Boletín de la Sociedad Zoológica del Uruguay, 2025

Vol. 34 (1): e34.1.8

ISSN 2393-6940

https://journal.szu.org.uy

DOI: https://doi.org/10.26462/34.1.8

ABSTRACT

The reproductive biology of Melanophryniscus

devincenzii was described here, based on morphometrics

and gonadal analysis of 86 individuals collected in the

Sertão Municipal Natural Park, Rio Grande do Sul, Brazil.

Individuals were captured both manually and using pitfall

traps. We identified 63 males and 23 females, with the sex

ratio biased towards males. Sexual dimorphism was

observed, with females being larger than males. No

significant correlation was found between female body

size and oocyte number, nor between male body size and

testicular mass, indicating that energetic investment may

be more focused on reproduction than on growth. All

analyzed females (except one) exhibited mature oocytes,

immature and atretic oocytes in their ovaries, suggesting

that females may be latent to reproductive event. This

reproductive latency benefit to those species that have

explosive reproductive events and depend on rainfall.

Keywords: Explosive reproduction, reproductive

investment, fecundity, fat bodies.

RESUMEN

Biología reproductiva de Melanophryniscus

devincenzii (Anura: Bufonidae) en el sur de Brasil. Se

describe la biología reproductiva de Melanophryniscus

devincenzii basada en el análisis morfométrico y gonadal

de 86 individuos colectados en el Parque Natural

Municipal de Sertão, Río Grande del Sur, Brasil. Los

individuos fueron capturados (manualmente y usando

trampas de caída). Identificamos 63 machos y 23

hembras. La proporción de sexos estuvo sesgada a favor

de los machos, y se comprobó la existencia de

dismorfismo sexual en la especie, con hembras más

grandes que los machos. No se encontraron relaciones

significativas entre el tamaño de las hembras y el número

de oocitos; ni entre el tamaño de los machos y la masa

testicular, indicando que la inversión energética estaría

centrada más en la reproducción que en el crecimiento.

Todas las hembras analizadas (con la excepción de una)

presentaron oocitos maduros, acompañados de oocitos

inmaduros y atrésicos en sus ovarios, evidenciando que

las hembras estarían siempre preparadas para el evento

reproductivo, lo cual resultaría beneficioso considerando

que los eventos reproductivos del tipo explosivo

dependen de las precipitaciones.

Palabras Clave: Reproducción explosiva, inversión

reproductiva, fecundidad, cuerpos grasos.

INTRODUCTION

Amphibians exhibit a wide range of reproductive

modes, which include variations in egg characteristics,

oviposition sites, and the presence of parental care

(Duellman & Trueb, 1986; Haddad & Prado, 2005). The

reproduction of several anuran species is particularly

influenced by abiotic variables. Temperature and

precipitation are the primary climatic factors affecting

anuran reproductive cycles (Cardoso & Martins, 1985),

which can directly alter the timing and duration of the

reproductive period (Lajmanovich, 2000). According to

Wells (1977), there are two temporal reproductive

strategies in anurans: prolonged and explosive

breeding. However, these strategies represent the

extremes of a continuum along which most species are

distributed. Anurans with explosive reproduction

concentrate their reproductive activity within hours or

days, with males and females migrating synchronously

to breeding sites. In prolonged breeders, the activity

period extends over several weeks or months; the

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

REPRODUCTIVE BIOLOGY OF Melanophryniscus devincenzii (ANURA: BUFONIDAE) IN

SOUTHERN BRAZIL

1 1 2 1

Schaiani V. Bortolini , Gisela Pereira * , Noeli Zanella , Raúl Maneyro

1 Laboratorio de Herpetología, Instituto de Ecología y Ciencias Ambientales. Facultad de Ciencias, Universidad

de la República, Iguá 4225, Montevideo 11400, Uruguay.

2 Instituto da Saúde, Campus I, Universidade de Passo Fundo, Bairro São José. CEP 99001-970, Passo

Fundo, RS, Brazil.

* Corresponding author: gpereira@fcien.edu.uy

Fecha de recepción: 28 de octubre de 2024

Fecha de aceptación: 16 de diciembre de 2024

BORTOLINI et al.

arrival of both sexes is asynchronous, and females

may choose males (Wells, 1977; Wells, 2007).

Reproduction in amphibians may be influenced by

other factors, such as the energy allocated to fat bodies

(FB) (Jørgensen, 1992). Fat bodies serves as a crucial

nutritional source during periods of drought or

hibernation (Saidapur & Hoque, 1996; Hillman et al.,

2009). These structures are located in the gonads of

both males and females and are involved in the

metabolism of gonadal processes (particularly lipid

synthesis and storage), as well as in folliculogenesis

and yolk production (Mendez-Tepepa et al., 2023).

Previous studies have demonstrated an inverse

relationship between FB abundance and gonadal

development in anurans, confirming that these

structures serve as energy reserves for reproduction

(Díaz-Páez & Ortiz, 2001; Pereira & Maneyro, 2012;

Pereira et al., 2015). In female anurans that do not

exhibit parental care, reproductive success largely

depends on the total energy stored in the oocytes

(Grafe et al., 1992). Additionally, the quality of their

offspring is directly affected by the amount of energy

stored in the fat bodies (Díaz-Paéz & Ortiz, 2001),

which supports oocyte development (Mendez-Tepepa

et al., 2023).

Melanophryniscus Gallardo, 1961 is a

monophyletic genus belonging to the family Bufonidae,

distributed across Brazil (from Espírito Santo, Minas

Gerais, and Mato Grosso do Sul to Rio Grande do Sul),

Bolivia (with one species reaching the inter-Andean

valleys in the south), central and northern Argentina

(from Buenos Aires to Jujuy), Paraguay, and Uruguay

(Frost, 2024). Currently, the genus comprises 31

species, commonly known as “South American red-

bellied toads” (Frost, 2024). The first study on

reproductive behavior in this genus was carried out on

Melanophryniscus stelzneri, which determined an

explosive reproduction (Fernández, 1927). Patterns of

reproductive activity were studied on different species

of the Genus Melanoprhyniscus. For instance,

reproductive activity was reported to M. rubriventris

(Vaira et al., 2005) and M. diabolicus (Cairo et al.,

2008); while oviposition site selection was examined in

M. rubriventris and M. stelzneri (Goldberg et al., 2006;

Pereyra et al., 2011); and calling activity was studied in

males of M. stelzneri (Pereyra et al., 2016); and M.

cupreuscapularis (Duré et al., 2015). More recent

investigations have focused on the, reproductive

biology and use of microhabitats in M. montevidensis

(Pereira & Maneyro, 2016; 2018), the reproductive

traits of M. atroluteus, M. devincenzii, and M. krauczuki

(Marangoni & Baldo, 2023); as well as the sexual

dimorphism, and fertility aspects of Melanophryniscus

fulvoguttatus (Carrillo et al., 2024) There are few

studies on the natural history of M. devincenzii (i.e.,

Bortolini et al., 2013). Despite these advancements,

more studies about natural history of M. devincenzii are

nedded, to better understand the variation among

poplations.

It is worth noting that several of these studies

demonstrated a skewed sex ratio in favor of males

during reproductive events in species of the genus

Melanophryniscus (Bustos Singer & Gutiérrez, 1997;

Vaira, 2005; Cairo, 2013; Pereira & Maneyro, 2018;

Marangoni & Baldo, 2023), as well as the existence of

sexual dimorphism, with females being larger than

males (Pereira & Maneyro, 2018; Carrillo et al., 2024).

Regarding the relationship between fecundity (F)

and/or oocyte size with female size, the results are

varied within the genus. For example, in species such

as M. diabolicus and M. montevidensis, fecundity was

not correlated with female size (Cairo et al., 2008;

Pereira & Maneyro, 2018). Opposite results were

reported for M. fulvoguttatus, where fecundity was

positively correlated with female size (Carrillo et al.,

2024). On the other hand, in M. atroluteus, a negative

correlation was found between oocyte size and female

size, while in M. krauczuki, this relationship was

positive (Marangoni & Baldo, 2023). Finally, it is

important to highlight that species with explosive

reproductive dynamics tend to maintain a constant lipid

reserve for the potential reproductive event (e.g.,

Valdez & Maneyro, 2016), which was demonstrated in

M. montevidensis (Pereira & Maneyro, 2018).

The aim of this study was to describe the

reproductive parameters of a Melanophryniscus

devincenzii (Anura: Bufonidae) population from the

Sertão Municipal Natural Park (PNMS), Rio Grande do

Sul, Brazil, by analyzing the reproductive investment of

females and the potential associations between fat

body reserves and the reproductive activity of both

males and females.

MATERIAL AND METHODS

Study Species

Melanophryniscus devincenzii (Klappenbach,

1968) (Fig. 1), commonly known as Devincenzi's toad

or Rivera's toad, is a small-sized species characterized

by a frontal swelling (Maneyro & Carreira, 2016). It is

included in the phenetic group M. tumifrons, which

comprises eight species with a restricted distribution

across Paraguay, Uruguay, Argentina, and Brazil

(Caramaschi & Cruz, 2002; Brusquetti & Lavilla, 2006;

Airaldi et al., 2009). Some references regarding the

natural history attributed to M. aff. devincenzii should

currently be considered as M. devincenzii (Langone &

Lavilla, 2024). In Brazil, it is found in the northern and

central-western regions of the state of Rio Grande do

Sul (Bastos et al., 2023). This species is primarily diur-

nal, although it may also be active at night, exhibiting

explosive breeding dynamics associated with heavy

rainfall and high humidity (Maneyro & Carreira, 2016).

The species is not considered threatened at a local

(Bastos et al., 2023) or global scale (IUCN, 2023).

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

3Reproductive biology of Melanophryniscus devincenzii

Study site and individual collection

Individuals of M. devincenzii were captured using

manual capture and dry pitfall traps with drift fences

(Cechin & Martins, 2000), between 2005 and 2011,

from the Municipal Natural Park of Sertão (PNMS), Rio

Grande do Sul, Brazil (28º02'31" S, 52º13'28" W) (Fig.

2). Four trap arrays were located, each consisting of 8

plastic containers placed 10 m apart and connected by

a one-m-high drift fence. These traps were checked

three times a week. Additionally, active searches for

specimens were conducted during both day and night.

All captured individuals are deposited in the Amphibian

Collection of the University of Passo Fundo (CAUPF).

We sexed all specimens, males were considered

reproductively active if they had a developed vocal sac,

or if they were found vocalizing or in amplexus at the

study site.

Laboratory Work

Individuals of M. devincenzii were sexed via

gonadal analysis. Before dissection, body mass (BM,

in g) and snout-vent length (SVL, in mm) were

recorded. In females, the gonads were removed to

measure ovarian mass (OM) using a digital balance

accurate to 0.001 g. Oocytes were subsequently

classified by developmental stage as mature,

immature (translucent), or atretic, following the criteria

of Camargo et al. (2005) and Pereira & Maneyro

(2012). Females were considered mature if they had

mature oocytes in their ovaries, with the smallest

female carrying mature oocytes serving as a reference

to distinguish mature from immature females.

Fecundity (F = number of mature oocytes) was

recorded for both ovaries. Reproductive investment

(RI) in reproductive females was then calculated as the

ratio of F to SVL (RI = F/SVL) (Carrillo et al., 2024). The

Kruskal-Wallis test was used to study the variation in

the IR of females among reproductive events. To

explore potential relationships between SVL and F, as

well as between SVL and OM, linear regression

analyses were performed on these variables.

The size (in mm) of the smallest male found in

reproductive activity was used as a threshold to

distinguish between mature and immature individuals.

Testicular mass (TM, in g) of each individual was

measured as an indicator of testicular activity in mature

Fig. 1. Melanophryniscus devincenzii individual in its natural habitat. Photo: Raúl Maneyro.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

BORTOLINI et al.

males (de Oliveira et al., 2007; Bortolini et al., 2018).

The relationship between SVL and TM was analyzed

through linear regression of these variables, and

monthly variation in TM of mature males was examined

using the non-parametric Kruskal-Wallis test.

Additionally, to determine the existence of sexual size

dimorphism in the species, SVL and BM were

compared between sexes using the non-parametric

Mann-Whitney U test. To assess differences in the

frequency of captured males and females, the sex ratio

(SR = number of males / number of females) was

calculated. Finally, to examine the association between

the reproductive activity of M. devincenzii and fat

accumulation, fat bodies (FB) in both males and

females were categorized based on abundance

(absent, small, intermediate, and large) (Pereira &

Maneyro, 2012).

RESULTS

A total of 86 individuals of M. devincenzii were

captured and analyzed, comprising 23 females and 63

males. Three instances of breeding activity were

recorded for the species at the study site, where males

were observed calling, amplectant pairs, and egg

clutches in lotic water bodies and along stream

margins. Peaks in capture numbers corresponded to

these three breeding events that occurred in

September 2009 (N=8 individuals), December 2010

(N=37), and July 2011 (N=22) (Fig. 3). During these

events, males were more abundant than females (Fig.

3), and the sex ratio was skewed in favor of males

(SR=3.6 in December 2010; and SR=2.7 in July 2011).

The exception was in September 2009, where the sex

ratio was balanced (SR=1). The mean SVL for males

was 23.2 ± 1.7 (range: 17.7–27.4) mm, while the mean

body mass (BM) was 1.4 ± 0.4 (range: 0.5–2.2) g. For

females, these values were 27.5 ± 2.1 (range:

23.1–32.5) mm and 2.2 ± 0.6 (range: 1.3–3.7) g. The

smallest female with mature oocytes measured 24.9

mm, and the smallest male found in reproductive

activity measured 20.45 mm. All individuals found

below these reference values for each sex were

considered immature. A total of 22 mature females and

60 mature males were identified. Significant sexual

size dimorphism was found in M. devincenzii, as

females exhibited a greater SVL (U=72; p<0.05) (Fig.

4a) and greater BM (U=180.5; p<0.05) (Fig. 4b) than

males.

The mean ovary mass (OM) in females was 0.25 ±

0.25 (range: 0.01–0.92) g. Of the 23 females analyzed,

22 exhibited all three types of oocytes in their ovaries:

mature, immature, and atretic. The mean fecundity (F)

was 126.5 ± 109.4 (range: 12–313) oocytes. No

significant relationship was found between log SVL

and log F (R²=0.08; p=0.19; Fig. 5a); however, a

significant positive relationship was found between

log SVL and log OM (R²=0.19; p=0.04) (Fig. 5b). The

10 10

mean RI was 4.5 ± 3.8 (0.5–12.2) (Fig. 6), and no

significant variation was observed in this parameter

across different breeding events (H=5.55; p=0.06). For

Fig. 2. Collection site of Melanophryniscus devincenzii individuals. The black circle represents the location of the Municipal

Natural Park of Sertão (PNMS) in Rio Grande do Sul.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

5Reproductive biology of Melanophryniscus devincenzii

males, the mean testis mass (TM) was 0.004 ± 0.002

(range: 0.0007–0.001) g. No significant relationship

was found between log SVL and log TM (R²=0.0001;

10 10

p=0.93) (Fig. 7), and there was no significant variation

in TM across months (H=7.69; p=0.05).

Finally, 60.9% of the females analyzed did not

display fat bodies (FB). In December 2010 and July

2011 (during reproductive activity events), females

exhibited FB across different abundance categories

(Fig. 8). In contrast, 83.3% of the males analyzed

presented FB. Males found in reproductive activity

exhibited FB, with the "abundant" category

predominating in December 2010 and July 2011 (Fig.

9).

DISCUSSION

This study recorded three reproductive activity

events for Melanophryniscus devinzenzii: in

September 2009, December 2010, and July 2011. The

species exhibits an explosive reproductive pattern,

breeding in semi-permanent marshes in open

environments following heavy rainfall. Reproductive

events for this species have been reported at the study

site following precipitation exceeding 60 mm (Bortolini

et al., 2011). Reproductive activity associated with

intense rainfall has been observed in most species of

the genus, including M. stelzneri (Bustos Singer &

Gutierrez, 1997), M. montevidensis (Prigioni & Garrido,

1989; Pereira & Maneyro, 2018), M. rubiventris (Vaira,

Fig. 3. Variation in the absolute frequency of Melanophryniscus devincenzii individuals observed during

breeding activity at the study site.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

Fig. 4. a. Variation in snout-vent length (SVL, mm) and b. variation in body mass (BM, g), of males and females of M.

devincenzii.

BORTOLINI et al.

2005), M. diabolicus (Cairo et al., 2008), and M.

cambaraensis (Santos & Grant, 2011). During breeding

events, clutches and exotrophic tadpoles were

observed in flowing water bodies, indicating that the

species exhibits reproductive mode 2 (Haddad &

Prado, 2005). Egg deposition in lotic water bodies has

also been observed in Uruguayan populations of M.

devincenzii (Maneyro, 2008).

Throughout the study, 73.3% of all individuals

analyzed were males. Additionally, in two of the three

recorded reproductive events, male abundance was

higher than that of females. Similar patterns have been

reported in Uruguayan populations of M.

montevidensis, another species with an explosive

breeding strategy, where a sex ratios 3:1 or higher were

recorded during reproductive events (Pereira &

Maneyro, 2018).

Male-biased sex ratio during reproduction are

common feature in explosive breeders, as documented

by Wells (2007) and have been demonstrated in

several species of the genus Melanophryniscus, such

as M. stelzneri, M. rubriventris y M. diabolicus; (Bustos

Singer & Gutiérrez, 1997; Vaira, 2005; Cairo, 2013).

Furthermore, sexual size dimorphism was confirmed in

M. devincenzii, with females being significantly larger

and heavier than males. This pattern is consistent with

observations in most anuran species (Shine, 1979),

where larger females may produce larger clutches and

oocytes of greater size (Prado et al., 2000; Camargo et

al., 2008). However, Wells (2007) suggests that there

are no differences in the sizes of amplectant males and

females in explosive breeding dynamics. The results of

this study contradict this assertion, since sexual

dimorfism is evidente. However, this female biased

Fig. 6. Frequency distribution of Reproductive Investment (RI) in females of M. devincenzii.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

Fig. 5. a. Linear regression model between log SVL (mm) and log F (number of oocytes); and b. linear regression model

10 10

between log SVL (mm) and log OM (g) in females of M. devincenzii.

10 10

a b

size dimorphism can not be associated to female

fecundity. A possible explanation could be that M.

devincenzii males initiate reproductive activity at

smaller body sizes, allowing them to participate in

multiple reproductive events, thus maximizing their

reproductive success. This strategy would be

advantageous considering that explosive breeding

events are unpredictable due to their high dependence

on rainfall.

All analyzed females (with the exception of one),

including those collected outside reproductive events,

exhibited mature oocytes in their ovaries,

accompanied by immature oocytes in various stages

and atretic oocytes. This finding suggests that the

females are prepared for potential reproductive events

by consistently maintaining a reserve of mature

oocytes. The recorded fecundity for the females was

comparable to that reported in other species of

Melanophryniscus, such as M. diabolicus (Cairo et al.,

2008), M. cambaraensis (Caorsi, 2011), and M.

montevidensis (Pereira & Maneyro, 2018). No

significant relationship was found between fecundity

and body size, indicating that females allocate their

energy resources to reproduction rather than growth in

order to maximise future reproductive success, as

observed in species with prolonged reproductive

periods (Camargo et al., 2005; Pereira & Maneyro,

2012). Consistent with this, several females exhibited

values close to the mean and high reproductive

indices, previously reported in species exhibiting

explosive reproduction (Pereira & Maneyro, 2018).

Contrary to expectations, a significant relationship was

Fig. 7. Linear regression model between log SVL (mm) and log TM (g) in males of M. devincenzii.

10 10

Fig. 8. Temporal variation in fat body abundance in females of M. devincenzii.

Reproductive biology of Melanophryniscus devincenzii

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

found between ovarian mass and female size.

However, it is crucial to note that ovarian mass is

influenced not only by mature oocytes but also by the

immature and atretic oocytes observed in the analyzed

females. In males, size did not correlate with testicular

mass, and no monthly variation in this parameter was

recorded, which is expected, as monthly variation in

both gonadal volume and mass serves as indicators of

prolonged reproductive activity (de Oliveira et al., 2007;

Pereira & Maneyro, 2015).

Fat bodies can be used as energy reserves for

gametogenesis; they are generally scarce during the

spawning period and may decrease as oocytes

develop (Frost, 1983; Jørgensen, 1992). In species

with seasonal reproductive activity, an inverse

relationship between fat body abundance and gonadal

development has been reported, suggesting the

utilization of fat bodies as energy reserves for

reproduction (Díaz-Páez & Ortiz, 2001; Pereira &

Maneyro, 2012; Pereira et al., 2015). In the present

study, over 80% of the analyzed males exhibited fat

bodies, indicating that they consistently maintain a lipid

reserve that would enable increased testicular activity

and calling in response to a potential reproductive

event. In contrast, the analyzed females predominantly

lacked fat bodies. However, females engaged in

reproductive activity displayed fat bodies in various

categories of abundance (with the exception of

September 2009), which is consistent with an

explosive reproductive pattern. Our results indicate

that the species exhibits some of the expected

characteristics for anuran explosive breeders;

however, it should be important to analyze a greater

number of females to determine whether the dynamics

of fat bodies are consistent with this reproductive

strategy.

ACKNOWLEDGMENT

We thank to the County of Sertão for the license to

the use of area, the University of Passo Fundo for the

logistics, and the academics of the Biological Sciences

Course and Herpeto Laboratory for assistance in field

activities. We also thank ICMBio/ SISBIO for the

license (n. 26826-1/2011).

REFERENCES

Airaldi, K., Baldo, J.D., & Lavilla, E.O. (2009).

Amphibia, Anura, Bufonidae, Melanophryniscus

devincenzii: First record for Paraguay and

geographic distribution map.

Bastos, R.P.; Martins, M.R.; Guidorizzi, C.E.; Andrade,

S.P.; Bataus, Y.S.L.; Faria, D.C.C.; … Zank, C.

(2023). Melanophryniscus devincenzii. Sistema

de Avaliação do Risco de Extinção da

Biodiversidade - SALVE. Disponível em: https://

salve.icmbio.gov.br Digital Object Identifier

(DOI): https://doi.org/10.37002/salve.ficha.

17803.2 - Acesso em: 11 de set. de 2024.

Bortolini, S.V., Maneyro, R., Coppes, F.A., & Zanella, N.

(2013). Diet of Melanophryniscus devincenzii

(Anura: Bufonidae) from Parque Municipal de

Sertão, Rio Grande do Sul, Brazil. The

Fig. 9. Temporal variation in fat body abundance in males of M. devincenzii.

BORTOLINI et al.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

Herpetological Journal, 23(2), 115-119.

Bortolini, S.V., Gonçalves, F.A., & Zanella, N. (2018).

Reproductive aspects of a population of

Physalaemus gracilis (BOULENGER 1883)

(Anura: Leptodactylidae) from south Brazil.

Neotropical Biology and Conservation, 13(1),

37.

Brusquetti, F., & Lavilla, E.O. (2006). Lista comentada

de los anfibios de Paraguay. Cuadernos de

herpetología, 20.

Bustos Singer, R., & Gutiérrez, M. (1997).

Reproducción y desarrollo larval del sapo enano

Melanophryniscus stelzneri stelzneri

(Weyemberg, 1875) (Anura: Bufonidae).

Cuadernos de Herpetología, 11.

Cairo, S.L., Zalba, S.M., & Úbeda, C.A. (2008).

Reproductive behaviour of Melanophryniscus

sp. from Sierra de la Ventana (Buenos Aires,

Argentina). South American Journal of

Herpetology, 3(1), 10-14.

Cairo, S.L., Zalba, S.M., & Úbeda, C.A. (2013).

Reproductive pattern in the southernmost

populations of South American redbelly toads.

Journal of Natural History, 47(31-32), 2125-

2134.

Camargo, A., Naya, D.E., Canavero, A., da Rosa, I.,

Maneyro, R., & Naya, D.A. (2005). Seasonal

activity and the body size-fecundity relationship

in a population of Physalaemus gracilis

(Boulenger, 1883) (Anura, Leptodactylidae)

from Uruguay. In Annales Zoologici Fennici (pp.

513-521). Finnish Zoological and Botanical

Publishing Board.

Camargo, A., Sarroca, M., & Maneyro, R. (2008).

Reproductive effort and the egg number vs. size

trade-off in Physalaemus frogs (Anura:

Leiuperidae). Acta oecologica, 34(2), 163-171.

Caorsi VZ. (2011). Comportamento reprodutivo de

Melanophryniscus cambaraensis (Anura:

Bufonidae) na Floresta Nacional de São

Francisco de Paula, Rio Grande do Sul, Brasil

[undergraduate thesis]. [Porto Alegre (RS)]:

Universidade Federal de Rio Grande do Sul.

Caramaschi, U., & Cruz, C.A.G. (2002). Taxonomic

status of Atelopus pachyrhynus Miranda-

R i b e i r o , 1 9 2 0 , r e d e s c r i p t i o n o f

Melanophryniscus tumifrons (Boulenger, 1905),

and descriptions of two new species of

Melanophryniscus from the state of Santa

Catarina, Brazil (Amphibia, Anura, Bufonidae).

Arquivos do Museu Nacional, 60(4), 303-314.

Cardoso, A.J., & Martins, J.E. (1985). Diversidade de

anuros durante o turno de vocalizações, em

comunidade neotropical. Papéis avulsos de

Zoologia, 36, 279-285.

Carrillo, J.F., Rocha, P.C., Cabral, A.A.F., Firme,

M.S.F., Serafim, L.L., Mori, K.Y., & Santana, D.J.

(2024). Diet, sexual dimorphism, and fertility

aspects of Melanophryniscus fulvoguttatus

(Mertens, 1937) from Central-East Brazil.

Herpetozoa, 37, 141-147.

Cechin, S.Z., & Martins, M. (2000). Eficiência de

armadilhas de queda (pitfall traps) em

amostragens de anfíbios e répteis no Brasil.

Revista brasileira de zoologia, 17, 729-740.

de Oliveira, E.F., Feio, R.N., & da Matta, S.L.P. (2007).

Aspectos reprodutivos de Dendropsophus

minutus (Peters, 1872) no município de Viçosa,

Minas Gerais. Revista Ceres, 54(313), 231-239.

Díaz-Páez, H., & Ortiz, J.C. (2001). The reproductive

cycle of Pleurodema thaul (Anura,

Leptodactylidae) in central Chile. Amphibia-

Reptilia, 22(4), 431-445.

Duellman, W.E., & Trueb, L. (1986). Biology of

amphibians. Baltimore, the John Hopkins

University Press.

Duré, M.I., Schaefer, E.F., & Kehr, A.I. (2015). Acoustic

r e p e r t o i r e o f M e l a n o p h r y n i s c u s

cupreuscapularis (Céspedez and Álvarez 2000)

(Anura: Bufonidae): advertisement, encounter,

and release calls. Journal of Herpetology, 49(1),

53-59.

Fernández, K. (1927). Reproducción de batracios

argentinos (segunda parte). Boletin Acad. Nac.

de Ciencias en Cordoba, 29, 271-328.

Frost, J.S. (1983). Comparative feeding and breeding

strategies of a sympatric pair of leopard frogs

(Rana pipiens complex). Journal of

Experimental Zoology, 225(1), 135-140.

Frost, D.R. (2024). Amphibian Species of the World: an

Online Reference. Version 6.2 (Date of access).

Elec t ronic Data b ase a ccessi ble a t

https://amphibiansoftheworld.amnh.org/index.p

hp. American Museum of Natural History, New

York, USA. doi.org/10.5531/db.vz.0001.

Goldberg, F.J., Quinzio, S., & Vaira, M. (2006).

Oviposition-site selection by the toad

Melanophryniscus rubriventris in an

unpredictable environment in Argentina.

Canadian Journal of Zoology, 84(5), 699-705.

Grafe, T.U., Schmuck, R., & Linsenmair, K.E. (1992).

Reproductive energetics of the African reed

frogs, Hyperolius viridiflavus and Hyperolius

marmoratus. Physiological Zoology, 65(1), 153-

171.

Haddad, C.F., & Prado, C.P. (2005). Reproductive

modes in frogs and their unexpected diversity in

the Atlantic Forest of Brazil. BioScience, 55(3),

207-217.

Hammer, Ø., Harper, D.A., & Ryan, P.D. (2001). PAST:

Paleontological statistics software package for

education and data analysis. Palaeontologia

electronica, 4(1), 9.

Hillman, S.S., Withers, P.C., Drewes, R.C., & Hillyard,

S.D. (2009). Ecological and environmental

physiology of amphibians. Oxford University

Press.

IUCN SSC Amphibian Specialist Group. (2023).

9Reproductive biology of Melanophryniscus devincenzii

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8

Melanophryniscus devincenzii. The IUCN Red

List of Threaten e d Spec i e s 2023 :

e.T54819A101423809. https://dx.doi.org/10.

2305/IUCN.UK.2023-1.RLTS.T54819A101423

809.en. Accessed on 08 October 2024.

Jørgensen, C. (1992). Growth and reproduction.

Environmental physiology of the amphibians.

Lajmanovich, R.C. (2000). Interpretación ecológica de

una comunidad larvaria de anfibios anuros.

Interciencia, 25(2), 71-79.

Langone, J.A., & Lavilla, E.O. (2024). Una revisión

histórica de los nombres aplicados a las

especies del género Melanophryniscus

Gallerdo, 1961 (Amphibia: Anura: BUfonidae).

Comunicaciones Zoológicas del Museo

Nacional de Historia Natural. 13(209), 1-139.

Maneyro, R. (2008). Checklist of anurans (Amphibia,

Anura) from “Campo del Abasto” and

surroundings, Rivera department, Uruguay.

Boletín de la Sociedad Zoológica del Uruguay,

17, 34-41.

Maneyro, R., & Carreira, S. (2016). Guía de anfibios del

Uruguay. Segunda Edición. Ediciones de la

Fuga.

Marangoni, F., & Baldo, J.D. (2023). Life-history traits

of three syntopic species of the South American

redbelly t o a ds (Anura : Bufonidae:

Melanophryniscus) from the Atlantic Forest of

Argentina.

Méndez-Tepepa, M., Morales-Cruz, C., García-Nieto,

E., & Anaya-Hernández, A. (2023). A review of

the reproductive system in anuran amphibians.

Zoological Letters, 9(1), 3.

Pereira, G., & Maneyro, R. (2012). Size-fecundity

relationships and reproductive investment in

females of Physalaemus riograndensis

Milstead, 1960 (Anura, Leiuperidae) in Uruguay.

The Herpetological Journal, 22(3), 145-150.

Pereira, G., Abadie, M., & Maneyro, R. (2015).

Reproductive pattern and dynamics of fat

bodies in males of a Uruguayan population of

Physalaemus riograndensis (Anura,

Leptodactylidae). Studies on Neotropical Fauna

and Environment, 50(3), 213-220.

Pereira, G., & Maneyro, R. (2016). Use of reproductive

m i c r o h a b i t a t by Mel a n o p h r yn i s cu s

montevidensis (Anura: Bufonidae) from

Uruguay. Zoological Science, 33(4), 337-344.

Pereira, G., & Maneyro, R. (2018). Reproductive

biology of Melanophryniscus montevidensis

(Anura: Bufonidae) from Uruguay: reproductive

effort, fecundity, sex ratio and sexual size

dimorphism. Studies on Neotropical Fauna and

Environment, 53(1), 10-21.

Pereyra, L., Lescano, J., & Leynaud, G. (2011).

Breeding-site selection by red-belly toads,

Melanophryniscus stelzneri (Anura: Bufonidae),

in Sierras of Córdoba, Argentina. Amphibia-

Reptilia, 32(1), 105-112.

Pereyra, L.C., Akmentins, M.S., Sanabria, E.A., &

Vaira, M. (2016). Diurnal? Calling activity

patterns reveal nocturnal habits in the

apo sema tic toad Mel anop hryn iscu s

rubriventris. Canadian Journal of Zoology,

94(7), 497-503.

Prado, C.P.A., Uetanabaro, M., & Lopes, F.S. (2000).

Reproductive strategies of Leptodactylus

chaquensis and L. podicipinus in the Pantanal,

Brazil. Journal of Herpetology, 135-139.

Prigioni, C.M., & Garrido, R.R. (1989). Algunas

observaciones sobre la reproducción de

Melanophryniscus stelzneri montevidensis

(Anura, Bufonidae). Boletín de la Sociedad

Zoológica del Uruguay, 5, 13-14.

Saidapur, S.K., & Hoque, B. (1996). Long-term effects

of ovariectomy on abdominal fat body and body

masses in the frog Rana tigrina during the

recrudescent phase. Journal of Herpetology,

30(1), 70-73.

Santos, R.R., & Grant, T. (2011). Diel pattern of

migration in a poisonous toad from Brazil and

the evolution of chemical defenses in diurnal

amphibians. Evolutionary Ecology, 25, 249-258.

Shine, R. (1979). Sexual selection and sexual

dimorphism in the Amphibia. Copeia, 297-306.

Vaira, M. (2005). Annual variation of breeding patterns

of the toad, Melanophryniscus rubriventris

(Vellard, 1947). Amphibia-Reptilia, 26(2), 193-

199.

Valdez, V., & Maneyro, R. (2016). Reproductive biology

of Odontophrynus americanus females

(Duméril & Bribon, 1 8 4 1 ) (Anura,

Cycloramphidae) from Uruguay. Pan-American

Journal of Aquatic Sciences, 11(3), 188-197.

Wells, K.D. (1977). The social behaviour of anuran

amphibians. Animal Behaviour, 25, 666-693.

Wells K.D. (2019). The ecology and behavior of

amphibians. University of Chicago Press.

Zar, J.H. (2010). Biostatistical Analysis, Fifth Edition,

Prentice Hall.

Section Editors:

Anita Aisenberg, Macarena González,

Carolina Rojas-Buffet

BORTOLINI et al.

Bol. Soc. Zool. Uruguay (2ª época). 2025. ISSN 2393-6940Vol. 34 (1): e34.1.8