Population ecology
The development of population ecology owes much to demography and actuarial life tables. Population ecology is important in conservation biology, especially in the development of population viability analysis which makes it possible to predict the long-term probability of a species persisting in a given habitat patch. Although population ecology is a subfield of biology, it provides interesting problems for mathematicians and statisticians who work in population dynamics.
Fundamentals
| Term | Definition |
| Species population | All individuals of a species. |
| Metapopulation | A set of spatially disjunct populations, among which there is some immigration. |
| Population | A group of conspecific individuals that is demographically, genetically, or spatially disjunct from other groups of individuals. |
| Aggregation | A spatially clustered group of individuals. |
| Deme | A group of individuals more genetically similar to each other than to other individuals, usually with some degree of spatial isolation as well. |
| Local population | A group of individuals within an investigator-delimited area smaller than the geographic range of the species and often within a population. A local population could be a disjunct population as well. |
| Subpopulation | An arbitrary spatially delimited subset of individuals from within a population. |
The most fundamental law of population ecology is Thomas Malthus' exponential law of population growth.
A population will grow exponentially as long as the environment experienced by all individuals in the population remains constant.
This principle in population ecology provides the basis for formulating predictive theories and tests that follow:
Simplified population models usually start with four key variables including death, birth, immigration, and emigration. Mathematical models used to calculate changes in population demographics and evolution hold the assumption of no external influence. Models can be more mathematically complex where "...several competing hypotheses are simultaneously confronted with the data." For example, in a closed system where immigration and emigration does not take place, the rate of change in the number of individuals in a population can be described as:
where N is the total number of individuals in the population, B is the raw number of births, D is the raw number of deaths, b and d are the per capita rates of birth and death respectively, and r is the per capita average number of surviving offspring each individual has. This formula can be read as the rate of change in the population is equal to births minus deaths.
Using these techniques, Malthus' population principle of growth was later transformed into a mathematical model known as the :
where N is the biomass density, a is the maximum per-capita rate of change, and K is the carrying capacity of the population. The formula can be read as follows: the rate of change in the population is equal to growth that is limited by carrying capacity . From these basic mathematical principles the discipline of population ecology expands into a field of investigation that queries the demographics of real populations and tests these results against the statistical models. The field of population ecology often uses data on life history and matrix algebra to develop projection matrices on fecundity and survivorship. This information is used for managing wildlife stocks and setting harvest quotas
Geometric populations
The population model below can be manipulated to mathematically infer certain properties of geometric populations. A population with a size that increases geometrically is a population where generations of reproduction do not overlap. In each generation there is an effective population size denoted as Ne which constitutes the number of individuals in the population that are able to reproduce and will reproduce in any reproductive generation in concern. In the population model below it is assumed that N is the effective population size.Assumption 01: Ne = N
Nt+1 = Nt + Bt + It - Dt - Et
| Term | Definition |
| Nt+1 | Population size in the generation after generation t. This may be the current generation or the next generation depending on the situation in which the population model is used. |
| Nt | Population size in generation t. |
| Bt | Sum of births in the population between generations t and t+1. Also known as raw birth rate. |
| It | Sum of immigrants moving into the population between generations t and t+1. Also known as raw immigration rate. |
| Dt | Sum of deaths in the population between generations t and t+1. Also known as raw death rate. |
| Et | Sum of emigrants moving out of the population between generations t and t+1. Also known as raw emigration rate. |
from reproduction. Exponential populations grow without designated periods for reproduction. Reproduction is a continuous process and generations of reproduction overlap. This graph illustrates two hypothetical populations - one population growing periodically and the other population growing continuously. The populations in the graph have a doubling time of 1 year. The populations in the graph are hypothetical. In reality, the doubling times differ between populations.
Assumption 02: There is no migration to or from the population
It = Et = 0
Nt+1 = Nt + Bt - Dt
The raw birth and death rates are related to the per capita birth and death rates:
Bt = bt × Nt
Dt = dt × Nt
bt = Bt / Nt
dt = Dt / Nt
| Term | Definition |
| bt | Per capita birth rate. |
| dt | Per capita death rate. |
Therefore:
Nt+1 = Nt + -
Assumption 03: bt and dt are constant.
Nt+1 = Nt + -
| Term | Definition |
| b | Constant per capita birth rate. |
| d | Constant per capita death rate. |
Take the term Nt out of the brackets.
Nt+1 = Nt + Nt
b - d = R
Nt+1 = Nt + RNt
Nt+1 =
Take the term Nt out of the brackets again.
Nt+1 = Nt
1 + R = λ
| Term | Definition |
| λ | Finite rate of increase. |
Nt+1 = λNt
| At t+1 | Nt+1 = λNt |
| At t+2 | Nt+2 = λNt+1 = λλNt = λ2Nt |
| At t+3 | Nt+3 = λNt+2 = λλNt+1 = λλλNt = λ3Nt |
| At t+4 | Nt+4 = λNt+3 = λλNt+2 = λλλNt+1 = λλλλNt = λ4Nt |
| At t+5 | Nt+5 = λNt+4 = λλNt+3 = λλλNt+2 = λλλλNt+1 = λλλλλNt = λ5Nt |
Therefore:
Nt+1 = λtNt
| Term | Definition |
| λt | Finite rate of increase raised to the power of the number of generations (e.g. for t+2 → λ2, for t+1 → λ1 = λ, and for t → λ0 = 1 |
Doubling time of geometric populations
The doubling time of a population is the time required for the population to grow to twice its size. We can calculate the doubling time of a geometric population using the equation: Nt+1 = λtNt by exploiting our knowledge of the fact that the population is twice its size after the doubling time.2Ntd = λtd × Nt
λtd = 2Ntd / Nt
λtd = 2
The doubling time can be found by taking logarithms. For instance:
td × log2 = log2
log2 = 1
td × log2 = 1
td = 1 / log2
Or:
td × ln = ln
td = ln / ln
td = 0.693... / ln
Therefore:
td = 1 / log2 = 0.693... / ln
Half-life of geometric populations
The half-life of a population is the time taken for the population to decline to half its size. We can calculate the half-life of a geometric population using the equation: Nt+1 = λtNt by exploiting our knowledge of the fact that the population is half its size after a half-life.0.5Nt1/2 = λt1/2 × Nt
λt1/2 = 0.5Nt1/2 / Nt
λt1/2 = 0.5
The half-life can be calculated by taking logarithms.
t1/2 = 1 / log0.5 = ln / ln
Geometric (R) and finite (λ) growth constants
Geometric (R) growth constant
R = b - dNt+1 = Nt + RNt
Nt+1 - Nt = RNt
Nt+1 - Nt = ΔN
| Term | Definition |
| ΔN | Change in population size between two generations. |
ΔN = RNt
ΔN/Nt = R
Finite (λ) growth constant
1 + R = λNt+1 = λNt
λ = Nt+1 / Nt
Mathematical relationship between geometric and exponential populations
In geometric populations, R and λ represent growth constants. In exponential populations however, the intrinsic growth rate, also known as intrinsic rate of increase is the relevant growth constant. Since generations of reproduction in a geometric population do not overlap but do in an exponential population, geometric and exponential populations are usually considered to be mutually exclusive. However, geometric constants and exponential constants share the mathematical relationship below.The growth equation for exponential populations is
Nt = N0ert
| Term | Definition |
| e | Euler's number - A universal constant often applicable in exponential equations. |
| r | intrinsic growth rate - also known as intrinsic rate of increase. |
was the mathematician who established the universal constant 2.71828... also known as Euler's number or e.
Assumption: Nt ' = Nt '.
Therefore:
N0ert = N0λt
N0 cancels on both sides.
N0ert / N0 = λt
ert = λt
Take the natural logarithms of the equation. Using natural logarithms instead of base 10 or base 2 logarithms simplifies the final equation as ln = 1.
rt × ln = t × ln
| Term | Definition |
| ln | natural logarithm - in other words ln = loge = x = the power that e needs to be raised to to give the answer y. In this case, e1 = e therefore ln = 1. |
rt × 1 = t × ln
rt = t × ln
t cancels on both sides.
rt / t = ln
The results:
r = ln
and
er = λ
r/K selection
An important concept in population ecology is the r/K selection theory. The first variable is r and the second variable is K.An r-selected species is one that has high rates of fecundity, low levels of parental investment in the young, and high rates of mortality before individuals reach maturity. Evolution favors productivity in r-selected species. In contrast, a K-selected species has low rates of fecundity, high levels of parental investment in the young, and low rates of mortality as individuals mature. Evolution in K-selected species favors efficiency in the conversion of more resources into fewer offspring.
Metapopulation
Populations are also studied and conceptualized through the "metapopulation" concept. The metapopulation concept was introduced in 1969:"as a population of populations which go extinct locally and recolonize."Metapopulation ecology is a simplified model of the landscape into patches of varying levels of quality. Patches are either occupied or they are not. Migrants moving among the patches are structured into metapopulations either as sources or sinks. Source patches are productive sites that generate a seasonal supply of migrants to other patch locations. Sink patches are unproductive sites that only receive migrants. In metapopulation terminology there are emigrants and immigrants. Metapopulation models examine patch dynamics over time to answer questions about spatial and demographic ecology. An important concept in metapopulation ecology is the rescue effect, where small patches of lower quality are maintained by a seasonal influx of new immigrants. Metapopulation structure evolves from year to year, where some patches are sinks, such as dry years, and become sources when conditions are more favorable. Ecologists utilize a mixture of computer models and field studies to explain metapopulation structure.
History
The older term, autecology, refers to roughly the same field of study as population ecology. It derives from the division of ecology into autecology—the study of individual species in relation to the environment—and synecology—the study of groups of organisms in relation to the environment—or community ecology. Odum considered that synecology should be divided into population ecology, community ecology, and ecosystem ecology, defining autecology as essentially "species ecology." However, for some time biologists have recognized that the more significant level of organization of a species is a population, because at this level the species gene pool is most coherent. In fact, Odum regarded "autecology" as no longer a "present tendency" in ecology, although included "species ecology"—studies emphasizing life history and behavior as adaptations to the environment of individual organisms or species—as one of four subdivisions of ecology.Journals
The first journal publication of the Society of Population Ecology, titled Population Ecology was released in 1952.Scientific articles on population ecology can also be found in the Journal of Animal Ecology, Oikos and other journals.