Morphological traits affect escape behaviour of the Balearic lizards
(Podarcis lilfordi)
Dror Hawlena
1,*
, Valentín Pérez-Mellado
2
, William E. Cooper Jr.
3
Abstract. Escape theory predicts that a prey should start escaping (flight initiation distance = FID) from a predator when the
costs of fleeing and the cost staying are equal or until future fitness is maximized. Consequently, prey escape performances
and current reproductive asset can affect FID. We tested effects of body condition, morphology, and whether the tail was
regenerated or original on FID in the Balearic lizard (Podarcis lilfordi) by ourselves simulating predators. Lizards with better
body condition had longer FID and lizards with longer intact tails had shorter FID. Lizards with regenerated tail presented
shorter FID than lizards with intact tails. These results suggest that impaired escape performance is counterbalanced by fitness
costs of tail regeneration or by alteration of escape behaviour. The weak association between morphology, body condition
and FID suggest that escape performances and asset protection have relatively small effect on P. lilfordi escape decisions.
Keywords: antipredatory behaviour, escape, flight initiation distance, predation risk, refuge, Squamata.
Escape theory predicts that a prey should
start to escape from an approaching predator
when the costs of fleeing (primarily opportu-
nity costs) and not fleeing (due to predation
risk) are equal (Ydenberg and Dill, 1986) or
until expected fitness is maximized (Cooper
and Frederick, 2007). Flight initiation distance
(= FID, distance between predator and prey
when escape starts) is predicted to be shorter
when cost of escaping increases or risk of
delaying escape decreases. These predictions
have been verified in various taxa (reviewed
by Stankowich and Blumstein, 2005) for many
environmental factors affecting risk, includ-
ing predator approach speed (Cooper, 1997a;
Cooper et al., 2003), number and relative attack
angles of predators (Cooper, Pérez-Mellado and
Hawlena, 2007) proximity of refuge (Cooper,
1997b), and weather conditions (Rand, 1964;
Hertz, Huey and Nevo, 1982), as well as fac-
tors affecting the costs of fleeing such as loss
of feeding and social opportunities (Cooper,
1 - School of Forestry & Environmental Studies, Yale Uni-
versity, 370 Prospect St., New Haven, CT 06511, USA
2 - Departamento de Biología Animal, Universidad de Sala-
manca, 37071 Salamanca, Spain
3 - Department of Biology, Indiana University Purdue Uni-
versity Fort Wayne, Fort Wayne, IN 46805, USA
*
Corresponding author; e-mail: dror.hawlena@yale.edu
1999, 2000b; Diaz-Uriarte, 1999; Martín, López
and Cooper, 2003a; Cooper, Pérez-Mellado and
Hawlena, 2006; Cooper and Wilson, 2007).
Risks and costs can vary as a consequence of
variation in morphological traits and body con-
dition of the prey (Stankowich and Blumstein,
2005). Prey that can escape faster or more effi-
ciently have lower cost of not fleeing when at a
given distance from a predator, but slower max-
imum speed might be associated with a change
in escape tactics toward greater reliance of cryp-
sis due to immobility to avoid being detected
and attacked (Cooper et al., 1990). Faster speed
alone would predict shorter predicted FID due
to lower risk at a given distance, but a switch
to greater crypsis would predict shorter FID. In
lizards running speed increases as body size and
hind limb length increase; tail length is also ex-
pected to affect speed (Warner and Shine, 2006;
Cooper and Wilson, 2008). Consequently in the
absence of change in tactics, escape theory pre-
dicts that individuals having greater body length
(snout-vent length = SVL) and longer hind
limbs will have shorter FID than conspecifics
with shorter SVL and limbs.
Effects of body condition on FID are diffi-
cult to predict because body condition may af-
fect both escape ability and initial fitness. Poor
body condition associated with decreased speed
© Koninklijke Brill NV, Leiden, 2009. Amphibia-Reptilia 30 (2009): 587-592
Also available online - www.brill.nl/amre