Adaptive significance of food income in European
snakes: body size is related to prey energetics
MARCO A. L. ZUFFI
1
*, SARA FORNASIERO
1
, RICCARDO PICCHIOTTI
1
, PIERA POLI
2
and MARCELLO MELE
2
1
Museo di Storia Naturale e del Territorio, Università di Pisa, via Roma 79, I-56011, Calci (Pisa),
Italy
2
Dipartimento di Agronomia e gestione dell’Agroecosistema, Università di Pisa, via San Michele degli
Scalzi 5, I-56121 Pisa, Italy
Received 7 September 2009; revised 1 December 2009; accepted for publication 1 December 2009
Feeding strategies and diet patterns have been extensively investigated in vertebrates and, more specifically, in
snakes. Although it has been hypothesized that prey species may differ in terms of energy content, almost no
theoretical or practical study has been carried out to determine actual nutritional values of the common prey types
of wild snakes. Our model taxa were a selection of widely distributed and well known European snake species,
which have all been studied in depth: approximately 76% of their diet is composed of mammals, reptiles, and
insects. We therefore selected a single model species for each of these categories and proceeded with the analyses.
Nutritional values were determined using a standard procedure: lizards and mice were richer in proteins than
insects (crickets); insects and mice were richer in lipids than lizards, and mice and crickets have a higher energy
content than lizards; lizards were rich in ashes. We then applied our experimental results to a selected sample of
European terrestrial snakes (11 populations, ten species, seven genera, two families) characterized by different
body size (50–160 cm total length) and reproductive strategies (oviparous versus viviparous), aiming to correlate
these parameters with patterns of energy income. A direct relationship was found between body mass/body length
ratio (BCI, body condition index) and meal energetics: the higher the BCI, the higher was the metabolic
requirement, whereas BCI was independent of species or of reproductive system effect. Large-sized snakes thus
need a highly diversified and more energy-rich diet than smaller snakes, supporting previous hypotheses. The
simple applicability of this method could be of valuable support in further comparative research work, reducing
experimental costs and stimulating further ecological, behavioural, and, possibly, phylogenetic comparisons.
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 307–317.
ADDITIONAL KEYWORDS: Colubridae – comparative analysis – diet – energy balance – Serpentes –
Viperidae.
INTRODUCTION
The energetics of food income, patterns of energy
consumption and field metabolic rates among verte-
brates have been reported at great length (Nagy,
2005; Codron et al., 2007), and food quality (Lanszki,
Molnár & Molnár, 2006), food abundance, and choice
of selected prey items (Estades, Vukasovic &
Tomasevic, 2008) have been studied in depth. Energy
requirements and subsequent dietary patterns have
often been related to sex, behavioural patterns, sea-
sonal variation, or to body size (Kohler et al., 2008;
Jiang et al., 2009).
The consequences of different experimental food
intake treatments, as well as their resulting effects
on growth patterns, physiology of development, and
reproduction, have recently received specific attention
in amphibians and in reptiles (Cox & Secor, 2007;
Secor, Wooten & Cox, 2007). Indeed, analyses on daily
energetic metabolism have greatly enhanced our
overall knowledge on the metabolic turnover in a
number of reptiles (Avery, 1971, 1978; Gallagher
*Corresponding author. E-mail: marcoz@museo.unipi.it
Biological Journal of the Linnean Society, 2010, 100, 307–317. With 1 figure
© 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 307–317 307