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Keystone Species Concept and General Type of Keystone Species1

By Tedi Setiadi2

The ecological term keystone species has been recognized since four decades ago.
Paine (1969) known as the first author describe the term following his study of the rocky
intertidal zone of Pacific Ocean. This concept triggered debate amongst ecologist especially
concerning the operational definition and the usage of the concept in ecology (Davic 2003).
Recent debate on the definition of keystone species involving several authors (De Leo and
Levin 1997, Khanina 1998, Piraino and Fanelli 1999) appears in Conservation Ecology
journal. In this communication, I describe the definition of keystone species and discuss the
importance of keystone species in maintaining communities and their ecological processes.

Definition of Keystone Species

The keystone species phenomenon was coined for the very first time by Paine (1966)
based on his experimental in intertidal ecosystem. However, by that time he did not use the
term keystone species. The term of keystone species was introduced three years later (Paine
1969). Keystone species is defined as a species of high trophic status whose activities exert a
disproportionate influence on the pattern of species diversity in a community (Paine 1969).
The term was originally applied to keystone predator defined as a species that feeds
preferentially on the dominant competitor among its prey species, such that the keystone
predators feeding prevents the dominant prey from excluding other species, and therefore


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maintains a higher species diversity in the system than in the keystones absence (Paine
1969). Paine (1969) encompassed specific ecosystem process which is top-down regulation
of species diversity, competitive interactions, and community persistence.
The Paines keystone species concepts (1969) become very popular in ecological
literature, the literature was cited more than 92 publications from 1970 to 1989 (Mills et al.
1993). The concept has been studied, modified, even though critiqued. Morover many
publications on keystone species have been neglected the principal idea of Paines work
which is the keystone species concept focused on food-web on the ecosystem (Davic 2003).
Thus, many species have been afforded Keystone Species status because they are so-called
keystone prey, competitors, mutualist, dispersers, pollinators, earth-movers, habitat
modifiers, engineers, hosts, processors, plant resources, dominant trees, etc. (Davic 2003).
Based on the application of the concept, there are two hallmarks of keystone: first, their
presence is crucial in maintaining the organization and diversity of their ecological
communities; and second, it is implicit that these species are exceptional, relative to the rest
of the community, in their importance (Mills et al. 1993).
Power et al (1996) summarized three important point on the keystone species concept,
which are: keystone species have been demonstrated occur in major ecosystem throughout
the world; keystone species are not always as top trophic level; keystone species can exert
effect toward several mechanisms i.e. consumption, mutualism, competition, dispersal,
pollination, disease, and by modifying habitat and its abiotic factors.
One of major debate to the keystone concept appeared in 1993, when Mills et al.
argue the ambiguous nature of its definition. Mills et al. (1993) argue that the term is broadly
applied, poorly defined and nonspecific meaning. This made it hard to identify exactly which
species should designated as having a keystone role in a community (Mills et al. 1993).

In order to response the critique to the keystone concept, new operational definition
was proposed by a group of ecologist known as Keystone Cops (Paine 1995) for the very
first time. The operational definition is based on the proportional biomass of species in
ecosystems in relation to their community importance, specifically the definition of keystone
species is a species whose effect on ecosystems in disproportionately large relative to its low
biomass in the community as a whole (Power et al. 1996). This definition distinguishes
keystone species from dominant species-whose effect to the ecosystem is large because the
species is very abundant. However, this operational definition has limited operational value
for conservation ecology because several thing, such as the difficulties to distinguishes
keystone species with dominant species, the difficulties to measure the biomass (Kotliar
2000, Davic 2003).
Davic (2003) developed another operational definition of keystone species which
considered the interaction between species and its effect on species diversity and competition
is large relative to its biomass dominance within a functional group.

This operational

definition links the community importance of keystone species to a specific ecosystem

process, e.g., the regulation of species diversity, within functional group at lower trophic
levels that are structured by competition for limited resource.

Type of Keystone Species and Its Role in Ecosystem

Mills et al. (1993) summarized several case studies related to keystone concept based
on utilisation of the concept into five types of keystone, which are: Keystone Predator;
Keystone Prey; Keystone Mutualist; Keystone Hosts; and Keystone Modifier. The
categorization is presented in Table 1.

Keystone Predator
The best example of this type of keystone species would the work of Paine (1969) on
the phenomenon of keystone species in the rocky intertidal system. The publication of his
work was the origin of keystone species concept. Paine (1969) found out the significant
changes in population density, species composition, and overt appearance was produced by
the removal of dominant carnivore, the starfish Pisaster ochraceus. The starfish play role as
controller of mussels Mytilus californianus population to a size that allowed the rest of the
species in the ecosystem still exist. Once the starfish were removed from the system, a
reduction in biodiversity of eight to 15 species was observed (Paine 1966).
Another example would be sea otter Enhydra lutris in rocky shore ecosystems (Estes
and Duggins 1995, Estes and Palmisano 1974). Sea otters have very strong cascading impacts
on the structure and functioning of rocky shore ecosystems despite their relatively low
abundance. Their disproportionate impacts stem from their voracious appetite for sea
urchins, herbivorous invertebrates whose feeding on kelps in turn determine whether rocky
bottoms will become dominated by large kelps (in the absence of urchins) versus by crustose
calcified algae (in the presence of urchins). The kelp beds facilitated by sea otter predation on
urchins offer both abundant food and physical structure in which fishes and other animals can
grow and escape their predators, whereas the crustose algal pavements that develop in the
absence of sea otters leave the bottom with no structure and little plant biomass to support
other animals.

Keystone Prey
The concept of keystone prey has been demonstrated by the research of Holt (1977 in
Mills et al 1993). Basically, certain prey in ecosystem which is able to maintain their

population in the face of predation can affect community structure by sustaining the density
of a predator, thus reducing the density of other prey. Holt suggested removal of keystone
species would increase species diversity in the community. However, another publication by
Noy-Meir (1981) suggested differently to Holts which is the removal of keystone prey
would decrease species diversity. The occurrence of keystone prey will allow the more
sensitive prey to coexist, since the predator switches to the keystone prey when numbers of
other species are low, then sensitive prey that otherwise would have been driven to extinction
may coexist in the presence of the predator-tolerant keystone prey (Noy-Meir, 1981).
Keystone Mutualist
Mills et al (1993) stated some species have been considered to be keystone because
they are critical to mutualistic relationship. This statement was derived from the publication
of Gilbert (1980) which introduced the term mobile links to describe animal that are
significant factors in the persistence of several plant species which, in turn, support otherwise
separate food webs. Gilbert (1980) described mobile link pollinators and seed dispersers as
keystone species.

Keystone Host
In this category, the plant species play role as keystone species. Mills et al (1993)
considered two way interaction between keystone mutualist (mobile link) and plant as the
host. Included in this group are those plants that support generalist pollinators and those fruit
dispersers that are considered critical mobile links (Gilbert 1980 in Mills et al. 1993).
Another author gives specific example of keystone plants such as palm nuts, figs, nectar
(Terborgh 1986 in Mills et al. 1993), and Didymopanax (Worthington 1982 in Power et al.

1996) to be keystone resources because they are critical to tropical forest nectar or fruit

Keystone Modifier
Mills et al. (1993) underlined that certain species has the ability to modified the
habitat feature and affects the survival of many other species. The species with such ability is
considered as keystone species. One of famous example in this category is the Canadan
beaver Castor Canadensis (Garibaldi and Turner 2004). The beavers damming of streams
creates ponds, which in turn serve as habitat for numerous other species from ducks to
aquatic plants and insects (Naiman et al. 1986, Pollock et al. 1995 in Garibaldi and Turner
2004) and the dams alter hydrology, biogeochemistry, and productivity on a wide scale
(Naiman et al. 1986).
Table 1. Categories of presumed keystones and the effects of their effective removal from a
system (after Mills et al. 1993)

Effect of removal


Increase in one or several predators/ consumers/ competitors, which

subsequently extirpates several prey/ competitor species


Other species more sensitive to predation may become extinct; predator

population may crash


Extirpation of dependent animals, potentially including pollinators and

seed dipersers


Failure of reproduction and recruitment in certain plants, with potential

subsequent losses


Loss of structures/ materials that affect habitat type and energy flow;
disappearance of species dependent on particular successional habitats
and resources

Many publications have been demonstrated discussion of keystone species concept
since its first introduction in late of 60s by T.R. Paine. Ongoing debates amongst ecologist
whose concern to this topic show how popular, perhaps important, this concept is. Paine
(1969) defined keystone species based on food web system in community specifically on top
predator. Afterward, the definition now expanded to more broadened sense including all level
trophic in the ecosystem. To clarify its concept in order to implement it for conservation
purpose, Power et al (1996) developed the operational definition of keystone species concept.
Based on the literature review by Mills et al. (1993) by interpreting the keystone
species term referred to, there are five categories of keystone species. These are: keystone
predator, keystone prey, keystone mutualist, keystone host, and keystone modifier. These
types of keystone species have been demonstrated very well on literatures.

References Cited
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