Behavioural Brain Research 173 (2006) 315–319
Short communication
Lateralized righting behavior in the tortoise (Testudo hermanni)
Gionata Stancher
a
, Elena Clara
a
, Lucia Regolin
a,∗
, Giorgio Vallortigara
b
a
Department of General Psychology, University of Padova, Via Venezia 8, 35131 Padova, Italy
b
Department of Psychology, B.R.A.I.N. Centre for Neuroscience, University of Trieste, Trieste, Italy
Received 28 April 2006; received in revised form 9 June 2006; accepted 13 June 2006
Available online 1 August 2006
Abstract
Lateralization of brain and behaviour at the population level has been documented in all vertebrate classes. Research was mostly carried out on
mammalian and avian species, the least investigated class with this regards being the Reptilia, with studies concentrating on lateralized aggressive
behaviour in lizards. No research has been carried out on lateralization in the Chelonian order. We investigated the presence of motor asymmetries
in the tortoise Testudo hermanni, using the righting response (i.e. the animal is positioned upside-down and the left/right side to which it uprights is
observed), a procedure already employed to assess behavioural lateralization in amphibians. The ability of righting has a particularly high adaptive
value in tortoises, as in case of overturning, and consequent exposure to sunrays, changes in body temperature and difficulties in respiration could
occur leading to serious conditions. Thirty-four tortoises underwent a series of righting tests in a standardized apparatus, 15 tortoises were also
retested 10 months later. A bias at the individual as well as at the population level was found for preferentially turning on the right side. Consistency
of responses at retest was also observed. The results are discussed with reference to the implications for the evolution of brain lateralization in
vertebrates.
© 2006 Elsevier B.V. All rights reserved.
Keywords: Righting response; Tortoise; Testudo hermanni; Motor asymmetry; Behavioural lateralization
A shared general pattern of lateralization can be found for
most vertebrate species [16,24,25]. Asymmetries in the central
nervous system could manifest themselves as lateral biases in
overt behaviour; motor lateralization is attested in all vertebrate
classes and has been investigated in several amphibian and rep-
tilian species [13,15,23].
Male lizards are more likely to attack conspecifics located on
their left side [5,6,10,11]. In these studies a general tendency was
found to respond (either to approach or to attack) to conspecifics
when they appeared on the left side. Responses to the right side
took longer than those to the left side and, for example, were used
to select food objects and to respond to stimuli after allocating
them to categories rather than on an individual basis [14].
Studies on newborn and adult amphibians have taken advan-
tage of their spontaneous righting response when overturned
[2,3,15]. According to these studies individual and population
biases do exist with a consistent favourite righting preferred side
in 70–80% of the cases.
∗
Corresponding author. Tel.: +39 049 8276650; fax: +39 049 8276600.
E-mail address: lucia.regolin@unipd.it (L. Regolin).
The righting response was also used in youngsters of Chely-
dra serpentina as a measure of fitness [19], it correlates with
temperature of incubation [7], but up to now it was never used
as a measure of lateralized behaviour.
We investigated the presence of lateralized bias in right-
ing responses in two subspecies of the Mediterranean tor-
toise Testudo hermanni [9]. T. h. ssp. hermanni and T. h. ssp.
boettgeri were classified as subspecies on the base of morpho-
logical, genetic, and biogeographical considerations. The two
subspecies are currently distributed to the west (ssp. hermanni)
and east (ssp. boettgeri) side of the Adriatic Sea, hence an
allopatric speciation process is assumed [26]. Modern popula-
tions are still reproductively isolated even though, during the last
10,000 years, with the end of the last glaciation, their habitats
progressively extended to the North.
Righting is a highly adaptive and efficiently performed
response for tortoises, being it of vital importance as, when
overturned, they might experience rather serious difficulties
at breathing and at body temperature control. Overturning
can occur in the natural environment during male fights [4],
and, occasionally, when climbing/walking through the uneven
grounds typical of Mediterranean habitat. The special adap-
0166-4328/$ – see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.bbr.2006.06.023