen Richards
2
1 Museum für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstr. 43, 10115 Berlin, Germany
2 Herpetology Department, South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia
http://zoobank.org/E134B166-6A06-41F7-B1C6-ACC8A503C2C2
Corresponding author: Rainer Günther (rainer.guenther@mfn-berlin.de)
Abstract
We describe a striking new species of the microhylid frog genus Mantophryne from
Woodlark Island in Milne Bay Province, Papua New Guinea. It is most similar to M.
lateralis but is distinguished from that species by its more slender body, longer shanks,
larger discs on the toes, and unique advertisement call. Most known specimens had, in
life, a striking golden tan mid-dorsum bordered by broad blackish dorsolateral bands. The
new species is currently known only from the rainforests of Woodlark Island, where males
call from elevated perches up to 4 m above the ground from climbing Freycinetia plants,
from crevices and hollows in elevated limestone outcrops, and from tree buttresses and on
top of fallen logs on the forest oor. It is the most arboreal member of this predominantly
terrestrial genus discovered to date.
Key Words
Frog
new species
taxonomy
bioacoustics
New Guinea
Introduction
The genus name Mantophryne was coined by Boulenger
(1897) to accommodate a single frog species (lateralis),
“similar to Xenorhina but with large eyes and ranoid
habit”, from the east of New Guinea. The name Man-
tophryne lateralis was retained by Méhely (1901) and
Vogt (1911) but treated as a synonym of Hylophorbus
rufescens Macleay, 1878, by Fry (1913) and Van Kamp-
en (1923). Parker (1934) stated that the ve type speci-
mens of Mantophryne lateralis were not all conspecic
and that the name bearing syntype must be allocated to
Asterophrys rufescens Parker, 1934 (he treated Hylo-
phorbus as a synonym of Asterophrys Tschudi, 1838).
In his revision of the subfamily Asterophryinae Günther,
1858, Zweifel (1972) included lateralis in the genus
Phrynomantis Peters, 1867, described a similar species
as Phrynomantis infulata, and also allocated Asterophrys
louisiadensis Parker, 1934 to this genus. However the
genus name Phrynomantis was subsequently shown by
Received 26 December 2015
Accepted 7 April 2016
Published 11 May 2016
Academic editor:
Johannes Penner
Dubois (1988) to be preoccupied by an African taxon
and it was therefore replaced by Callulops Boulenger,
1888. Zweifel (1972) had also resurrected the genus
name Hylophorbus mainly because members of that ge-
nus have an eleutherognathine jaw, in contrast to a sym-
phygnathine jaw in members of the genus Phrynomantis
(Callulops). On the basis of osteological and myological
studies Burton (1986) then resurrected the name Man-
tophryne where he accommodated the symphygnathine
species infulata, lateralis and louisiadensis. These three
species were later complemented by M. axanthogaster
Kraus & Allison, 2009.
In a recent paper Oliver et al. (2013) studied phylo-
genetic relationships between the closely related genera
Mantophryne, Hylophorbus and Pherohapsis Zweifel,
1972. They used three mitochondrial and three nuclear
genes and conrmed M. lateralis, M. louisiadensis and
M. axanthogaster as congeneric; the monotypic Phero-
hapsis menziesi Zweifel, 1972 is also nested within Man-
tophryne and was transferred to that genus; M. infulata is
Zoosyst. Evol. 92 (1) 2016, 111–118 | DOI 10.3897/zse.92.7629
Copyright Takafumi Nakano. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits
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Günther, R. & Richards, S. : Description of a new Mantophryne species
112
nested within Hylophorbus and was included in that ge-
nus as H. infulatus. Accordingly Mantophryne at present
contains four species and quite a number of undescribed
forms (Oliver et al. 2013, Frost 2015).
During eld work on Woodlark Island, Papua New
Guinea (Fig. 1) in April 2011 three strikingly coloured
male frogs were encountered that, on the basis of their
overall morphology and the symphygnathine condition of
the maxillary bones belong to the genus Mantophryne but
cannot be assigned to any of the known species. We de-
scribe and illustrate the new species here.
Material and methods
Frogs were located at night by tracking their advertise-
ment calls, and all specimens were photographed in life
prior to preservation. Tissue probes from liver were taken
from two of the three collected specimens and stored in
about 96% ethanol to enable DNA sequencing. All speci-
mens were xed in 10% formalin and later transferred to
75% ethanol for permanent storage. Measurements were
taken with a digital calliper (> 10 mm) or with a binocular
dissecting microscope tted with an ocular micrometer (<
10 mm) to the nearest 0.1 mm.
SUL snout-urostyle length: from tip of snout to distal
tip of urostyle-bone; SUL is subject to lower mea-
surement error than the traditionally used snout-
vent length (SVL) (R. Günther, pers. obs.) so we
have used it here. However both measurements are
very similar. We therefore directly compare SUL
measurements reported here with SVL measu-
rements of congeners presented in the literature;
TL tibia length: external distance between knee and
ankle;
TaL length of tarsus: external distance between tarsal
and ankle joints held at a right angle;
T4L length of fourth toe: from tip of toe to proximal
end of inner metatarsal tubercle;
T4D transverse diameter of disc of fourth toe;
F3L length of third nger from tip to proximal margin
of central palmar tubercle;
F3D transverse diameter of disc of third nger;
F1D transverse diameter of disc of rst nger;
T1D transverse diameter of disc of rst toe;
HL head length: from tip of snout to posterior margin
of tympanum;
HW head width, taken in the widest point;
SL snout length: from an imaginary line that connects
the centres of eyes to tip of snout;
END distance from anterior corner of orbital opening to
centre of naris;
IND internarial distance between centres of external
nares;
ED eye diameter: from anterior to posterior corner of
orbital opening;
TyD horizontal diameter of tympanum.
Figure 1. Map of New Guinea showing location of Woodlark Island, type locality of Mantophryne insignis.

Zoosyst. Evol. 92 (1) 2016, 111–118
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113
Advertisement calls were recorded with a Marantz PMD-
660 digital recorder and a Sennheiser ME66 shotgun micro-
phone and analysed with Avisoft-SAS Lab Pro software.
Specimens are stored in the collection of the South
Australian Museum, Adelaide (SAMA), the Museum für
Naturkunde Berlin (ZMB) and prospectively the Papua
New Guinea National Museum (PNGNM), Port Moresby.
Our information concerning features of Mantophryne
species was taken from original descriptions (Boulenger
1897; Parker 1934; Zweifel 1972; Kraus and Allison
2009), recompiled treatises (Parker 1934; Zweifel 1972;
Burton 1986, Menzies 2006; Oliver et al. 2013) and our
own observations including direct examination of eight
Mantophryne lateralis (SAMA R69327–34) specimens
from mainland New Guinea.
Results
Mantophryne insignis sp. n.
http://zoobank.org/A66147A7-C0C9-443E-85B0-9A1589D415B8
Holotype. SAMA R69237 (eld number = FN SJR
13920); adult male, collected on the slopes of Talpos
Mountain, Woodlark Island (Fig 1), Milne Bay Province,
Papua New Guinea (9°09.364’S, 152°46.495’E, 180 m
a.s.l.) on 15.IV.2011 by S.J. Richards.
Paratypes. ZMB 83181 (FN SJR 13923), same data
as for holotype, and FN SJR 13932 (to be deposited in
the PNG National Museum), Upper Muniai Creek, Wood-
lark Island, Milne Bay Province, Papua New Guinea
(9°07.502’S,
152
°
44.902’E, 30 m a.s.l.).
Diagnosis. The new species is assigned to the genus
Mantophryne on the basis of the following characters:
body slender, circum-marginal grooves on all nger and
toe discs, those on the toes wider than those of ngers;
symphygnathine condition of the maxillary bones (ante-
rior processes of the maxillary bones not fused but almost
in contact and joined by a ligament); and two protuber-
ances on chin. It differs from all hitherto known conge-
ners by its 1) relatively long legs (TL/SUL 0.55–0.57 vs.
<0.51 in males of all congeners; Menzies 2006, Kraus
and Allison 2009), 2) more expanded terminal discs on
the toes, 3) smooth dorsum, 4) advertisement calls con-
sisting of 38–52 notes and lasting 4.6–6.5 s, and 5) dis-
tinct colouration normally comprising a uniformly gold-
en tan dorsum, broad blackish dorsolateral bands edged
below with a narrow, slightly undulating white stripe and
large, distinct dark brown blotches each encircled by a
white border, on the abdomen.
Description of the holotype. In life mid-dorsal
band and dorsal surfaces of thighs uniform golden tan,
dorsal surfaces of shanks yellow, posterior of thighs, ex-
tending partly to dorsal surfaces, orange-red; supra-can-
thal stripe, extending onto upper eyelids, yellowish; dor-
solateral band blackish, ventral boundary of this band
delineated by narrow white stripe; axillary region yel-
lowish and inguinal region orange-red. Lateral surfaces
Figure 2. Holotype of Mantophryne insignis sp. n., dorsolateral view in life.

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Günther, R. & Richards, S. : Description of a new Mantophryne species
114
Figure 3. Holotype of Mantophryne insignis sp. n., ventral view in life.
of shanks and dorsal surfaces of tarsi dark brown, their
ventral parts mottled with dark brown. Iris silvery with
a few irregular dark lines; anterior and posterior portions
of iris more strongly pigmented by dark lines and with an
orange hue (Fig. 2). Ground colour of all ventral surfac-
es whitish. Throat and chest covered by variably intense
brown pigmentation and scattered large dark-brown spots
especially along the chin and on the bases of the forelegs.
Abdomen and lower areas of anks are covered by a pat-
tern of very conspicuous dark-brown blotches, each en-
circled by a narrow white line; areas between the spots
are less densely pigmented than the throat and chest;
brown spots on inferior thighs are smaller than on abdo-
men and not encircled by white lines (Fig. 3).
Colouration of the preserved holotype: Dorsal sur-
faces of head, shanks and mid-dorsum uniform light
grey; dorsal surfaces of thighs with faint brown mot-
tling; those of upper arm more strongly mottled with
brown; anterior of lower arm and dorsal surfaces of
hands, tarsi and metatarsi irregularly brown; conspi-
cuous large dark brown spots edged with a white line
extend to posterior of lower arm and anterior tarsus
and metatarsus; a relatively wide whitish supra-canthal
stripe extends from snout tip to middle of eyelid. The
broad dorsolateral bands begin at the posterior edge of
the orbital opening, are widest on middle of anks, and
end with a small tip at cloacal opening. Ground colour
of all ventral surfaces is off-white. Mottling is same co-
lour as in life.
Measurements of the holotype are listed in Table 1.
There is a longitudinal incision in the abdomen. Head
longer than broad (HL/HW 1.06). Snout truncate, with
only a slight narrowing at the tip in dorsal view and
protruding in lateral view. Canthus rostralis rounded,
straight anterior to eyes before bending laterally above
the nares; loreal region at, nares directed laterally, close
to end of snout and not visible from above; distance be-
tween nares greater than distance between eye and naris
(END/IND 0.86). Supratympanic skin fold scarcely pro-
nounced, tympanic annulus clearly visible; horizontal di-
ameter of tympanum more than half that of eye (TyD/ED
0.55). Pupil horizontally oval. Tongue very broad, long
and free laterally and posteriorly, its posterior margin not
notched. Anterior prepharyngeal ridge with three lobes
and posterior ridge with ten denticles. Vocal slits small
and near angle of jaws. No webbing between ngers; one
well developed subarticular tubercle on nger I and II
and two well developed tubercles on ngers III and IV;
three less prominent metacarpal tubercles; no other pal-
mar tubercles; all ngers bear small but distinct, grooved
discs; disc of third nger clearly smaller than that of
fourth toe (F3D/T4D 0.63); relative length of ngers
III>IV>II>I (Fig. 4). Legs long and slender (TL/SUL
0.57). Toe discs clearly broader than those of ngers and
all with distinct circum-marginal grooves; no webbing
between toes; one well-developed subarticular tubercle
on toe I and toe II, two prominent subarticular tubercles
on toes III, V and three on toe IV; clearly expressed inner

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115
metatarsal tubercle, no outer one; relative length of toes
IV>III>V>II>I (Fig. 5). All dorsal and ventral surfaces
of legs, body and head smooth except two inconspicuous
)( -shaped longitudinal ridges beginning between eyes
and reaching to occiput. Two small chin protuberances,
clearly visible in the living specimen, disappeared in
preservative. There are 3 faint, whitish tubercles on the
upper edge of the eyelid.
Variation in the type series. Body size (SUL) of
three adult males (including the holotype) varied only
slightly, from 35.0 mm to 36.2 mm, mean 35.5 mm, SD
(standard deviation) 0.61. Measurements of all types are
listed in Table 1. Deviations in colouration of the para-
types from the holotype are insignicant; all share the
striking colour pattern of a golden tan dorsum with broad,
blackish dorsolateral bands and a heavily spotted venter.
Additional variation. Three additional adult male
specimens of this species (BPBM 40135–7) collected on
Woodlark Island by F. Kraus are 34.1–35.3 mm SUL and
agree closely with the description of the type series in all
features except that the dorsal colouration of one of the
three specimens (BPBM 40135), including the mid-dorsal
band, is creamy tan rather than golden tan.
Vocalisation. The advertisement call of Mantophryne
insignis sp. n., recorded at an air temperature of 25 °C, is a
rattle of several seconds duration (Fig. 6).
Figure 4. Ventral view of the left hand of the holotype of Man-
tophryne insignis sp. n. in life.
Figure 5. Ventral view of the left foot of the holotype of Manto-
phryne insignis sp. n. in life.
Two complete calls of the holotype and two calls of
SJR 13932 were analysed. Call duration varied from
4.6 to 6.5 s, mean 5.9 s. Number of notes per call was
38–52, mean 47.5. Note repetition rate was from 8.0 to
8.3/s, mean 8.1 notes/s. Mean note duration of two calls
from the holotype was 53.1 ms, SD 15.4, range 30–78 ms,
n=104; mean internote interval duration was 72.9 ms, SD
12.7, range 41–153 ms, n=102. For technical reasons note
and internote length from the calls of the second speci-
men could not be measured exactly and therefore are not
considered here. Note and internote interval length and
amplitude of notes clearly increased during the course of
the call (compare Figs 7 and 8).
In three of the four calls the last internote interval is
clearly the longest and in one call the last but one inter-
val was the longest. All notes are composed of pulses, and
these mostly cluster into pulse-groups. The rst pulse of al-
most all notes is clearly separated from the following (clus-
tered) pulses. Frequencies scatter mainly from 1.0 to 3.5
kHz with dominant frequency at 2.0 kHz (Fig. 9). There are
no harmonics evident and no modulation of frequencies.

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Günther, R. & Richards, S. : Description of a new Mantophryne species
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Distribution and ecological remarks. The three
type specimens of M. insignis were detected by their
calls, which were uttered at night from hidden perch-
es 50–80 cm high in a limestone block, a tree buttress
and a fallen log, all in lowland rainforest (30–180 m
asl) in south-central Woodlark Island. However two of
three additional specimens found calling on Woodlark
Island by F. Kraus (pers. comm.) were approximately 4
m above the ground, in climbing pandanus (Freycinetia
sp.) plants. The third specimen was calling from under
a leaf on the forest oor. The slender body form, long
legs and expanded toe discs (relative to congeners) re-
ect the unusually arboreal habits of this Mantophryne
species. Given the uniformity of habitat across the is-
land, and the lack of major topographic relief, it is like-
ly that the species is widespread in lowland rainforest
on Woodlark Island. This species has not been reported
from any other islands in the region and may be endem-
ic to Woodlark.
Etymology. The name insignis is derived from the
Latin ‘insignis’ meaning remarkable or conspicuous, and
refers to the species’ distinctive colour pattern and unusu-
al (for the genus) ecology.
Comparison with other species. Mantophryne
lateralis, which is mainly distributed throughout the
lowlands of eastern New Guinea (Oliver et al. 2013), is
most si milar to the new species in having black lateral bands
and distinct spotting on the abdomen. Some specimens of
this species also have extremely smooth skin, approaching
the state in the new species. However it is larger than the
new species (males more than 40 mm SUL vs. less than
40 mm SUL in M. insignis), has shorter legs (SUL/TL
<0.51 vs. 0.55–0.57), smaller toe discs (T4D/SUL <0.034–
0.042 vs. 0.042–0.044), lacks the conspicuous golden tan
dorsum, and has very long advertisement calls (up to 30 s)
at 22–24 °C with a note duration of about 200 ms vs. calls
of about 6 s and a mean note length of about 50 ms in the
new species (Zweifel 1972, Menzies 2006).
With a snout-vent length up to 82 mm Mantophryne
louisiadensis is substantially larger than the new species;
it also has a broader head, a more robust habitus, lacks
dark lateral bands and does not exhibit the striking golden
tan middorsal area or clearly delimited brown spots on the
abdomen (Zweifel 1972, Kraus and Allison 2009).
Mantophryne axanthogaster male is also larger (> 40
mm SVL) than M. insignis and further differs from the
new species by its ecked (vs. uniform golden tan) dor-
sum, lack of dark lateral bands, and uniform grey venter
(vs. strongly spotted). Moreover, its advertisement call
contains 13–18 notes (vs. 38–52 notes) with a mean note
duration of 126 ms (vs. about 50 ms) and a mean note
repetition rate of 1.8 notes/s (vs. 8.1 notes/s) at 26.5 °C
(Kraus and Allison 2009).
With a snout-vent length of 25–31 mm Mantophryne
menziesi is smaller than the new species, lacks dark later-
al bands and a blotched venter, and has a grey- brown (vs.
golden tan) dorsum. Its advertisement call is also differ-
ent, with notes having a length of about 200 ms (tempera-
ture not available) (vs. 50 ms). Moreover in M. menziesi
the squamosal and frontoparietal bones meet to form an
arch over the prootic region, a character which is unique
for asterophryine microhylid frogs.
Hylophorbus infulatus (until recently Mantophryne
infulata) is similar to M. insignis sp. n. in many body
proportions. The species differ, however, in their internarial
spacing. The ratio END/IND of 20 specimens of H.
infulatus ranges from 0.73–0.84 (Zweifel 1972) whereas
three specimens of the new species have values of 0.86–
0.91. Moreover, the species differ in their colouration. H.
infulatus has an inconspicuous brown mid-dorsum with
some darker markings (vs. uniform golden tan mid-dorsum
in most M. insignis), the upper margin of its blackish
dorsolateral band is poorly dened (vs. well dened
Table 1. Body measurements and body ratios of the type series
of Mantophryne insignis sp. n. Reg-No=registration numbers;
FN are the eld numbers of Stephen Richards (SJR). SAMA
R69237 is the holotype, all three specimens are adult males. All
measurements are in mm; SD=standard deviation; all other ab-
breviations are explained in “Materials and methods”.
Reg-No
FN
SAMA
R69237
SJR
13920
ZMB
83181
SJR
13923
PNGNM
SJR
13932
Mean±SD
SUL 36.2 35.0 35.4 35.5±0.61
TL 20.7 19.8 19.5
TaL 12.2 12.0 11.5
T4L 20.1 19.1 18.3
T4D 1.6 1.5 1.5
F3L 10.7 9.2 9.1
F3D 1.0 1.0 1.0
T1D 1.1 1.0 1.0
F1D 0.8 0.7 0.7
HL 12.5 12.2 12.3
HW 11.8 11.5 11.6
END 3.0 2.9 3.1
IND 3.5 3.2 3.4
ED 3.8 4.1 4.0
TyD 2.1 2.5 2.3
SL 5.5 6.0 5.6
TL/SUL 0.57 0.57 0.55 0.56±0.01
TaL/SUL 0.33 0.34 0.32 0.33±0.01
T4L/SUL 0.56 0.55 0.52 0.54±0.02
T4D/SUL 0.044 0.043 0.042 0.043±0.001
F3L/SUL 0.30 0.26 0.26 0.27±0.02
F3D/SUL 0.022 0.029 0.028 0.026±0.004
T4D/F3D 1.60 1.50 1.50 1.53±0.06
F1D/SUL 0.022 0.020 0.020 0.021±0.001
T1D/F1D 1.38 1.43 1.43 1.41±0.03
HL/SUL 0.35 0.35 0.35 0.35±0.00
HW/SUL 0.33 0.33 0.33 0.33±0.00
HL/HW 1.06 1.06 1.06 1.06±0.00
END/IND 0.86 0.91 0.91 0.89±0.03
ED/SUL 0.105 0.117 0.113 0.112±0.006
TyD/SUL 0.058 0.071 0.065 0.065±0.007
TyD/ED 0.55 0.61 0.58 0.58±0.03
SL/SUL 0.152 0.171 0.158 0.160±0.009

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Figure 6. Wave form of a complete advertisement call of the holotype of Mantophryne insignis sp. n. with 52 notes.
Figure 7. Wave form (above) and spectrogram (below) of a sequence of 15 notes from the rst part of an advertisement call of the
holotype of Mantophryne insignis sp. n.
Figure 8. Wave form (above) and spectrogram (below) of a sequence of 9 notes from the last part of an advertisement call of the
holotype of Mantophryne insignis sp. n.

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in M. insignis) and its ventral surfaces are mottled with
diffuse ecks (vs. covered with well-dened dark brown
blotches) (Zweifel 1972).
Acknowledgements
Field work on Woodlark Island was supported by Wood-
lark Mining Limited and SJR is particularly grateful to
George and Eleanor Clapp of WML’s Environment De-
partment for their assistance and support. Francis Crome
and Daniel Moriarty also assisted the second author’s
work in various ways. The PNG National Research In-
stitute and Department of Environment and Conservation
approved the second author’s research visa, and the export
of specimens respectively. Carolyn Kovach and Mark
Hutchinson provided assistance at the South Australian
Museum, and Lisa Capon produced the map. Fred Kraus
(University of Michigan) kindly provided eld observa-
tions on the new species’ habitat use, and a photograph
of an additional specimen from Woodlark Island and we
are also grateful to Allen Allison (Bishop Museum) who
provided some measurements of this material.
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Figure 9. Power spectrum of an advertisement call of Manto-
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