- Functional traits and phylogeny - 381
Journal of Vegetation Science 19: 381-392, 2008
doi: 10.3170/2008-8-18378, published online 10 March 2008
© IAVS; Opulus Press Uppsala.
Functional traits and phylogeny: What is the main ecological process
determining species assemblage in roadside plant communities?
Valladares, F.
1,2*
; Tena, D.
1,5
; Matesanz, S.
1,6
; Bochet, E.
3,7
; Balaguer, L.
4,8
;
Costa-Tenorio, M.
4,9
; Tormo, J.
3,10
; García-Fayos, P.
3,11
1
Instituto de Recursos Naturales. Centro de Ciencias Medioambientales, CSIC. Serrano 115 Bis, ES-28006 Madrid, Spain
2
Departamento de Biología y Geologia, Escuela Superior de Ciencias Experimentales y Tecnológicas, Universidad Rey
Juan Carlos, c/ Tulipán s/n, ES-28933 Móstoles, Spain;
3
Centro de Investigaciones sobre desertiicación (CSIC, Universidad Valencia, Generalitat Valenciana), Camí de la
Marjal s/n, ES-46470 Albal, Valencia, Spain;
4
Departamento de Biología Vegetal I, Facultad de Biología, Universidad
Complutense de Madrid, C/ Jose Antonio Novais s/n, ES-280040, Madrid, Spain;
5
E-mail tiotena@hotmail.com;
6
E-mail silvia@ccma.csic.es;
7
E-mail esther.bochet@uv.es;
8
E-mail balaguer@bio.ucm.es;
9
E-mail mct@bio.ucm.es;
10
E-mail jaume.tormo@uv.es;
11
E-mail patricio.garcia-fayos@uv.es;
*
Corresponding author; Fax +34 913940800; E-mail valladares@ccma.csic.es
Abstract
Question: What is the main ecological process determining spe-
cies assemblage in roadside herbaceous plant communities?
Location: Roadside slopes (roadcuts and embankments) in the
south (Málaga, mesic Mediterranean) and east (Valencia, dry
and continental) of the Iberian Peninsula.
Methods: We identiied 417 plant species, from which we
selected the 331 most abundant (within the 70th abundance
percentile) at each site. We compiled information on 28 func-
tional traits and on the biogeographic range of each of these
331 species. We quantiied the phylogenetic signal of each
trait for the species of each community and determined the
number of functional convergences or divergences over the
phylogenetic tree for each of the four situations (roadcuts and
embankments in the two sites).
Results: There was a signiicant phylogenetic signal in many
traits, being positive in Valencia embankments and negative in
Valencia roadcuts with almost no signal in any type of slope in
Málaga. Each trait was signiicantly correlated with 20% - 35%
of all other traits but correlation coeficients were low. No sig-
niicant phylogenetic signal was found for the species’ distribu-
tion range in any of the four communities studied, which might
be the consequence of the complex mixture of biogeographic
origins of the species found in these communities.
Conclusion: The lack of a phylogenetic signal in most traits
in Málaga, a climatically favourable locality, suggests that
competitive exclusion was the main process involved in the
assemblage of these communities. The signiicant and either
positive or negative phylogenetic signal (in embankments and
roadcuts respectively), the latter coupled with a signiicant
number of functionally convergent nodes in the phylogenetic
tree, suggests that environmental iltering is the most likely
process involved in the harsh locality of Valencia.
Keywords: Environmental gradient; Mediterranean; Phyloge-
netic signal; Plant community; Road slope.
Abbreviation: AOT = Analysis of traits.
Introduction
We have little knowledge of the mechanisms deter-
mining the assemblage of the species that make up a
community and, although it has been known for some
time now that neither plants nor their functional traits are
distributed randomly (Raunkiær 1934; Diamond 1975),
numerous studies are still attempting to determine the
processes that give rise to community structure (e.g.
Keddy & Weiher 2001; Temperton & Hobbs 2004). From
an ecological point of view, two main hypotheses might
account for the non-random distribution of a group of
species and their functional traits (Tofts & Silvertown
2000). On one hand, the classical theory of competitive
exclusion states that competition is directly proportional
to the similarity among species, which brings co-existing
species to maximise the difference for a given functional
trait (Johansson & Keddy 1991). This trait is, in turn,
distributed among the species of the community in an
overdispersed manner (i.e. exhibiting phenotypic repul-
sion) regarding a null hypothesis of random distribution
of species and functional traits (Armbruster et al. 1994;
Wilson & Watkins 1994; Wilson & Gitay 1995). On
the other hand, community structure might result from
the inluence of environmental ilters and only those
species possessing speciic attributes would persist in
the community (Montalvo et al. 1991; Fernández-Alés
et al. 1993; Díaz & Cabido 1997; Díaz et al. 2001). In
this case, the species would be more similar than what
would be expected by chance and a given trait would be
clustered (i.e. exhibiting phenotypic attraction) regarding
a null hypothesis of randomness. Neither hypothesis is
exclusive, so that competitive exclusion and environmen-
tal ilters concur, although their relative importance can
differ depending on environmental conditions (Weiher &