Acta Herpetologica 10(1): 67-72, 2015

ISSN 1827-9635 (print) © Firenze University Press 
ISSN 1827-9643 (online) www.fupress.com/ah

DOI: 10.13128/Acta_Herpetol-15547

Re-description of the external morphology of Phyllomedusa iheringii 
Boulenger, 1885 larvae (Anura: Hylidae), with comments on the external 
morphology of tadpoles of the P. burmeisteri group

Samanta Iop¹,*, Victor Mendes Lipinski¹, Bruno Madalozzo¹, Franciele Pereira Maragno¹, Sonia Zanini 
Cechin¹, Tiago Gomes Dos Santos²

¹ Universidade Federal de Santa Maria.  Programa de Pós Graduação em Biodiversidade Animal – Centro de Ciências Naturais e Exa-
tas. Prédio 17, sala 1140 - D, Cidade Universitária Camobi, Km 29 CEP: 97105-900 - Santa Maria, RS – Brasil. *Corresponding author. 
E-mail: samantaiop@yahoo.com
² Universidade Federal do Pampa, Campus São Gabriel. Av. Antônio Trilha, 1847 CEP: 97300-000 – São Gabriel, RS – Brasil

Submitted on 2015, 12th February; revised on 2015, 28th April; accepted on 2015, 30th April 
Editor: Sebastiano Salvidio

Abstract. Phyllomedusa iheringii is a leaf frog endemic to the Uruguayan Savanna, which reproduces in water bod-
ies in open areas. Here, based on the lack of some informative characteristics observed on the first description of 
this species, we re-describe the larval external morphology of P. iheringii from localities in Brazil and Uruguay, and 
compared them with other species from the P. burmeisteri group. The tadpoles of P. iheringii belong to the suspension-
rasper guild. The body length corresponds to approximately one-third of the total length. The body is pyriform in dor-
sal view and laterally triangular. The snout is rounded in a dorsolateral view. The spiracle is single and almost ventral, 
not forming a free tube, and sinistral. The opening of vent tube is dextral. The oral disc is anteroventral and anteriorly 
directed, with single ventral emargination. Marginal papillae uniseriate, interrupted in a wide dorsal gap, and with 
pointed tips. The labial tooth row formula is 2(2)/3(1). The third lower row is four times shorter than the others. Fur-
thermore, the tadpoles of P. iheringii showed morphological differences in relation to other species of the P. burmeis-
teri group, this species seems to be smaller in average and have fewer marginal papillae in the oral discs.

Keywords. Tadpole morphology, Phyllomedusinae, Uruguayan Savanna ecoregion.

The genus Phyllomedusa currently has 30 species and 
is widely distributed in the Neotropics, occurring from 
Panama, the Pacific slopes of Colombia, South America 
east of the Andes, and southwards to northern Argenti-
na and Uruguay (Frost, 2014). Species of this genus are 
distinguished from other Neotropical hylids by their con-
trasting colour patterns, vertical pupils, and an arboreal 
reproductive mode (Caramaschi, 2006). In Brazil, 23 spe-
cies of Phyllomedusa are recorded and all have external 
larval morphology known (Segalla et al., 2012). Never-
theless, larval internal oral features and chondrocranium 
are poorly known for most species. Similarly, the larval 
morphology still requires to be described or re-described, 

because some previous descriptions raise doubts concern-
ing the identification of some structures, as is the case of 
P. iheringii.

The group of Phyllomedusa burmeisteri comprises 
five species (P. bahiana, P. burmeisteri, P. distincta, P. iher-
ingii, and P. tetraploidea) that share similarities in exter-
nal morphology, vocalization, and cytogenetic features, 
and possess a complex pattern of strips on the thighs as 
a diagnostic characteristic (Pombal Jr. and Haddad, 1992; 
Silva-Filho and Juncá, 2006; Faivovich et al., 2010). Spe-
cies of the P. burmeisteri group show a parapatric or 
allopatric distribution throughout the Atlantic Rainforest 
and replace each other throughout the geographic distri-



68 Samanta Iop et alii

bution of the group (Brunes et al., 2010). Phyllomedusa 
iheringii is the only species of the group that occurs in 
open areas and is endemic to the Uruguayan Savanna, 
occurring in Uruguay and southern of the Brazilian state 
of Rio Grande do Sul (Olson et al., 2001; Brunes et al., 
2010; Frost, 2014).

Phyllomedusa iheringii reproduces in both lentic or 
lotic water bodies (but in backwater zones), and uses 
marginal vegetation for both vocalization and spawning 
(Maneyro and Carreira, 2012). Males call whilst perched 
on vegetation and the eggs are laid in leaf nests. Tadpoles 
are exotrophic and drop into lentic water to complete 
their development (mode 24; Haddad and Prado, 2005). 
The first description of the P. iheringii tadpole was pro-
vided by de Sá and Gerhau (1983) was based on a sin-
gle individual hatched from on single egg mass, from one 
single locality. This first effort to describe the species, lack 
of information concerning the sizes, lenght, and posi-
tions of some body structures treated as diagnostic char-
acteristics. The dataset provided may not represent with 
precision the morphological intraspecific variation of the 
species within its distributional range and may have lim-
ited the understanding of intraspecific variation as well 
as inter-specific comparisons with other species of the 
group. Here, we re-describe the external morphology of 
the tadpole of P. iheringii, accounting for intraspecific 
variation and providing comparisons with other species 
of the P. burmeisteri group. In addition, we present data 
on natural history.

The tadpoles of Phyllomedusa iheringii were col-
lected at four sites in the Uruguayan Savanna: Encruzil-
hada do Sul (30°33’50.7’’ S, 52°34’13.1’’ W, 414 m a.s.l.; 
about 100 km from the type locality; ZUFSM 8166), 
Santa Maria (29°49’25.3” S, 53°37’19.3” W, 229 m a.s.l.; 
ZUFSM8831), and São Sepé (30°15’27.5’’ S, 53°34’52.4’’ 
W, 346 m a.s.l.; ZUFSM 8832-3) in Rio Grande do Sul, 
Brazil; and in Cerro Chapeu, Rivera, Uruguay (30°57’11’’ 
S,  55°28’22’’ W, 257 m a.s.l.; RML 2448). The larvae were 
collected and immediately fixed in a formalin 10% solu-
tion in field.

The re-description was based on eight specimens 
of  stage  37 (Gosner, 1960) (seven individuals from São 
Sepé and one from Rivera),  from which 21 morphologi-
cal measurements were taken which according to  McDi-
armid and Altig (1999), and Altig (2007) and Lavilla and 
Scrocchi (1986). To allow  comparison  within  P. iheringii 
and with other species from P. burmeisteri  group,  we also 
took the same measures of other 32 individuals (stages 
26-31, 34-36; Gosner, 1960) (Table 1). Tadpole measure-
ments were made using a stereomicroscope (0.01 mm 
precision), except for TL, which was measured with a 
digital caliper (0.01 mm precision) (Table 2).

External morphology

The tadpoles of Phyllomedusa iheringii belong to 
the suspension-rasper guild (McDiarmid and Altig 
1999; Both et al., 2011), and have an elongated body 
(BH/BW: 0.92; Tab. 1). The body length corresponds 
to approximately one-third of the total length (BL/TL: 
0.34). The body is piriform in dorsal view and triangu-
lar in lateral view (Fig. 1A, B). The snout is rounded in 
a dorsolateral view. The nostrils are rounded and dor-
solateral (ND: 0.25  ±  0.04). The internostril distance is 
larger than the distance between the nostrils and the 
snout (IND: 3.60  ±  0.22; NSD: 1.44  ±  0.27). Eyes are lat-
eral (ED: 2.15  ±  0.19). The interorbital distance is greater 
than the distance between the eyes and the snout (IOD: 
7.75  ±  0.41; ESD: 6.16  ±  0.35). The spiracle is sinistral, 
single and almost ventral, not forming a free tube. The 
opening of vent tube is dextral. The tail is pointed, with 
flagellum, and longer and taller than the body (TAL/
TL: 0.66; MTH/BH: 1.33). The tail musculature is high-
er than wide (TMH/TMW: 1.16). The dorsal fin origi-
nates anterior of dorsal tail-body junction, the ventral fin 
is higher than the dorsal fin (VMH: 5.93  ±  1.01; DMH: 
1.74  ±  0.24). The oral disc is anteroventral and anteriorly 
directed, not emarginate. Marginal papillae uniseriate, 
interrupted in a wide dorsal gap, and with pointed tips. 
Submarginal papillae laterally aggregate in the oral disc, 
not forming rows. The upper jaw sheath is curved, but 
flat in the middle, not serrated and its width is approxi-
mately seven times greater than high (UJSW/UJSH: 7.86). 
The lower jaw has a ‘v’ shape, not serrated and approxi-
mately six times longer than high (LLJ/HLJ: 6.06). The 
labial tooth row formula is 2(2)/3(1) and the third lower 
row is four times shorter than the others (Fig. 1C).

Coloration

Usually, the dorsum of the body and tail varies from 
yellowish to greenish. The lateral portion of the body is 
grey-violet, with brown pigmented areas in the middle 
of fin. The fins are translucent and have small, uniformly 
distributed melanophores. The intestine region is blu-
ish in lateral and ventral views. There are two darkened 
blotches on the head, between the eyes and the nostrils. 
When preserved in formalin (10%), the tadpoles are uni-
formly yellowish or greyish.

Variation

The tadpoles of Phyllomedusa iheringii do not have 
ontogenetic variation, though geographic variation 



69The tadpole of Phyllomedusa iheringii 

occurs. Such variation may be the result of different envi-
ronmental conditions (temperature, pH), food availability 
or presence of predators among water bodies. Although 

tadpoles from the three localities are morphologically 
similar, some of them have small variations (Table 1). 
Indeed, some individuals in different developmental stag-

Table 1. Morphological measurements of 40 tadpoles of Phyllomedusa iheringii from different localities of the Uruguayan Savana. Mean, 
standard deviation and range (mm) are shown. The stage and sample size are in parentheses. BH – body height, BL – body length, BW – 
body width, DMH – dorsal fin height, ED – eye diameter, ESD – eye–snout distance, HLJ –lower jaw sheath height, IND – inter-narial dis-
tance, IOD – interorbital distance, LLJ –lower jaw sheath length, MTH – maximum tail height, ND – nostril diameter, NSD – nostril–snout 
distance, OD – oral disc diameter, TAL – tail length, TL – total length, TMH – tail musculature height, TMW – tail musculature width, 
UJSH – upper jaw sheath height, UJSW – upper jaw sheath width, VMH – ventral fin height.

Encruzilhada do Sul
(stage 27, n = 1)

Santa Maria
(stage 27, n = 3)

São Sepé
(stage 26-31, n = 18)

São Sepé
(stage 34-36, n = 7)

Rivera
(stage 36, n = 3)

São Sepé and Rivera
(stage 37, n = 8)

BH 5.60
4.80 ± 0.12 

(4.67 – 5.60)
6.45 ± 0.71

(4.74 – 7.41)
7.07 ± 0.94

(8.52 - 6.04)
9.77 ± 0.93

(9.16 – 10.84)
7.74 ± 0.64

(8.35 – 6.41)

BL 8.60
6.85 ± 0.14 

(6.72 – 8.60)
11.48 ± 1.54

(8.85 – 13.86)
14.78 ± 1.26

(16.79 – 13.43)
16.05 ± 1.24

(14.77 – 17.25)
15.02 ± 1.86

(16.51 – 10.95)

BW 5.08
4.45 ± 0.10

 (4.39 – 5.08)
6.87 ± 0.69

(5.30 – 7.83)
8.25 ± 0.81

(9.23 – 6.94)
8.69 ± 0.33

(8.31 – 8.89)
8.42 ± 0.76

(9.39 – 7.32)

DMH 1.15
0.95 ± 0.07

 (0.87 – 1.15)
1.27 ± 0.14

(1.00 – 1.28)
1.59 ± 0.14

(1.76 – 1.34)
1.66 ± 0.12

(1.56 – 1.80)
1.74 ± 0.24
(2.06 – 1.4)

ED 1.60
1.37 ± 0.03

 (1.29 – 1.60)
2.03 ± 0.35

(1.29 – 2.42)
2.55 ± 0.27

(2.74 - 2.09)
2.10 ± 0.14

(2.02 – 2.26)
2.50 ± 0.19

(2.77 – 2.18)

ESD 3.47
3.00 ± 0.16

 (2.87 – 3.47)
4.73 ± 0.65

(3.54 – 5.85)
5.97 ± 0.51

(6.58 – 5.25)
5.52 ± 0.37

(5.26 – 5.94)
6.16 ± 0.35
(6.9 – 5.75)

HLJ 0.13
0.10 ± 0.01

 (0.09 – 0.13)
0.14 ± 0.05

(0.10 – 0.29)
0.16 ± 0.06
(0.3 – 0.11)

0.15 ± 0.03
(0.13 – 0.19)

0.15 ± 0.04
(0.19 – 0.1)

IND 2.00
1.80 ± 0.02

(1.78 – 2.00)
2.95 ± 0.29

(2.32 – 3.44)
3.40 ± 0.22

(3.08 - 3.18)
3.61 ± 0.11

(3.52 – 3.73)
3.60 ± 0.22

(3.84 – 3.22)

IOD 3.88
3.69 ± 0.03

(3.65 – 3.88)
5.73 ± 0.7

(4.42 – 6.63)
7.36 ± 0.79

(8.61  –  6.02)
7.09 ± 0.53

(6.48  –  7.46)
7.75 ± 0.41

(8.32  –  7.2)

LLJ 0.58
0.58 ± 0.08

(0.54 – 0.67)
0.73 ± 0.18

(0.54 – 1.28)
0.90 ± 0.14

(1.2  –  0.81)
1.21 ± 0.25

(0.93  –  1.42)
0.91 ± 0.09

(1.05  –  0.77)

MTH 6.92
5.50 ± 0.13

(3.72 – 6.92)
7.78 ± 1.29

(3.72 – 9.15)
9.88 ± 0.70

(10.48  –  8.6)
9.06 ± 1.30

(8.06  –  10.53)
10.33 ± 1.26

(12.22  –  8.38)

ND 0.13
0.14 ± 0.03 

(0.11 – 0.16)
0.20 ± 0.06

(0.13 – 0.33)
0.23 ± 0.04

(0.23  –  0.18)
0.21 ± 0.02

(0.20  –  0.23)
0.25 ± 0.04

(0.31  –  0.21)

NSD 0.92
0.56 ± 0.24

(0.40 – 0.92)
1.09 ± 0.25

(0.75 – 1.54)
1.49 ± 0.24

(1.74  –  1.5)
1.47 ± 0.07

(1.41  –  1.55)
1.44 ± 0.27

(1.85  –  1.1)

OD 1.28
1.49 ± 0.21

(1.25  –  1.62)
2.58 ± 0.50

(1.77 – 4.15)
3.21 ± 0.62

(4.57  –  2.79)
3.41 ± 0.68

(2.89  –  4.18)
3.12 ± 0.18

(3.32  –  2.82)

TAL 15.70
15.58 ± 1.02  

(14.41 – 16.26)
23 ± 3.12

(17.02 – 29.73)
28.21 ± 3.35

(32.78  –  23.64)
23.53 ± 0.86

(22.72  –  24.44)
29.61 ± 2.06

(32.28  –  26.98)

TL 24.30
22.43 ± 1.13  

(21.13 – 24.30)
34.48 ± 4.01

(26.11 – 40.2)
42.99 ± 4.41

(48.93  –  37.07)
39.58 ± 2.10

(37.49  –  41.69)
44.63 ± 2.55

(47.73  –  40.88)

TMH 2.85
2.21 ± 0.18

(2.09 – 2.85)
3.63 ± 0.73
(2.32 – 4.5)

4.70 ± 0.56
(5.02  –  3.74)

3.11 ± 0.14
(3.00  –  3.27)

4.52 ± 0.50
(4.86  –  3.3)

TMW 1.99
1.52 ± 0.24

(1.30 – 1.99)
3.02 ± 0.56

(2.02 – 3.82)
3.99 ± 0.58

(4.5  –  3.03)
2.43 ± 0.24

(2.20  –  2.67)
3.88 ± 0.59

(4.79  –  2.85)

UJSH 0.16
0.12 ± 0.03

(0.10 – 0.16)
0.23 ± 0.07

(0.14 – 0.46)
0.44 ± 0.53

(0.36  –  0.210)
0.31 ± 0.06

(0.24  –  0.36)
0.22 ± 0.04

(0.29  –  0.18)

UJSW 0.99
0.84 ± 0.01

(0.22 – 0.99)
1.39 ± 0.31

(0.97 – 2.38)
1.54 ± 0.68

(2.53  –  1.58)
1.95 ± 0.36

(1.68  –  2.36)
1.73 ± 0.07

(1.82  –  1.63)

VMH 3.33
2.63 ± 0.14

(2.30 – 3.33)
4.02 ± 0.56

(2.80 – 4.88)
5.05 ± 0.42

(5.26  –  4.92)
4.87 ± 1.17

(3.75  –  6.09)
5.93 ± 1.01

(7.55  –  4.08)



70 Samanta Iop et alii

es (25, 26, 31, 36 and 37) have an oral formula of 2(2)/3, 
instead of the usual formula of 2(2)/3(1), and the third 
lower row in some cases, not linked to a specific devel-

opmental stage, is much smaller in comparison with the 
other ones. The marginal papillae can be light white or 
have several melanophores. Most individuals have a few 
blotches on the body and tail, but many others had fewer 
of these markings, which were less dark. Some individu-
als from São Sepé had melanophores on the ventral fins 
and blueish intestines. However, specimens from Encru-
zilhada do Sul, Santa Maria, and Cerro Chapeu had no 
melanophores on the ventral fins, and the intestine 
ranged from brown to black. 

Natural history

Tadpoles of Phyllomedusa iheringii occurred through-
out the spring and summer of the austral region (from 
September to March), and inhabit deeper and non-vege-
tated areas of ponds, pools of streamlets, and small dams 
used in cattle raising. At least, larvae of other 13 anuran 
species occurred syntopically with larvae of Phyllome-
dusa iheringii at the studied waterbodies: Dendropso-
phus minutus, Elachistocleis bicolor, Hypsiboas pulchellus, 
Leptodactylus gracilis, L. latrans, Odontophrynus ameri-
canus, Physalaemus cuvieri, P. gracilis, P. henselii, Pseu-
dopaludicola falcipes, Scinax fuscovarius, S. granulatus, 
and S. uruguayus. Larvae usually forage near the water 
surface (head up, tilted upwards at 45º), but dive quickly 
when disturbed (without forming schools). On two occa-
sions, we observed predation on tadpoles of P. iheringii 
in a temporary pond at São Sepé: in the first incident 
(December 2003), a tadpole was devoured by a beetle 
larva (Dytiscidae) early in the night, near the water sur-

Table 2. Comparison of external morphological characteristics between tadpoles from the Phyllomedusa burmeisteri species group. *The 
original description uses the stage VIII of Rugh (1951). Stages were assessed according to Gosner (1960).

Species Stage Oral formula Ventral gap Marginal papillae Reference

Phyllomedusa bahiana Lutz, 1925 34–36 2(2)/3(1) Present
Mostly uniserial; biserial at 
emargination

Silva – Filho and Juncá, 2006

Phyllomedusa burmeisteri 
Boulenger, 1882

35 2(2)/3(1) Absent
Two series, with four or more 
series on the labial emargination

Cruz, 1982

Phyllomedusa distincta A. Lutz in 
B. Lutz, 1950

37 2(2)/3(1) Absent
Two series, with a few 
scattered papillae on the labial 
emargination

Cruz, 1982

Phyllomedusa iheringii Boulenger, 
1885

28* 2(2)/3 Absent Uniserial de Sá and Gerdau, 1983

Phyllomedusa iheringii Boulenger, 
1885

37
2(2)/3(1)

2(2)/3
Absent

Uniserial, with submarginal 
papillae laterally aggregate in the 
oral disc, not forming rows

Present study

Phyllomedusa tetraploidea Pombal 
Jr. and Haddad, 1992

37 2(2)/3(1) Absent
Uniserial, with numerous series 
on the emargination

Pombal Jr. and Haddad, 1992

*The original description uses the stage VIII of Rugh (1951). Stages were assessed according to Gosner (1960).

Fig. 1. Tadpole of Phyllomedusa iheringii (museum catalog ZUFSM 
10015) at Gosner stage 36: (A) dorsal view (scale = 10 mm); (B) lat-
eral view (scale  =  10 mm); (C) oral disc (scale  =  1 mm). Drawnigs 
by B.M. and T.G.S



71The tadpole of Phyllomedusa iheringii 

face; the second occasion (November 2011) occurred in 
the afternoon, when a Great Kiskadee bird (Pitangus sul-
furatus, Tyrannidae) repeatedly plunged into the water to 
catch and eat several tadpoles.

The morphological characteristics presented here for 
Phyllomedusa iheringii are similar to those reported by de 
Sá and Gerhau (1983). However, we found variation in 
the labial tooth row formula (Table 2), the color of papil-
lae, body, tail, fins, and the intestine (which ranges from 
black to blue). The variation in coloration is probably 
related to the structural characteristics of the pond and to 
avoid predation and competitors (Thibaudeau and Altig, 
2012). Indeed, experiments among tadpoles of several 
species in the presence of predators exhibit differences in 
coloration and morphology (McCollum and Leimberger, 
1997; Van Buskirk and McCollum, 2000; Touchon and 
Warkentin, 2008). Tadpoles assessed here were collected 
in ponds with distinct structural variables and subject to 
different predation pressures, and thus we suggest that 
future studies could test the adaptive value of tadpole col-
oring on different pressures of predators and competitors. 

Several characteristics of the Phyllomedusa iheringii 
tadpole are shared with other species of the P. burmeis-
teri group, such as the triangular shape of the body in 
dorsal view, ventral fin higher than the dorsal fin, and 
the oral formula 2(2)/3(1), with a reduced third row of 
lower teeth and dorsal gap (Cruz, 1982; de Sá and Ger-
hau, 1983; Pombal Jr. and Haddad, 1992; Silva-Filho and 
Juncá, 2006). However, the tadpole of P. iheringii is dis-
tinguished from other species of the P. burmeisteri group 
by fewer marginal papillae in the oral disc (only one 
series), whereas P. burmeisteri has two rows of papillae 
in the anterior and posterior portion of the oral disc, and 
four series on the labial emargination (Cruz, 1982); P. 
bahiana has one row in the anterior and posterior region 
and a double row on the labial emargination (Silva-Filho 
and Juncá, 2006); P. distincta has double series (Cruz, 
1982); P. tetraploidea has one row, but with numerous 
rows on the labial emargination (Pombal Jr. and Haddad, 
1992), and P. bahiana is the only species in the group that 
has a ventral gap of marginal papillae (Silva-Filho and 
Juncá, 2006). In summary, the tadpole of P. iheringii is 
distinct from other species of the P. burmeisteri group by 
the oral disc, mainly by the number of marginal papillae 
and the labial emargination.

The species belonging to the genus Phyllomedusa 
possess complex polypeptides on the skin that can be 
toxic and cause irritation, which might potentially pro-
tect the species against predation (Duellman and Trueb, 
1986; Caramaschi and Cruz, 2002). However, several spe-
cies of vertebrates and invertebrates prey on eggs, larvae, 
and adults of Phyllomedusa (Castanho, 1996; Feltrim and 

Cechin, 2000; Toledo et al., 2005; Dias et al., 2012). Here, 
we add two predators of larvae of this genus (a Great 
Kiskadee bird and a beetle larva). Until now, odonate 
naiads, belostomatid bugs, spiders and fishes were identi-
fied as predators of tadpoles of Phyllomedusa (Azevedo-
Ramos et al., 1992; Gascon, 1992; Schmidt and Amézqui-
ta, 2001; Santos-Silva et al., 2013)

Tadpoles of Phyllomedusa iheringii are similar to 
those of other species of P. burmeisteri group, but smaller 
in average and with fewer marginal papillae in the oral 
disc (not emarginated). Variation (between populations 
and individuals) on larvae of P. iheringii was primarily 
related to size, the oral disc formula (and the size of the 
third lower row) and the color pattern of the intestines 
and fins. In this sense, we believe our results add impor-
tant information about the external morphology and 
natural history of P. iheringii, an endemic species of the 
Uruguayan Savanna ecoregion. We suggest future studies 
to focus on frog species that are relatively common in the 
Neotropics, such as P. iheringii, but that lack basic infor-
mation as aspects of behavior, reproductive biology, and 
morphological descriptions of tadpoles. 

ACKNOWLEDGEMENTS

We are grateful to Raúl Maneyro Landó for pro-
viding specimens from Uruguay. S.I, V.M.L, B.M and 
F.P.M. thank CAPES, for the doctoral fellowships. T.G.S 
and S.Z.C. are grateful to CNPq for research fellowships 
(proc. 307352/2013-7 and 304929/2012-3, respectively). 
Collecting permits (#29509 and 29508) were provided by 
the Instituto Chico Mendes de Conservação da Biodiver-
sidade (ICMBio). We thank to the anonymous reviewers 
for their comments and suggestions on the manuscript.

REFERENCES

Altig, R. (2007): A primer for the morphology of anuran 
tadpoles. Herpetol. Conserv. Biol. 2: 71-74.

Azevedo-Ramos, C., Van Sluys, M.,  Magnusson, W.E.  
(1992): Influence of  tadpole movement on predation 
by odonate naiads. J. Herpetol. 26: 335-338.

Both, C., Melo, A.S., Cechin, S.Z., Hartz, S.M. (2011): 
Tadpole co-occurrence in ponds: When do guilds and 
time matter? Acta Oecol. 37: 140-145. 

Brunes, T.O., Sequeira, F., Haddad, C.F.B., Alexandrino, J. 
(2010): Gene and species trees of a Neotropical group 
of treefrogs: genetic diversification in the Brazilian 
Atlantic Forest and the origin of a polyploid species. 
Mol. Phyl. Evol. 57: 1120 -1133. 



72 Samanta Iop et alii

Caramaschi, U. (2006): Redefinição do grupo de Phyl-
lomedusa hypochondrialis, com redescrição de P. meg-
acephala (Miranda-Ribeiro, 1926), revalidação de P. 
azurea Cope, 1862 e descrição de uma nova espécie 
(Amphibia, Anura, Hylidae). Arq. Mus. Nac. Rio de 
Janeiro 64: 159-179. 

Caramaschi, U., Cruz, C.A.G. (2002): Phyllomedusa: 
posição taxonômica, hábitos e biologia (Amphibia, 
Anura, Hylidae ). Phyllomedusa 1: 5-10. 

Castanho, L.M. (1996): Phyllomedusa distincta (Leaf-
Frog). Predation. Herpetol. Rev. 27: 141. 

Cruz, C.A.G. (1982): Conceituação de grupos de espécies 
de Phyllomedusinae brasileiras com base em carac-
teres larvários (Amphibia, Anura, Hylidae). Arq. Univ. 
Fed. Rur. Rio de Janeiro 5: 147-171.

de Sá, R., Gerhau, A. (1983): Observaciones sobre la bio-
logia de Phyllomedusa iheringii Boulenger 1885 (Anu-
ra, Hylidae). Bol. Soc. Zool. Urug. 1: 44-49. 

Dias, T.M., Maragno, F.P., Santos, T.G., Cechin, S.Z. 
(2012): Phyllomedusa iheringii: egg predation. Herpe-
tol. Bull. 121: 38-40.

Duellman, W.E., Trueb, L. (1986): Biology of amphibians. 
McGraw-Hill, Baltimore and London.

Faivovich, J., Haddad, C.F.B., Baêta, D., Jungfer, K-H, 
Álvares, G.F.R., Brandão, R.A., Sheil, C., Barrientos, 
L.S., BarrioAmorós, C.L., Cruz, C.A.G., Wheeler, W.C. 
(2010): The phylogenetic relationships of the charis-
matic poster frogs, Phyllomedusinae (Anura, Hyli-
dae). Cladistics 26: 227-261.

Feltrim, A.C., Cechin, S.Z. (2000): Helicops infrataeniatus. 
Diet. Herpetol. Rev. 31: 46. 

Frost, D.R. (2014): Amphibian species of the world: an 
online reference. Version 5.6 (9 January 2013). Elec-
tronic database accessible at <http://research.amnh.
org/herpetology/amphibia/index.html>. American 
Museum of Natural History, New York, USA.

Gascon, C. (1992): Aquatic  predators  and  tadpole  prey  
in central  Amazonia:  field  data  and experimental  
manipulations. Ecology 73: 971-980.

Gosner, K.L. (1960): A simplified table for staging anuran 
embryos and larvae with notes on identification. Her-
petologica 16: 183-190. 

Haddad, C.F.B., Prado, C.P.A. (2005): Reproductive 
modes in frogs and their unexpected diversity in the 
Atlantic Forest of Brazil. BioScience 55: 207-217. 

Lavilla, E.O., Scrocchi, G.J. (1986): Morfometría larval de 
los géneros de Telmatobiinae (Anura: Leptodactyli-
dae) de Argentina y Chile. Physis 44: 39-43.

Maneyro, R., Carreira, S. (2012): Guía de anfíbios del 
Uruguay. Ediciones de la Fuga, Montevideo. 

McCollum, S.A., Leimberger, J.D. (1997): Predator-
induced morphological changes in an amphibian: pre-

dation by dragonflies affects tadpole shape and color. 
Oecologia 109: 615-621.

McDiarmid, R.W., Altig, R. (1999): Tadpoles: The biol-
ogy af anuran larvae. The University of Chicago Press, 
Chicago. 

Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Bur-
gess, N.D., Powell, G.V.N., Underwood, E.C., D’amico, 
J. A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, 
C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, 
J.F., Wettengel, W.W., Hedao, P., Kassem, K.R. (2001): 
Terrestrial ecoregions of the world: a new map of life 
on earth. BioScience 51: 933-938. 

Pombal Jr., J.P., Haddad, C.F.B. (1992): Espécies de Phyl-
lomedusa do grupo burmeisteri do Brasil oriental, 
com descrição de uma espécie nova (Amphibia, Hyli-
dae). Rev. Brasil. Biol. 52: 217-229. 

Rugh, R.  (1951): The frog. Its reproduction and develop-
ment. The Blakiston Company, Philadelphia. 

Santos-Silva, C.R, Santos, E.D.S., Gomes, L., Ferrari, S.F. 
(2013): Predation of a Phyllomedusa nordestina tad-
pole (Anura, Hylidae) by a fishing spider, Thaumasia 
sp. (Areneae, Pisauridae), in a temporary pond in the 
Raso da Catarina, Bahia, Brazil. Herpetol. Notes 6: 
193-194.

Schmidt, B., Amézquita, A. (2001): Predator-induced 
behavioural responses: tadpoles of the neotropical 
frog Phyllomedusa tarsius do not respond to all preda-
tors. Herpetol. J. 11: 9-15.

Segalla, M.V., Caramaschi, U.C., Carlos, A.G., Garcia, 
P.C.A., Grant, T., Haddad, C.F.B., Langone, J. (2012): 
Brazilian amphibians: list of species. Electronic data-
base accessible at <http://www.sbherpetologia.org.br>. 
Sociedade Brasileira de Herpetologia, São Paulo, Bra-
zil.

Silva-Filho, I.S.N., Juncá, F.A. (2006): Evidence of full 
species status of the neotropical leaf-frog Phyllome-
dusa burmeisteri bahiana (A. Lutz, 1925) (Amphibia, 
Anura, Hylidae). Zootaxa 1113: 51-64. 

Thibaudeau, G., Altig, R. (2012): Coloration of anuran 
tadpoles (Amphibia): Development, dynamics, func-
tion, and hypotheses. ISRN Zoology 2012: 1-16.

Toledo, L.F., Woehl Jr., G., Woehl, E.N., Prado, C.P.A. 
(2005): Scinax nasicus, Hyla albomarginara, Hyla bis-
choffi and Phyllomedusa distincta (Tree frogs): avian 
predation. Herpetol. Bull. 92: 31-32. 

Touchon, J.C., Warkentin, K.M. (2008): Fish and dragon-
fly nymph predators induce opposite shifts in color 
and morphology of tadpoles. Oikos 117: 634-640.

Van Buskirk, J., McCollum, S.A. (2000). Functional 
mechanisms of an inducible defence in tadpoles: mor-
phology and behaviour influence mortality risk from 
predation. J. Evol. Biol. 13: 336-347.


	Acta Herpetologica
	Vol. 10, n. 1 - June 2015
	Firenze University Press
	Obituary: Valery K. Eremchenko (1949-2014)
	Leo J. Borkin1, Tatjana Dujsebayeva2, Roberto Sindaco3, Matthias Stöck4
	Haplotype variation in founders of the Mauremys annamensis population kept in European Zoos 
	Barbora Somerová1, Ivan Rehák2,*, Petr Velenský2, Klára Palupčíková1, Tomáš Protiva1, Daniel Frynta1
	Reproductive ecology of Sichuan digging frogs (Microhylidae: Kaloula rugifera)
	Wei Chen1,*, Lina Ren2, Dujuan He2, Ying Wang2, David Pike3
	Toxic effects of carbaryl on the histology of testes of Bufotes variabilis (Anura: Bufonidae)
	Özlem Çakici
	Basal frequency of micronuclei and hematological parameters in the Side-necked Turtle, Phrynops hilarii (Duméril & Bibron, 1835)
	María A. Latorre 1,2,*,#; Evelyn C. López González1,2, #; Pablo A. Siroski1,2,3; Gisela L. Poletta1,2,4
	Into a box interiors: clutch size variation and resource allocation in the European pond turtle
	Marco. A.L. Zuffi1,*, Simonetta Citi2, Elena Foschi1, Francesca Marsiglia1, Eva Martelli1
	Where to “Rock”? Choice of retreat sites by a gecko in a semi-arid habitat
	Andreia Penado1,2, Ricardo Rocha3,4,*, Marta Sampaio3, Vanessa Gil3, Bruno M. Carreira3, Rui Rebelo3
	Age structure, growth and longevity in the common toad, Rhinella arenarum, from Argentina 
	Clarisa de L. Bionda1,2,*, Silvia Kost 4, Nancy E. Salas1, Rafael C. Lajmanovich3, Ulrich Sinsch4, Adolfo L. Martino1
	On a putative type specimen of Pleurodema bibroni Tschudi, 1838 from Chile (Anura: Leptodactylidae)
	Daiana Paola Ferraro
	Re-description of the external morphology of Phyllomedusa iheringii Boulenger, 1885 larvae (Anura: Hylidae), with comments on the external morphology of tadpoles of the P. burmeisteri group
	Samanta Iop¹, Victor Mendes Lipinski¹, Bruno Madalozzo¹, Franciele Pereira Maragno¹, Sonia Zanini Cechin¹, Tiago Gomes Dos Santos²
	Book Review: 
	Harold Heatwole, John W. Wilkinson (Eds). Amphibians Biology. Volume 11 - Status of conservation and decline of Amphibians. Eastern Hemisphere. Part 4 . Southern Europe and Turkey
	Sebastiano Salvidio
	Book Review: 
	Antonio Romano. Atlante degli anfibi del Parco Nazionale del Cilento Vallo di Diano e Alburni - Distribuzione, biologia, ecologia e conservazione
	Sebastiano Salvidio
	ACTA HERPETOLOGICA
	Journal of the Societas Herpetologica Italica