Ecological facilitation describes species interactions that benefit at least
one of the participants and cause harm to neither. Facilitations can be
categorized as mutualisms, in which both species benefit, or commensalisms,
in which one species benefits and the other is unaffected. Much of classic
ecological theory (e.g., natural
selection, niche separation, metapopulation
dynamics) has focused on negative interactions such as predation and competition, but positive interactions
(facilitation) are receiving increasing focus in ecological research.[1][2][3][4][5]
This article addresses both the mechanisms of facilitation and the increasing
information available concerning the impacts of facilitation on community
ecology.
Categories
There are two basic categories of facilitative interactions:
- A mutualism is an interaction between species that is beneficial to both. A familiar example of a mutualism is the relationship between flowering plants and their pollinators. The plant benefits from the spread of pollen between flowers, while the pollinator receives some form of nourishment, either from nectar or the pollen itself.
- A commensalism is an interaction in which one species benefits and the other species is unaffected. Epiphytes (plants growing on other plants, usually trees) have a commensal relationship with their host plant because the epiphyte benefits in some way (e.g., by escaping competition with terrestrial plants or by gaining greater access to sunlight) while the host plant is apparently unaffected.
Strict categorization, however, is not possible for some
complex species interactions. For example, seed germination
and survival in harsh environments is often higher under so-called nurse plants
than on open ground. A nurse plant is one with an established canopy, beneath
which germination and survival are more likely due to increased shade, soil
moisture, and nutrients. Thus, the relationship between seedlings and their
nurse plants is commensal. However, as the seedlings grow into established
plants, they are likely to compete with their former benefactors for resources.
Mechanisms
The beneficial effects of species on one another are
realized in various ways, including refuge from physical stress, predation, and
competition, improved resource availability, and transport.
Refuge from physical stress
Facilitation may act by reducing the negative impacts of a
stressful environment. As described above, nurse plants facilitate seed
germination and survival by alleviating stressful environmental conditions. A
similar interaction occurs between the red alga Chondrus
crispus and the canopy-forming seaweed Fucus in intertidal
sites of southern New England, USA. The alga survives higher in the
intertidal zone—where temperature and desiccation stresses are greater—only
when the seaweed is present because the canopy of the seaweed offers protection
from those stresses. The previous examples describe facilitation of individuals
or of single species, but there are also instances of a single facilitator
species mediating some community-wide stress, such as disturbance. An example of such
“whole-community” facilitation is substrate stabilization of cobble beach plant
communities in Rhode Island, USA, by smooth cordgrass (Spartina alterniflora). Large beds of
cordgrass buffer wave action, thus allowing the establishment and persistence
of a community of less disturbance-tolerant annual
and perennial plants below the high-water mark.
In general, facilitation is more likely to occur in
physically stressful environments than in favorable environments, where
competition may be the most important interaction among species. This can also
occur in a single habitat containing a gradient from low to high stress. For
example, along a New England, USA, salt marsh tidal gradient, a presence of black
needle rush (Juncus gerardi) increased the fitness of marsh elder
(Iva) shrubs in lower elevations, where soil salinity was
higher. The rush shaded the soil, which decreased evapotranspiration, and in turn decreased soil
salinity. However, at higher elevations where soil salinity was lower, marsh
elder fitness was decreased in the presence of the rush, due to increased
competition for resources. Thus, the nature of species interactions may shift
with environmental conditions.
Refuge from predation
Another mechanism of facilitation is a reduced risk of being
eaten. Nurse plants, for example, not only reduce abiotic stress, but may also physically impede herbivory of seedlings
growing under them. In both terrestrial and marine environments, herbivory of
palatable species is reduced when they occur with unpalatable species. These
“associational refuges” may occur when unpalatable species physically shield
the palatable species, or when herbivores are “confused” by the inhibitory cues
of the unpalatable species. Herbivory can also reduce predation of the
herbivore, as in the case of the red-ridged clinging crab
(Mithrax forceps) along the North
Carolina, USA, coastline. This crab species takes refuge in the branches of
the compact Ivory Bush Coral (Oculina arbuscula) and
feeds on seaweed in the vicinity of the coral. The reduced competition with
seaweed enhances coral growth, which in turn provides more refuge for the crab.
A similar case is that of the interaction between swollen-thorn acacia trees (Acacia spp.)
and certain ants (Pseudomyrmex spp.) in Central
America. The acacia provides nourishment and protection (inside hollow
thorns) to the ant in return for defense against herbivores. In contrast, a
different type of facilitation between ants and sap-feeding insects may
increase plant predation. By consuming sap, plant pests such as aphids produce a
sugar-rich waste product called honeydew, which is consumed by ants in
exchange for protection of the sap-feeders against predation.
Refuge from competition
Another potential benefit of facilitation is insulation from
competitive interactions. Like the now familiar example of nurse plants in
harsh environments, nurse logs in a forest are sites of increased seed
germination and seedling survival because the raised substrate of a log frees
seedlings from competition with plants and mosses on the forest floor. The
crab-coral interaction described above is also an example of refuge from
competition, since the herbivory of crabs on seaweed reduces competition
between coral and seaweed. Similarly, herbivory by sea urchins (Strongylocentrotus droebachiensis)
on kelps (Laminaria
spp.) can protect mussels
(Modiolus modiolus) from overgrowth by kelps competing for space in the subtidal
zone of the Gulf of Maine, USA.
Improved resource availability
Facilitation can increase access to limiting resources such
as light, water, and nutrients for interacting species. For example, epiphytic
plants often receive more direct sunlight in the canopies of their host plants
than they would on the ground. Also, nurse plants increase the amount of water
available to seedlings in dry habitats because of reduced evapotranspiration
beneath the shade of nurse plant canopies. However, the most familiar examples
of increased access to resources through facilitation are the mutualistic
transfers of nutrients between symbiotic organisms. A symbiosis is
a prolonged, close association between organisms, and some examples of
mutualistic symbioses include:
Associations between a host species and a microbe
living in the host’s digestive tract, wherein the host provides habitat
and nourishment to the microbe in exchange for digestive services. For example,
termites
receive nourishment from cellulose digested by microbes inhabiting their gut.
Associations between fungi and algae, wherein the
fungus receives nutrients from the alga, and the alga is protected from harsh
conditions causing desiccation.
Associations between reef-building corals and photosynthetic
algae called zooxanthellae, wherein the zooxanthellae provide
nutrition to the corals in the form of photosynthate, in exchange for nitrogen
in coral waste products.
Associations between fungi and plant roots, wherein the
fungus facilitates nutrient uptake (particularly nitrogen) by the plant in
exchange for carbon in the form of sugars from the plant root. There is a
parallel example in marine environments of sponges on the roots of mangroves,
with a relationship analogous to that of mycorrhizae and terrestrial plants.
Transport
The movement by animals of items involved in plant
reproduction is usually a mutualistic association. Pollinators may increase
plant reproductive success by reducing pollen waste, increasing dispersal of pollen, and increasing the
probability of sexual reproduction at low population density. In return, the pollinator
receives nourishment in the form of nectar or pollen. Animals may also disperse
the seed or fruit of plants, either by eating it (in which case they receive
the benefit of nourishment) or by passive transport, such as seeds sticking to
fur or feathers.
Community Effects
Although facilitation is often studied at the level of
individual species interactions, the effects of facilitation are often
observable at the scale of the community, including impacts to spatial
structure, diversity, and invasibility.
Spatial structure
Many facilitative interactions directly affect the
distribution of species. As discussed above, transport of plant propagules by
animal dispersers can increase colonization rates of more distant sites, which
may impact the distribution and population dynamics of the plant species.
Facilitation most often affects distribution by simply making it possible for a
species to occur in a site where some environmental stress would otherwise
prohibit growth of that species. This is apparent in whole-community
facilitation by a foundation species, such as sediment stabilization in cobble
beach plant communities by smooth cordgrass. A facilitating species may also
help drive the progression from one ecosystem type to another, as mesquite
apparently does in the grasslands of the Rio
Grande Plains. As a nitrogen-fixing tree, mesquite establishes more readily
than other species on nutrient-poor soils, and following establishment,
mesquite acts as a nurse plant for seedlings of other species. Thus, mesquite
facilitates the dynamic spatial shift from grassland to savanna to woodland across
the habitat.
Diversity
Facilitation affects community diversity (defined in this
context as the number of species in the community) by altering competitive
interactions. For example, intertidal mussels increase total community species
diversity by displacing competitive large sessile species such as seaweed and barnacles.
Although the mussels decrease diversity of primary space holders (i.e., large
sessile species), a larger number of invertebrate
species are associated with mussel beds than with other primary space holders,
so total species diversity is higher when mussels are present.The effect of
facilitation on diversity could also be reversed, if the facilitation creates a
competitive dominance that excludes more species than it permits.
Invasibility
Facilitation of non-native species, either by native species
or other non-native species, may increase the invasibility of a community, or
the ease with which non-native species become established in a community. In an
examination of 254 published studies of introduced species, 22 of 190 interactions
studied between introduced species in the studies were facilitative. It is
worth noting that 128 of the 190 examined interactions were predator-prey relationships of a
single plant-eating insect reported in a single study, which may have
overemphasized the importance of negative interactions. Introduced plants are
also facilitated by native pollinators, dispersers, and mycorrhizae.
Thus, positive interactions must be considered in any attempt to understand the
invasibility of a community.
Conclusion
Facilitation is a significant ecological process that
produces community-level effects through individual interactions. By improving
dispersal, increasing access to resources, and providing protection from
stress, predation, and competition, facilitation can impact community
structure, diversity, and invasibility. Incorporation of facilitation into such
classic theories as natural selection and niche separation should be a goal of
current and future ecologists. This will require further research into the
mechanisms of facilitation at the level of individuals, and the impacts of
facilitation at the level of population, community, and ecosystem. Continued
study of positive species interactions will serve to improve our understanding
of processes and application of theories.
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