Passive and Active Defense in
Toads: The Parotoid Macroglands
in Rhinella marina and
Rhaebo guttatus
PEDRO L. MAILHO‐FONTANA
1
,
MARTA M. ANTONIAZZI
1
, LUÍS F. TOLEDO
2
,
VANESSA K. VERDADE
3
, JULIANA M. SCIANI
1
,
KATIA C. BARBARO
1
, DANIEL C. PIMENTA
1
,
MIGUEL T. RODRIGUES
4
, AND CARLOS JARED
1
*
1
Instituto Butantan, São Paulo, Brazil
2
Museu de Zoologia, Universidade Estadual de Campinas, Campinas, Brazil
3
Centro de Ciências Naturais e Humanas, Universidade Federal ABC, Santo André, Brazil
4
Departamento de Zoologia, Universidade de São Paulo, São Paulo, Brazil
The skin of amphibians is characterized by the presence of mucous
glands, mainly associated with protection against desiccation, and
granular glands, or poison glands, associated with chemical
defense against predators and microorganisms (Toledo and
Jared, '95; Hillman et al., 2009).
In some species the poison glands are grouped into large and
very conspicuous protuberances, such as the parotoid macroglands
ABSTRACT
Amphibians have many skin poison glands used in passive defense, in which the aggressor causes its
own poisoning when biting prey. In some amphibians the skin glands accumulate in certain regions
forming macroglands, such as the parotoids of toads. We have discovered that the toad Rhaebo
guttatus is able to squirt jets of poison towards the aggressor, contradicting the typical amphibian
defense. We studied the R. guttatus chemical defense, comparing it with Rhinella marina,a
sympatric species showing typical toad passive defense. We found that only in R. guttatus the
parotoid is adhered to the scapula and do not have a calcified dermal layer. In addition, in this
species, the plugs obstructing the glandular ducts are more fragile when compared to R. marina. As
a consequence, the manual pressure necessary to extract the poison from the parotoid is twice as
high in R. marina when compared to that used in R. guttatus. Compared to R. marina, the poison of
R. guttatus is less lethal, induces edema and provokes nociception four times more intense. We
concluded that the ability of R. guttatus to voluntary squirt poison is directly related to its
stereotyped defensive behavior, together with the peculiar morphological characteristics of its
parotoids. Since R. guttatus poison is practically not lethal, it is possibly directed to predators'
learning, causing disturbing effects such as pain and edema. The unique mechanism of defense of R.
guttatus may mistakenly justify the popular myth that toads, in general, squirt poison into people's
eyes. J. Exp. Zool. 9999A: XX–XX, 2013.
©
2013 Wiley Periodicals, Inc.
How to cite this article: Mailho‐Fontana PL, Antoniazzi MM, Toledo LF, Verdade VK, Sciani JM,
Barbaro KC, Pimenta DC, Rodrigues MT, Jared C. 2013. Passive and active defense in toads: The
parotoid macroglands in Rhinella marina and Rhaebo guttatus. J. Exp. Zool. 9999:1–13.
J. Exp. Zool.
9999A:1–13, 2013
Grant sponsor: CAPES and CNPq‐INCTTox—Brazilian Federal Government.
Correspondence to: Carlos Jared, Cell Biology Laboratory, Instituto
Butantan, Av. Vital Brasil 1500, CEP 05503‐000 São Paulo, Brazil.
E‐mail: jared@butantan.gov.br
Received 8 July 2013; Revised 6 September 2013; Accepted 13
September 2013
DOI: 10.1002/jez.1838
Published online XX Month Year in Wiley Online Library
(wileyonlinelibrary.com).
RESEARCH ARTICLE
©
2013 WILEY PERIODICALS, INC.