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A New Species Of The Rhinella Margaritifera Group (Anura: Bufonidae) From

Brazilian Pantanal

Article in Zootaxa · January 2010

DOI: 10.5281/zenodo.193265

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 Zootaxa 2339: 57–68 (2010) ISSN 1175-5326 (print edition)
www.mapress.com / zootaxa/ Article ZOOTAXA
Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

A new species of the Rhinella margaritifera group (Anura: Bufonidae) from

Brazilian Pantanal

ROBSON W. ÁVILA1,4, ANDRÉ PANSONATO2 & CHRISTINE STRÜSSMANN2,.3

1
Departamento de Parasitologia, Instituto de Biociências, Universidade Estadual Paulista, Distrito de Rubião Jr., s/nº, CEP 18618-
000, Botucatu, SP, Brazil
2
Pós-Graduação em Ecologia e Conservação da Biodiversidade, Universidade Federal de Mato Grosso, Av. Fernando Corrêa da
Costa, s/nº, CEP 78060-900, Cuiabá, MT, Brazil
3
Departamento de Ciências Básicas e Produção Animal, Faculdade de Agronomia e Medicina Veterinária, Universidade Federal de
Mato Grosso, Av. Fernando Corrêa da Costa, s/nº, CEP 78060-900, Cuiabá, MT, Brazil
4
Corresponding author. E-mail: robsonavila@gmail.com

Abstract

We describe adult morphology, advertisement call and some natural history traits of a new species of toad from the
Pantanal, western Brazil. Rhinella paraguayensis sp. nov. belongs to the Rhinella margaritifera group, and is
characterized by medium size, snout rounded in dorsal view, with a vertical apical ridge, supraorbital crests weakly
developed, parietal crest not well developed, postorbital crest prominent, presence of a dorsolateral line of tubercles,
tympanum evident, bony protrusions at angle of jaws, absence of vertebral apophyses and of projections on upper
eyelids, and parotoid glands small. The new species is distinguished from other species of the group by geographic
distribution and by the use of floating mats of vegetation as reproductive site.

Key words: morphology, advertisement call, Rhinella margaritifera group, Pantanal

Introduction

Species of the Rhinella margaritifera group are medium-sized toads, inhabitants of forest leaf litter in Central
and South America (Frost 2009). This group remains problematic taxonomically in spite of recent efforts for
elucidating the many fuzzy species boundaries. The difficulty lies principally in the cryptic morphological
diversity and a lack of diagnostic characteristics for distinguishing the species (Caramaschi & Niemeyer
2003; Caramaschi & Pombal 2006; Fouquet et al. 2007b; Lima et al. 2007). According to Fouquet et al.
(2007b) 14 species are currently recognized: R. acutirostris (Spix), R. alata (Thominot), R. castaneotica
(Caldwell), R. dapsilis (Myers & Carvalho), R. hoogmoedi Caramaschi & Pombal, R. lescurei Fouquet,
Gaucher, Blanc & Velez-Rodriguez, R. magnussoni Lima, Menin & Araújo, R. margaritifera (Laurenti), R.
martyi Fouquet, Gaucher, Blanc & Velez-Rodriguez, R. proboscidea (Spix), R. roqueana (Melin),R.
sclerocephala (Mijares-Urrutia & Arends), R. scitula (Caramaschi & Niemeyer), and Rhinella stanlaii
(Lötters & Köhler). Out of the 14 species, 12 occur in Amazonia, while R. hoogmoedi occurs in the Atlantic
forest and R. scitula in gallery forests in the Cerrado (Caramaschi & Niemeyer 2003; Caramaschi & Pombal
2006; Lima et al. 2007). Herein, we describe adults, advertisement calls, and provide information on natural
history of a new species belonging to this group and found in the Pantanal, western Brazil.

Material and methods

We examined 20 specimens housed in the Coleção Zoológica de Vertebrados of the Universidade Federal de
Mato Grosso (UFMT, Cuiabá, Mato Grosso State, Brazil). Specimens in the type-series were collected in the

Accepted by M. Vences: 5 Dec. 2009; published: 20 Jan. 2010 57

Parque Nacional do Pantanal, in the extreme southern portion of Poconé municipality, Mato Grosso state,
western Brazil. To infer on the distribution of the species, we examined additional specimens housed both in
UFMT and in the Coleção Zoológica de Referência do Campus de Corumbá (CEUCH; Campo Grande, Mato
Grosso do Sul, Brazil). Specimens from similar species examined are listed in the Appendix. Measurements in
12 specimens (seven adult males and five adult females) were taken with digital calipers to the nearest 0.1 mm
and follow Caramaschi and Pombal (2006). The following measurements were made: SVL (snout-vent
length); HL (head length); HW (head width); IND (internarial distance); END (eye to nostril distance); ED
(eye diameter); UEW (upper eyelid width); IOD (interorbital distance); POCL (postorbital crest length); HTD
(horizontal tympanum diameter); VTD (vertical tympanum diameter); PGL (parotoid gland length); HAL
(hand length); THL (thigh length); TL (tibia length); FL (foot length).
Vocalizations were recorded with a YOGA EM 9600 microphone and a Sony TCM 50000EV tape
recorder. The calls of two males were recorded: two from Poconé municipality in 2001 (holotype, UFMT
7430), and one from Cáceres municipality in March 2002 (UFMT 2112) by Christine Strüssmann. Tape
recordings were digitized at 44.1 kHz, resolution of 16 bits (FFT 1024) with Cool Edit 96 (™Syntrillium).
Terminology for acoustic parameters follows Martins (2003) and Heyer et al. (1990).

Results

Rhinella paraguayensis, sp. nov.

(Figures 1, 2A,3)

Holotype. UFMT 7430, adult male (Figure 1), collected on 15 May 2001 by Christine Strüssmann at “Parque
Nacional do Pantanal” (17º51'12”S; 57º24'39”W), Poconé municipality, Mato Grosso state, Brazil.

FIGURE 1. Holotype of Rhinella paraguayensis sp. nov. (UFMT 7430): A. dorsal view; B. ventral view; C. profile of
head; D. ventral view of right hand; E.ventral view of right foot.

58 · Zootaxa 2339 © 2010 Magnolia Press ÁVILA ET AL.

FIGURE 2. A) living male of Rhinella paraguayensis sp. nov. observed at Paraguay river margin in 22 December 2006.
Photo by Robson W. Ávila; B) living male of Rhinella scitula collected at Serra da Bodoquena. Photo by Masao
Uetanabaro; C) preserved specimens of Rhinella paraguayensis sp. nov. (right) and Rhinella scitula (left).

Paratypes. UFMT 4268, 4270 (adult males), UFMT 4272, 4274, 4276 (adult females), 11–12 May 2001,
by Tami Mott, Raul Vieira, Christine Strüssmann; UFMT 7145 (adult male) and 7150 (adult female), 11–15
September 2001, by Christine Strüssmann and Vinicius T. Carvalho; UFMT 4269 (adult male) and 4271
(adult female), 27 March 2001, by Rogério Rossi, Ana P. Carmignotto and Mário M. Rollo Junior; UFMT
7056 and 7062 (juveniles), 13–22 September 2001, by Vinicius T. Carvalho, all from the same locality as
holotype. UFMT 046 (juvenile), May 1995, by C. Strüssmann and UFMT 171 and 172 (juveniles), 20 August
1982, by J. C. Dalponte from “Base de Pesquisa da Fauna do Pantanal, IBAMA”, km 111 of “Rodovia
Transpantaneira" (17°10’S, 57°00”W), Poconé, Mato Grosso state, Brazil. UFMT 2112 (adult male), 02
March 2002, by Christine Strüssmann in Cáceres municipality (15°47’31”S, 57°35’28”), Mato Grosso state,

A NEW RHINELLA FROM PANTANAL Zootaxa 2339 © 2010 Magnolia Press · 59

Brazil. UFMT 2103 and 2109 (juveniles), 27 February 2002, by Christine Strüssmann, Vanda L. Ferreira,
Marcos A. Carvalho, Marcelo Gordo, Robson W. Ávila and Guilherme Mourão in Cachoeirinha River, Porto
Estrela municipality (15°73’48”S, 57°31’90”), Mato Grosso state, Brazil. CEUCH 1851 and 1866 (adult
males), 15 May 2005, by Robson W. Ávila, in Baía Vermelha (18°28’27”S, 57°27’08”W), Corumbá
municipality, Mato Grosso do Sul state, Brazil.
Diagnosis: A medium sized species belonging to the Rhinella margaritifera group based on the following
morphological characters: presence of cephalic crests, distinct tympanum and paratoid glands, lateral row of
tubercles and dorsal “dead-leaf” pattern (sensu Vélez-Rodrigues 2005; Hoogmoed 1990). Rhinella
paraguayensis sp. nov. is characterized by (1) adult males 42.3–52.6 mm SVL , adult females 51–53.3 mm
SVL, (2) snout rounded in dorsal and lateral view, with a vertical apical ridge that gives it a nearly mucronate
aspect, (3) canthal and antorbital crests not raised, (4) supraorbital crests weakly developed, (5) parietal crest
not developed, (6) postorbital crest prominent, (7) vocal slits present in males, (8) tympanum evident, (9)
presence of bony protrusions at angle of jaws, (10) parotoid glands small, (11) absence of projections on upper
eyelids, (12) presence of a dorsolateral line of tubercles, from the external border of the paratoid gland to the
groin, (13) vertebral apophyses absent, (14) tarsal fold absent, (15) when the legs are adpressed to the body,
heel don’t reach posterior margin of the tympanum, (16) toes about three quarters webbed.
Rhinella paraguayensis sp. nov. is distinguished from R. acutirostris and R. alata by supraorbital crest
less developed (supraorbital crest well developed in R. acutirostris and R. alata; Leavitt 1933; Hoogmoed
1986); from R. castaneotica by supratympanic crests developed, greater SVL, presence of a dorsolateral line
of tubercles, and tympanum evident (SVL 30.9–36.8 mm in males, 33.8–42.6 mm in females, dorsolateral line
of tubercles absent, and tympanum not evident in R. castaneotica; Caldwell 1991 and examined specimens
listed in Appendix); from R. dapsilis by the snout not developed in proboscis and the presence of a
dorsolateral line of tubercles (snout forming a proboscis and absence of a dorsolateral line of tubercles in R.
dapsilis; Myers & Carvalho 1945); from R. hoogmoedi by the rounded snout in lateral view, smaller tigh, tibia
and foot (snout nearly acute, thigh 15.7–25.8 mm, 15.1–24.5 mm and foot22.4–35.2 mm in R. hoogmoedi;
Caramaschi & Pombal 2006); from R. margaritifera by smaller SVL in females, less developed supraorbital
crest and absence of vertebral apophyses (46.1–72.9 mm SVL in females, supraorbital crests hypertrophied
and vertebral apophyses present in R. margaritifera; Hoogmoed 1989, 1990 and examined specimens listed in
Appendix); from R. roqueana by the presence of a dorsolateral line of tubercles and absence of vertebral
apophyses (absence of dorsolateral line of tubercles and presence of vertebral apophyses in R. roqueana);
from R. proboscidea by the rounded snout and the presence of a dorsolateral line of tubercles (absence of
dorsolateral line of tubercles and; snout pointed extending far beyond nostrils in R. proboscidea; Hoogmoed
1986, 1990); from R. scitula by the larger SVL, rounded snout in lateral view, well developed and larger
postorbital crest, and more granulose skin on dorsum (33.8–46.1 mm in males, 45.9–50.5 mm in females,
subacuminate snout in lateral view, postorbital crest not developed with 2.8–4.4 mm length and smoother skin
on dorsum in R. scitula – Figure 2; Caramaschi & Niemeyer 2003, and examined specimens listed in
appendix); from R. magnussoni by the greater SVL and the snout rounded in dorsal view (SVL 36.0–45.3 mm
in males, 43.1–53.3 mm in females and snout pointed in dorsal view, in R. magnussoni; Lima et al. 2007);
from R. martyi by smaller SVL, absence of vertebral apophyses and less developed cephalic crest (SVL 55.3 ±
5.8 mm in males, 64.7 ± 3.4 mm in females, presence of vertebral apophyses and prominent cephalic crests in
R. martyi; Fouquet et al. 2007b); from R. lescurei by greater SVL, developed postorbital crest and rounded
snout (SVL 34.6 mm in males and 43.7 mm in females, postorbital crest not developed and snout pointed in R.
lescurei; Fouquet et al. 2007b); from R. sclerocephala by smaller SVL, and absence of vertebral apophyses
(SVL 55.4–67.3 mm in males and 68.5–77.4 mm in females, and vertebral apophyses present in R.
sclerocephala; Mijares-Urrutia & Arends 2001). Finally, R. paraguayensis sp. nov. differed from R. stanlaii
by smaller SVL in females, and supraorbital crest not developed (SVL 57.2–59.4 mm in females, and
hypertrophied supraorbital crest in R. stanlaii; Lötters & Köhler 2000).
Description of the holotype. UFMT 7430 (Figure 1). Snout vent length 50.4 mm; head as wide (17.1
mm) as long (17.2 mm) at the angle of jaws. Other measurements are presented in Table 1. Snout rounded,
both in lateral and dorsal view, with a distinct fleshy vertical apical ridge that gives it a nearly mucronate

60 · Zootaxa 2339 © 2010 Magnolia Press ÁVILA ET AL.

aspect. Cephalic crests weakly developed, except from postorbital crests; paratoid glands small, elliptical;
nostrils lateral, protuberant; distance from eye to nostril slightly larger than eye diameter and than internarial
distance; upper eyelid width about 90% than eye diameter. Presence of a bony knob at the lower angle of jaws.
Tympanum distinct, large, elliptical; its horizontal diameter representing 75% of the vertical diameter and of
the upper eyelid width. A dorsolateral line of tubercles extend from the posterior border of the paratoid gland
to the groin. Vertebral apophyses absent. Vocal slits present, lateral to the tongue. Skin on dorsal and ventral
surfaces granulose, with scattered tubercles. Forearms and forelimbs robust, covered with small tubercles of
the same size of those in the dorsum. A line of small tubercles along the external border of the forearms;
fingers free, with tips slightly enlarged; length of fingers 4<2<1<3. Palmar tubercle oval, large, smooth;
thenar tubercle oval, about one third of the palmar tubercle; subarticular and supranumerary tubercles present.

TABLE 1. Measurements (mm) of holotype and of specimens in the type series of Rhinella paraguayensis sp. nov.
Means are ± 1 standard deviation, ranges are in parentheses. Abbreviations are defined in the material and methods
section.
Character Holotype Male (n = 6) Female (n = 5)
SVL 50.4 47.5 ± 3.5 (42.3–52.6) 52.4 ± 1.0 (51–53.3)
HL 17.2 16.6 ± 1.3 (14–18.1) 17.8 ± 0.9 (16.9–18.9)
HW 17.1 18.0 ± 1.4 (16.6–20.1) 17.8 ± 2.0 (15.5–20.2)
IND 4.0 3.1 ± 0.5 (3.2–3.5) 3.3 ± 0.2 (2.9–3.5)
END 4.1 4.8 ± 0.6 (3.8–5.6) 4.7 ± 1.0 (3.5–6)
ED 4.0 4.6 ± 0.6 (4–5.4) 4.7 ± 0.5 (4–5.3)
UEW 3.7 3.4 ± 0.2 (3–3.7) 3.2 ± 0.6 (2.4–3.9)
IOD 16 14.6 ± 1.1 (13–16.1) 15.5 ± 0.5 (15.2–16)
POCL 5.2 4.7 ± 0.5 (4.2–5.2) 5.6 ± 0.3 (5.3–6)
HTD 2.8 2.4 ± 0.6 (1.6–3.1) 2.7 ± 0.6 (1.9–3.3)
VTD 3.7 3.1 ± 0.4 (2.5–3.7) 3.2 ± 0.4 (2.8–3.7)
PGL 9.2 7.8 ± 1.3 (6–9.2) 7.9 ± 1.3 (6.8–9.9)
HAL 14.2 13.3 ± 1.0 (12–14.3) 14.0 ± 0.5 (13.5–14.8)
THL 13 11.1 ± 1.6 (8.8–13) 13.1 ± 2.0 (11–15.5)
TL 18 16.3 ± 1.6 (14.7–19) 17.9 ± 1.8 (16.5–20.3)
FL 22.3 24.9 ± 2.0 (22.3–28) 26.4 ± 2.9 (21.6–29.2)

Hindlimbs robust, thigh length smaller than tibia length, thigh+tibia length approximately 62% of the
SVL; tarsus+foot length longer than tibia and thigh length; a line of tubercles on external and internal borders
of tarsus. Inner metatarsal tubercle oval, two times larger than the outer metatarsal tubercle; subarticular and
supernumerary tubercles present, small, round. Length of toes 1<5<2<3<4, webbing formula: I 1-2 II 1-3 III
1-2 ½ IV 2 ½-1V.
Colour in preservative: dorsum light brown, dark brown on legs and head; a yellowish vertebral line from
the snout to cloaca, surrounded by three black triangular spots at midbody and a pair of triangular black spots
at the level of the parietal crest. Venter uniform brown; throat, hand, foot and legs dark brown; an irregular
dark brown band in the forearm. Color in life (Figure 2A) similar to that in preservative. Iris gold, with black
reticulations.
Variation within type series. The type series include 20 specimens (6 adult males, 5 adult females and 9
juveniles). Dorsal coloration varies a lot among individuals in the type series (Figure 3), with the fine
vertebral line present in 75% of the individuals and only one individual (holotype) have a wide vertebral line.
Few specimens don’t have the triangular spots in the midbody. The ventral coloration is also variable, from
the uniform brown to spotted bellies.

A NEW RHINELLA FROM PANTANAL Zootaxa 2339 © 2010 Magnolia Press · 61

FIGURE 3. Variation in dorsal and ventral color pattern of Rhinella paraguayensis sp. nov.

62 · Zootaxa 2339 © 2010 Magnolia Press ÁVILA ET AL.

FIGURE 4. Oscillogram (A–B) and audio spectrogram (C) of the advertisement calls of Rhinella paraguayensis sp. nov.
(UFMT 7430), recorded on 15 May 2001 in the Parque Nacional do Pantanal , Poconé municipality, Mato Grosso state,
Brazil (18:00 h, air temperature 21,5ºC).

Advertisement call. The call of the new species is composed by 18.4 ± 8.7 call series (9–30 calls, n=5),
with duration of call series of 16.5 ± 7.2s (n=5), emitted at regular intervals of 10.4 ± 0.8s (n= 5). The call of
R. paraguayensis sp. nov. is composed by mean 6.5 ± 0.65 notes (5–8 notes, n=77) , with a mean duration of
0.3s ± 0.04s (0.2–0.42, n=82), followed by intercall intervals of 0.5s ± 0.015s (0.22–0.98, n=82). The average
rate of call per minute was 39.3 ± 6.4. The mean duration of note was 0.03s ± 0.01s (0.01–0.07, n=163),
followed by inter-notes intervals of 0.02s ± 0.01s (0.01–0.04, n=163). The average rate of notes per minute
was 504.8 ± 113 (262.4–792.4). Frequency ranges from 618.9–1771.8 Hz (n=125); mean dominant frequency
was 1438.7 ± 70.5 Hz (1113.7–1568.5, n=125), displayed between the fourth and fifth note (Figure 4).
Natural history. In the Upper Paraguay River Basin, individuals of Rhinella paraguayensis sp. nov.
inhabit the leaf litter of gallery forests, and of forests established over paleo-levees throughout the Pantanal
wetlands. Riparian systems are used as reproductive sites by individuals in this new species (Figure 5), which
apparently presents a unimodal pattern of reproduction, as shown below.
During the peak of the floods, in April-May, which are coincident with the beginning of the dry season,
the Paraguay River discharges increased (Barros et al. 2004), facilitating the movements of dense stands of
macrophytes (particularly the water hyacinth Eichhornia azurea; Figure 5). These macrophyte stands offer
both vocalization perches for calling males and lentic habitats for the deposition of egg clutches, as well as
protection for the tadpoles. Additionally, the displacement of macrophyte stands downriver may facilitate the
transport and dispersal of tadpoles, juveniles and also adults of R. paraguayensis sp. nov., as already reported
for several amphibian species using macrophytes as rafts in others river systems (Hoogmoed, 1993; Schiesari
et al. 2003).
Observations of calling males occurred on seven occasions. In 10 June 1989 a male calling from nude
ground at “Rodovia Transpantaneira”, Poconé, Mato Grosso. In April and May 2001, males of the new
species were observed calling on floating stems and leaves of E. azurea at “Parque Nacional do Pantanal”,
Poconé municipality, Mato Grosso. From 4 March 2002 males were observed calling (a male was recorded
and used in the call description) in a flooded swampy areas in the left margin of the Sepotuba River, Cáceres
municipality, Mato Grosso. Between 14–17 May 2002 several males of R. paraguayensis sp. nov. were
observed calling on nude ground in the Paraguay River banks, approximately 0.5 m from the river margin, in

A NEW RHINELLA FROM PANTANAL Zootaxa 2339 © 2010 Magnolia Press · 63

sites where the availability of floating mats of E. azurea was low, in the Serra do Amolar, Corumbá
municipality, Mato Grosso do Sul. In October 2002, males were observed calling from riparian systems of the
Paraguay River, in the Serra do Amolar, Corumbá municipality, Mato Grosso do Sul. In 11 and 12 August
2003, during the dry season, several males of the new species were observed calling on floating stems and
leaves of E. azurea, as far as 2 m from the Paraguay River bank. River width in this locality, near Acurizal
ranch (17º49'51”S; 57º32'53”W), is near 100 m. Vocalizations were heard from around 15:00 h, in full sun, to
21:00h, with full moon.

FIGURE 5. A. General view of a riparian system along the margins of the Paraguay River near Acurizal Ranch
(17°49’51” S; 57°33’06” W), in the western border of the Brazilian Pantanal, Poconé municipality, Mato Grosso state.
This kind of riparian system, formed by gallery forests and dense stands of macrophytes anchored along the margins of
river, is used by individuals of Rhinella paraguayensis sp. nov. as a reproductive site, in the peak of the dry season.

Therefore, calling activities of Rhinella paraguayensis sp. nov. in the Pantanal were recorded from March
to October and seems to be correlated with the Paraguay River flooding cycle .
Rhinella paraguayensis sp. nov. is usually abundant in the localities where the species was already
recorded. In a field study in the Parque Nacional do Pantanal, it was the third most abundant among the 13
anuran species recorded by pitfall trapping (http://www.ibama.gov.br/siucweb/unidades/parna/
planos_de_manejo/78/index.htm).
Distribution. The new species occurs along the western drainages of the Paraguay River, in the Brazilian
states of Mato Grosso and Mato Grosso do Sul (Figure 7).
Etymology. The specific epithet is an adjective, derived from the type locality of the new species: the
Paraguay River basin.

64 · Zootaxa 2339 © 2010 Magnolia Press ÁVILA ET AL.

FIGURE 6. Mean monthly discharges of the Paraguay River near Acurizal Ranch from 1968–2003, in the western
border of the Brazilian Pantanal. Bar indicates the calling period based on our observations. Data were taken from ANA
– Agência Nacional de Águas. Sistema de Informações Hidrológicas. Available at http://hidroweb.ana.gov.br/. Accessed
25 May 2009.

FIGURE 7. Geographical distribution map of Rhinella paraguayensis sp. nov (triangles) and R. scitula (circles). Star =
type locality of Rhinella paraguayensis sp. nov and square = type locality of R. scitula.

A NEW RHINELLA FROM PANTANAL Zootaxa 2339 © 2010 Magnolia Press · 65

TABLE 2. Measurements of acoustic parameters for males of Rhinella paraguayensis sp. nov. Values are mean ±
standard deviation, range in parentheses.
UFMT 7430 UFMT 2112 Mean
Call series 19.3 ± 9.3 17 ± 11.3 18.4 ± 8.7
(13–30) (9–25) (9–30)
Call series duration 16.3 ± 9.1 16.9 ± 6.4 16.5 ± 7.2
Intervals call series 10.6 ± 0.9 10.1 ± 0.4 10.4 ± 0.8
Call notes 6.5 ± 0.6 6.3 ± 0.7 6.5 ± 0.6
(5–7) (5–8) (5–8)
Call duration (s) 0.33 ± 0.20 0.27 ± 0.03 0.30 ± 0.04
(0.26–0.42) (0.2–0.32) (0.2–0.42)
Intercall interval (s) 0.48 ± 0.16 0.5 ± 0.15 0.5 ± 0.15
(0.22–0.98) (0.38–0.97) (0.22–0.98)
Call minute 39.3 ± 10 35.7 ± 16 39.3 ± 6.4
Note duration (s) 0.03 ± 0.01 0.02 ± 0.01 0.03 ± 0.01
(0.02–0.07) (0.01–0.04) (0.01–0.07)
Inter–notes interval (s) 0.02 ± 0.01 0.02 ± 0.01 0.02 ± 0.01
(0.01–0.04) (0.01–0.04) (0.01–0.04)
Notes minute 499.8 ± 118.7 515.3 ± 102 504.8 ± 113
(262.4–792.4) (323.4–728.3 (262.4–792.4)
Frequency ranges (Hz) 615.3 – 1813.7 647.5 – 1750.4 618.9 – 1771.8
n= 68 n= 56 n=125
Dominant frequency (Hz) 1411.9 ± 73.8 1470.6 ± 50.9 1438.7 ± 70.5
(1113.7–1482.9) (1315.8–1568.5) (1113.7–1568.5)

Discussion

The Rhinella margaritifera complex remains as one of the major problems in South American anuran
taxonomy (see Fouquet et al. 2007a). There are several undescribed species, and the cryptic morphology
amongst representatives of the group turns difficult the systematic arrangement even of the species already
described (Hoogmoed 1989, 1990; Lima et al. 2007). Nevertheless, recent studies have demonstrated that
genetic and acoustic differences can be useful tools for differentiating species in this group (Lima et al. 2007;
Fouquet et al. 2007b).
The dominant frequency in calls of R. paraguayensis sp. nov. is similar to what was recorded in Rhinella
sp. clade E (1407 Hz), and higher than in R. martyi (1169 Hz), R. lescurei (1161 Hz), and R. margaritifera
(1265 Hz) (Fouquet et al. 2007b) and in a representative of the Rhinella margaritifera complex in Bolivia
(1332 Hz; De la Riva et al. 1996). Dominant frequency is considerably lower in R. paraguayensis sp. nov.
than in R. castaneotica (1650 Hz; Köhler & Lötters 1999). Frequency range is lower than in R. magnussoni
(1600–2820 Hz; Lima et al. 2007), R. castaneotica (900–2600 Hz; Köhler & Lötters 1999), and R.
proboscidea (1620–3200 Hz; Zimmerman & Bogart 1988) and similar to the Bolivian populations assigned to
the Rhinella margaritifera complex (500–1500; De la Riva et al. 1996). Note duration was lower than R.
magnussoni (0.06 ± 0.01 s); Lima et al. 2007) and R. proboscidea (0.12 s; Zimmerman & Bogart 1988).
Moreover, there are clear ecological differences between some of the species in the Rhinella margaritifera
species group, including geographic distribution, reproductive habitat, and vocalization microhabitat (Lima et
al. 2007). Species in the R. margaritifera complex are distributed mainly in the Amazonia Biome, with a few
species occurring in other forested (e.g. Atlantic Forest – R. hoogmoedi Caramaschi & Pombal 2006) or
opened ecosystems (e.g. Cerrado (Brazilian savanna) – R. scitula Caramaschi & Niemeyer 2003; Pantanal –
R. paraguayensis sp. nov., present study; Figure 7).

66 · Zootaxa 2339 © 2010 Magnolia Press ÁVILA ET AL.

 The calling and reproductive site of R. paraguayensis sp. nov. (Leaves of floating macrophyte or in the
ground of river margins) contrasts with several species in the R. margaritifera complex. For instance, males of
R. magnussoni call from leaf litter or bellow fallen leaves distant from streams or terrestrial ponds, with egg
deposition in fallen tree trunks (Lima et al. 2007); in R. castaneotica, calling and egg deposition site occurs in
the empty fruit capsules of Brazilian nut trees (Caldwell 1991); males of R. proboscidea use shallow puddles
bordering small streams or temporary pools (Zimmerman & Bogart 1988, Menin et al. 2006); in R.
margaritifera, males call from the leaf litter surrounding permanent pools, swamps and artificial ponds (Wells
1979), while males of Rhinella sp. use shallow pools on top of large flat boulder with concave surface in the
middle of creek beds (Hoogmoed and Ávila-Pires 1991).
The calling site of the new species bears resemblance with that of R. sclerocephala, whose males were
observed calling perched on leaf axis of vegetation (Mijares-Urrutia & Arends 2001); of R. hoogmoedi,
observed perched in tree trunks at 60 cm from the ground (Pombal & Gordo 2004), and of R. lescurei, which
calls on dead trunks, vines and roots distant about 10 m from slowly running water (Fouquet et al. 2007b).
Males calling on the ground near river margins were also observed in R. scitula and R. hoogmoedi, although
these species were recorded using small streams inside riparian forest until now (Caramaschi & Niemeyer
2003; Pombal & Gordo 2004), instead of the margins of larger rivers, as observed in R. paraguayensis sp.
nov.
In spite of several recent contributions, much work is still needed to evaluate species composition, as well
as understanding the geographical distribution and ecological diversity of R. margaritifera complex.

Acknowledgements

We thank PROBIO, Ecotrópica, IBAMA, for partial financial support to field expeditions; José Augusto
Ferraz de Lima, for the arrangements to work at the Pantanal National Park; Pierre Girard for the help to
obtain discharges data for Amolar and Ricardo A. Kawashita Ribeiro for preparing the distribution map.
Masao Uetanabaro for providing photographs of live specimen of R. scitula and Franco L. Souza for gently
provided voucher specimens of R. scitula.

References

Barros, V., Chamorro, L., Coronel, G. & Baez, J. (2004) The Major Discharge Events in the Paraguay River: Magnitudes,
Source Regions, and Climate Forcings. Journal of Hydrometeorology, 5(6), 1161–1170.
Caldwell, J.P. (1991) A new species of toad in the genus Bufo from Pará, Brasil, with an unusual breeding site. Papéis
Avulsos de Zoologia, 37(26), 389–400.
Caramaschi, U. & Niemeyer, H. (2003) Nova espécie do complexo de Bufo margaritifer (Laurenti, 1768) do Estado do
Mato Grosso do Sul, Brasil (Amphibia, Anura, Bufonidae). Boletim do Museu Nacional, Nova Série, Rio de Janeiro,
501, 1–16.
Caramaschi, U. & Pombal Jr, J.P. (2006) A new species of Rhinella Fitzinger, 1826 from the Atlantic rain forest, eastern
Brazil (Amphibia, Anura, Bufonidae). Papéis Avulsos de Zoologia, 46(23), 251–259.
De la Riva, I., Bosch, J. & Márquez, R. (1996) Advertisement calls of two Bolivian toads (Anura: Bufonidae: Bufo).
Herpetological Journal, 6, 59–61.
Fouquet, A., Vences, M., Salducci, M.-D., Meyer, A., Marty, C., Blanc, M. & Gilles, A. (2007a) Revealling cryptic
diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera
species groups. Molecular Phylogenetics and Evolution, 43, 567–582.
Fouquet, A., Gaucher, P., Blanc, M. & Velez-Rodriguez, C.M. (2007b) Description of two new species of Rhinella
(Anura: Bufonidae) from the lowlands of the Guiana shield. Zootaxa, 1663, 17–32.
Frost, D.R. (2009) Amphibian Species of the World: An online reference. Version 5.3, American Museum of Natural
History, New York. Available from: http://research.amnh.org/herpetology/amphibia/index.php. (accessed 5
September, 2009).
Heyer, W.R., Rand, A.S., Cruz, C.A.G., Peixoto, O.L. & Nelson, C.E. (1990) Frogs of Boracéia. Arquivos de Biologia,
31, 231–410.
Hoogmoed, M.S. (1986) Biosystematic studies of the Bufo “typhonius” group. A preliminary progress report. In: Rocek,

A NEW RHINELLA FROM PANTANAL Zootaxa 2339 © 2010 Magnolia Press · 67

 Z. (Ed.), Studies in Herpetology. Charles University, Prague, pp. 147–150.
Hoogmoed, M.S. (1989) South American bufonids (Amphibia: Anura: Bufonidae), an enigma for taxonomists. In:
Fontanet, X. & Horta, N. (Eds.), Treballs d’Ictiologia i Herpetologia. Barcelona, pp. 167–180.
Hoogmoed, M.S. (1990) Biosystematics of South American Bufonidae, with special reference to the Bufo “typhonius”
group. In: Peters, G. & Hutterer, R. (Eds), Vertebrates in the Tropics. Museum Alexander Koening, Bonn, pp. 113–
123.
Hoogmoed, M.S. (1993) The Herpetofauna of Floating Meadows. 199–213p. In: Ouboter, P.E. (Ed, Freshwater
Ecossystems of Suriname. Kluwer Academic Publishers. pp 199–213.
Hoogmoed, M.S. & Avila-Pires, T.C.S. (1991) Annotated checklist of the herpetofauna of Petit Saut, Sinnamary river,
French Guiana. Zoologische Mededelingen, 65, 53–88.
Köhler, J. & Lötters, S. (1999) Annotated list of amphibian records from the Departamento Pando, Bolivia, with
description of some advertisement calls. Bonner Zoologische Beitraege, 48, 259–273.
Leavitt, B.B. (1933) On three races of Bufo typhonius. Copeia, 1933, 7–8.
Lima, A.P., Menin, M. & Araújo, M.C. (2007) A new species of Rhinella (Anura: Bufonidae) from Brazilian Amazon.
Zootaxa, 1663, 1–15.
Lötters, S. & Köhler, J. (2000) A new toad of the Bufo typhonius complex from humid montane forests of Bolivia.
Spixiana, 23(3), 293–303.
Martins, I.A. & Jim, J. (2003) Bioacoustic analysis of advertisement call in Hyla nana and Hyla sanborni in Botucatu,
São Paulo, Brazil. Brazilian Journal of Biology, 63, 507–516.
Menin, M., Rodrigues, D.J. & Lima, A.P. (2006). The tadpole of Rhinella proboscidea (Anura: Bufonidae) with notes on
adult reproductive behavior. Zootaxa, 1258, 47–56.
Mijares-Urritia, A. & Arends, A. (2001) A new toad of the Bufo margaritifer complex (Amphibia: Bufonidae) from
northwestern Venezuela. Herpetologica, 57(4), 523–531.
Myers, G.S. & Carvalho, A.L. (1945) Notes on some new or little-known Brazilian amphibians, with an examination of
the history of the plata salamander, Ensatina platensis. Boletim do Museu Nacional do Rio de Janeiro, Nova Série,
Zoologia, 35, 1–24+18 Plates.
Pombal Jr., J.P. & Gordo, M. (2004) Anfíbios anuros da Juréia. In: Marques, O.A.V. & Duleba, W. (Eds), Estação
Ecológica Juréia-Itatins: Ambiente físico, flora e fauna. Holos Editora, Ribeirão Preto, pp. 243–256.
Schiesari, L., Zuanon, J., Azevedo-Ramos, C., Garcia, M., Gordo, M., Messias, M. & Vieira, E.M. (2003) Macrophyte
rafts as dispersal vectors for fishes and amphibians in the lower Solimões River, Central Amazon. Journal of
Tropical Ecology, 19(3), 333–336.
Vélez-Rodriguez, C.M. & Ruiz-C, P.M. (2002) A new species of Bufo (Anura: Bufonidae) from Colombia.
Herpetologica, 58, 453–462
Wells, K.D. (1979) Reproductive behavior and male mating success in a Neotropical toad, Bufo typhonius. Biotropica,
11(4), 301–307.
Zimmerman, B.L. & Bogart, J.P. (1988) Ecology and calls of four species of Amazonian forest frogs. Journal of
Herpetology, 22(1), 97–108.

Appendix. Additional specimens examined.

Rhinella castaneotica: BRAZIL: Mato Grosso: Colniza: UFMT 6772, 6781, 6802, 8063
Rhinella margaritifera: BRAZIL: Mato Grosso: Colniza: UFMT 6786, 6801
Rhinella scitula: BRAZIL: Mato Grosso do Sul: Bodoquena: UFMT 9853-9864

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