OIKOS 100: 223–231.

2003

MINI- Minireviews provides an opportunity to summarize existing knowledge of selected

ecological areas, with special emphasis on current topics where rapid and significant
REVIEW advances are occurring. Reviews should be concise and not too wide-ranging. All key
references should be cited. A summary is required.

Guilds or functional groups: does it matter?

Jacques Blondel

Blondel, J. 2003. Guilds or functional groups: does it matter? – Oikos 100: 223– 231.

Although most researchers use the terms ‘‘guild’’ and ‘‘functional group’’ more or
less synonymously, these two concepts bear different meanings. The guild concept
refers primarily to the mechanisms of resource sharing by species in a competitive
context whereas the functional groups concept is concerned with how a resource or
any other ecological component is processed by different species to provide a specific
ecosystem service or function. In many cases but not necessarily all, the two concepts
are the two ‘‘faces’’ or ‘‘sides’’ of the same coin: the sharing by species of a similar
resource is the guild facet (structural), while the ecosystem processes these species
eventually perform through resource exploitation is the functional group facet. The
two concepts differ in that competitive relationships within groups of species are not
the focus of the functional group approach, exactly as processes or functions are not
the focus of the guild approach. A group of species can be considered either as a
guild or a functional group depending on the question addressed. Guild and
functional group membership is independent of phylogenetic relationships but be-
cause species tend to share similar life history traits and adaptations through
common evolutionary history, guild and functional group associates are often closely
related. The concept of guild has had broader application in animal studies than in
plant studies, whereas the reverse is true for the concept of functional group. Recent
methodological advances to objectively partition species into guilds and functional
groups, taking into consideration the most relevant characters or traits for delineat-
ing them, provide the means to construct an operational framework for making in
situ and ex situ experiments that are urgently needed for a better understanding of
the role of species in ecosystem functioning, especially in relation to global change
concerns.

J. Blondel, CEFE-CNRS, F-34293 Montpellier cedex 5, France ( jacques.blondel@

cefe.cnrs-mop.fr).

In a seminal paper published more than 40 years ago,
George Evelyn Hutchinson (1959) paved the way to a
new approach in understanding community organisa-
tion and structure. At a time when ecologists were
much interested in mechanisms responsible for species
coexistence he developed the view that the boundaries
of realised niches are set by competition for limited
resources. Hutchinson was not directly concerned with
guilds or functional groups, not even citing these terms,
but he implicitly assumed that groups of species sharing

Accepted 19 August 2002

Copyright © OIKOS 2003
ISSN 0030-1299

OIKOS 100:2 (2003) 223

similar resources are common in nature because ‘‘a
complex trophic organisation of a community is more
stable than a simpler one’’ (Hutchinson 1959, p. 155).
The main question was: what sets limits to the number
of species coexisting at a given trophic level, or to cite
Hutchinson himself, ‘‘why are there not more different
kinds of animals?’’. His approach on species coexistence
was based on morphological differentiation of traits of
importance for resource acquisition, assuming that the
spacing out of niches is reflected in the degree of
morphological divergence between species exploiting
the same resource dimension. Hutchinson empirically
found from many examples in vertebrates and inverte-
brates that adjacent species tend to exhibit regular
differences in body size or in the size of feeding struc-
tures with size ratios being approximately 2.0 for body
mass and 1.3 (the cubic root of 2.0) for length of
feeding structures. This Hutchinson’s ratio rule together
with the assumption that species are ecologically unique
was the cornerstone of the widely accepted theories of
the niche and of dynamic equilibrium which have been
a starting point for hundreds of studies on the structure
of communities (Cody and Diamond 1975, Diamond
and Case 1986, Putman 1994 and references therein).
This classical approach to community ecology led to
the notion that communities are built from groups of
species that share certain similarities, either ecological
or phylogenetic. Hence the need arose to split commu-
nities into groupings of lower rank on the basis of
similarity in resource utilisation. In this context the
term ‘‘guild’’ was first utilised by Root (1967) and then
the parallel term ‘‘functional groups’’ by Cummins
(1974). Simberloff and Dayan (1991) reviewed the guild
concept and provided some insight into its relationship
to functional groups, concluding that most researchers
actually use them more or less synonymously with a
preference for guild because ‘‘the metaphor of the guild
seemed more elegant’’. Steneck (2001) also remarked
that ‘‘depending on the application, guilds can be syn-
onymous with functional groups’’.
Interest in the concepts of guilds and functional
groups has increased in recent years in the context of
current concerns about the effects of global changes,
especially biodiversity decline, CO2 increase, climate
warming, and various ecosystem disturbances including
invasions, on such issues as community dynamics and
ecosystem functioning (Walker 1992, Körner 1993,
Prentice et al. 1993, Diaz 1995, Chapin et al. 1996,
Bond 1997, Lavorel et al. 1997, Smith et al. 1997,
Wilson 1999 and references therein). In this paper, I
shall discuss, mainly from a zoological point of view,
how the two terms have been or should be considered,
focusing mostly on vertebrates and emphasising what
differentiates them (Table 1).
Guilds as arenas for deciphering mechanisms
of coexistence and resource sharing
The term ‘‘guild’’ was used many decades ago by plant
geographers (Schimper 1903) and plant ecologists
(Clements 1905), but Root (1967), who is a zoologist,
first formally defined the term as ‘‘a group of species
that exploit the same class of environmental resources
in a similar way. This term groups together species
without regard to taxonomic position, that overlap
significantly in their niche requirements’’. The rationale
is that the ecological relationships among guild associ-
ates are moulded by competition for limited resources,
which is by no means surprising considering the im-
petus given at the time to interspecific competition by
Hutchinson (1959) and the work of MacArthur and his

Table 1. Attributes, characteristics and fields of application of guilds and functional groups.

Guilds Functional groups

Definition based on Similarity in resource sharing Similarity in ecosystem function

What they are Permanent or temporary structural subunits Permanent or temporary assemblages of species
of communities; depend on resources performing a same ecosystem process
What they do Exploit resources in a similar way Process the same resources or habitat features
(e.g. soil) for an ecosystem service
How they are described Structural criteria Process-oriented criteria
Questions addressed Structure at the community scale, arenas for Processes at the ecosystem scale, arenas for
deciphering differences (coexistence) in an deciphering similarities (redundancy) for
evolutionary context achieving ecosystems functions
Predictions from removal Changes in the abundance of the remaining No response if redundancy (compensation);
experiments species (compensation) change in ecosystem response if not
Intra-group interspecific Competition between species in a ‘‘species Similarities between species in a functional
relationships packing’’ context context
Inter-group interspecific More competition within guilds than between Not relevant
relationships guilds
Taxonomic scale Often within-taxon assemblages Often across-taxon assemblages

224 OIKOS 100:2 (2003)

colleagues (Hutchinson and MacArthur 1959,
MacArthur and Levins 1967) for explaining the struc-
ture of communities (Brown 1981). This means that
guilds are assumed to be coevolved entities and ‘‘arenas
of intense interspecific competition’’ (Pianka 1980, Jak-
sić and Medel 1990).
However, if interspecific competition has for long
been considered as the main force driving species pack-
ing along resource gradients, challenging views have
been proposed on several counts e.g. by Jaksić (1981),
Jaksić et al. (1993) and Wiens (1989, 1993, see also
Chesson and Case 1986). Wiens (1993) predicts that
niche overlap among species varies according to re-
source abundance with less competition and more over-
lap in resource use when resource is abundant than
when it is rare, a prediction that had been supported by
Schoener (1982) in his review on competition studies.
Root did not consider any functionality in the guild
concept which, as he wrote, is ‘‘the most evocative and
succinct term for groups of species having similar ex-
ploitation patterns’’ (Root 1967, p. 336). The kind of
resource used to define guilds, e.g. food, habitat struc-
tures, nest-sites, roost-sites, hosts, or whatever else
shows that ecological processes are not involved in the
definition and utilisation of the term because the guild
concept only focuses on similarity in resource sharing
irrespective of the ecological consequences (i.e. func-
tion) of resource sharing (Table 1). Therefore any guild
designations which are not based on resource sharing
but on other characters such as demographic or be-
havioural traits, violate the definition of the guild (Sim-
berloff and Dayan 1991). The guild concept became a
standard currency among community ecologists in the
golden age of community ecology in the sixties and
seventies (Cody and Diamond 1975, Brown 1981, Dia-
mond and Case 1986, Simberloff and Dayan 1991), and
generated two kinds of studies (Table 1): (1) those that
described, organised and classified the species that com-
pose communities in operational and logical subunits
(Cody 1974, Holmes et al. 1979), and (2) those whose
purpose was to explain the ecological and evolutionary
mechanisms that permit coexistence of species along
resource gradients (species packing, Landres and Mac-
Mahon 1980, Case et al. 1983, Dayan et al. 1989, 1990,
McNaughton 1993). Segregation along resource gradi-
ents of guild associates has been primarily based on
differences in morphological traits closely associated to
feeding techniques and often assumed to be generated
through character displacement.
A same species may belong to different guilds during
its life as a result of ontogenetic changes in resource
use, a point which has not been much considered in
studies of guild structure within communities although
some authors (MacNally 1983, Simberloff and Dayan
1991, Gerking 1994) mentioned that ontogenetic shifts
could modify the guild structure of species assemblages.
Ontogenetic changes in resource use have been reported
OIKOS 100:2 (2003)
in many groups of animals, invertebrates of course with
completely different life forms in larval and adult
stages, but also fishes (Werner and Gilliam 1984).
Theoretically guild membership is independent of
phylogenetic relationship but since closely related spe-
cies likely share similar life history traits and adapta-
tions to resources and habitats, members of a guild are
often closely related. Few attempts have been made to
include in the same guild distantly related species,
especially multi-class groupings, in part because of the
practical difficulty of deciphering interspecific interac-
tions among very different species. This explains why
guilds and most functional groups have been investi-
gated within single-taxon assemblages, such as bird,
lizard, or small mammal assemblages (Jaksić 1981).
Exceptions are the studies of Jaksić and Medel (1990)
and Jaksić et al. (1993) on the guild structure of a three
class-predatory assemblage including reptiles, mammals
and birds in California, and of granivorous communi-
ties of rodents, birds and ants in desert ecosystems
(Brown et al. 1979).
If the concept of guild had much success in animal
studies, it has not been very popular in plants, for
several reasons. Perhaps most importantly the notion of
a resource that can be exploited ‘‘in a similar way’’ is
not obvious for plants, and very difficult to categorise
and quantify (Simberloff and Dayan 1991, de Kroon
and Olff 1995). This explains that attempts to define
plant guilds resulted in classifications which did not
differ much from the classical life-form approach
(Raunkiaer 1934) or other traditional systems of clas-
sification. In recent years, however, the term has been
applied to investigate the responses of potentially com-
peting plant species to ecosystem disturbance (Lavorel
et al. 1997, Wilson 1999).
From guilds to functional groups
The guild approach has sometimes been considered as a
starting point for understanding the role of guild asso-
ciates in communities (Jaksić 1981). Therefore, one
step further has been to explore whether guild associ-
ates have similar functional roles in spite of their eco-
logical uniqueness. If so, this would imply that different
communities are constructed with similar kinds of
guilds. This rationale, which makes a link between
guilds and functional groups, led to the proposition
that guilds are fundamental units of communities. This
conceptual extension addresses the two fundamental
collective attributes of species assemblages, namely the
sharing by species of similar resources that are ex-
ploited in a similar way (the guild), and the ecosystem
processes they eventually perform through resource ex-
ploitation (the functional group). Depending on the
question addressed, the same suite of species can be
225
considered either as a guild or as a functional group, as
explained in Table 1. It should be noted that the guild
concept only refers to relationships between guild asso-
ciates for resource acquisition, whereas the functional
group concept refers to a wide range of potential
ecosystem functions such as biochemical cycles, resis-
tance to invasion or to fire, resource acquisition, water
uptake, storage of resources, defence against herbivory,
pollination, seed dispersal, or any physical processes
such as ecosystem engineering, disturbance and biotur-
bation. When the resource considered is food, which is
the case in the vast majority of guild studies in animals,
guild associates are by necessity the predators of the
resource they exploit, so that traits of interest are
related to food acquisition whereas in the functional
group concept, any kind of trait can theoretically be
involved depending on the function considered. Such
explicit differences between the two concepts should
help avoid the confusion between ‘‘functional’’ or ‘‘ef-
fect groups’’ and ‘‘response groups’’ (Lavorel and Gar-
nier 2002). For example, Gilbert (1980) proposed a
scheme to show how the bulk of neotropical diversity is
organised by chemical mosaics and mutualism. Key
elements of the food web organisation include ‘‘mobile
links groups’’ i.e. suites of species that perform such
important functions as pollination (hummingbirds,
bats, moths and bees), seed dispersal (birds and bats),
and plant defence (ants). Species involved in each of
these functions belong to the same effect group but, as
guild associates sharing the same resource, which is
either nectar (pollinators) of fruits (seed dispersers),
they also belong to the same response group.
Functional groups as clusters of players
engaged in the same ecosystem processes
Cummins (1974) emphasised the ‘‘…need to identify
functional groups of organisms, at least partially inde-
pendent of traditional taxonomic determinations, in
order to address important process-oriented ecosystem
questions’’ thereby defining the term ‘‘functional
group’’ which, historically, has been introduced after
the guild concept. Most authors, e.g. Körner (1993) and
Hobbs et al. (1995), gave somewhat similar definitions,
focusing on the ‘‘similar roles or processes’’ assumed to
be achieved by functional groups associates. Thus spe-
cies within a functional group are by definition ecologi-
cally equivalent and provide some degree of
redundancy to the system. The term is definitely pro-
cess-oriented and has been widely used in this sense,
especially in invertebrate communities (Hawkins and
MacMahon 1989, Table 1). It also bears a slightly
different meaning in plant ecology by designating spe-
cies with similar responses to environmental factors, the
so-called ‘‘response groups’’ (Lavorel et al. 1997, La-
vorel and Garnier 2002).
226
The main conceptual differences between guilds and
functional groups are tentatively summarised in Table
1. The two concepts differ in that competitive relation-
ships within groups are not included in the definition of
the functional group concept, exactly as processes or
functions are not included in the definition of the guild
concept. A same group of species can be considered
either as a guild or a functional group depending on the
question addressed. Guilds are concerned with identify-
ing species that share a common resource and determin-
ing how this resource is shared among them, the
‘‘similar way’’ of the Root’s definition, whereas func-
tional groups are concerned with how the resource is
processed by different species in such a way that it
provides an ecosystem function, the ‘‘similar role’’ of
the functional group definition. In the guild approach
the subject is the species which exploit resources for
themselves whereas in the functional group approach
the subject is the resource which, so to speak, utilises
mediators, i.e. members of a functional group, for being
processed (Table 1). As noted by MacMahon et al.
(1981), ‘‘…it does not matter whether an organism
removes a tree leaf for nesting material, for food, or as
a substrate to grow fungi which in turn are eaten; the
leaf is gone and the leaf users belong to a common
guild’’. The different leaf consumers admittedly belong
to the same guild since they share the same resource
(which is actually questionable because the leaf resource
is rarely limited), but the functional response, if any, is
quite different depending on how and for which pur-
pose the leaf has been removed and processed. From
his studies of Serengeti savannahs, McNaughton (1993)
demonstrated a relationship between grassland diversity
and the grazing pressure of herds of four species of
ungulates (Thompson’s gazelles, African buffaloes,
plain zebras, and blue wildebeest). These four species
belong to the same guild because they share the same
resource for which they potentially compete, but the
point which makes them at the same time a functional
group is that the grazing pressure of the four species
combined has an effect on an important ecosystem
attribute, namely plant species diversity. The guild ap-
proach has been much more popular than the func-
tional group approach in vertebrates whereas the
opposite is true for invertebrates, micro-organisms and
plants.
As for the guild concept, the term functional group
does not imply any phylogenetic relationships among
the species although several species in a functional
group are likely to be closely related. Examples of
multi-taxon functional groups are seed dispersers in
desert ecosystems which include ants, birds and rodents
(Brown et al. 1979) or pollinators of Lauraceae which
include hummingbirds, bats and moths (Gilbert 1980).
Yet anatomical, morphological or physiological traits
are not the cornerstone of functional groupings (Ste-
neck 2001), perhaps because multi-taxon identification
OIKOS 100:2 (2003)
of functional groups is extremely difficult, mostly for
practical reasons. How can one sample and analyse by
the same approach distantly related taxa that differ in
morphology, physiology, behaviour, and demography,
but perform similar functions, such as for example
saproxylic beetles, nematodes and fungi which are en-
gaged in the same process of recycling dead wood?
How can one classify and assign in functional groups
(or guilds) animals which change their diet seasonally,
or which have a completely different biology depending
on whether they are in larval of adult stages (amphibi-
ans, many invertebrates)?
Convergence in guild structure and ecosystem
functioning
The statement of May and Seger (1986) ‘‘…the view
that there exist ecological roles is supported by the
observation that ecological communities often bear
striking resemblance to each other though their con-
stituent species have very different evolutionary
origins’’ opened the door to the idea that guilds and
functional groups should be repeatable in space and
time, irrespective of which species compose them.
Repeatability in space is a basic assumption of the
convergence paradigm (Mooney 1977, Orians and Sol-
brig 1977, Cody and Mooney 1978). The rationale of
convergence is that partitioning of communities into
groups of species with morphological resemblances (i.e.
guilds) should be similar in different regions with simi-
lar environmental conditions, whatever the phyloge-
netic relatedness of the constituting species (Hawkins
and MacMahon 1989). Terborgh and Robinson (1986)
explained the many efforts to demonstrate morphologi-
cal convergence as reflecting attempts to demonstrate
that similar morphologies are involved in the achieve-
ment of the same ecosystem functions. In this line of
reasoning various studies compared the structure of
parallel guilds in different regions, e.g. in birds (Recher
1969, Cody 1973, 1975, Cody and Mooney 1978, Rick-
lefs and Travis 1980). However, a recurrent problem in
these kinds of studies, which has been only recently
more or less satisfactorily solved, was the arbitrary
threshold of ecological overlap chosen to define guilds,
with the risk of ad hoc interpretation of guild conver-
gence (Blondel et al. 1984). Orians and Paine (1983)
who discussed many of the problems of convergence
studies were skeptical about convergence, arguing that
the resemblance between communities is often more
parsimoniously explained by similar physiological re-
sponses of individual species to similar environments.
The high degree of similarity between communities
probably reflects phylogenetic constraints and the lim-
ited number of adaptive zones or adaptive solutions to
common environmental conditions more than evolu-
OIKOS 100:2 (2003)
tionary convergence, especially in vertebrates such as
birds or fishes, which are distributed in the three dimen-
sions of space. Perhaps one of the most convincing
examples of convergent guilds and functional groups is
that of nectar-feeding birds, which belong to distantly
related groups living in different regions: sunbirds (Nec-
tariniidae) in the paleotropics, honeycreepers
(Drepanididae) on the Hawaiian Islands, hummingbirds
(Trochilidae) in the New World, and honeyeaters
(Meliphagidae) in Australasia. All these birds exploit
the same resource in a similar way which involves a
high specialisation of bill and tongue to extract nectar
from long tubular flowers. In this particular example
convergence can be extended to the astonishing resem-
blance of flying characteristics and mouthparts among
the pollinating butterflies, moths, bats, and birds.
Repeatability of guilds in time should also be demon-
strable if guilds are fundamental units or basic ‘‘build-
ing blocks’’ of communities. One example of such
‘‘guild signatures’’ (Karr 1980) in time has been pro-
vided by a complete guild turnover realised from exper-
imental defaunation of small mangrove cays off the
coast of Florida (Simberloff and Wilson 1969).
Analysing the trophic organisation of invertebrate com-
munities in seven ‘‘trophic classes’’ before experimental
defaunation and after recolonisation, Heatwole and
Levins (1972) found a striking similarity in the appor-
tionment of species in the different trophic classes,
which by the way are also functional groups, despite
large taxonomic differences and heterogeneity among
the islands. The trophic structure of communities was
re-established with different species belonging to the
same trophic levels, indicating some constancy of guild
structure in time.
Making links between guilds and functional
groups
In many pollination and seed dispersal processes plants
and animals are engaged in more or less sophisticated
mutualistic interactions (McKey 1975, Wheelwright and
Orians 1982, Herrera 1995). These systems help us to
make a link between the way resources are partitioned,
which is a guild matter, and the fate of the resource
itself, which is a functional group matter. Murray et al.
(1987) reported on a variety of plant species, often quite
distantly related, that are pollinated by two guilds of
hummingbirds. Plant species are only guild associates
because they share the same resource (pollinators) but
hummingbirds are at the same time guild and func-
tional group members because in addition to sharing
the same resource from the plants (nectar), they also
achieve the process of pollination.
In an insightful paper on the relationships between
plants and birds that disperse their propagules, Ferry
227
(1984) considered the coevolutionary processes from
the points of view of both the plants and the birds.
Bird-dispersed seeds belong to several groups that cor-
respond to different evolutionary strategies. Fruits can
be caught or stuck in the plumage or fur of a bird or a
mammal (epizoochory), berries can be swallowed and
then the seeds regurgitated or defecated at some dis-
tance (endozoochory), or else dry fruit seeds can be
cached to be found again some time later with ‘‘forgot-
ten’’ caches being eventually available for germination
(synzoochory). Whereas epizoochory is a passive, non-
functional process from the animal’s point of view
because there is no reward for dispersing the seeds, the
two other strategies result in the same functional pro-
cess but require two different bird guilds because the
‘‘subject resource’’ is exploited through different ways
that all together result in processing the same ecosystem
function. From the birds’ point of view, there are only
two guilds because only fruits of endozoochorous
plants (berries, drupes) and those of synzoochorous
plants (dry seeds) are a dietary resource for them.
Plants engaged in close mutualistic interactions with
dispersing animals evolved a syndrome of ‘‘zoochory’’,
i.e. they evolved similar or complementary fruit charac-
ters such as size, shape, colour and nutritive value of
the fruits (Herrera 1995). All the plants that are dis-
persed by the same set of species belong to the same
guild because they potentially compete for the same
shared resource, i.e. birds. All bird species that share
the same food resource, for examples berries, thus
belonging to a same guild, evolved similar morphologi-
cal, physiological, digestive and behavioural traits al-
lowing them to exploit this resource (Herrera 1995).
With respect to participating in seed dispersal, the
different assemblages of birds are not only guilds but
also functional groups. In the context of a guild analy-
sis, Snow (1971) classified seed-eating birds as ‘‘legiti-
mate’’ and ‘‘illegitimate’’ seed-eaters. The former
participate in the process of dispersal by regurgitating
or defecating the intact seed which will eventually ger-
minate, thus belonging to a functional group. In con-
trast, the latter destroy the seed by eating it. Both
legitimate and illegitimate animals can belong to the
same guild but only the former belong to the seed-dis-
persing functional group.
Other good examples of functional groups are earth-
worms, which include several species assemblages con-
tributing differently to major soil ecological processes
(Lavelle 1997), and dung beetles, whose function is to
recycle the non-digestive part of the green food of large
herbivores. The 135 or so species of dung beetles in the
Mediterranean region have been assigned to three main
groups depending on how they process and use the
dung (Lumaret 1995). The ‘‘dweller species’’ (e.g.
Aphodius) live inside the dung, the ‘‘tunneller species’’
(e.g. Bubas) bury pieces of faecal material under the
dung itself down to a depth of ca 0.5 – 1 m, and finally
228
the ‘‘roller species’’ (e.g. Scarabaeus) make a ball of
dung within which females lay their eggs and roll it
over several metres until they find a soil suitable for
burying it. All dung beetles exploit the same resource,
hence they belong to the same guild, but the different
ways of processing this resource result in different
decomposition processes (e.g. shallow vs deep soil cy-
cling of nutriments), hence different functional groups.
How to define guild and functional group
membership?
Perhaps the most difficult problem of the guild/func-
tional group endeavour, which presumably explains
why the two concepts have fallen into disuse (Salt
1979), is that species partitioning into well-defined
groups has too often been done on rather arbitrary
grounds. Many studies relied more on taxonomy and
intuition to partition a priori species into guilds and
functional groups than on sound quantitative methods,
and comparisons of species groupings to those expected
under null hypotheses have been rare. Defining guilds
and functional groups without objective procedures,
and using them as convenient terms for describing
groups of taxonomically related syntopic species, car-
ries the risk of making these concepts no more than
empty buzzwords. I will not enter into the methodolog-
ical aspects of defining guild and functional group
membership because this would be beyond the scope of
this essay. I will only make one point about interspe-
cific competition which is central in the guild approach.
The most rigorous means to properly assign species
into guilds is to test whether guild associates are in
exploitative competition, past or present (MacNally
1983). From his studies on New Guinea birds, Dia-
mond (1975) developed the theme of competitively de-
termined assembly rules governing the structure and
size of guilds. The underlying assumption is that
‘‘through diffuse competition, the component species of
a community are selected, and coadjusted in their
niches and abundances, so as to fit with each other and
to resist invasion’’ (Diamond 1975, p. 342). Accord-
ingly some combinations of species are permissible and
resist invaders whereas others are forbidden. But this
approach was challenged, and degenerated in a hot
debate, mostly because species combinations had not
been compared to null hypotheses and competition had
not been demonstrated but just inferred from compara-
tive studies (Connor and Simberloff 1979, Diamond
and Gilpin 1982, Gilpin and Diamond 1984). However,
from an analysis of many guild studies, MacNally
(1983) actually demonstrated that exploitative competi-
tion usually occurs among guild members, especially
when guilds include a moderate to small number of
species. As already explained in this paper, Jaksić et al.
OIKOS 100:2 (2003)
(1993) and Wiens (1989, 1993) pointed out that inter-
specific competition is not necessarily a prerequisite for
the emergence of guild structure. Opportunistic use of
temporarily superabundant resources may set the stage
for the same emergent phenomenon. Demonstrating
current morphological responses through character dis-
placement within guild members would disabuse the
‘‘ghost of competition past’’ (Connell 1980) whereby
the extinction of competitors does not change the pat-
terns of current resource sharing which resulted from
past coevolutionary specialisation. Such morphological
responses have been elegantly demonstrated in some
studies, for example in Darwin’s finches by Schluter et
al. (1985) and in carnivorous mammals (canids,
mustelids, viverrids, and felids) by Dayan et al. (1989,
1990).
Much effort has been made recently to objectively
establish guild and functional group membership by
Jaksić and Medel (1990) and Pillar (1999) who gave
means to determine cut-off points in cluster analysis
from suitable computer algorithms. A plethora of more
or less sophisticated statistical multivariate procedures
for guild assignment among which cluster analyses,
principal component analyses, canonical correlations,
Monte Carlo techniques, bootstrap procedures have
been commonly used (Jaksić and Medel 1990, Sim-
berloff and Dayan 1991, Muñoz and Ojeda 1998, Pillar
1999).
Conclusion
Guild members share a same resource and by doing so
may eventually become involuntary partners for achiev-
ing an ecosystem function, whereas functional groups
include by definition partners that are engaged in run-
ning a same ecosystem process. Guilds and functional
groups may be the two facets of the same coin (struc-
tural vs functional) but this is not always the case
because resource use does not necessarily affect the way
an ecosystem good or service is provided or performed.
However, the difference between the two terms is not
always reflected in the studies that actually divide spe-
cies among guilds and functional groups. This is pre-
sumably because resource partitioning often (but not
always) links species with ecosystem functioning
(Loreau et al. 2001) so that, to some degree, the re-
source partitioning that guilds rely on is often relevant
for ecosystem functioning. To make character-based
guilds and functional groups more operational, rigor-
ous criteria should be used to select those species-spe-
cific traits that are the most relevant in resource sharing
(guilds) or in processing a function. To reach this goal
in the functional group approach, the identification of
well-defined target ecosystem processes is important
because the breakdown of communities into functional
OIKOS 100:2 (2003)
groups depends on the process considered. In situ and
ex situ experiments should help answer the kind of
questions that are relevant to the functional group
approach, e.g. whether functional group associates are
equivalent or redundant (Walker 1992), play a role of
‘‘insurance’’ for the maintenance of ecosystem func-
tions, ‘‘plug leaks’’ in the efficiency of resource
turnover, or provide immunity against pathogens or
disturbances. Another interesting question to be experi-
mentally investigated is whether some species are struc-
tural with regards to ecosystem processes (keystone
species) whereas others are interstitial. Consequences of
species extinctions on ecosystem functioning will be
quite different whether the former or the latter are lost.
The distinction between guilds and functional groups
should certainly be made clearer and more operational
by carefully relating these terms to groups of species
depending on what they really do in nature, either
sharing a same resource for themselves as subjects, or
using a same resource that makes them associates in
running a same function (functional groups). Making
the guild/functional group framework more operational
is important in the scope of the current concern about
the effects of global change on ecosystem structure and
function.
Acknowledgements – I am most grateful to Eric Garnier and
Robert E. Ricklefs for most useful comments and suggestions.
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