Basic and Applied Ecology 13 (2012) 109–115

INVITED VIEWS IN BASIC AND APPLIED ECOLOGY

Imagine a world without seed dispersers: A review of threats,
consequences and future directions
Nina Farwig∗ , Dana G. Berens
Dept. Ecology – Conservation Ecology, University of Marburg, Karl-von-Frisch-Str. 8, 35043 Marburg, Germany

Received 29 July 2011; accepted 18 February 2012

Abstract
Seed dispersing animals, ranging from small insects to large mammals, provide a crucial service for a large number of plant
species worldwide. However, a decline in dispersers due to direct and indirect threats leads to disruptions of seed dispersal
processes. As disperser species are differently susceptible to these threats, consequences for ecosystems are hard to predict.
Impacts range from hampered regeneration of plant species to shifts in communities and a decline in ecosystem function. Here,
we review these threats as well as expected consequences for communities and for the entire ecosystem. We further introduce
options to protect dispersers and consider future research directions.

Zusammenfassung
Von Insekten bis hin zu Großsäugern spielen Tiere weltweit eine wichtige Rolle als Samenausbreiter für zahlreiche
Pflanzenarten. Jedoch führen direkte und indirekte Gefährdungen zum Rückgang von Samenausbreitern und den von ihnen
abhängigen Prozessen. Da Ausbreiterarten unterschiedlich störungsanfällig sind, ist es schwierig, Konsequenzen für das gesamte
Ökosystem vorherzusagen. Bisherige Auswirkungen reichen von verminderter Regeneration verschiedener Arten über Ver-
schiebungen in Pflanzengemeinschaften bis hin zu einem Rückgang an Ökosystemfunktion. Wir fassen die Gefährdungen sowie
deren Konsequenzen für Gemeinschaften und Ökosysteme zusammen. Außerdem geben wir einen Überblick über mögliche
Optionen zum Schutz von Samenausbreitern sowie über zukünftige vielversprechende Forschungsfragen.
© 2012 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Keywords: Animal–plant interaction; Environmental change; Functional consequences; Dynamics of species communities; Human impact;
Loss of biodiversity; Seed dispersal; Specialization

∗ Corresponding author. Tel.: +49 6421 2823478; fax: +49 6421 2823387.

E-mail address: farwig@biologie.uni-marburg.de (N. Farwig).

1439-1791/$ – see front matter © 2012 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.baae.2012.02.006
110 N. Farwig, D.G. Berens / Basic and Applied Ecology 13 (2012) 109–115
Introduction
The dependence of plants on animals as dispersal vectors
varies across the globe. In tropical regions, up to 90% of
plant species are dispersed by animals (Howe & Smallwood
1982). The importance of animals as seed dispersers is less
pronounced in temperate regions, varying from 0% to 60%
(Willson, Rice, & Westoby 1990). Still, even in biomes such
as the arctic tundra, animal seed dispersal plays an impor-
tant role in shaping plant communities (Bruun, Lundgren, &
Philipp 2008).
Seed dispersal by animals has both ecological and
evolutionary consequences for biodiversity and ecosystem
function. In the following, we briefly introduce the broad
spectrum of animals acting as seed dispersers and focus on the
relevance of this mutualistic interaction for species, commu-
nities and ecosystems. Further, we review factors leading to
disperser loss as well as observed consequences. Last, we dis-
cuss options to protect dispersers and consider future research
directions that might help us understand the dynamics of this
biotic interaction.
Who disperses seeds?
A broad range of animals are involved in seed dispersal pro-
cesses, ranging from small insects to large vertebrates. The
majority of dispersers are mammals, birds and ants (Stiles
2000). Other, less common, but nonetheless interesting vec-
tors are annelids (Eisenhauer & Scheu 2008), slugs (Türke
et al. 2010), beetles (De Vega, Arista, Ortiz, Herrera, &
Talavera 2011), grasshoppers (Duthie, Gibbs, & Burns 2006),
amphibians (Da Silva & De Britto-Pereira 2006), reptiles
(Traveset & Riera 2005), or fish (Galetti, Donatti, Pizo, &
Giacomini 2008). Animals transport seeds externally, e.g.
attached to their fur or as cachers (e.g. Fischer, Poschlod,
& Beinlich 1996; Gómez 2003), or internally as food items
(Howe & Westley 1988).
Dispersing animals carry seeds away from the mother
plant. While ants transport seeds over a distance of a few
meters (Gomez & Espadaler 1998), migratory birds may
transport seeds over several hundreds of kilometers (Clausen,
Nolet, Fox, & Klaassen 2002). Thus, animals provide seed
dispersal on a local, regional, or even transcontinental scale.
Going along with the variety of seed dispersers, dispersal
modes and extents of dispersal, animals differ in their quan-
tity, quality and effectiveness of dispersal services provided
to plants (Schupp 1993; Schupp, Jordano, & Gomez 2010).
What is the relevance of animal-mediated
seed dispersal?
Seed dispersal by animals involves a great diversity of ani-
mal and plant lineages and is an essential component in the
formation and maintenance of biodiversity as it has led to
morphological, physiological and behavioral adaptations in
both plants and animals (Herrera & Pellmyr 2002).
Seed dispersal provides benefits for both animals and
plants. Fruit-eating animals profit by dispersing seeds in the
form of nutrition (Howe & Westley 1988). From the plant’s
perspective, dispersal provides escape from high density of
competing siblings, from natural enemies in the vicinity of
the parent trees and enables the colonization of vacant recruit-
ment sites (Howe & Smallwood 1982). Especially directed
dispersal to most suitable habitats for plant recruitment is
largely dependent on the movement behavior of dispers-
ing animals and has pivotal implications for communities
(Wenny & Levey 1998; Gómez 2003). Further, seed disper-
sal is vital for the maintenance of genetic diversity and for
the adaptation potential of plants to micro-environmental or
broad-scale ecological conditions (Hamrick, Murawski, &
Nason 1993).
Seed dispersal is thus vital for the functioning of ecosys-
tems (Fig. 1). Animal seed dispersal facilitates natural
regeneration of plant species that are used for timber, fuel-
wood or medicinal purposes (Wenny et al. 2011). Further,
it is a key process for natural restoration of degraded areas
worldwide, thereby directly benefiting humans by reducing
management costs (Wunderle 1997; Sekercioglu, Daily, &
Ehrlich 2004; Mendez, Garcia, Maestre, & Escudero 2008).
Consequently, it significantly influences the carbon storage
potential of different habitats and, in the long-term, global
change (Laurance et al. 2006; Brodie & Gibbs 2009).
What are the main threats to seed
dispersers?
The global loss of biodiversity does not spare animals act-
ing as seed dispersers. Birds, as one of the major groups of
dispersers, are highly threatened, with 13% of species being
prone to extinction within the next 100 years (Sodhi, Liow, &
Bazzaz 2004). Directly and indirectly, human activities are
the main drivers of disperser loss. Among the major direct
threats are hunting for meat, ornaments or medicinal purposes
(e.g. Wright et al. 2000), and poisoning for pest control (e.g.
Brake & Smith 2005). Both hunting and poisoning target at
large species potentially acting as long-distance dispersers
(Wright 2003).
The range of human activities leading indirectly to the loss
of seed dispersers is overwhelming. Deforestation through
conversion into agricultural land and commercial logging
entails habitat loss for a wide range of species, especially in
tropical forests (Achard et al. 2002). Also other forms of habi-
tat alteration, e.g. fragmentation, intensification of farmland
production or urban expansion pose a threat to seed dis-
perser communities worldwide (Sekercioglu & Sodhi 2007;
Tscharntke et al. 2008). Increasing land-use intensity, e.g.
by enhanced use of pesticides and fertilizer, has led to the
 N. Farwig, D.G. Berens / Basic and Applied Ecology 13 (2012) 109–115 111

altered services

Ecosystem functionality
e.g.
- ecosystem goods
- natural restoration
Human activities feedback loops - carbon storage feedback loops

Direct
e.g.
- hunting Seed disperser Plant communities
- poisoning threatening changing interactions e.g.
communities
e.g. - genetic composition
Indirect - unique dispersers - community shifts
e.g. - large-bodied species - regeneration potential
- habitat alteration - functional diversity feedback loops
- land-use
- invasive species

Other trophic
communities
e.g.
- microorganisms
- herbivores
- predators

Fig. 1. Schematic diagram depicting the impact of human activities on seed dispersers and potential consequences for communities and
ecosystems: direct and indirect threats alter different aspects of seed disperser communities, thereby changing mutualistic interactions with
consequences for various facets of plant communities. These modifications may entail negative feedback loops within communities and
cascading effects on other trophic communities. In the long-term they imperil ecosystem functionality as well as human wellbeing.
impoverishment of farmland biodiversity, thereby also affect- vulnerable to hunting (Corlett 2007). As larger animals are
ing frugivorous animals (Freemark & Kirk 2001). Further, more affected by persecution than small-sized animals, plant
human-induced species invasions can lead to the disruption of species depending on their dispersal services are particularly
plant–animal interactions that have developed over evolution- threatened (Galetti, Donatti, Pires, Guimarães, & Jordano
ary timescales, which often entails a loss of native dispersers 2006; Peres & Palacios 2007).
(Traveset & Richardson 2006 and references therein). To Overall niche breadth is probably the most important
make things worse, drivers of disperser loss often act syn- determinant of a species’ tolerance to anthropogenic threats
ergistically: habitat loss and degradation entail a number of (Swihart et al. 2003). Specialized species are more sensitive
secondary threats, as they facilitate the colonization of inva- than generalist species, as flexibility in resource use allows a
sive species, the spread of diseases, and provide easier access species to respond to environmental changes (Swihart et al.
for hunters (Sodhi et al. 2004; Wotton & Kelly 2011). 2003; Colles, Liow, & Prinzing 2009). Especially habitat
specialization is closely linked to extinction risk (Owens &
Bennet 2000).
Which dispersers suffer most?
Certainly, not all species are equally susceptible to What would a world without dispersers look
extinction. A suite of traits may determine a species’ capa- like?
bility of adaptation to environmental changes, ranging from
physiological, morphological, and ecological to behavioral As expected, several studies show that loss of dispersers
attributes (Swihart, Gehring, Kolozsvary, & Nupp 2003). impairs interactions between plants and their mutualistic
More vagile birds show a higher level of persistence in East partners and subsequent dynamics in plant recruitment (e.g.
African forest fragments than less mobile species (Lens, Cordeiro & Howe 2003; Forget & Jansen 2007; Kirika,
Van Dongen, Norris, Githiru, & Matthysen 2002). Large Bleher, Böhning-Gaese, Chira, & Farwig 2008; Kirika,
body sizes usually imply low reproductive rates, leading to Farwig, & Böhning-Gaese 2008). Changes in the disperser
a low ability to recover from population declines (Owens & community lead to altered spatial recruitment patterns of
Bennet 2000; Safi & Kerth 2004). Orangutans, for example, affected species, e.g. if dispersal becomes restricted to the
reach sexual maturity comparatively late and have long inter- canopy radius of a species (Bond & Slingsby 1984; Christian
birth intervals, which makes their populations particularly 2001). Changes are most pronounced for species depending
112 N. Farwig, D.G. Berens / Basic and Applied Ecology 13 (2012) 109–115
on unique dispersal services: while highly specialized and
unique disperser species are most likely to go extinct, the
probability of others taking their role is low (Sekercioglu
et al. 2004). Non-random directed dispersal by Muntjac deer
to open sites has been shown to enhance the regeneration of
a tropical tree, a service that is not provided by other dis-
persers (Brodie, Helmy, Brockelman, & Maron 2009). Often
large-seeded plants fail to regenerate due to hunting of key
vertebrate dispersers (Vanthomme, Bellé, & Forget 2010).
Campos-Arceiz and Blake (2011) could show that the Asian
tapir is not able to replace elephants and rhinos in their func-
tion as dispersers of large-seeded plants as tapirs mainly act
as seed predators of these species. Wotton and Kelly (2011)
state that dispersal of five large-seeded tree species in New
Zealand is depending mainly on the New Zealand pigeon,
a seed disperser whose numbers have decreased dramati-
cally since human arrival on the island. This makes these
tree species especially susceptible to regeneration failure.
Changes in seed dispersal processes further influence the
genetic make-up of plant populations. Reduced genetic vari-
ability results in diminished adaptation potential (Keller &
Waller 2002). Moreover, reduced gene flow among plant
populations enhances the impact of inbreeding and genetic
drift and influences the genetic structure of plant populations
(Aldrich, Hamrick, Chavarriage, & Kochert 1998; Wang,
Leong, Smith, & Sork 2007; Farwig, Braun, & Böhning-
Gaese 2008).
Besides affecting the abundance and genetic template of
single species, loss of seed dispersers modifies diversity and
dynamics of plant communities. Thereby, the rich assemblage
of species dispersed by large vertebrates may be substituted
by fewer species dispersed abiotically or by small animals
(Terborgh et al. 2008). In consequence, the loss of disperser
species entails a decline in plant diversity. Ultimately, ongo-
ing co-extinction of species of both taxa may lead to a
downward spiral of diversity in the whole ecosystem (Bond
1995).
Consequently, the loss of dispersers imperils the function-
ing of ecosystems (Fig. 1). Reduced dispersal rates entail in
the long-term diminished population sizes of valuable timber
species and non-timber forest products (Wenny et al. 2011).
So far, research on the economical implications of seed dis-
perser loss is still scarce. However, increasing management
costs are expected if, due to the loss of animal dispersers,
the potential for natural regeneration as well as restoration
of degraded sites is missing. Thus, losing dispersers might
in the future even reduce the important carbon sink potential
especially of forest ecosystems (Brodie & Gibbs 2009).
How can we save seed dispersal?
Management approaches for the conservation of seed
dispersal should focus on protecting the functionality of
ecosystems. This implies to protect plant and animal species
that are involved in this mutualistic interaction as well as
the management of high-quality habitats at different spatial
scales (Telleria, Ramirez, & Perez-Tris 2005). The protection
of charismatic dispersers such as elephants or tapirs involves
the conservation of large contiguous habitats and simultane-
ously preserves other less conspicuous dispersers (Roberge &
Angelstam 2004). Similarly, the protection of keystone seed
dispersers, e.g. flying foxes on tropical islands, preserves an
important function for the entire ecosystem (Cox, Elmqvist,
Pierson, & Rainey 1991). Strategies that control illegal hunt-
ing, e.g. wildlife farming, have been successfully employed to
protect large-bodied dispersers (Nogueira & Nogueira-Filho
2011). Another management option is the systematic control
of competitive invasive species, which could preserve vital
indigenous disperser–plant interactions (Bellingham et al.
2010).
The protection of high-quality habitat comprises a holis-
tic community-wide approach. At a local scale, a mosaic of
high-quality habitats such as fragments with a high avail-
ability of food resources and nesting sites can maintain
high numbers of seed dispersers (e.g. Anzures-Dadda &
Manson 2007; Garcia, Zamora, & Amico 2010). Notably,
the conservation of keystone resources such as figs could
be a successful conservation tool in modified landscapes
(Terborgh 1986). Increasing connectivity through corridors
or isolated trees enhances the movement of animals within
fragmented landscapes and in turn preserves their regener-
ation potential (Berens, Farwig, Schaab, & Böhning-Gaese
2008; Neuschulz, Botzat, & Farwig 2011). Thus, even the
conservation of small habitat patches seems to provide great
potential for conservation of ecosystem functionality if the
budget for large reserves is not available.
What needs to be studied?
Until now, the impact of disperser loss on entire seed dis-
persal communities has rarely been studied (Reid & Armesto
2011). For the majority of dispersers, a community-wide
assessment through experimental setups is hardly feasi-
ble. Thus, analyses of interaction networks are a promising
approach to understand long-term consequences at the com-
munity scale. Habitat loss, hunting of seed dispersers or
invasive species could lead to a lower diversity of interac-
tions, higher specialization and ultimately lower robustness
in plant–disperser networks. Thus, a comparison of networks
along experimental and real-world gradients of different
types of anthropogenic impact (large vs. small fragments,
invaded vs. non-invaded, hunted vs. non-hunted sites, etc.)
could reveal the sensitivity of mutualistic communities to
changes in disperser assemblages. Other approaches could
aim at investigating changes in specialization of interaction
networks during the fruiting season to reveal whether early
or late fruiting plants are more prone to disruption.
Further, the frequently large number of plant–disperser
interactions limits our understanding at a community scale.
One approach to deal with this complexity is to get away
 N. Farwig, D.G. Berens / Basic and Applied Ecology 13 (2012) 109–115
from focusing on the identity of species, but to consider func-
tional traits that determine the role species play in disperser
networks (Dennis & Westcott 2006). For example, the level
of disturbance tolerance of communities could be assessed
by analyzing the diversity of functional traits related to suc-
cessful seed dispersal (e.g. gape width) in relation to habitat
degradation. Studies addressing the plasticity of species and
traits could provide information on the adaptive potential
of dispersers in changing environments. Here, aspects of
changes in dietary specialization of dispersers or predictabil-
ity of interactions across temporal and spatial scales hold
great promise for future studies (Levey & Benkman 1999).
Understanding the evolution of traits that may extend a
species niche following environmental alteration would allow
making predictions for future changes (Holt & Gomulkiewicz
2004). Thereby, considering phylogeny in disperser commu-
nities may be helpful to determine the degree to which related
species are able to replace each other in their disperser func-
tion (Rezende, Lavabre, Guimarães, Jordano, & Bascompte
2007). Studying functional and phylogenetic diversity of dis-
perser communities along an altitudinal or latitudinal gradient
may be helpful to determine the degree to which related
species are able to substitute each other in their disperser
function in the face of climate change (Rezende et al. 2007).
Such approaches would help to set conservation priorities by
identifying keystone dispersers that play an overriding role
in interaction networks.
While few studies have investigated the effect of dis-
perser loss on mutualistic communities, even less studies
tackle cascading effects on other trophic levels (Fig. 1). The
loss of dispersers may not only directly influence dependent
plant species, but may indirectly also impact seed predators
or herbivores. As studying trophic cascade effects requires
intensive sampling, experimental setups could be a promis-
ing approach here (see Wotton & Kelly 2011 for an elegant
example). One could test if dispersal limitation equally affects
seed predation and seedling recruitment rates of differently
sized seed species, for example, to deduce the species’ sus-
ceptibility toward the loss of dispersers. Further, studying
feedback loops, where the loss of one seed disperser leads to
the joint loss of others, poses a challenge for future research.
Acknowledgments
We thank Teja Tscharntke for his kind invitation to write
this invited view and Jörg Albrecht, Alexandra Botzat, Eike
Neuschulz, Sascha Rösner and three anonymous reviewers
for helpful comments on the manuscript. The authors are
supported by the Robert Bosch Stiftung.
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