Introduction to Population Ecology Olalere Shittu Dept. of Zoology University of Ilorin, Nigeria Introduction Population ecology is the study of populations (especially population abundance) and how they change over time. Crucial to this study are the various interactions between a population and its resources. A population can decline because it lacks resources or it can decline because it is prey to another species that is increasing in numbers. Populations are limited by their resources in its capacity to grow; the maximum population abundance (for a given species) an environment can sustain is called the carrying capacity. Introduction As a population approaches its carrying capacity, overcrowding signifies that there are less resources for the individuals in the population and this results in a reduction in birth rate. A population with these features is said to be density dependent. Most populations are density dependent to some extent, but some grow (almost) exponentially and these are, in effect, density independent. Ecological models that focus on a single species and the relevant carrying capacity are single species models. Alternatively, multispecies or community models focus on the interactions of specific species. Characteristics of Population Population characteristics are based on the individual characteristics of the individuals within the population. Population is described in terms of size, age, and structure. Along with the population’s physical attributes, factors which affect the population includes mortality from competition and predation. Population dynamics is the study of changes in populations through time. Demography is the study of the characteristics of populations. It provides a mathematical description of how those characteristics change over time. Demographics can include any statistical factors that influence population growth or decline, but several parameters are particularly important: population size, density, age structure, Population Size oThis is defined as the number of individuals present in a subjectively designated geographic range. oThe most fundamental demographic parameter is the number of individuals within a population. Despite the simplicity in its concept, locating all individuals during a census (a full count of every individual) is nearly impossible, so ecologists usually estimate population size by counting individuals within a small sample area and extrapolating that sample to the larger population. Regardless of the challenges in measuring population size, it is an important characteristic of a population with significant implications for the dynamics of the population as a whole. Population Size Populations display distinctive behaviors based on their size, viz: Small populations face a greater risk of extinction. Individuals in these populations can have a hard time finding quality mates so, fewer individuals mate and those that do risk inbreeding. Additionally, individuals in small population are more susceptible to random deaths. Events like fire, floods, and disease have a greater chance of killing all individuals in the population. Large populations experience their own problems. As they approach the maximum sustainable population size (carrying capacity), large populations show characteristic behavior. Population Size Populations nearing their carrying capacity experience the following difficulties: 1) They face greater competition for resources, 2) Shifts in predator-prey relationships, and 3) Lowered fecundity. If the population grows too large, it may begin to exceed the carrying capacity of the environment and degrade available habitat (Figure 1). Population Size Fig. 1: Graph showing population growth over time Exponential growth.: This means that the population grows very quickly over a short amount of time, and on a graph looks like the letter 'J'. Logistic growth: Due to limited resources, the environment cannot continue to support EG, therefore the growth of the population begins to level out and takes the shape of an 'S'. Carrying capacity: When population growth is restricted and the size of the population becomes stable, denoted by K . Population Size Fluctuations in population growth In theory, populations grow quickly, reach K, and level out. In reality, populations tend to fluctuate around K, and there are several ways this might occur in nature. Carrying capacity fluctuations can be chaotic. These are erratic fluctuations around K, often due to environmental factors that have an immediate impact on population size, e.g. disease and natural disasters. Fluctuations around K can also be cyclical (stable limit cycle). This type of oscillation around K is different from chaotic because it is regular and produces a normal pattern for the population. Population Size Damped oscillations are fluctuations of population size above and below K that lessen with time. The population will eventually reach a stable limit, and the fluctuations will become minimal. Logistic Growth Model The growth curve of a population, which is always limited by one or more factors in the environment, is expressed by the logistic growth equation: The growth rate of the population (dN/dt) equals its rate of increase (r X N, the No. of individuals present at any one time), adjusted for the amount of resources available. The adjustment is made by multiplying rN by the fraction of K still unused (K minus N, divided by K). As N increases (the population grows in size), the fraction by which r is multiplied (the remaining resources) becomes smaller and smaller, and the rate of increase of the population declines. Logistic Growth Model In mathematical terms, as N approaches K, the rate of population growth (dN/dt) begins to slow, reaching 0 when N = K (blue line in the figure). a b Density dependence in song sparrow: Reproductive success decreases (a) and mortality rates increases (b) as population size increases. Population Size Population Density (Complete description of size) The size of a population in relation to the amount of space that it occupies. Density is usually expressed as the number of individuals per unit area or volume. For example: the number of crows per square kilometre or the number of plankton per litre. Like all population properties, density is a dynamic characteristic that changes over time as individuals are added to or removed from the population. Closely related species of Gannet birds will maintain very different densities. Birth and immigration: The influx of new individuals from other areas can increase a population's density, while death and emigration; the movement of individuals out of a population to other areas can decrease its density Population Density Similar to population size, population density displays distinctive characteristics at both high and low values. Density-dependent factors, including competition, predation, migration and disease, intensify within populations as density increases. In contrast, densityindependent factors, such as weather, fire regimes, and flooding, impact populations regardless of their specific densities Population Density Similar to population size, population density displays distinctive characteristics at both high and low values. Density-dependent factors, including competition, predation, migration and disease, intensify within populations as density increases. In contrast, densityindependent factors, such as weather, fire regimes, and flooding, impact populations regardless of their specific densities. The unit of population density differ in different populations. It is generally expressed as the number of individuals, or the population biomass (weight basis) per unit area or volume. Larger organisms like trees may be expressed as 600 trees per hectare, whereas smaller ones like phytoplanktons as 3million cells per cubic metre of water. In terms of weight, it may be 100 pounds of fish per hectare of water surface. Population Density Patterns of dispersion of organisms in nature differs, hence it becomes important to distinguish between crude density and specific (ecological) density. 1) Crude density is the density (number or biomass) per unit total space 2) Specific or ecological or economic density is the density (number or biomass) per unit of habitat space i.e. available area or volume that can actually be colonized by the population. Broadly, population density is the total number of species within some natural habitat. Mathematically:D = n/a / t . Where D = density, n = number of individuals, a = area, t = the time unit. Population Density Fecundity: This is the potential reproductive capacity of a an organism or population measured by the number of gametes or asexual propagules As age structure suggests, some individuals within a population have a greater impact on population-level processes, such as growth. Fecundity describes the number of offspring an individual or a population is able to produce during a given period of time. In demographic studies, fecundity is calculated in age-specific birth rates, which may be expressed as the number of births per unit of time, the number of births per female per unit of time, or the number of births per 1,000 individuals per unit of time. Maximum (or physiological) fecundity is the theoretical maximum number of offspring produced in a population assuming no ecological constraints. Fecundity However, since every ecosystem implements constraints on its populations, ecologists prefer to measure realized (or ecological) fecundity, which is the observed number of offspring produced in a population under actual environmental conditions. While maximum fecundity is a constant for populations, realized fecundity varies over time based on the size, density, and age structure of the population. External conditions, such as food and habitat availability, can also influence fecundity. Density-dependent regulation provides a negative feedback if the population grows too large, by reducing birth rates and halting population growth through a host of mechanisms Fecundity In white-footed mice, for example, populations regulate their reproductive rate via a stress hormone. As population densities increase, so do aggressive interactions between individuals (even when food and shelter are unlimited). High population densities lead to frequent aggressive encounters, triggering a stress syndrome in which hormonal changes delay sexual maturation, cause reproductive organs to shrink, and depress the immune system Natality This broadly covers the production of new individuals by any organism. These new individuals are born, hatched, germinated, arise by division etc. In human population, natality rate means birth rate. Two types of natality rate, viz: 1. Maximum (absolute or potential or physiological) natality is the theoretical maximum production of new individuals under ideal conditions (i.e. no ecological limiting factors, reproduction being limited only by physiological factors) and is a constant for a given population. 2. Ecological or realized natality refers to population increase under an actual, existing specific condition. Natality Natality can be expressed as Nn/t = Absolute natality (B) Nn/Nt = Specific natality rate (b) (natality rate per unit population) Where, N = the initial number of organisms n = the new individuals in the population t = the time. Mortality (Death rate) Population size may decrease as a result of emigration or death. It may be expressed as percentage, numbers per thousand dying per year. Mortality may take any of the following forms:- 1. Minimum mortality: It is also called specific or potential mortality, and represents the theoretical minimum loss under ideal or nonlimiting conditions e.g. death as a result of old age or decline in physiology 2. Ecological or realized mortality: this is the actual loss of an individual under a given environmental condition. Mortality varies with population and environmental conditions. Mortality Mortality can be expressed mathematically as deaths per time: A birth ratio (100 X Births/deaths) = Vital index The important consideration is the members who survives. Thus survival rates are generally expressed by survival curves (loss rates) 3. Survivorship: It involves direct counting of individuals per unit time or intervals. It is calculated as: Survival rate = m1/ m0 X 100 Where m0 = number of marked individuals at the beginning and m1 = number of surviving individuals at the time interval. Three main survivor curves 1. Convex curves (Type 1): Minimal mortality recorded at the beginning i.e. young and adult stages. Individuals that have a high probability of surviving through early and middle life but have a rapid decline in the number of individuals surviving into late life. 2. Diagonal curves (Type II): Shows a roughly constant mortality rate for the species through its entire life. This means that the individual's chance of dying is independent of their age. Type II survivorship curves are plotted as a diagonal line going downward on a graph. 3. Concave curves (Type III): It depicts species where few individuals will live to adulthood and die as they get older because the greatest mortality for these individuals is experienced early in life. This type of survivorship curve is drawn as a concave curve on a graph. Types of Survival curves Life span and population Information about life span and age are important factors in characterizing a population and predicting its course. How many people are below, within or beyond the reproductive age? How long is the life span? Which age group are most vulnerable to die? Concept of r-selected Organisms adapted to survive in unstable environments are referred to as r-selected. r-selected organisms’ live in settings where population levels are well below the maximum number that the environment can support i.e. the carrying capacity, there numbers are growing exponentially at the maximum rate at which the population can increase if resources are unlimited. Organisms that are r-selected tend to be small, short lived, and opportunistic, and to grow through irregular boom-and-burst population cycles. Examples include insects, annual plants, bacteria, frogs and rats etc. Species considered pests typically are r-selected organisms that are capable of rapid growth when environmental conditions are favorable. K-selected Organisms Organisms adapted to survive in stable environments are referred to as K-selected. This is because they live in environments in which the number of individuals is at or near the environment's carrying capacity (often abbreviated as K). K-selected species are typically larger, grow more slowly, have fewer offspring and spend more time parenting them. Examples include large mammals, birds, and longlived plants such as redwood trees. K-selected species are more prone to extinction than r-selected species because they mature later in life and have fewer offspring with longer gestation times. Biotic Potential Each population has the inherent power to grow. When the environment is unlimited (Space, food, other organisms not exerting a limiting effect), the specific growth rate (i.e. the population growth rate par individual) becomes constant and maximum for the existing conditions. The value of the growth rate under these favourable conditions is maximum, is characteristic of a particular population age structure, and is a single index of the inherent power of a population to grow Index r is the difference between the instantaneous specific natality rate (rate per time per individual) and the instantaneous specific death rate (r = b – d). Age Structure Not all individuals contribute equally to a population. Occasionally, researchers find it useful to characterize the different contributions made by different individuals. First, individuals are sorted into agespecific categories called cohorts, such as "juveniles" or "sub-adults". Researchers then create a profile of the size and age structures of the cohorts to determine the reproductive potential of that population, in order to estimate current and future growth. Usually, a rapidly expanding population will have larger reproductive cohorts, stable populations show a more even distribution of age classes, and rapidly declining populations have large older cohorts. Spatial Distribution (Dispersion) The spatial pattern in which individuals are dispersed within a given area is that population’s distribution, which may vary with time and available resources. • There are three major types of spatial distributions: – Clumped – Uniform – Random Spatial Distribution (Dispersion) Clumped distribution – includes family and social groups • Examples: elephant herds, wolf packs, prides of lions, flocks of birds, and schools of fish. • Advantages: – Provides many eyes to can search for local food sources. – Confuses predators with sheer numbers. – Cooperation for hunting more effectively. Spatial Distribution (Dispersion) Clumped distribution – includes family and social groups • Examples: elephant herds, wolf packs, prides of lions, flocks of birds, and schools of fish. • Advantages: – Provides many eyes to can search for local food sources. – Confuses predators with sheer numbers. – Cooperation for hunting more effectively. Spatial Distribution (Dispersion) Uniform distribution – constant distance maintained between individuals; common among territorial animals defending scarce resources or defending breeding territories. • Examples: iguanas, shorebirds, tawny owls • Advantage: a uniform distribution helps ensure adequate resources for each individual. Random distribution - Rare, exhibited by individuals that do not form social groups; occurs when resources are not scarce enough to require territorial spacing or cooperative behavior. • Examples: Trees and other plants in rain forests. Spatial Distribution (Dispersion) Uniform distribution – constant distance maintained between individuals; common among territorial animals defending scarce resources or defending breeding territories. • Examples: iguanas, shorebirds, tawny owls • Advantage: a uniform distribution helps ensure adequate resources for each individual. Random distribution - Rare, exhibited by individuals that do not form social groups; occurs when resources are not scarce enough to require territorial spacing or cooperative behavior. • Examples: Trees and other plants in rain forests.