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Two new treefrogs of the Boophis
rappiodes group from eastern
Madagascar (Amphibia Mantellidae)
M. Vences a & F. Glaw b
a Institute for Biodiversity and Ecosystem Dynamics, Zoological
Museum, University of Amsterdam , PO Box 94766, 1090 , GT
Amsterdam , The Netherlands E-mail:
b Zoologische Staatssammlung , Münchhausenstr. 21, D-81247 ,
München , Germany E-mail:
Published online: 30 Jul 2012.
To cite this article: M. Vences & F. Glaw (2002) Two new treefrogs of the Boophis rappiodes
group from eastern Madagascar (Amphibia Mantellidae), Tropical Zoology, 15:1, 141-163, DOI:
10.1080/03946975.2002.10531171
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Tropical Zoology 15: 141-163, 2002
Two new treefrogs of the Boophis rappiodes group
from eastern Madagascar (Amphibia Mantellidae)
1
2
M. VENCES and F. GLAW
1 Institute for Biodiversity and Ecosystem Dynamics, Zoological Museum, University
of Amsterdam, PO Box 94766, 1090GT Amsterdam, The Netherlands
(E-mail: m.vences@t-online.de)
2
Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 München, Germany
(E-mail: Frank.Glaw@zsm.mwn.de)
Received 21 March 2002, accepted 22 May 2002
Two new sibling species of Boophis Tschudi 1838 are described from Anda-
sibe in central-eastern Madagascar. Both are small greenish treefrogs with a
translucent ventral skin and without lateral fringes along lower arm and tarsus,
and are thereby assignable to the phenetic B. rappiodes group. Boophis bottae n.
sp. is morphologically similar to B. rappiodes (Ahl 1928) and occurs syntopically
with this species. It strongly differs from its sibling by advertisement calls (long
trill notes instead of two-pulse notes), and by a reddish-brown dorsal pattern
which does not fade soon in ethanol, often covering the entire back (vs a red
pattern that largely fades in ethanol in B. rappiodes). Boophis tasymena n. sp. is
similar to B. erythrodactylus (Guibé 1953) but differs in advertisement calls
(notes composed of two instead of four-seven pulses) and lack of red colour on
tips of fingers and toes. A lectotype is designated for B. erythrodactylus. The dis-
covery of the two new species in addition to the revised distributional informa-
tion for B. rappiodes, B. erythrodactylus and B. viridis Blommers-Schlösser 1979
confirms that mid-elevational central-eastern Madagascar is the centre of diver-
sity for many Malagasy amphibian groups but has a relatively low degree of
endemism. DNA sequence divergence was high within each pair of sibling
species (6-7% in a fragment of the mitochondrial 16S rRNA gene), suggesting
that their reproductive isolation was not a recent event and probably predates
the Pleistocene.
KEY WORDS: Amphibia, Anura, Mantellidae, Boophis viridis, Boophis rappiodes,
Boophis erythrodactylus, Boophis bottae n. sp., Boophis tasymena n.
sp., Madagascar, advertisement calls, distribution.
Introduction
Materials and methods
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M. Vences and F. Glaw
Species accounts
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Boophis viridis Blommers-Schlösser 1979
Boophis rappiodes (Ahl 1928)
Boophis bottae n. sp.
Boophis erythrodactylus (Guibé 1953)
Boophis tasymena n. sp.
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Discussion
Acknowledgements
References
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INTRODUCTION
With currently more than 45 species, the treefrog genus Boophis is one of the
most speciose groups of Malagasy anurans. It is part of an endemic frog radiation
(RICHARDS et al. 2000) which has recently been defined as the family Mantellidae
(VENCES & GLAW 2001). Boophis was first recognized as a natural unit and divided into
seven species groups by BLOMMERS-SCHLÖSSER (1979). Later, BLOMMERS-SCHLÖSSER &
BLANC (1991) assigned all known species in the genus to these groups, while GLAW &
VENCES (1994) proposed some modifications to the grouping. According to the molec-
ular data of RICHARDS et al. (2000), who studied representatives of five of the seven
species groups (sensu GLAW & VENCES 1994), Boophis is a monophyletic assemblage.
One characteristic pattern in Boophis is the high proportion of sibling species
which are well defined by advertisement calls and partly by colouration, but virtu-
ally indistinguishable by external morphology. This pattern has received particular
attention by BLOMMERS-SCHLÖSSER (1979) in her treatment of the Boophis rappiodes
group. She assigned two described species (B. rappiodes and B. erythrodactylus) to
this group and described two new taxa (B. mandraka and B. viridis). She found all
these species in central-eastern Madagascar, at her collecting localities Mandraka
and Andasibe, and demonstrated close syntopy of two species (B. erythrodactylus,
B. mandraka Blommers-Schlösser 1979) at one site (Mandraka). All species of the
B. rappiodes group are small, basically greenish-coloured treefrogs with a translu-
cent venter in life, reminiscent of the Neotropical Glass frogs of the family Cen-
trolenidae. When preserved, the often diagnostic colour pattern is lost: the greenish
colour changes first to pale yellow and then to whitish. If present, the red pattern
often becomes brownish or fades to white. Although a few morphological charac-
ters are known to distinguish some of the species as defined by BLOMMERS-SCHLÖSSER
(1979) (e.g., the position of the nostrils, which are closer to the eye in B. mandraka
but closer to the snout tip in the other species), in general the B. rappiodes group
exemplifies the difficulties in distinguishing sibling Malagasy frog species better
than most other species assemblages (paralleled, however, by the B. luteus group;
e.g., ANDREONE 1993, ANDREONE et al. 1995).
The recent intensive survey work carried out in several parts of eastern Mada-
gascar revealed that the species inventory of Madagascar’s herpetofauna is far from
being complete (GLAW & VENCES 2000). Several of the newly discovered forms are
assignable to the B. rappiodes group. In the present paper, we describe two of these
as new species, partly revise the two species B. rappiodes and B. erythrodactylus,
and provide new information about the distribution and advertisement calls of B.
viridis. A revision of the remaining representative of the group, B. mandraka, and
two new sibling species will be published elsewhere.

Two new treefrogs from Madagascar
143
MATERIALS AND METHODS
Specimens were collected at night, mainly by locating calling males with the aid of
electric torches. They were euthanised using chlorobutanol, fixed in 90% ethanol and pre-
served in 70% ethanol. Samples of femur muscle were preserved in pure 90% ethanol; DNA
was extracted from these tissue samples, and a fragment of the mitochondrial 16S rRNA gene
(560 nucleotides) was amplified and sequenced using protocols given in VENCES et al. (2000).
Morphological measurements of the preserved specimens were taken by the senior
author with callipers to the nearest 0.1 mm: SVL (snout-vent length), HW (head width), HL
(head length), ED (horizontal eye diameter), END (eye-nostril distance), NSD (nostril-snout
tip distance), NND (nostril-nostril distance), TD (horizontal tympanum diameter), HAL (hand
length), FORL (forelimb length), HIL (hindlimb length), FOL (foot length), FOTL (foot length
including tarsus). The webbing formula is given according to BLOMMERS-SCHLÖSSER (1979).
Vocalizations were recorded using portable tape recorders with an external microphone
(Vivanco EM 238) and were analyzed with the MEDAV sound analyzing system Spektro 3.2.
Temporal measurements are given as range, with mean standard deviation, and number of
temporal units measured, in parentheses.
Institutional abbreviations used are as follows: MNHN (Muséum National d’Histoire
Naturelle, Paris); UADBA (Université d’Antananarivo, Département de Biologie Animale)
(numbers given refer to the fieldnumbers of different collectors; UADBA-MICET, collection of
the MICET team; UADBA-RD, collection of D. Rakotomalala; UADBA-MV, collection of M.
Vences; UADBA-FG/MV, collection of F. Glaw and M. Vences); ZFMK (Zoologisches
Forschungsinstitut und Museum Alexander Koenig, Bonn); ZMA (Zoölogisch Museum Ams-
terdam); ZMB (Zoologisches Museum der Universität, Berlin); ZSM (Zoologische
Staatssammlung München). Coordinates and altitudes of localities are given according to
GLAW et al. (2001) or given at first mention.
DNA sequences were submitted to public databases; EMBL/Genbank accession num-
bers are the following (voucher specimens in parentheses): Boophis bottae n. sp. (ZSM
344/2000), AJ314817; Boophis erythrodactylus (ZSM 324/2000), AJ314814; Boophis rappiodes
(ZSM 347/2000 and UADBA-FG/MV 2000.59), AJ314815 and AJ314816; Boophis tasymena n.
sp. (ZFMK 62888), AF215339; Boophis viridis (ZSM 338/2000), AJ314818.
SPECIES ACCOUNTS
Boophis viridis Blommers-Schlösser 1979
Boophis viridis BLOMMERS-SCHLÖSSER 1979. Name-bearing type: holotype ZMA 7100A, adult male,
collected by R. Blommers-Schlösser on 14 November 1972. Type locality: “near Perinet (high-
road R.N. 2 at km 142)” according to original description (Perinet being the old French name
of Andasibe). Other types: paratypes ZMA 7100B, two adult males.
Identity and diagnosis. See Table 1 for morphometric measurements. A mem-
ber of the Boophis rappiodes group as recognizable by relatively small size (SVL 29-
31 mm in males, 32-35 mm in females; Table 1 and BLOMMERS-SCHLÖSSER 1979),
greenish and slightly translucent dorsal colouration (during the day; at night more
reddish), translucent venter (inner organs can be clearly seen through the skin in
live specimens) and absence of lateral fringes along lower arm and tarsus (Fig. 1).
Distinguishable from all other species of the group by the larger size of males
(below 27 mm in all other species), characteristic iris colouration (inner iris area

144
M. Vences and F. Glaw

Two new treefrogs from Madagascar
145

146
M. Vences and F. Glaw
brown, outer iris area blue), and by advertisement calls (unharmonious pulsed
notes instead of pulsed click or trill notes of more harmonious structure).
Material examined. UADBA-MICET 331, 335, 344, 347 (Marotreho forest, 6 km S
Ranomafana, 21°18.14’S, 47°27.4’E, 910
m elevation); UADBA-MICET 260-262, 304
(Ankopakopaka forest, 12 km NW Ikongo, 21°49.39’S, 47°20.20’E, 645 m elevation); UADBA-
MV 2001.186, 2001.188-189 (Andasibe); ZFMK 53620, 60012-60014 and 62211 (Andasibe);
ZSM 338/2000 and 675/2001 (Andasibe).
Natural history and advertisement calls. A few observations were reported by
BLOMMERS-SCHLÖSSER (1979) and GLAW & VENCES (1994). According to published
data and observations made in 1994-1996 and 2000-2001 at Andasibe, males general-
ly call at night from leaves at 1-2 m above the ground, along (or up to 10 m away
from) slow-moving brooks and streams, at forest edges. However, on 28 December
1994, during dry weather conditions, calling males were sitting on trees up to 5 m
above the ground. Calling activity was recognized in the months October, November,
December, and March. One pair in axillary amplexus (the female being ZFMK 62211)
was observed on 31 January 1996 and laid 154 eggs in captivity. In contrast to the
pale-yellowish eggs of B. rappiodes, B. erythrodactylus, and B. bottae, the eggs of B.
viridis were dark brown. Another pair in amplexus was observed on 10 March 1996.
Advertisement calls were briefly described by GLAW & VENCES (1994); we here
provide a more detailed description. Calls (recorded on 28 December 1994, in the
evening around 19:00 hr) consisted of two note types. Notes of type 1 (Fig. 2) con-
sisted of up to 40 pulses (pulse repetition rate about 135/sec) and had a duration of
131-295 msec (213
41 msec, n = 15). They were sometimes arranged in series,
with intervals of 347-755 msec (546 129 msec, n = 10) between notes. The fre-
quency ranged between 2150 and 3300 Hz. Each note was emitted as one expira-
tion. Note type 2 generally consisted of three pulses (less frequently up to five puls-
es) and had a duration of 42-74 msec (53 11 msec, n = 9). The intensity of the first
pulse was usually lower than that of the following pulses. This note type was often
emitted in short series of 2-3 notes; inter-note interval duration was 347-484 msec
(420 59 msec, n = 6). When both note types were combined (e.g., a note of type 1
followed by a series of notes of type 2), the note of type 1 was usually more intense.
Distribution. The species was originally known only from the type locality,
Andasibe in central-eastern Madagascar, but was recorded from the Andringitra
massif at 700-720 m altitude by RAXWORTHY & NUSSBAUM (1996). Recent collections
by teams of the University of Antananarivo in the Ranomafana and Ikongo areas
yielded one specimen (UADBA-MICET 331) which clearly can be assigned to B.
viridis (as well as several additional specimens probably belonging to the species:
UADBA-MICET 260-262, 304, 335, 344, 347).
Boophis rappiodes (Ahl 1928)
Rhacophorus rappiodes AHL 1928. Name-bearing type: holotype ZMB 30540, collected by Braun. Type
locality: “Ankoraka, Sahambendrana (Zentral-Madagaskar)” according to the original descrip-
tion; corresponding to Akkoraka near Sahembendrana according to BLOMMERS-SCHLÖSSER &
BLANC (1991). Other types: none.
Identity. During fieldwork in Andasibe, central-eastern Madagascar, we noted
that B. rappiodes as defined by BLOMMERS-SCHLÖSSER (1979), BLOMMERS-SCHLÖSSER

Two new treefrogs from Madagascar
147
Fig. 1. — Boophis viridis, adult specimen from Andasibe.
Fig. 2. — Sonagram and oscillogram of a note of type 1 of Boophis viridis, recorded at Andasibe.

148
M. Vences and F. Glaw
& BLANC (1991) and GLAW & VENCES (1994) was actually a mix of two different
forms. One form had extensive reddish-brown markings on its greenish back, which
in preservative changed into a persistent brownish pattern, and a call that included
long trills, sometimes preceded and followed by click notes. Specimens of the sec-
ond form often had less extensive and usually bright red markings on the greenish
back (Fig. 3), which in ethanol faded after a few days or weeks, and had calls con-
sisting of short click series only. The two forms were observed syntopically at Anda-
sibe, while at Nahampoana in south-eastern Madagascar only the second form was
found and trill calls were not heard. DNA sequences (560 nucleotides of the 16S
rRNA gene) differed in the two forms; 38 substitutions and 3 indels were found, cor-
responding to a total pairwise sequence divergence of 7.3%. Altogether, the evidence
is overwhelming that the two forms are reproductively isolated and constitute two
separate species. The holotype of Boophis rappiodes is in rather good state of preser-
vation (despite the contrary statement of GUIBÉ 1978) and clearly shows remains of
a light dorsolateral band on each side (presumably yellow in life) running from the
nostril over the eye along the supratympanic fold, fading in the shoulder region.
Except for a fine dark spotting all over the dorsum, no other pattern is recognizable.
This agrees exactly with specimens from Nahampoana (ZFMK 53621-53622) which
are clearly attributable to the form with little dark pigment on the back. We there-
fore conclude that this form must be considered as Boophis rappiodes (Ahl 1928),
while the species with more dark pigment on the back will be described below.
Diagnosis. See Table 1 for morphometric measurements. A member of the
Boophis rappiodes group as recognizable by small size (males 20-25 mm, females
30-34 mm), greenish and slightly translucent dorsal colouration, translucent venter
(inner organs can be clearly seen through the skin in live specimens) and absence
of lateral fringes along lower arm and tarsus (Fig. 3). Distinguished from Boophis
mandraka by nostril position (closer to snout tip vs slightly closer to eye), iris
colour (absence of distinct reticulations on a light beige iris), usual presence of red
markings on the back (vs generally absent) and continuation of dorsolateral stripes
from eye to supratympanic fold (vs restricted to snout tip-eye). Distinguished from
B. viridis by smaller size and presence (vs absence) of yellowish dorsolateral
stripes. For distinction from B. erythrodactylus and the two new species described
herein, see below.
Material examined. UADBA-FG/MV 2000.59 (Andasibe); ZFMK 53621-53622 (Naham-
poana); ZFMK 53623-53625 and 59869 (Andasibe); ZFMK 62278 (Vohiparara); ZMB 30450
(holotype; Ankoraka); ZSM 347/2000 and 676/2001 (Andasibe).
Natural history and advertisement call. Calling males were observed at night
2-3 m high in the vegetation along a large, slow-moving stream at Andasibe (some-
times more than 10 m from the water), and about 1 m high in the vegetation along
a rather large and noisy brook at Nahampoana, at forest edges or outside primary
forest. At Nahampoana, advertisement calls were recorded on 4 January 1992 at
18:30 hr (air temperature about 25 °C). Each note consisted of two pulses; duration
of inter-note intervals was 2800-5400 msec. Duration of single pulses was 13-21
msec (n = 6), duration of inter-pulse intervals was 40-45 msec (n = 3). The frequen-
cy was between 2500 and 3500 Hz. Similar calls were recorded at Andasibe (from
specimen ZSM 347/2000; Fig. 4). The second pulse had generally a higher intensity
and longer duration than the first pulse. Duration of the first pulse of a note was

Two new treefrogs from Madagascar
149
Fig. 3. — Boophis rappiodes (ZFMK 59869), adult female from Andasibe.
Fig. 4. — Sonagram and oscillogram of a call of Boophis rappiodes, recorded from specimen
ZSM347/2000 at Andasibe.

150
M. Vences and F. Glaw
9-29 msec (14 7 msec, n = 9); duration of inter-pulse intervals was 42-62 msec (55
6 msec, n = 9); duration of the second pulse was 11-27 msec (20 7 msec, n = 9).
The frequency ranged between 2500 and 3400 Hz.
Distribution. Besides the holotype from (1) Ankoraka and specimens collected
by us at (2) Andasibe, (3) Nahampoana, and (4) Vohiparara, BLOMMERS-SCHLÖSSER
& BLANC (1991) recorded B. rappiodes from three additional localities, Mandraka,
Moramanga and Anosibe. These records are probably based on ZMA 7161 and
7168, tadpoles and young from Mandraka, and ZMA 7166, tadpoles “collected along
the road from Moramanga to Anosibe at km 27” (BLOMMERS-SCHLÖSSER 1979). As it
is not known at present how the tadpoles and young of B. rappiodes can be distin-
guished from those of B. bottae n. sp. (see below), we consider these sites as in
need of confirmation. Also the identity of B. rappiodes records from Marojejy,
Anjanaharibe-Sud and Tsararano in north-eastern Madagascar (RAXWORTHY et al.
1998, ANDREONE et al. 2000, RASELIMANANA et al. 2000) remains to be revised.
Boophis bottae n. sp.
Identity. This is the B. rappiodes-like species with distinct dark pigment on the
back and trill calls (see above). Calls of this species were attributed to B. rappiodes
by GLAW & VENCES (1994).
Diagnosis. A member of the Boophis rappiodes group as recognizable by small
size (males 21-24 mm, one female 35 mm), greenish and slightly translucent dorsal
colouration, translucent venter (inner organs can be clearly seen through the skin
in live specimens) and absence of lateral fringes along lower arm and tarsus. Dis-
tinguished from B. mandraka by nostril position (closer to snout tip than to eye vs
slightly closer to eye), and iris colour (absence of distinct reticulations on a light
beige iris). Distinguished from B. viridis by smaller size, iris colouration, and pres-
ence (vs absence) of yellowish dorsolateral stripes. Distinguished from B. erythro-
dactylus by lack of reddish colour of fingertips, and from B. erythrodactylus and B.
tasymena n. sp. (see below) by lack of regular pattern of red dorsal spots. By mor-
phology and colouration, B. bottae n. sp. is most similar to B. rappiodes. It is distin-
guished, however, by a more extensive dark pattern on the dorsum. This difference
is most distinct in living or freshly preserved specimens; while the dorsal pattern in
B. rappiodes is intensely red, and remains red in preservative before it eventually
fades, the pattern in B. bottae is reddish-brown in life and becomes persistently
dark brown in preservative, often covering almost the entire dorsum (Fig. 5). Addi-
tionally, B. bottae is easily distinguished by its advertisement calls as no other
species of the B. rappiodes group emits long trill calls composed of double click
notes (Fig. 6).
Holotype. ZSM 678/2001, adult male, collected by M. Vences and D. Vieites on 16 Feb-
ruary 2001 close to Andasibe (at a bridge on the road between the National Road 2 and the
Andasibe village), central-eastern Madagascar 18°56’S, 48°25’E, ca 900 m elevation.
Paratypes. UADBA-MV 2000.194 and 2000.196, ZSM 679/2001, three adult males, same
collection data as holotype; UADBA-FG/MV 2000.60 and ZSM 344/2000, two adult males, col-
lected by F. Glaw and M. Vences on 9 February 2000 at the type locality; ZFMK 60015-60016,
two adult males, collected by F. Glaw on 28 December 1994 at Andasibe; ZFMK 62220-62221,
one adult male and one adult female, collected by F. Glaw on 9 February 1996 at Andasibe.

Two new treefrogs from Madagascar
151
Fig. 5. — Boophis bottae n. sp. (paratype ZFMK 60016), adult male from Andasibe.
Fig. 6. — Sonagram and oscillogram of a section of a call of Boophis bottae n. sp. (series of notes
of type 1), recorded at Andasibe.

152
M. Vences and F. Glaw
Further material. UADBA-RD 1724 and 1732 (Vinanitelo, 15.5 km SE Vohitrafeno,
21°46.5’S, 47°20.8’E, 1100 m elevation); UADBA-MICET 19-21 (Mandriandry forest, 4.4 km
SW Tolongoina, 21°35.3’S, 47°29.1’E, 750 m elevation); UADBA-MICET 144 and 147 (Amba-
haka forest, 9 km NW Ambatofotsy, 21°44.2’S, 47°24.5’E, 750 m elevation). As no bioacoustic
data of these specimens are known, we do not include them in the type series; they agree,
however, with the type material in morphology and colouration.
Description of the holotype. SVL 24.2 mm. For measurements, see Table 1.
Body slender; head wider than long, much wider than body; snout rounded in dor-
sal and lateral views, nostrils directed dorsolaterally, slightly protuberant, nearer to
tip of snout than to eye; canthus rostralis moderately distinct, curved; loreal region
concave; tympanum distinct, rounded, 41% of eye diameter; supratympanic fold
rather indistinct and straight; tongue ovoid, distinctly bifid posteriorly; vomerine
teeth distinct, in two elongate aggregations, positioned posterolateral to choanae;
choanae rounded. Arms slender, subarticular tubercles single; metacarpal tubercles
not recognizable; fingers webbed; webbing formula 1 (1), 2i (1.5), 2e (1), 3i (2.25),
3e (1), 4 (1); relative length of fingers 1 < 2 < 4 < 3, finger 2 distinctly shorter than
finger 4; finger disks distinctly enlarged; unpigmented nuptial pads recognizable on
the inner side of first finger. Hindlimbs slender; tibiotarsal articulation reaches
beyond snout tip when hindlimb is adpressed along the body; lateral metatarsalia
separated by the webbing; inner metatarsal tubercle recognizable, no outer
metatarsal tubercle; webbing formula between toes 1 (0), 2i (0.5), 2e (0), 3i (0.5), 3e
(0), 4i (1.5), 4e (1), 5 (0.25). Skin on the upper surface smooth; ventral skin smooth
on throat where the presence of a vocal sac is clearly recognizable, slightly granu-
lar on belly; no distinct enlarged tubercles in the cloacal region.
After 5 months in preservative, the basic colour is beige-whitish. No further
pigments are present on the flanks, while the whole dorsal side is irregularly cov-
ered with more or less intense dark pigment. This dark colour is interrupted by a
pattern of small pigmentless spots. The area above the eyes is beige to reddish. The
dorsal side of the legs is covered by few small dark pigment cells. The venter is
unpigmented, the inner organs are visible through the translucent belly skin.
Variation. The paratypes agree largely in morphology and colouration with the
holotype. The dark dorsal pattern is strongly expressed in the single female speci-
men (ZFMK 62221) and some males (ZFMK 62220), while in some other males
(ZFMK 60016; Fig. 5) it is less distinct and sometimes relatively faint (ZFMK
60015). A dark inter-ocular stripe or band is always visible, and the dark pattern is
always more intense on the posterior back. Two light dorsolateral bands are visible
in most preserved specimens.
Colour in life. Based on colour slides of several paratype specimens (Fig. 5).
The ground dorsal colour, including limbs, webbing and finger and toe disks is
light greenish, more yellowish towards the flanks. Thin yellow dorsolateral stripes
run from behind the eye to at least the forelimb insertion, sometimes along the
whole body. Additionally, the dorsal surface has a pattern consisting of four general
pigment types, but presence and extent of all of them is very variable. First, a red-
brownish pattern is generally present above the eyes and as an inter-ocular band,
and often as spots and larger markings on the posterior back. Second, a black pat-
tern is often seen as very fine regular spotting on the back (probably consisting of
single melanophores); often, the blackish pigment forms larger and more intense
spots (probably dense layers of melanophores) at the center of the red-brownish

Two new treefrogs from Madagascar
153
markings, i.e. above and between the eyes. Third, a pattern of bright red dorsal
spots is often present; although sometimes intermediate states between these spots
and the red-brown markings are observed, generally the red colour appears similar
to that observed in B. rappiodes. Fourth, small dorsal yellow spots are often also
observed. Few markings are present on fore- and hindlimbs. The venter is translu-
cent. The iris is beige with a turquoise outer iris area and a brownish central mark-
ing in its lower part; the iris periphery is blue.
Etymology. Dedicated to Ursula Bott, ZFMK, in recognition of her invaluable
help during the past 10 years.
Natural history and advertisement calls. Calling males were regularly observed
during the night at Andasibe, 1-3 m high in the vegetation along a slow-moving
stream, in part syntopically with B. rappiodes. Advertisement calls recorded from
ZFMK 62220 at Andasibe on 9 February 1996 (20:30 hr) at 23 °C air temperature
had a complex structure. Note type 1 was a double click which corresponded to
one expiration and was emitted in long trill series of up to 2800 msec (note repeti-
tion rate 8-9/sec). Each note consisted of one longer and one shorter click pulse
(Fig. 6). Duration of the first, longer pulse was 21-27 msec (22 2 msec, n = 15),
duration of the second pulse was 7-11 msec (9 1 msec, n = 15), inter-pulse inter-
val duration was 26-29 msec (27 1 msec, n = 15). Inter-note intervals in trill calls
had a duration of 48-70 msec (54 5 msec, n = 13). The frequency of the long puls-
es was 4050-4450 Hz with a dominant frequency of 4150-4350 Hz, the frequency of
the short pulses was 3600-4900 Hz with a dominant frequency of 4200-4500 Hz.
Notes of type 2 generally consisted of three pulses (the first pulse having a lower
intensity) and were usually emitted at the end of one trill call with a note repetition
rate of about 2/sec. Pulse duration was 7-28 msec (17 9 msec, n = 5) for the first
pulse, 21-26 msec (24 2 msec, n = 5) for the second pulse, and 20-27 msec (24
3 msec, n = 5) for the third pulse. Inter-pulse interval duration was 28-40 msec (33
6 msec, n = 5) between first and second pulse and 14-25 msec (20 4 msec, n =
5) between second and third pulse. Calls recorded on 27 February 1996 at
Vohiparara had an almost identical structure; the duration differences between the
two pulses of note type 1 were less distinct. The vocal sac was single subgular and
remained partly inflated between calls.
Distribution. The species is known from (1) the type locality Andasibe, and
from (2) the Ranomafana area. The latter locality is corroborated by our own call
recordings and photographs from Vohiparara, and by UADBA specimens.
Boophis erythrodactylus (Guibé 1953)
Hyperolius erythrodactylus GUIBÉ 1953. Name-bearing type: lectotype by present designation, MNHN
1994.1469 (originally numbered 1953.171A), adult male, collected in September 1952 by R.
Paulian. Type locality: “forêt de Mahajeby, près de Morafenobe, Ouest de Madagascar” accord-
ing to the original description. Other types: paralectotypes, 1953.171, adult male, and
1994.1470-1471 (originally numbered 1953.171B-C), two subadult specimens, with same col-
lecting data as lectotype.

154
M. Vences and F. Glaw
Remarks. According to GUIBÉ (1953), the original syntype series of Hyperolius
erythrodactylus consisted of three males and one female. We could determine the
sex reliably in two (male) specimens only (MNHN 1953.171 and 1994.1469). These
still show distinctly the many light (originally red; see GUIBÉ 1953) spots scattered
on the dorsum, and a few very faint dark minute spots above the eyes. In contrast,
in the subadults, the light spotted pattern is not recognizable, while the black pig-
ment forms a faint interocular band on the dorsal head surface. Whether these two
specimens are actually conspecific with the lectotype cannot be reliably assessed.
Considering the identification of a sibling species described below as B. tasymena
n. sp., it is important to stabilize the name B. erythrodactylus by designation of a
lectotype. Of the two males, MNHN 1994.1469 was in a better state of preservation
upon examination in May 2001, and we therefore choose it as lectotype.
Identity. Boophis erythrodactylus was defined by GUIBÉ (1953), BLOMMERS-
SCHLÖSSER (1979) and BLOMMERS-SCHLÖSSER & BLANC (1991) largely by its charac-
teristic colour pattern, with numerous red spots of similar size (whitish in ethanol)
on a greenish back. During recent fieldwork, we observed that populations of frogs
with this character differed in advertisement calls. While specimens from Mandra-
ka, on which the observations of BLOMMERS-SCHLÖSSER (1979) were based, emitted
notes composed of 4-7 pulses of rather low frequency (below 3500 Hz), those from
other sites emitted notes composed of two or three click pulses of higher frequency
(above 3500 Hz). DNA sequences (547 nucleotides of the 16S rRNA gene) of one
specimen of each form (from Mandraka and Andasibe, respectively) differed from
each other in 32 substitutions and 2 indels, corresponding to a total pairwise
sequence divergence of 6.1%. The high genetic differentiation and consistent call
differences indicate that the two forms correspond to distinct sibling species. Mor-
phologically, we could not detect any significant difference between them, but the
Mandraka specimens always had red tips of fingers and toes which were absent in
the other individuals. According to GUIBÉ (1953), who examined the type specimens
of B. erythrodactylus shortly after preservation, intense red colour was present on
“l’extrémité des trois doigts externes, ainsi que celle des trois orteils externes”. The
red colour on 2-3 external fingers and toes was also recognizable in May 2001 in
the ZFMK and ZSM specimens from Mandraka examined (see below), 1-7 years
after their fixation and preservation in ethanol. The presence of red colour on some
external or all fingertips of specimens from Mandraka was also mentioned by
BLOMMERS-SCHLÖSSER (1979). In contrast, specimens from Andasibe, Mantady and
An’Ala (all with two-click calls) had no trace of red colour on any finger- or toe-
tips. We therefore consider the species present at Mandraka (Fig. 7) as conspecific
with the types of B. erythrodactylus from Mahajeby forest. Although the latter local-
ity has been said to be in western Madagascar, it appears to be rather on the west-
ern slopes of the central plateau of Madagascar, and at least two species occurring
at this site [Mantidactylus opiparis (Peracca 1893) and a new species of the M.
albofrenatus group; VENCES & GLAW in press] are common in forests above 1000 m
elevation such as the one at Mandraka (ca 1200 m).
Diagnosis. A member of the Boophis rappiodes group as recognizable by small
size (males 20-25 mm, female 32-33 mm; Table 1 and BLOMMERS-SCHLÖSSER 1979),
greenish and slightly translucent dorsal colouration, translucent venter (inner
organs can be clearly seen through the skin in live specimens) and absence of later-
al fringes along lower arm and tarsus. Distinguished from Boophis mandraka by

Two new treefrogs from Madagascar
155
nostril position (closer to snout tip than to eye vs slightly closer to eye), and iris
colour (absence of distinct reticulations on a light beige iris). Distinguished from B.
viridis by smaller size. Distinguished from all species of the B. rappiodes group
except for B. tasymena n. sp. (see below for a distinction from that species) by a
regular dorsal pattern of similarly-sized red spots and red colour of (at least the
external) tips of fingers and toes (Fig. 7).
Material examined. MNHN 1953.171 and 1994.1469-1471 (lectotype and three paralecto-
types from Mahajeby forest); UADBA-MV 2001.206-207 (Mandraka); ZFMK 59813-59814
(Mandraka); ZSM 342/2000 and 677/2001 (Mandraka).
Distribution. The species is at present only known from (1) the type locality
Mahajeby and (2) Mandraka. BLOMMERS-SCHLÖSSER & BLANC (1991) mentioned
three other localities for the species, namely Andasibe (as Perinet), Manjakatompo
and Vondrozo. The distributional information of these authors was almost exclu-
sively based on the ZMA and MNHN holdings which were revised by us. The
Manjakatompo locality was based on tadpoles (ZMA 7205-7206; BLOMMERS-
SCHLÖSSER 1979) which most probably correspond to the species Boophis ankara-
tra Andreone 1993 (GLAW & VENCES 1994 and unpublished data). The Andasibe
locality was based on two males (ZMA 6929) collected along the “highroad R.N. 2
at km 142, alt 1100 m” (BLOMMERS-SCHLÖSSER 1979). Unfortunately, in the ZMA
material, “the red colour has disappeared completely ..., probably because the col-
lection was exposed to the light for some time” (BLOMMERS-SCHLÖSSER 1979).
Therefore it is not possible, at present, to ascertain whether the specimens labeled
ZMA 6929 are to be assigned to B. erythrodactylus or to B. tasymena n. sp. (see
below). As the latter species is common in the Andasibe region, we consider the
presence of B. erythrodactylus as in need of confirmation, although the syntopic
occurrence of both taxa cannot be excluded. The locality Vondrozo is based on
MNHN 1930.432. This specimen is in rather good state of preservation, but the
colour pattern has faded completely. A reliable determination of this specimen
(see Table 1 for measurements) is not possible, and we therefore propose to con-
sider also the Vondrozo locality for B. erythrodactylus as in need of confirmation.
As to the presence of B. erythrodactylus at Ranomafana (ANDREONE 1994), we sup-
pose that this record was based on observations of the very similar B. tasymena n.
sp. (see below) which is common in this area.
Natural history and advertisement calls. Calling males were observed about 2-3
m high in the vegetation along broad, fast-flowing streams, syntopically with B.
mandraka (see also BLOMMERS-SCHLÖSSER 1979). Calls from Mandraka (recorded on
9 February 1994, 21:45 hr, at 23 °C air temperature) were series of usually 4-7 click
pulses (sometimes up to 14 clicks) (Fig. 8). Duration of each pulse was 9-15 msec,
pulse repetition rate was 9-10/sec. Inter-pulse interval duration was 99-109 msec.
The frequency was between 1850 and 3500 msec, and often increased towards the
last pulses of a call.
Boophis tasymena n. sp.
Identity. A B. erythrodactylus-like species with two- or three-pulse calls of high
frequency, and without red tips on fingers and toes (see above).

156
M. Vences and F. Glaw
Fig. 7. — Boophis erythrodactylus, adult male from Mandraka (specimen preserved in UADBA col-
lection).
Fig. 8. — Sonagram and oscillogram of a call of Boophis erythrodactylus, recorded at Mandraka.

Two new treefrogs from Madagascar
157
Diagnosis. A member of the Boophis rappiodes group as recognizable by
small size (males 21-23 mm, one female 32 mm; Table 1), greenish and slightly
translucent dorsal colouration, translucent venter (inner organs can be clearly
seen through the skin in live specimens) and absence of lateral fringes along
lower arm and tarsus. Distinguished from Boophis mandraka by nostril position
(closer to snout tip than to eye vs slightly closer to eye), and iris colour (absence
of distinct reticulations on a light beige iris). Distinguished from B. viridis by
smaller size. Distinguished from all species of the B. rappiodes group except for B.
erythrodactylus by a regular dorsal pattern of similarly-sized red spots. The species
is most similar to B. erythrodactylus, but differs in advertisement calls (2-3 click
pulses vs 4-7 pulses), in the lack of red colour on the tips of fingers and toes (vs
presence at least on the external fingers and toes; Fig. 9), and possibly in a small-
er mean size of males (SVL 21-23 mm vs 24-25 mm in the samples analyzed by
us; Table 1). It furthermore shows important genetic differences from B. erythro-
dactylus (see Identity section of that species) which fully support its status as a
distinct species.
Holotype. ZSM 1085/2001 (originally ZFMK 62224), adult male, collected by F. Glaw on
4 February 1996 at Andasibe, central-eastern Madagascar, 18°56’S, 48°25’E, ca 900 m eleva-
tion (Fig. 9).
Paratypes. ZFMK 62225, adult male, same collecting data as holotype; ZFMK 62243-
62244, two adult males, collected by F. Glaw on 10 February 1996 at Mantady; ZFMK 62262
and 62267, two adult males, collected by F. Glaw on 3 February 1996 at An’Ala; ZFMK 62888,
adult female, collected by F. Glaw, D. Rakotomalala and F. Ranaivojaona on 9-11 March 1996
at Andasibe.
Description of the holotype. SVL 22.7 mm. For measurements, see Table 1.
Body slender; head wider than long, much wider than body; snout moderately trun-
cated in dorsal and lateral views; nostrils directed laterally, slightly protuberant,
nearer to tip of snout than to eye; canthus rostralis relatively indistinct, curved;
loreal region slightly concave; tympanum distinct, rounded, 57% of eye diameter;
supratympanic fold indistinct, relatively straight; tongue ovoid, distinctly bifid pos-
teriorly; rudiments of vomerine teeth can be recognized posterolateral to choanae;
choanae rounded. Arms slender, subarticular tubercles single; metacarpal tubercles
not recognizable; fingers webbed; webbing formula 1 (1), 2i (1.25), 2e (1), 3i (2.25),
3e (2), 4 (1); relative length of fingers 1 < 2 < 4 < 3, finger 2 distinctly shorter than
finger 4; finger disks distinctly enlarged; unpigmented nuptial pads on inner side of
first finger. Hindlimbs slender; tibiotarsal articulation reaches snout tip when
hindlimb is adpressed along the body; lateral metatarsalia separated by the web-
bing; inner metatarsal tubercle present, outer metatarsal tubercle absent; webbing
formula between toes 1 (0), 2i (0.5), 2e (0), 3i (1.25), 3e (0.5), 4i (2), 4e (2), 5 (0.5).
Skin on the upper surface smooth; ventral skin smooth on throat, slightly granular
on belly; no distinct enlarged tubercles in the cloacal region.
After more than 5 years in preservative, the main colouration is a dirty
yellow-white. The whole dorsal area, including fore- and hindlimbs, is stippled with
small, distinct red spots, all of more or less similar size. Large, intense red mark-
ings are present above the eyes, and are connected by a discontinuous red inter-
ocular stripe. The centre of these supraocular markings is covered by blackish pig-
ment. No red colour is present on the tips of fingers and toes. The ventral side is
completely unpigmented.

158
M. Vences and F. Glaw
Fig. 9. — Boophis tasymena n. sp. (holotype ZSM 1085/2001), adult male from Andasibe.
Fig. 10. — Sonagram and oscillogram of a call of Boophis tasymena n. sp., recorded at Andasibe.

Two new treefrogs from Madagascar
159
Variation. The paratypes agree well in morphology and colouration with the
holotype. Most males (in preservative) have red patches above the eyes and a red
inter-ocular line of different width and intensity. Generally, also some dark pigment
is present at these places. In some specimens, three (ZFMK 62262) or four (ZFMK
62225) regularly spaced black spots are present on the tibia. Especially ZFMK
62225 is also characterized by two symmetric dark brown markings on the central
back. The single female (ZFMK 62888) is characterized by the absence of any trace
of supraocular or inter-ocular markings; the sole pattern in this specimen is the
fine regular red spotting all over the dorsal surface. No trace of red colour on the
tips of fingers or toes is visible in any specimen.
Colour in life. Based on colour slides of the holotype and of several paratype
specimens. The ground colour is greenish, including the limbs, webbing and finger
and toe disks. Sometimes, the latter are of a more intense green than the limbs, but
never show any red colour. The ventral side is translucent. Yellow dorsolateral lines
run from the eye to the forelimb insertion. Dorsally two further pigment types are
present to a variable extent: the bright red spotting; reddish-brown markings above
the eyes and as an inter-ocular transversal band, but seldom as symmetric mark-
ings on the back. The iris is beige with a bright light blue outer area and a brown-
ish central marking in its lower part; the iris periphery is blue.
Etymology. Derived from the Malagasy words tasy (pattern, spot) and mena
(red) and referring to the characteristic red spotted pattern. The name is used as an
invariable noun standing in apposition to the generic name.
Natural history and advertisement calls. Calling males were found at night sit-
ting 1-3 m high in trees and shrubs along broad brooks, sometimes at distances of
up to 20 m from the water. Calls recorded at Andasibe on 4 February 1996 at 19:30
hr (24 °C air temperature) and An’Ala on 3 February 1996 at 18:30 hr (23 °C air
temperature) were very similar. They usually consisted of two pulses (Fig. 10).
Pulse duration was 10-24 msec (17 3 msec, n = 46), duration of inter-pulse inter-
vals was 71-97 msec (81 8 msec, n = 23). The frequency ranged between 3850 and
6100 Hz, the dominant frequency was 5000-5500 Hz. Calls recorded at Vohiparara
on 3 March 1996 regularly contained also three-pulse notes. Pulse duration was
5-17 msec (7 3 msec, n = 17), duration of inter-pulse intervals was 63-90 msec (77
9 msec, n = 11). The frequency was 3700-5400 Hz, the dominant frequency 4250-
4900 Hz. The vocal sac was single subgular, and each two- or three-pulse note was
emitted as one expiration. The vocal sac did not remain inflated between calls, but
was slightly inflated before the start of each call.
Distribution. Besides (1) the type locality Andasibe and the localities of
paratypes, (2) Mantady and (3) An’Ala, the species is also known from (4)
Vohiparara. The latter locality is corroborated by field observations and call record-
ings by F. Glaw (see call descriptions above).
DISCUSSION
In addition to the characters used here for distinguishing among the species
of the B. rappiodes group, several others have been used in the literature and will

160
M. Vences and F. Glaw
therefore be discussed in the following. BLOMMERS-SCHLÖSSER (1979) distinguished
B. viridis by a strongly protruding nostril (vs slightly protruding in the other
species). We do not consider this character to be very useful, especially as B.
viridis is easily diagnosed by its size and general appearance. The more strongly
protruding nostrils of B. viridis are recognizable in most individuals, but at least
in one of our specimens, ZFMK 60013, the nostril state did not differ from other
species of the group.
BLOMMERS-SCHLÖSSER (1979) also distinguished B. rappiodes by its shorter
hand length (ratio HAL/SVL 0.26 vs 0.33 in the other species). In our sample
(Table 1), mean relative hand length of B. rappiodes males was actually the lowest
(0.299), but also the values of B. viridis (0.300), B. bottae (0.305) and B. tasymena
males (0.308) were low, with largely overlapping ranges. Although relative hand
length may be of some use to distinguish particular species (e.g., in our sample,
B. mandraka males had a high mean value of 0.340), it is not a very useful diag-
nostic character according to our data.
In contrast, relative tympanum diameter, as used by BLOMMERS-SCHLÖSSER
(1979) to characterize B. erythrodactylus, appears useful also according to our
data set. Instead of the ratio TD/ED as used by BLOMMERS-SCHLÖSSER (1979), we
preferred the ratio TD/SVL, as eye diameter can strongly depend on the state of
fixation and preservation of the specimens. Considering males only, the highest
mean values, 0.071 and 0.070, corresponded to B. erythrodactylus and its sibling
B. tasymena, while the other species of the group ranged between 0.060 and
0.062. Even the minimum values of B. erythrodactylus (0.069) and B. tasymena
(0.067) were higher than those of any other species except for B. rappiodes in
which two individuals attained 0.068-0.069. We therefore confirm that relative
tympanum diameter in B. erythrodactylus and B. tasymena is larger than in the
other species of the group.
A further difference within the B. rappiodes group is relative female size. All
species agree with the general trend that Boophis females are distinctly larger
than males (in contrast to many species of the mantelline genus Mantidactylus).
However, the relative mean male/female size ratio of B. viridis (87%) is distinctly
higher than in the other species of the group (66-73% in B. rappiodes, B. bottae,
B. erythrodactylus and B. tasymena). As female size is similar in B. viridis and the
other species (Table 1), this indicates an evolutionary trend of male size differenti-
ation independently of female size. It may be hypothesized that the derived state
is the large male size with low sexual dimorphism in B. viridis. This species
occurs syntopically with three other species of the B. rappiodes group at Andasibe
(B. rappiodes, B. bottae and B. tasymena) and is bioacoustically the most divergent
species of the group. It is well known that the frequency of vocalizations is
inversely correlated with individual size (DUELLMAN & TRUEB 1986). It is therefore
possible that the size diversification of this species has been influenced by factors
of bioacoustic competition with other syntopic species of the B. rappiodes group.
An alternative explanation for the relatively enlarged size of B. viridis males may
be intraspecific male combats (SHINE 1979) which, however, have not been
observed in this species.
The discovery and description of two additional species of the B. rappiodes
group indicates that the centre of endemism of the group is in central-eastern
Madagascar. Two further new species of the group, siblings to B. mandraka, were
also discovered in this region (this complex of species will be revised elsewhere).
According to the distribution maps of BLOMMERS-SCHLÖSSER & BLANC (1991), only

Two new treefrogs from Madagascar
161
one dubious record of B. mandraka was then known from the Châines Anosyennes
in the extreme south-east. The corresponding vouchers, MNHN 1973.1100-1101
and 1973.1103-1105, have distinct fringes along the lateral edge of the lower arm
and tarsus, which are absent in the B. rappiodes group but typical of the B. luteus
group (BLOMMERS-SCHLÖSSER & BLANC 1991); these specimens may actually belong
to Boophis andohahela Andreone et al. 1995, B. albipunctatus Glaw & Thiesmeier
1993, or B. sibilans Glaw & Thiesmeier 1993. Thus, the record of B. rappiodes in
Nahampoana (GLAW & VENCES 1994) was the first of a species of this group in far
south-eastern Madagascar. Recent surveys recorded species of the group from the
north-east: B. mandraka and B. rappiodes from Marojejy, Anjanaharibe-Sud,
Tsararano, and B. mandraka from Ambolokopatrika (RAXWORTHY et al. 1998,
ANDREONE et al. 2000, RASELIMANANA et al. 2000). However, at these northern and
southern sites, only one or two species were found, while the Andasibe-Mandraka
area in central-eastern Madagascar harbours at least eight species.
The elevational distribution of the group is largely confined to mid-
elevations (700-1200 m). At north-eastern sites, B. rappiodes and B. mandraka
were found between 500 and 1560 m (RAXWORTHY et al. 1998, ANDREONE et al.
2000, RASELIMANANA et al. 2000), while the south-eastern Nahampoana site, located
at ca 300 m above sea level, is an exception. However, other typical mid-elevation-
al species, such as Boophis luteus (Boulenger 1882), also reach lower elevations at
Nahampoana.
Altogether, the new distributional and taxonomic information confirms that
the high faunal diversity in mid-elevational central-eastern Madagascar (LEES 1996)
is not a sampling artefact. On the other hand, the degree of endemism in this
region may be lower than previously thought, and species thought to be exclusive
to the environments of Andasibe are increasingly recorded also from other sites.
According to our data, all species revised herein are known from at least one addi-
tional site, mostly in south-eastern Madagascar (Ranomafana area). The genetic
differentiation (in the 16S rRNA gene; Table 2) of Boophis tasymena to B. erythro-
dactylus and B. bottae to B. rappiodes (6-7%) is of a similar order of magnitude as,
for instance, that of B. erythrodactylus and B. tasymena to B. rappiodes (8%); it is
lower than between B. viridis and the other species (9-10%), but still much higher
than would be expected from the similar morphology of these sibling species. In a
molecular phylogenetic analysis (VENCES et al. 2002), B. bottae was the sister
species of B. rappiodes, and B. tasymena the sister species of B. erythrodactylus, as
would be expected from the similarities in colouration and morphology. When the
Table 2.
Genetic differences between species of the B. rappiodes group. The values are total pairwise diver-
gences in per cent (above diagonal) and total number of substitutions including indels (below diag-
onal) in the analyzed 16S rRNA gene fragment (560 nucleotides).
B. viridis
B. rappiodes
B. bottae B. erythrodactylus B. tasymena
B. viridis
B. rappiodes
—
58
10.4%
—
9.8%
7.3%
9.3%
8.0%
9.8%
8.0%
B. bottae
B. erythrodactylus
B. tasymena
55
52
55
41
45
45
—
45
51
8.5%
—
34
9.1%
6.1%
—

162
M. Vences and F. Glaw
usual rate estimate of 0.6% pairwise sequence divergence per million years is
applied, the origin of these sibling species clearly predates the Pleistocene, as is
common in rainforest species (MORITZ et al. 2000).
ACKNOWLEDGEMENTS
We are grateful to Domoina Rakotomalala who assisted in the field and during exami-
nation of UADBA specimens; to Fara Ranaivojaona and David Vieites for their help in the
field. Wolfgang Böhme (Bonn), Rainer Günther (Berlin), Berthus van Tuijl (Amsterdam),
Alain Dubois and Annemarie Ohler (Paris) allowed examination of specimens held in their
care. We are indebted to the Malagasy authorities for research and export permits. The work
of both authors was made possible by a cooperation accord between the Zoologisches
Forschungsinstitut und Museum A. Koenig, Bonn and the Zoologische Staatssammlung
München, with the Departément de Biologie Animale, Université d’Antananarivo, and was
financially supported by the Deutscher Akademischer Austauschdienst (DAAD).
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