Habitat of Ants
Are Your Neighbors Good Enough?
Using Minnesotan Ants in a Biodiversity Model -- A spatial explicit approach to model ant biodiversity under local competition --
Background
Conservation Biology is an applied science. One goal of this integrated discipline
is to clarify the issue and seek the solution to preserve biodiversity, while most
natural communities contain a significant amount of species diversity which are endangered
by modernization and globalization of human development. The biodiversity of ecological
community provides an important understanding with a number of implications for ecosystem
functioning and stability. A great number of work has been done in both experimental
methods (field observation, field manipulation and laboratory experiments)
and theoretical methods, such as the analysis of mathematical
and computer-based models.
Since species diversity is accepted as one of the most important descriptive parameters
in an ecological community, the central question of Conservation Biology is raised:
Why are there so many kinds of organisms? We can view
the number of species in co-existence in communities as a complex function of local
spatial interaction (mainly competition of limited resource, with symbiosis, parasitism
and other interactions) under evolutionary selection pressure and adaptive radiations
with biogeographical barriers. MacArthur and Wilson (1967) examined how the rates
of colonization and extinction might influence the species number within islands.
With greatly over-simplification, larger area may contain more available niche to
support species diversity. The other factor to affect species abundance distribution
within community is succession time under ecological constraint, if we assume that
tropical ecosystems are older than the other ecosystems, and arctic environments
are more physically shaped than tropical climax communities. Productivity hypothesis
also suggested diversity may be a function of community productivity because primary
productivity provides the basis of resource (Brown 1973). Finally, the hypothesis
of spatial complexity claimed that diversity may be explained in terms of environmental
mosaics or spatial subdivision (Tilman 1994): the more complex and heterogeneous
an environment is, the more niche it may offer to support diverse fauna and flora.
However, the content of biodiversity can be enriched by the role of biotic
interactions (Sander 1968). It suggested the increase of biotic interactions
would promote diversity. For example, predation can maintain diversity within community,
even though predator can eliminate prey species. Interspecific competition also promotes
biodiversity by increasing specialization and decreasing niche width, which leads
to overlapping of co-existence or allows more species share or partition resource
before exclusion by severe competition. In addition, intraspecific competition can
also serve as a principal mechanism of evolutionary selection. It is generally essential
to consider both interspecific and intraspecific competitions shaping biodiversity
in community. From the non-spatial mathematical model of Lotka-Volterra model (Volterra
1926, Lotka 1932), coexistence to enhance biodiversity is only
possible if intraspecific competition is more important than interspecific competition.
This may be questionable under the spatial mechanism of co-existence, because local
selection pressure of intraspecific competition can react with interspecific neighborhood
in a spatial explicit sense. Local population maintains active range and determines
the adjacent impact toward the other adjacent species. To maintain biodiversity,
space and local distributed pattern are also important.
Objectives of Modeling Biodiversity
To examine the spatial roles of different competition, I choose ant fauna to study
biodiversity because these social insects have rich behavioral interactions in both
colony (intraspecific) and species (intraspecific) levels. Currently, the spatial
explanation of species diversity distribution within community is still needed to
expand. From classical ecological theory, the stable co-existing species are limited
by resource and environmental uncertainty. This poses a dilemma:
How does ant community maintain biodiversity spatially in a dynamic of interspecific
and intraspecific competitions? What is the implication to Conservation Biology
from this spatially interactive mechanism of ant biodiversity? Through the spatial
explicit simulation and modeling, I propose to study what is the trade-off effect
of ant biodiversity (coexistence) between interspecific and intraspecific competitions
under different levels of environmental resource.
My First Model Description (co-existing
competition model of ants)
This spatial explicit model is built with a grid system and each cell acts as a spatial
unit to observe ant species coexistence. Within site, there is a frame-based component
to allow multiple ant species to occupy a single patch with interspecific interaction.
Between sites, different ant species compete with adjacent neighborhood (interspecies),
or interact with different colonies of the same species (intercolonies or intraspecies).
The competition among different species and colonies can be gradient between exclusive
and non-exclusive levels, which depends on specific ant density and local resource
abundance.
Assumptions
1. Simple competitors between sites
From my pilot observation and data in first year (2000), when available food appears,
ants can be roughly classified as early arriver, late overwhelmer,
occasional patroller and cryptic species. I assume early ant arrivers come
from local patch because the spatial convenience leads to their rapid response. Within
one bait station, it may or may not have the other late overwhelmer species from
neighborhood to compete food. Thus the same ant species can be early arriver or late
overwhelmer, which based on the spatial location and distribution pattern of their
nest sites.
In my first stage of modeling, I only consider the first two competitors in grid
system. To simplify the ant interactions in the beginning of model building, I assume
occasional patroller and cryptic species are NOT significant important in competition.
Generally, it is true for some cryptic species, such as thief ant (Solenosis molesta
) can only steals little pieces of food without causing a strong competition.
I wish to point out here it may not be true for some large and aggressive ants. For
example, Camponotus noveboracensis is a beautiful red-black large carpenter
ant which travels for a long distance to patrol its active range. Even small number
of individuals can strongly attack the local abundant ants and cause severe mortality
of small ants like Tapinoma sessile and Lasius aliens. In addition,
it is needed to release this assumption in the future in order to consider all ant
species in the further biodiversity modeling.
2. Resolution is suitable to model the continuous space: One cell can carry multiple
species, but only one colony per species is allowed per cell.
3. Eliminate the complex polygeny (many queens in a colony) and polydomy (many nests
or entrances in a colony) situation: All different sites in grid system occupied
by the same ant species are treated for different colonies. This colony unit operates
intraspecific competition separately.
Output of Model Simulation
1. Species richness: plus other biodiversity index with different weight of relative
abundance (e.g. Shannon and Weaver index, Simpson index...etc.)
2. Species coexistence pattern: Where is the biodiversity hot spot to preserve? Why?
3. Species-abundance curve: Stable ecosystem or many endangered species?
Conclusion
It is a start to look at how the complex biodiversity system works in community scale
from an emphasis of competitive interaction. The knowledge and insight from this
model will help conservation biologists focus on the a number of key variables to
explain the phenomena of ant coexistence which is regulated by a series of unique
spatially interactive mechanisms. The model prediction will also allow us to apply
behavioral indicators to enrich diversity index, and develop
better estimates of " biodiversity " by concerning more biotic interaction
in a spatial ecological network.