Journal of Tropical Ecology (2012) 28:125–126. © Cambridge University Press 2011
doi:10.1017/S0266467411000587
SHORT COMMUNICATION
The snake head-shape signal: a reply to Valkonen & Mappes
Murilo Guimar ˜ aes
*,1
and Ricardo J. Sawaya†
*
Departamento de Biologia Animal, Programa de P ´ os Graduac ¸˜ ao em Ecologia, Instituto de Biologia, Caixa Postal 6109, Universidade Estadual de Campinas,
13083-970, Campinas, S ˜ ao Paulo, Brazil
† Departamento de Ci ˆ encias Biol ´ ogicas, Universidade Federal de S ˜ ao Paulo, Rua Prof. Artur Riedel, 275, 09972-270, Diadema, S ˜ ao Paulo, Brazil
(Accepted 25 September 2011)
Key Words: defence, head triangulation, mimicry, morphology, plasticine replicas, predation, snakes
It has already been suggested that snake head
triangulation might be related to mimicry of the head
shape of vipers (Greene & McDiarmid 2005, and
references therein). Until very recently, this hypothesis
has never been experimentally tested. We first tested the
hypothesis of snakes’ head shape as a dangerous signal
to predators by use of plasticine models (Guimar˜ aes &
Sawaya 2011). We suggested in that study that shape
of the head does not confer advantage itself but may
work in synergy with a set of traits including colour and
behavioural displays that warn and discourage predator
attacks.
Shortly after our publication, Valkonen et al. (2011)
published an article in which they suggest triangular
head shape was a warning signal to predators, also
based on experiments using plasticine snakes. Valkonen
& Mappes (2012) argue: ‘In contrast to the findings of
Guimar˜ aes & Sawaya (2011), our results demonstrate
that the triangular head shape typical of vipers can act as
a warning signal to predators.’
We consider the debate on the evolutionary
significance of head shape and its possible associated
mimetic behaviour very productive. In order to further
advance that discussion, we need to clarify some
points discussed by Valkonen & Mappes (2012). The
combination of colour pattern and head shape used by
Valkonen et al. (2011) may have played an important
role in repelling predators. We think a more conservative
1
Corresponding author. Email: mu.guima@gmail.com
interpretation would be one very similar to ours: head
shape associated with colour pattern can act as a warning
signal to predators. Thus, we do not agree that their
findings are in contrast with ours, but in the same
direction.
However, an interesting point discussed by Valkonen
& Mappes (2012) is that attacks by mammalian and
avian predators must not be pooled together. It would
be better indeed to consider only avian predators in
order to ascertain that only visual signals are involved.
In our experiment (Guimar˜ aes & Sawaya 2011) we
pooled attacks by mammalian and avian predators. We
recorded 17 and 11 attacks to oval and triangular-shaped
heads, respectively, pooling avian and mammalian
predators, which was not significantly different.
Considering separately the two types of predators in
our experiments (omitted from our publication), we
recorded for avian predators 11 and seven attacks to
oval and triangular-shaped heads, respectively; and for
mammalian predators six and four attacks, respectively.
Those results would not change our conclusions, even
when considering only avian predators (χ
2
= 0.89, df = 1,
P = 0.35).
More studies are necessary to better understand the
evolutionary role of head shape of snakes as visual
cues to predators and its possible related mimetic
complexes (see Greene & McDiarmid 2005 for a review,
complementary questions and future prospects in snake
mimicry). However, we believe head triangulation must
be considered a complementary warning signal to
predators, acting within a set of traits in the defensive
behaviour of snakes.