Ecological Modelling 251 (2013) 32–43
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Ecological Modelling
jo u r n al hom ep age : www.elsevier.com/locate/ecolmodel
Trait contributions to fish community assembly emerge from trophic
interactions in an individual-based model
Henrique C. Giacomini
a,∗
, Donald L. DeAngelis
b
, Joel C. Trexler
c
, Miguel Petrere Jr.
a,1
a
Departamento de Ecologia, Universidade Estadual Paulista “Júlio de Mesquita Filho”, Campus de Rio Claro, Avenida 24-A, 1515, CEP 13506-900, Rio Claro, SP, Brazil
b
U.S. Geological Survey, Department of Biology, University of Miami, 271 Cox Science Center, 1301 Memorial Drive, Coral Gables, FL 33146, USA
c
Department of Biological Sciences, Florida International University, 3000 NE 151 St, North Miami, FL 33181, USA
a r t i c l e i n f o
Article history:
Received 28 May 2012
Received in revised form
11 November 2012
Accepted 4 December 2012
Available online 9 January 2013
Keywords:
Community dynamics
Fish diversity
Life history
Environmental gradients
Tradeoffs
Priority effects
a b s t r a c t
Community ecology seeks to understand and predict the characteristics of communities that can develop
under different environmental conditions, but most theory has been built on analytical models that are
limited in the diversity of species traits that can be considered simultaneously. We address that limi-
tation with an individual-based model to simulate assembly of fish communities characterized by life
history and trophic interactions with multiple physiological tradeoffs as constraints on species perfor-
mance. Simulation experiments were carried out to evaluate the distribution of 6 life history and 4 feeding
traits along gradients of resource productivity and prey accessibility. These experiments revealed that
traits differ greatly in importance for species sorting along the gradients. Body growth rate emerged as
a key factor distinguishing community types and defining patterns of community stability and coexis-
tence, followed by egg size and maximum body size. Dominance by fast-growing, relatively large, and
fecund species occurred more frequently in cases where functional responses were saturated (i.e. high
productivity and/or prey accessibility). Such dominance was associated with large biomass fluctuations
and priority effects, which prevented richness from increasing with productivity and may have limited
selection on secondary traits, such as spawning strategies and relative size at maturation. Our results
illustrate that the distribution of species traits and the consequences for community dynamics are inti-
mately linked and strictly dependent on how the benefits and costs of these traits are balanced across
different conditions.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
Community assembly is the process by which immigrants sur-
vive to form a local assemblage of coexisting species. This process
is dynamic, with the resultant community at any point in time
depending on the previous balance between successful invasions
and deterministic or stochastic extinctions (Drake, 1990; Tilman,
2004). Assembly studies – and community studies in general –
are much more interesting when functional traits are explicitly
accounted for, assuming they are relevant to species adjustment
to environmental conditions and to resource partitioning, working
as a signature of niche differentiation and structuring by inter-
actions (McGill et al., 2006). Accordingly, compelling arguments
have been made for a trait-based approach in community ecology
∗
Corresponding author. Present address: Department of Ecology and Evolution-
ary Biology, University of Toronto, Office RW 520B, 25 Harbord St., Toronto, ON,
Canada M5S 3G5. Tel.: +1 416 978 7338; fax: +1 416 978 5878.
E-mail address: hgiacomini@gmail.com (H.C. Giacomini).
1
Present address: UNISANTA - Mestrado em Ecologia, Rua Doutor Oswaldo Cruz,
266, Boqueirão, 11045-907 Santos, SP, Brazil.
(McGill et al., 2006; Mims et al., 2010). Many trait-based stud-
ies of animal community assembly have searched for evidence of
regular niche spacing among co-occurring species (i.e. trait overdis-
persion), based on a variety of morphological traits (Gotelli and
Graves, 1996). But trait overdispersion is not expected to occur in
studies encompassing large spatial scales or very heterogeneous
landscapes (Weiher and Keddy, 1995), in which case the search for
trait-environment associations is the usual and most interesting
aspect of assembly theory.
For fish, especially in freshwater, the trilateral classification of
Winemiller and Rose (1992) among opportunistic, periodic and
equilibrium strategies has been a good predictor of general associ-
ations between species life history and environmental conditions
(Mims et al., 2010; Mims and Olden, 2012). In this case, and in many
empirical investigations, habitat predictability and the strength of
biotic interactions are the main factors driving selection of life
history traits such as fecundity, life span, and offspring size (e.g.
Reznick and Endler, 1982; Roff, 1992; Lamouroux et al., 2002;
Mims and Olden, 2012). For studies interested in the outcomes
of biotic interactions, as in the present paper, two key gradients
influencing species traits and diversity are resource productiv-
ity and predation pressure (Sih et al., 1985; Rosenzweig, 1995;
0304-3800/$ – see front matter © 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.ecolmodel.2012.12.003