Studies on Neotropical Fauna and Environment

Vol. 44, No. 2, August 2009, 93–107

ORIGINAL ARTICLE
NNFE

A new species of Adenomera (Amphibia: Anura: Leptodactylidae) from the Atlantic

rain forest in Santa Catarina, southern Brazil†
Axel Kweta*, Josefina Steinerb & Anne Zillikensb,c
Studies on Neotropical Fauna and Environment

a
Staatliches Museum für Naturkunde, Abt. Zoologie, Stuttgart, Germany; bDepartamento de Biologia Celular, Embriologia
e Genética, Centro de Ciências Biológicas, Universidade Federal de Santa Catarina, Florianópolis, SC, Brazil; cZoologisches
Institut, Universität Tübingen, Germany
(Received 15 February 2009; accepted 6 March 2009)

As part of the remarkable biodiversity of anurans in the southern range of the Brazilian Mata Atlântica, Adenomera
engelsi sp. nov. is described here as a new taxon occurring on the island of Santa Catarina and adjacent coastal
mainland. This litter-inhabiting frog is restricted to the Atlantic rain forest biome, living in primary and second-
ary woods and semi-open areas from sea level to about 900 m a.s.l., partly in sympatry with its congener A. arau-
caria. Both species are also referred to the Leptodactylus marmoratus group. Adenomera engelsi sp. nov. is a
medium-sized member of the genus (snout–vent length 20.9–22.7 mm in males), characterized by a brown dorsal
coloration with a maculated pattern of variable dark spots and blotches and a unique advertisement call, consist-
ing of single, unpulsed notes with duration of 95–160 ms, dominant frequency between 3500 and 4300 Hz, and
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note repetition rates of 10–24 notes/min.

Está descrita, como parte da enorme biodiversidade de anuros na região sul da Mata Atlântica brasileira,
Adenomera engelsi sp. nov. que ocorre na Ilha de Santa Catarina e no litoral adjacente. A espécie está restrita ao
bioma da Mata Atlântica, onde vive no folhiço em mata primária, secundária e áreas semi-abertas, desde o nível
do mar até 900 m, parcialmente em simpatria com A. araucaria. Ambas as espécies também são atribuidas ao
grupo Leptodactylus marmoratus. Adenomera engelsi sp. nov., de tamanho medio dentro do gênero (SVL 20.9–
22.7 mm em machos), é caraterizada pela coloração dorsal marrom, com manchas escuras de tamanho e forma
variável e um canto de anúncio distinto, que consiste de notas únicas e sem pulsos de 95–160 ms com frequência
dominante entre 3500–4300 Hz e taxa de repetição de 10–24 notas por minuto.
Keywords: advertisement call; Anura; biodiversity; Brazil Mata Atlântica; Leptodactylus marmoratus group

Introduction fauna. Invertebrate and vertebrate taxa, and within the

In the worldwide list of biodiversity hotspots, the Mata
Atlântica is ranked in top position (Myers et al. 2000;
UNESCO Natural Sciences 2007; Carnaval et al. 2009)
due to the species richness and high endemism rates.
As over 90% of the former coastal rain forests have
been cleared, the flora and fauna is highly endangered
today. The biome has the status of a biosphere reserve
and in 1988 was declared a national heritage site by the
Federal Constitution of Brazil. Recently the govern-
ment released a plan for protection and partial refor-
estation of areas with forest remnants (Presidência da
República Brasil 2008). Within the bi-national cooper-
ation program “Science and Technology for the Mata
Atlântica”, funded since 2002 by the CNPq in Brazil
and the BMBF in Germany, a project on internal
dynamics of rain forests was initiated in Santa Catarina
(Zillikens & Steiner 2007). Specificities of animal–plant
interactions are studied, including the diversity of the
latter anurans (Kwet et al. 2005), are subjects of this
long-term project, revealing numerous yet undescribed
species.
One example is the Neotropical frog genus Adeno-
mera Steindachner, 1867, revalidated by Heyer (1974)
for small leaf litter frogs of the Leptodactylus
marmoratus group. Recently, Frost et al. (2006) placed
Adenomera into synonymy with Leptodactylus, propos-
ing the subgenus Leptodactylus (Lithodytes) for the
former genera Adenomera and Lithodytes. However,
Almeida & Angulo (2006), Angulo & Reichle (2008),
and Fouquet et al. (2007) considered the statement of
this subgenus premature as the recommendation of
Frost et al. (2006) was based mainly on Heyer’s (1998)
study, in which the genus Adenomera was represented
by only one species with its taxonomic position not
fully resolved. Hence, Almeida & Angulo (2006) and
Angulo & Reichle (2008) referred to these frogs as the

†In memoriam Marcos Di-Bernardo.

*Corresponding author. Email: kwet.smns@naturkundemuseum-bw.de

ISSN 0165-0521 print/ISSN 1744-5140 online

© 2009 Taylor & Francis
DOI: 10.1080/01650520902901659
http://www.informaworld.com
 94 A. Kwet et al.
Leptodactylus marmoratus species group, a term previ-
ously used already by Heyer (1973). Although it is likely
that Adenomera is nested within a paraphyletic genus
Leptodactylus (Heyer 1998; Kokobum & Giaretta 2005;
A Angulo, personal communication), there are mor-
phological (e.g. Heyer 1974; de Sá et al. 2005), ethologi-
cal (Kokobum & Giaretta 2005), and bioacoustic data
(Kwet & Angulo 2002; Kwet 2007) supporting the idea
that all species of the marmoratus group may form a
natural unit with a single ancestral lineage. de Sá et al.
(2005), Fouquet et al. (2007), and Ponssa & Heyer
(2007) treated both Adenomera and Lithodytes as valid
genera, and molecular analyses (A Fouquet, personal
communication) seem to support the monophyly of
Adenomera as a sister group of the genus Leptodactylus.
Unlike Frost (2008), two recent species descriptions of
L. ajurauna (Berneck et al. 2008) and L. coca (Angulo &
Reichle 2008) and pending further studies, we prefer to
keep the genus Adenomera as a valid taxon (see also
Kwet 2007), comprising currently 13 recognized species
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(= Leptodactylus marmoratus group) of which six are
distributed in the Brazilian Atlantic rain forest domain
(Almeida & Angulo 2006; Kwet 2007; Berneck et al.
2008): A. ajurauna (Berneck, Costa & Garcia, 2008), A.
araucaria Kwet & Angulo, 2002, A. bokermanni (Heyer,
1973), A. marmorata Steindachner, 1867, A. nana
(Müller, 1922), and A. thomei (Almeida & Angulo,
2006).
Due to the cryptic morphology with broadly over-
lapping characteristics and high intraspecific variation
in dorsal pattern and coloration, most of these species
are difficult to identify (Kwet & Angulo 2002; Kok
et al. 2007). However, advertisement calls which play
an important role in species reproductive separation
by mate recognition have been suggested as a useful
means of resolving the systematics of this group (e.g.
Heyer 1984). In fact bioacoustic data have been fre-
quently used for successful species delimitation in pre-
vious studies (Kwet 2001; Kwet & Angulo 2002;
Angulo & Icochea 2003; Angulo et al. 2003; Almeida
& Angulo 2006; Kwet 2007).
Regarding the South Brazilian Atlantic rain forest,
advertisement calls recorded from different popula-
tions of Adenomera in the states of Rio Grande do Sul,
Santa Catarina and southern Paraná (Kwet & Angulo
2002; Kwet 2006, 2007) revealed great variation and
allowed the discrimination of at least five call types,
from males of presumably distinct species. Based on
considerable differences in bioacoustic parameters,
Kwet (2007) revalidated A. nana and suggested the
presence of A. araucaria and three additional, mor-
phologically similar (cryptic) species in this region.
Here we present morphological data for one of these
taxa occurring on the Atlantic island of Santa Cata-
rina and adjacent continental regions of eastern Santa
Catarina. This species, which is morphologically
similar to A. nana, was preliminarily named Adeno-
mera sp. 1 in Kwet (2006, 2007) and was recently
referred to as Leptodactylus sp. 1 grupo marmoratus
by Lucas (2008). It is described here as a new species.
Materials and methods
Adult frogs were collected from 2002 through 2004 at
various sites on the island of Santa Catarina and also
on the nearby mainland. Specimens were fixed and
stored in 70% ethanol and are housed in the Museu de
Ciências e Tecnologia da PUCRS, Porto Alegre, Rio
Grande do Sul (MCP). For morphological compari-
sons, material in the collections of Staatliches Museum
für Naturkunde Stuttgart (SMNS), National Museum
of Natural History, Washington, DC (USNM),
Museum für Naturkunde Berlin (formerly Zoolo-
gisches Museum Berlin, ZMB), Zoologisches Museum
Hamburg (ZMH), and Zoologische Staatssammlung
München (ZSM) was examined. The following meas-
urements were taken to the nearest 0.1 mm with dial
calipers: snout–vent length (SVL); head length (HL);
head width (HW); horizontal eye diameter (ED); hori-
zontal tympanum diameter (TD); eye–nostril distance,
from center of the nostril to the anterior edge of eye
(EN); interorbital distance between anterior edges of
eyes (IO); internarial distance between centers of nos-
trils (IN); thigh length (THL); shank length (TIL); foot
length from proximal edge of inner metatarsal eleva-
tion to tip of fourth toe (FL).
Advertisement calls from more than 20 Adenomera
populations in southern Brazil were recorded during
field studies from 2002 to 2006 using a Sony WM-
D6C tape recorder, a Sennheiser microphone system
K6 with ME66 module, and metal tapes (Sony Metal
XR-90). Bioacoustic parameters for these recordings,
which are referable to five different taxa, are analyzed
in Kwet (2006, 2007) and summarized here (Table 1).
A distribution map and precise recording data are
found in Kwet (2007). The following recordings from
eastern Santa Catarina, referred to as Adenomera sp. 1
in Kwet (2007), belong to the new species described in
this paper: AK15B08 (part) from Lagoa do Perí, Ilha
de Santa Catarina; AK15B12 from Praia dos Naufra-
gados, Ilha de Santa Catarina; AK20A05 from Porto
Belo; AK20B08 from Águas Mornas; AK21A07 from
Taquaras; AK22A19 from Lagoa do Perí, Ilha de
Santa Catarina; AK22B07 from São Pedro de Alcân-
tara; and AK23A04 from Santo Amaro da Imperatriz.
Single calls of the new species from three populations
in Porto Belo (AK20A05), Ilha de Santa Catarina
(AK22A19), and Águas Mornas (AK20B08) are fig-
ured here in comparison with calls of the most similar
congener A. nana from Blumenau (AK22B11),
 Studies on Neotropical Fauna and Environment 95

Table 1. Measurements in mm of Adenomera engelsi sp. nov.; type series, n = 12 males (including holotype MCP 6415).

MCP MCP MCP MCP MCP MCP MCP MCP MCP MCP MCP MCP
Males 6379 6415 6438 6439 6440 7704 7705 8255 8256 8266 8267 8268 Min–Max Mean SD

SVL 22.5 21.9 20.9 22.7 21.3 21.9 21.5 21.3 21.8 21.7 21.0 22.5 20.9–22.7 21.75 0.59
HL 7.9 7.2 7.3 8.1 7.6 7.9 7.6 7.6 7.6 7.7 7.2 8.0 7.2–8.1 7.64 0.30
HW 7.6 7.4 7.3 7.7 7.2 7.6 7.4 7.3 7.2 7.1 7.0 7.6 7.0–7.7 7.37 0.22
ED 1.7 1.8 1.7 2.0 1.9 2.0 1.9 1.9 2.0 2.2 2.2 2.2 1.7–2.2 1.96 0.18
TD 1.2 1.3 1.3 1.6 1.2 1.4 1.3 1.3 1.4 1.5 1.6 1.6 1.2–1.6 1,39 0.15
EN 2.0 2.1 2.0 2.0 2.0 2.0 2.0 2.1 2.1 2.0 2.0 2.1 2.0–2.1 2.03 0.05
IO 4.3 4.2 3.9 4.3 4.2 4.4 4.3 4.3 4.3 4.5 3.8 4.4 3.8–4.5 4.24 0.20
IN 2.2 2.4 2.4 2.1 2.4 2.2 2.3 2.3 2.3 2.4 2.2 2.2 2.1–2.4 2.28 0.10
THL 10.8 10.0 10.1 10.9 10.3 9.9 9.5 9.6 9.3 10.3 9.9 10.0 9.3–10.9 10.05 0.48
TIL 11.3 10.7 10.6 11.4 10.5 10.7 10.7 10.3 10.2 10.5 10.3 10.7 10.2–11.4 10.66 0.37
FL 12.3 12.2 12.4 13.2 11.9 11.9 11.9 11.3 12.3 11.9 11.0 12.0 11.0–13.2 12.03 0.55

Jaraguá do Sul (AK22A01), and Corupá (AK27A17). adult male; MCP 8213, 15 December 2004, munici-
Additionally, a series of consecutive notes of syntopi- pality of Santo Amaro da Imperatriz, subadult.
cal advertisement calls of the new species and A. arau-
caria is figured here from Lagoa do Perí, Ilha de Santa
Catarina (AK15B08). Acoustic analyses were per- Diagnosis
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formed with Cool Edit software and calls were digi-
tized at a sampling rate of 44.1 kHz (Hamming
windowing function set at a resolution of 1024 bands).
Results
Adenomera engelsi sp. nov. Kwet (Figures 1–3)
Adenomera sp. 1: Kwet 2006, 2007.
Leptodactylus sp. 1 grupo marmoratus: Lucas
2008, p. 53–54.
Type series
Holotype
MCP 6415, adult male, collected on 11 October 2002
by Axel Kwet, Anne Zillikens, and Tatiana Miranda,
near Praia dos Naufragados (Ilha de Santa Catarina),
municipality of Florianópolis, state of Santa Catarina,
Brazil (27°49′S, 48°34′W), about 50 m a.s.l.
Paratypes
MCP 6379, 19 October 2002, and 7704–05, 10
November 2003, Praia dos Naufragados (Ilha de
Santa Catarina), municipality of Florianópolis; MCP
6438–40, 13 October 2002, and MCP 8266–68, 2
December 2004, Morro do Perí (Ilha de Santa Cata-
rina), municipality of Florianópolis; MCP 8255–56,
11 December 2004, municipality of São Pedro de
Alcântara. All type specimens are adult males.
Referred specimens
MCP 6344, 10 October 2002, Córrego Grande (Ilha
de Santa Catarina), municipality of Florianópolis,
The new species is assigned to the genus Adenomera
(or Leptodactylus marmoratus species group sensu
Heyer 1973), based on small size, overall morphol-
ogy, and advertisement call features. Within the
genus, it is a robust, medium-sized member (SVL in
males 20.9–22.7 mm, mean 21.75 mm; see Table 1)
being characterized by (1) maculated dorsal pattern
consisting of variably sized, longitudinally arranged
spots and a distinct triangle in the interorbital region,
followed by a chevron-like blotch forming an hour-
glass-shaped figure; (2) presumed absence of distinct
color-pattern polymorphism (a longitudinally striped
morph with light mid-dorsal line and/or broad dorso-
lateral stripes is not known to date); (3) throat uni-
formly cream-colored without dark pigmentation; (4)
tip of the snout in males protruding and ending in a
small whitish shovel (more distinct than in its South
Brazilian congeners); (5) toe tips expanded and
slightly flattened, with small disks; (6) distinct gland
behind the jaw articulation; (7) presumed direct
development of larvae in foam nests, due to the lack
of water bodies near calling males in natural habitats;
(8) distinct and unique advertisement call consisting
of single unpulsed notes with duration of 95–160 ms,
dominant frequency between 3500 and 4300 Hz, and
repetition rates of 10–24 notes/min.
Compared with other species of Adenomera known
from the Brazilian Atlantic rain forest, A. engelsi sp.
nov. differs as follows: (1) from the morphologically
most similar A. nana (Figure 4A–E) by the larger size
(SVL in male A. nana 16.3–19.4 mm, mean 18.27 mm;
Kwet 2007), different advertisement call with lower
dominant frequency, more acuminate snout profile,
and dorsal surface with more spotted pattern, espe-
cially on the posterior part of the body; (2) from the
 96 A. Kwet et al.
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Figure 1. Adenomera engelsi sp. nov., holotype MCP 6415, adult male: (A) dorsal view; (B) ventral view; (C) lateral view; (D)
ventral view of right hand; (E) ventral view of right foot. Scale bars: 1 mm (D, E).
 Studies on Neotropical Fauna and Environment 97
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Figure 2. Adenomera engelsi sp. nov., variation in dorsal pattern of the preserved type series. First row from left to right: MCP
6438–40 and MCP 8266–68 from Morro do Perí, Ilha de Santa Catarina. Second row from left to right: MCP 6415 (holotype),
MCP 6379, and MCP 7704–05 from Praia dos Naufragados, Ilha de Santa Catarina; MCP 8255–56 from São Pedro de
Alcântara.

partly sympatrically occurring A. araucaria (Figure
4A, B, F–H) by the larger size (SVL in male A. arau-
caria 17.1–18.8 mm, mean 18.4 mm; Kwet & Angulo
2002), different advertisement call without pulses or
notable frequency modulation (call with severe, dis-
tinct frequency and amplitude modulations in A. arau-
caria; Kwet & Angulo 2002), absence of a
longitudinally striped morph with light dorsolateral
stripes, more acuminate snout profile, and toe tips
that are more expanded and partly disked (toe tips
rounded but not flattened or disked in A. araucaria);
(3) from A. bokermanni by the smaller size (maximum
SVL 25.1 mm in A. bokermanni, see Heyer 1973,
versus maximum SVL 22.7 mm in A. engelsi sp. nov.)
and expanded, flattened toe tips with small disks (nei-
ther expanded nor flattened in A. bokermanni); (4)
from A. marmorata by the more acuminate snout pro-
file, the distinct and strongly developed gland behind
the jaw articulation, and absence of a longitudinally
striped morph with light dorsolateral stripes; (5) from
A. ajurauna by the larger size (SVL 17.2–20 mm in A.
ajurauna), the whitish throat (dark brown in A.
ajurauna), and the lighter dorsal coloration with dis-
tinct, dark pattern (uniform pattern without dark
markings in A. ajurauna); (6) from A. thomei by the
expanded and flattened toe tips (neither expanded nor
flattened in A. thomei) and the presumed terrestrial
reproductive mode (aquatic mode in A. thomei; see
Almeida & Angulo 2006); (7) from A. diptyx from
adjacent Argentina and Paraguay by the absence of a
distinct light mid-dorsal line (present in this species)
and the presumed terrestrial reproductive mode
(aquatic mode in A. diptyx; see de la Riva 1995, 1996).
Description
Description of the holotype: adult male, MCP 6415
(Figures 1, 2A, B), measurements see Table 1. Body
robust. Head nearly as long as broad; outline of
snout slightly rounded, subelliptical to subacuminate
(Figure 1A, B); snout profile acuminate, tip of the
snout protruding and ending in a small but distinct,
whitish shovel (Figure 1C). Nostrils anterior,
directed dorsolaterally, closer to tip of the snout than
 98 A. Kwet et al.
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Figure 3. Adenomera engelsi sp. nov., variation in living specimens: (A) MCP 6414, holotype, dorsal; (B) MCP 6414, holo-
type, ventral; (C) MCP 8255, male, São Pedro de Alcântara; (D) MCP 8267, male, Ilha de Santa Catarina, Lagoa do Perí; (E)
MCP 8266, male, Ilha de Santa Catarina, Lagoa do Perí; (F) MCP 6344, male with unusual pattern, Ilha de Santa Catarina,
Córrego Grande; (G) MCP 6438, calling male, Ilha de Santa Catarina, Lagoa do Perí; (H) MCP 8213, subadult, Santo Amaro
da Imperatriz.
 Studies on Neotropical Fauna and Environment 99
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Figure 4. Comparison between three similar, partly sympatrically occurring species of Adenomera in Santa Catarina: (A) dorsal
views of A. araucaria (left), Ilha de Santa Catarina, Florianópolis; A. engelsi sp. nov. (middle), Ilha de Santa Catarina, Lagoa do
Perí; and A. nana (right), Jaragua do Sul; (B) ventral views of A. araucaria (left), Ilha de Santa Catarina, Florianópolis; A. engelsi
sp. nov. (middle), Ilha de Santa Catarina, Lagoa do Perí; and A. nana (right), Jaragua do Sul; (C) two males of A. nana from
Corupá with typical dorsal pattern; (D) male of A. nana from Blumenau with striped pattern; (E) male of A. nana from Blumenau
with less distinct pattern; (F) male of Adenomera araucaria from Pró-Mata, São Francisco de Paula; (G) male of Adenomera arau-
caria from Santo Amaro da Imperatriz; (H) male of Adenomera araucaria from São Pedro de Alcantâra with striped pattern.
 100 A. Kwet et al.
to the eye. Internarial distance about one-third of
head width. Tympanum distinct, its diameter two-
thirds of eye diameter. Supratympanic fold poorly
developed, highlighted with black from back of eye
to arm insertion. Canthus rostralis indistinct,
rounded, cranial crests absent; loreal region slightly
concave. Distinct, cream-coloured, oval gland at jaw
articulation. Vocal sac single, internal; paired, elon-
gate vocal slits. Vomerine teeth in two short trans-
verse series, posterior to choanae and separated from
each other by about two-thirds the length of one
tooth row. Tongue elongate-ovoid. Fingers relatively
short; finger lengths III > I > II > IV; fingers free,
without webbing or fringes, finger tips rounded,
slightly widened, but not expanded to disks (Figure
1D). Two large, ovoid metacarpal tubercles; size of
inner metacarpal tubercle about two-thirds of outer
metacarpal tubercle. Prominent, rounded subarticu-
lar tubercles on fingers, more pronounced on the
thumb. Nuptial asperities absent. Hindlegs short,
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thigh slightly longer than shank. Toe lengths IV > III
> V > II > I; toe tips bulbous, expanded and slightly
flattened to ungrooved discs; toes free, without web-
bing or fringes (Figure 1E). Two distinct, ovoid met-
atarsal tubercles and well-developed, rounded
subarticular tubercles; size of outer metatarsal tuber-
cle about two-thirds of inner metatarsal tubercle.
Distinct inner tarsal fold with a single row of small
white-tipped tubercles on crest running about two-
thirds of tarsus length. Tarsal and foot texture gran-
ular, with a profusion of small, but distinct white-
tipped tubercles. Upper shank and ventral surfaces
smooth, without perceptible tubercles. Dorsal tex-
ture slightly rugose ranging from smooth to granular
with some larger tubercles arranged in two longitudi-
nal, dorsolateral glandular folds extending from
behind the eye, over tympanum and shoulder, to
inguinal region. In the preserved specimen (Figure
1A), pustules and glandular folds are considerably
less marked than in life (Figure 3A).
Coloration in life
Dorsal coloration greyish-brown, with orange-brown
hints on snout and limbs, and with symmetrically
arranged dark patterns of marks and blotches (Figure
3A). A black triangular interorbital spot with whitish
margins and an irregular lighter-colored center is fol-
lowed by a pair of irregularly formed blotches between
arm insertions that are joined to a chevron-shaped
mark forming an hourglass-shaped figure. Several
smaller spots on dorsum, partly longitudinally
arranged into two dorsolateral lines running
discontinuously along the glandular folds from
behind the eyes to the inguinal region. Indistinct, light
mid-dorsal hairline stripe on posterior third of body,
between anus and sacrum. Distinct cream-colored
gland at jaw articulation. Two dark cross-bars on
each shank and on thigh; several dark spots on arms.
Ventral coloration whitish, immaculate (Figure 3B),
skin on throat with yellowish hue but without dark
pigments. Ventral surface of thigh greyish, ventrola-
terally dotted with small melanophores.
Coloration in preservative
In 70% ethanol, orange hint of dorsal coloration
turned to light greyish-brown, with the dark pattern
of spots, blotches and cross-bars becoming more dis-
tinct. Ventral coloration white, immaculate, without
yellowish or greyish hues.
Variation
Measurements were taken from the type series of 12
males (Table 1). According to these data, A. engelsi
sp. nov. can reach a maximum SVL of 22.7 mm in
males. Females could not be collected yet. In the hol-
otype, HW is a bit larger than HL, whereas in all
other specimens HL is slightly larger than HW. The
maculated dorsal coloration is variable (Figures 2, 3)
consisting of a conspicuous pattern of longitudinally
arranged, irregular, dark marks on orange-brown or
light to dark brown background. In most specimens
the dark triangular blotch in the interorbital region
presents one to three light small spots centrally, and
the following pair of dark dorsal blotches between
arm insertions is joined mid-dorsally forming a
chevron-shaped mark. In some specimens this
chevron-like blotch is in contact with the larger trian-
gular spot, enforcing the overall appearance of an
hourglass-shaped figure in the region between eyes
and shoulders. Contrary to A. araucaria and A. nana
(Kwet & Angulo 2002; Kwet 2007), where two differ-
ent dorsal patterns occur (striped pattern with two
broad, whitish, dorsolateral lines and pattern with-
out such longitudinal stripes), light dorsolateral
stripes are absent in all known specimens of A.
engelsi sp. nov. Although MCP 6438, MCP 6339,
MCP 6379, and the holotype present an indistinct,
whitish mid-dorsal hairline stripe from above the
vent to sacrum, most specimens are lacking this mid-
dorsal stripe. Dorsolateral glandular folds are varia-
bly developed in the type series. In summary, despite
evident similarity with the already-described conge-
ners, especially the partially sympatric or even syn-
topic species, quite a number of characters in size,
morphology, coloration, and calls substantiate the
bioacoustic discrimination of A. engelsi sp. nov. as
an own species.

Etymology
We are pleased to dedicate the new frog species to Prof.
Wolf Engels (University of Tübingen, Germany), in
recognition of his contributions to the knowledge of
the fauna and flora of southern Brazil, and for sup-
porting our herpetological research in Rio Grande do
Sul and Santa Catarina.
Distribution and conservation
The new species occurs in the subtropical Atlantic rain
forest biome of southern Brazil, living in primary and
secondary forests of the coastal mountain range, from
sea level to about 900 m a.s.l. To date, it is known only
from east of Santa Catarina state, i.e. from the island
of Ilha de Santa Catarina and adjacent continental
regions (mesoregion of Grande Florianópolis and
extreme south of adjacent mesoregion of Vale do
Itajaí). The western-most record is from Taquaras
(municipality of Rancho Queimado) and, based on
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call recordings without voucher specimens, the distri-
bution limit for A. engelsi sp. nov. to the north is near
Porto Belo, and to the south at Serra do Tabuleiro
(municipality of Santo Amaro da Imperatriz). To
date, there are no records of the new species from the
Araucaria highlands (mesoregion of Serrana) with
forests of Araucaria angustifolia where its congener
A. araucaria occurs. However, at many other locali-
ties, e.g. on the Ilha de Santa Catarina, A. engelsi sp.
nov. lives in sympatry with A. araucaria. The morpho-
logically most similar A. nana and two additional spe-
cies seem to have a parapatric distribution in
northeastern Santa Catarina with the lowland region
of Vale do Itajaí as a presumed geographic barrier.
Adenomera engelsi sp. nov. might be classified as
“Least Concern” according to the criteria established
by IUCN (2008). Although its extent of occurrence
may be less than 5000 km2, at present the new species
does not appear to be in decline. It is locally common
with presumed large population sizes occurring in
protected areas such as Parque Estadual da Serra do
Tabuleiro where the habitats are not threatened, and
it seems to be adaptable to habitat alterations.
Natural history
Adenomera engelsi sp. nov. is a ground-dweller
endemic to subtropical rain forests and semi-open
bush-land in the southeast of Santa Catarina. During
breeding time in the warm summer months, males
were observed from November to February calling on
the ground, usually hidden under fallen leaves or in
dense vegetation on the forest floor. The diurnal and
nocturnal activity of this species is strongly correlated
Studies on Neotropical Fauna and Environment 101
with rainfall, with increasing activity after heavy
showers. Calling generally starts in the afternoon with
increasing call frequency until the late evening, then
decreasing until around midnight. Eggs and larvae of
the new species were not observed, but the lack of
water bodies near calling males and in natural habitats
suggest the direct development of larvae in foam nests
as described for A. marmorata (Lutz 1947). However,
many details of the reproductive biology of A. engelsi
sp. nov. have to be clarified in future studies.
Advertisement call
The advertisement call of Adenomera engelsi sp.
nov. consists of single, unpulsed notes with durations
of 95–160 ms that are emitted at low calling rates of
10–24 notes/min (Table 2; Figures 5, 6). The call,
which has been already described as Adenomera sp. 1
in Kwet (2006, 2007), has a dominant frequency
between 3500 and 4300 Hz; the fundamental fre-
quency (first harmonic) lies at about 2000 Hz; other
harmonics are found between 5400 and 6400 Hz, at
around 8000 Hz, and at 10,000 Hz. The notes are
lacking significant amplitude modulation and present
only a weak frequency modulation, meaning that,
after initially rising, the frequency falls to about the
same level as at the beginning of the note.
In most localities, the advertisement calls of
Adenomera engelsi sp. nov. could be heard synchro-
nously with the calls of A. araucaria which was
described by Kwet & Angulo (2002) and Kwet (2007).
Both calls are figured here from an area of syntopy on
Ilha de Santa Catarina (Figure 6A, B). They show
similar note durations (95–160 ms versus 64–140 ms
in A. araucaria), but the call of A. engelsi sp. nov. has
a noticeable lower dominant frequency (3500–4300
Hz, versus 4200–5600 Hz) and a lower call rate, with
10–24 notes/min (26–72 notes/min in A. araucaria).
Additionally, the call of A. araucaria presents a dis-
tinct upward frequency modulation of 500–1200 Hz
(A. engelsi sp. nov. without notable frequency modu-
lation), and a severe amplitude modulation which
could take the form of 6–18 weak, irregular pulses
(A. engelsi sp. nov. with unpulsed notes or a weak
amplitude modulation in the form of two or three
indistinct pulses). These bioacoustic differences are
easily perceivable to the human ear.
On the other hand, the advertisement calls of
A. engelsi sp. nov. and A. nana (both described in Kwet
2007) sound similar and are figured here for direct com-
parisons from six different populations (Figure 5A, B).
Both calls are characterized by the absence of a distinct
frequency modulation and by the low calling rates
which are a bit higher in A. nana (10–24 notes/min in A.
engelsi sp. nov., versus 18–37 notes/min in A. nana). All
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102
A. Kwet et al.
Table 2. (A) Acoustic parameters from different populations of Adenomera engelsi sp. nov. and (B) comparison (condensed data) between Adenomera populations in Rio
Grande do Sul, Santa Catarina, and southern Paraná (Guaratuba) which are referable to five different taxa; for better comparisons only recordings obtained with air
temperatures between 19–25°C are included.

Air No. of calls Dominant Fundamental Other

temperature (no. of Call duration Pulses frequency frequency frequencies Frequency
Identification (°C) individuals) (ms) Calls/min per call (Hz) (Hz) (Hz) modulation

(A) Adenomera engelsi sp. n.

Taquaras 25.5 10 (1) 147.0 (134–155) 18–20 1 3780–3860 – 5700–5850 0–50
Águas Mornas 17 10 (1) 130.0 (122–145) 18 1 3460–4150 – 5400–6040 30–50
Santo Amaro 20 3 (1) 129.7 (123–133) 12–14 1 4130–4290 – 6200–6400 20–40
Ilha de Santa Catarina 22–23 20 (2) 115.6 (96–133) 18–24 1 3850–4240 – 6000–6240 0
São Pedro de Alcantara 17 10 (1) 138.3 (112–163) 10–12 1 3820–4050 – 5800–6200 0–20
Porto Belo 20.5 9 (1) 124.6 (118–134) 14 1 3970–4190 – 6020–6200 60
(B)
Adenomera araucaria 19–23.7 64 (7) 115.4 (64–220) 22–72 6–18 4200–5650 2200–2830 6990–7960 500–1200
Adenomera nana 19–25 78 (8) 95.3 (70–122) 18–30 1 4620–5440 2400–2700 6900–7900 0–80
Adenomera engelsi sp. n. 20–23 32 (4) 123.3 (96–134) 12–24 1 3850–4290 – 6000–6400 0–60
Adenomera bokermanni 24.5 10 (1) 71.6 (67.7–79.1) 93 1 2900–3050 1440–1610 5950–6120 (−150–0)
“Pirabeiraba”
Adenomera sp. 23 10 (1) 31.6 (30.6–32.3) 120 6 4000–4200 – – 300
“Guaratuba”

Note: Adenomera bokermanni “Pirabeiraba” shows negative values for frequency modulation because frequency is falling with time; numbers in brackets under call duration refer to ranges.
Data source: Kwet & Angulo (2002); Kwet (2006, 2007).
 Studies on Neotropical Fauna and Environment 103
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Figure 5. Comparison between the audiospectrograms (A) and oscillograms (B) of three notes of the advertisement call of
Adenomera engelsi sp. nov. (from left to right) from Porto Belo (AK20A05, 20.5°C air temperature), Ilha de Santa Catarina
(AK22A19, 22°C air temperature), and Águas Mornas (AK20B08, 17°C air temperature), compared with three notes from
specimens of Adenomera nana from Blumenau (AK22B11, 21°C air temperature), Jaraguá do Sul (AK22A01, 20°C air temper-
ature), and Corupá (AK27A17, 18.8 °C air temperature); see Kwet (2007) for precise recording data.

calls recorded from populations of A. nana showed
considerably higher dominant frequencies than those
from A. engelsi sp. nov. without overlapping parame-
ters (dominant frequency 4620–5440 Hz in A. nana,
versus 3500–4300 Hz in A. engelsi sp. nov.). In addi-
tion, the calls of A. nana generally present shorter call
durations (70–120 ms in A. nana, versus 95–160 ms in
A. engelsi sp. nov.) and higher call rates (18–30 notes/
min in A. nana, versus 10–24 notes/min in A. engelsi sp.
nov.). The harmonic structure in both calls is also dif-
ferent; in A. nana the fundamental frequency (first har-
monic) lies at about 2500 Hz (2000 Hz in A. engelsi sp.
nov.), harmonic frequency bands are found at about
7500 Hz (between 5400 and 6400 Hz in A. engelsi sp.
nov.) and at 10,000–10,500 Hz (two frequency bands at
8000 and at 10,000 Hz in A. engelsi sp. nov.).
Because in frogs male advertisement calls are
important cues for reproductive cooperation between
both sexes, clear differences in these courtship songs
indicate an established function as interspecific mat-
ing barriers. Female choice requires at least a sure
recognition of conspecific males (Ryan 2001). The
described differences in the male voices of closely
related Adenomera species verify in particular our
proposal of separating a new taxon A. engelsi sp. nov.
This is particularly evident in recordings from a site
of syntopic occurrence of two species where males of
both taxa called simultaneously. The species-specific
calls are clearly discerned (Figure 6A, B).
Discussion
The ingenious naturalist Johann Friedrich Theodor
Müller, known as Fritz Müller, observed and repeat-
edly described the anuran biodiversity of Santa Cat-
arina around Desterro (today Florianópolis) and
Blumenau during decades of research in the sub-
tropical rain forests, which at that time in the
 104 A. Kwet et al.
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Figure 6. Audiospectrogram (A) and oscillogram (B) of a series of consecutive notes of advertisement calls of Adenomera
engelsi sp. nov. (en) and syntopical A. araucaria (ar) from Lagoa do Perí, Ilha de Santa Catarina (AK15B08); recorded on 16
October 2002, 17:30 h, 22°C air temperature.
nineteenth century were nearly untouched. Müller
observed how ostracods (Elpidium) can move from
one bromeliad pool to another by attaching to frogs
and other plant visitors (Müller 1879), and he com-
municated many discoveries to Charles Darwin, as
documented in their long-lasting correspondence,
e.g. the observation of a marsupial frog (Flectono-
tus) near Blumenau (which he referred to as
Hylodes? sic), where the embryos developed in a
pouch on the mother’s back, or a “toadlet being
characterized by an unusual long tail” which he
found on humid stones near a cascate (presumably a
tadpole of Cycloramphus in late development stage
with hindlegs, shortly before metamorphosis (Zillig
1997, p. 214).
Despite severe deforestation, even today the
southern range of the Mata Atlântica houses a rich
anuran fauna. A first survey in the Araucaria forest
of the Pró-Mata reserve on the Araucaria Plateau in
the Serra Geral revealed 32 species of Anura in this
area of a mere 5000 ha (Kwet & Di-Bernardo 1999),
47 species when including surrounding areas (Kwet
2001), and actually 76 species when comprising the
whole anuran hotspot of northeastern Rio Grande
do Sul, an area of roughly 300 km2 (Kwet 2004). In
comparison, a recently published checklist com-
prises 84 species recorded for the entire state of Rio
Grande do Sul (Machado & Maltchik 2007), and in
the state of Santa Catarina the anuran biodiversity
at least totals 110 species (Lucas 2008). However,
since the taxonomy of many taxa requires a careful
revision, and more field studies will undoubtedly
result in the detection of new species, we are far
from a sound knowledge of the total biodiversity of
amphibians in this hotspot area which for Santa
Catarina may comprise over 140 species (Lucas
2008). In particular, the numerous cases of rather
similar species can be resolved only by application

of suitable discrimination approaches such as
molecular data or bioacoustic analyses.
This is meaningfully demonstrated in our presen-
tation of a new Adenomera species which required
consideration of quite a number of similar and pre-
sumably closely related congeners. In fact the extens-
ive morphological variation with broadly overlapping
characteristics in different populations of the small
leaf litter frogs, assigned to the genus Adenomera or
to the Leptodactylus marmoratus group, makes this
species complex a taxonomically very difficult group
(Heyer 1973, 1977, 1984). However, bioacoustic anal-
yses have been successfully used for clarifying the spe-
cies’ identity in different zoogeographic regions, such
as Peru (Angulo & Icochea 2003; Angulo et al. 2003),
French Guyana (Boistel et al. 2006), and Brazil (Kwet
& Angulo 2002; Almeida & Angulo 2006). Based on
call recordings, two or more Adenomera species could
be recognized at most study sites.
For the Brazilian Atlantic rain forest biome, Heyer
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(1973) reported two species occurring in sympatry, A.
marmorata and A. bokermanni, but these nominal spe-
cies have been later suspected to be composites of
three or more species (Heyer 1977, 1984). This
assumption was recently confirmed by the revalida-
tion of A. nana (Kwet 2007) and the descriptions of A.
araucaria (Kwet & Angulo 2002), A. thomei (Almeida
& Angulo 2006), A. ajurauna (Berneck et al. 2008),
and A. engelsi sp. nov. within this group. Such species
complexes presumably point at rather recent separa-
tion processes as described for several other species
groups, e.g. tree frogs of the Hypsiboas pulchellus
group (Kwet 2004, 2008; Garcia et al. 2007).
Kwet (2006, 2007; see summarized data in Table 2)
compared the advertisement calls from more than 20
different Adenomera populations in eastern and cent-
ral Santa Catarina and adjacent regions of Paraná
and Rio Grande do Sul. This bioacoustic analysis
revealed remarkable differences in temporal and spec-
tral characteristics depending on the recording local-
ity, and the data, along with subtle morphological
differences (Appendix 1), supported the occurrence of
five distinct taxonomic units in the Atlantic rain for-
est biome of southern Brazil. Besides A. araucaria, A.
engelsi sp. nov. and A. nana, two additional taxa seem
to occur in this region.
A call with uncertain identity recorded from Pir-
abeiraba, extreme northeastern Santa Catarina, by
Ronald W. Heyer which was analyzed as type I call in
Kwet & Angulo (2002) and own recordings (AK27B04)
obtained recently from Joinville near Pirabeiraba,
present the same spectral and temporal patterns, mean-
ing they are conspecific. The unpulsed call of this spe-
cies which was referred to as Adenomera sp. 2 in Kwet
(2007) showed very high call rates (between 90 and 160
Studies on Neotropical Fauna and Environment 105
calls/min), a weak frequency modulation (with negative
value because of the falling frequency) with low domi-
nant frequency of about 3000 Hz, and a distinct har-
monic structure (see Table 2). This call belongs to A.
bokermanni as suggested in Lucas (2008, p. 46) and
recently confrimed by unpublished recordings from J.
Zina (personal communication).
Another differing recording from Guaratuba,
southern Paraná (Table 2), was described as Adeno-
mera sp. 3 in Kwet (2006, 2007) and might belong to
an undescribed species. It is characterized by high
calling rates (about 120 notes/min), short note dura-
tions (30–35 ms), a dominant frequency of 4000–4200
Hz, and a characteristic upward frequency modula-
tion of about 300 Hz towards the end of the note.
Some of these five species of Adenomera seem to
have overlapping distribution ranges living partly in
sympatry or even in syntopy. The southernmost
member of this genus, A. araucaria, occurs from
northeastern Rio Grande do Sul to southern and
central-eastern Santa Catarina (Kwet & Angulo 2002;
Kwet 2007), whereas A. engelsi sp. nov. is restricted to
the island of Ilha de Santa Catarina and adjacent
continental regions. Both species occur sympatrically
on the island of Ilha de Santa Catarina and nearby
mainland areas. Adenomera nana is parapatrically dis-
tributed in northeastern Santa Catarina and adjacent
southern and southern-central Paraná (unpublished
recordings from R Lingnau and CE Conte, personal
communication). Call recordings from Joinville,
Pirabeiraba (SC), and Guaratuba (PR) suggest at
least two additional taxa in the border region between
Santa Catarina and Paraná, besides A. bokermanni a
presumably undescribed species. Further field studies
are needed for resolving the taxonomic status of these
populations.
Acknowledgements
This study is part of the project “Internal Dynamics
of Rain Forest: Specificity of Animal–Plant Interac-
tion” within the Brazilian–German program “Mata
Atlântica” in cooperation with the Brasilien-Zentrum
of the University of Tübingen, and we acknowledge
the financial support of BMBF (01LB0205) and
CNPq (690143/01-0). We would also like to thank
Wolf Engels, Andreas Schlüter, Rodrigo Lingnau,
Mirco Solé, and Tatiana Miranda for support and
valuable field assistance, and Sabine Heinle for help
with the figures. Ariadne Angulo, Mirco Solé, and
Andreas Schlüter reviewed an early draft of this man-
uscript. Glaucia F. Pontes (MCP), W. Ronald Heyer
(USNM), Rainer Günther (ZMB), Jakob Haller-
mann (ZMH) and Frank Glaw (ZSM) kindly allowed
access to collections under their care.
 106 A. Kwet et al.
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Appendix 1. Additional specimens examined
Adenomera araucaria – Brazil: Rio Grande do Sul: São Francisco
de Paula, CPCN Pró-Mata: MCP 2421 (holotype), MCP 1794,
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Studies on Neotropical Fauna and Environment 107
1849, 3208–09, 3463, 3672–73, 3676–77; Bom Jesus, Encruzil-
hada das Antas: MCP 3345–46. Santa Catarina: Ilha de Santa
Catarina: MCP 1340, 8125, 8262, 8285–86; Santo Amaro da
Imperatriz: MCP 8212.
Adenomera marmorata – Brazil: Rio de Janeiro: NMW 16453
(holotype). São Paulo: Salesópolis, Boracéia: MCP 6658–61,
USNM 209116–20.
Adenomera nana – Brazil: Santa Catarina: Blumenau: MCP
8717–20; Corupá (Rio Novo, Colonia Hansa): ZMB 66829–
844, 66950, 66976–67011, 67814–16, ZMH A01737–44,
A01746–50, ZSM 661/1920/3 (lectotype), ZSM 661/1920/1–2
(paralectotypes); Ilhota, Morro do Baú: MCP 1345–52;
Jaraguá do Sul: MCP 8149–50; Joinville: MCP 8633; Pirabeir-
aba: USNM 243737–39, 243741–42; São Bento do Sul: MCP
8751–55.
Adenomera sp. – Brazil: Paraná: Guaratuba: MCP 7688-89.