RESEARCH
PAPER
Trophic level scales positively with body
size in fishesgeb_579 1..10
Tamara N. Romanuk*, April Hayward† and Jeffrey A. Hutchings
Department of Biology, Dalhousie University,
1355 Oxford Street, Halifax, Nova Scotia,
Canada B3H 4J1
ABSTRACT
Aim The existence of a body size hierarchy across trophic connections is widely
accepted anecdotally and is a basic assumption of many food-web models. Despite
a strong theoretical basis, empirical evidence has been equivocal, and in general the
relationship between trophic level and body size is often found to be weak or
non-existent.
Location Global (aquatic).
Methods Using a global dataset for fishes (http://www.fishbase.org), we explored
the relationship between body size and trophic position for 8361 fishes in 57 orders.
Results Across all species, trophic position was positively related to maximum
length (r
2
= 0.194, b = 0.065, P < 0.0001), meaning that a one-level increase in
trophic level was associated with an increase in maximum length by a factor of 183.
On average, fishes in orders that showed significantly positive trophic level–body
size relations [mean = 51.6 cm 11.8 (95% confidence interval, CI)] were 86 cm
smaller than fishes in orders that showed no relation [mean = 137.1 cm 50.3
(95% CI), P < 0.01]. A separate slopes model ANCOVA revealed that maximum
length and trophic level were positively correlated for 47% (27 of 57) of orders, with
two more orders showing marginally non-significant positive relations; no signifi-
cant negative correlations were observed. The full model (order ¥ body size)
explained 37% of the variation between body size and trophic position (P <
0.0001).
Main conclusions Our results support recent models which suggest that trophic
level and body size should be positively correlated, and indicate that morphological
constraints associated with gape limitation may play a stronger role in determining
body size in smaller fishes. Differences among orders suggest that the nature of the
trophic level–body size relation may be contingent, in part, on evolutionary history.
Keywords
Allometry, body mass, fish, FishBase, gape limitation, metabolic theory of
ecology, scaling, size classes.
*Correspondence: Tamara N. Romanuk,
Department of Biology, Dalhousie University,
1355 Oxford Street, Halifax, Nova Scotia,
Canada B3H 4J1.
E-mail: tromanuk@gmail.com
†Present address: Department of Biology,
University of Florida, PO Box 118525,
Gainesville, FL 32611, USA.
INTRODUCTION
Body size is of paramount importance in ecological dynamics
(Elton, 1927; Peters, 1983; Calder, 1984; Schmidt-Nielsen, 1984;
Cohen et al., 1993; Brown et al., 2004; Marquet et al., 2005; Arim
et al., 2007; Carbone et al., 2007): it shapes the demand that
organisms place on their environment for energy and materials
(through its well known, if poorly understood, relationship with
metabolic processes; Brown et al., 2004) and plays an important
role in structuring both inter- and intra-specific interactions.
Among the most important interactions affected by body size
are consumer–resource interactions, as body size constrains the
range of prey sizes a predator can consume (Cohen et al., 1993;
Arim et al., 2007; Carbone et al., 2007; Hildrew et al., 2007). The
existence of a body size hierarchy across trophic connections is
widely accepted as a basic assumption of many food-web
models (Cohen et al., 1993; Williams & Martinez, 2000; Becker-
man et al., 2006; Petchey et al., 2008). Cohen et al. (1993), for
example, have suggested that body size provides a mechanistic
interpretation of the assumed trophic hierarchy of the cascade
Global Ecology and Biogeography, (Global Ecol. Biogeogr.) (2010)
© 2010 Blackwell Publishing Ltd DOI: 10.1111/j.1466-8238.2010.00579.x
www.blackwellpublishing.com/geb 1