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Lesson 1 of 5Population Dynamics
Unit 5 · Lesson 1 of 5

Population Characteristics and Sampling

Population size, density and dispersion patterns; estimating size with quadrats, transects and mark–recapture; natality, fecundity and mortality; life tables, survivorship curves, age structure and minimum viable population size.

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Key things to remember

  1. 1Density: (or in water); quadrat estimate
  2. 2Lincoln–Petersen mark–recapture: , assuming a closed population, no lost marks, random mixing and equal catchability.
  3. 3Dispersion: clumped (patchy resources, social groups), uniform (territoriality, competition, allelopathy), random (uniform resources, no interaction).
  4. 4Survivorship: Type I dies mostly old (humans), Type II constant rate (songbirds), Type III dies mostly young (oysters, cod).
  5. 5Fecundity is potential reproductive output; natality is the actual birth rate; mortality is the death rate.
  6. 6Minimum viable population: the smallest size that can survive demographic, environmental and genetic chance events over a set time.

Suppose a wildlife agency needs to know whether the moose in a northern Ontario management area are increasing or declining, so it can set hunting quotas. Nobody can count every moose in thousands of square kilometres of forest. The same problem faces anyone who studies a population, whether it is trilliums on a hillside, walleye in a lake or bacteria in a flask. Ecologists solve it by sampling: counting carefully in a part of the habitat and scaling up, or marking some animals and seeing how many turn up again. Once they have numbers, they also ask how the individuals are spread out, how many are born and die at each age, and whether the population is large enough to survive at all. Those measurements are the raw material for every model in this unit.

Size, density and dispersion

A population is a group of individuals of the same species living in the same area at the same time, able to interbreed. Its population size () is simply the number of individuals. Size alone can mislead: 500 white-tailed deer is a sparse population in a 1000 km² forest but a crowded one in a 5 km² park. Population density () corrects for this by dividing by the space occupied: individuals per unit area for land organisms (per m², per hectare, per km²) or per unit volume for aquatic and microscopic organisms (per litre, per mL). Crude density uses the total area; ecological density uses only the area of usable habitat, which is often more meaningful because a lake-dwelling loon cannot use the surrounding forest.

Density is population size divided by the area or volume the population occupies.
Clumpedresources in patches, or socialgroups (e.g. herds, fish schools,seeds falling near the parent)Uniformcompetition or territorialspacing (e.g. nesting penguins,creosote bushes)Randomresources spread evenly andlittle interaction (e.g. dandelionsfrom wind-blown seeds)
Fig 1Dispersion patterns: clumped individuals gather in groups, uniform individuals are evenly spaced, random individuals show no pattern.
Dispersion patternUsual causeExamples
Clumped (most common)Resources are patchy (water, food, shelter); social behaviour (herds, packs, schools); offspring stay near the parent (seeds that fall close, clonal roots).Caribou herds, wolf packs, schooling fish, trembling aspen clones, mushrooms on a rotting log
Uniform (even spacing)Individuals repel each other: territorial defence, competition for a scarce resource such as water, or chemicals that inhibit neighbours (allelopathy).Nesting northern gannets on Bonaventure Island, Quebec; creosote bushes in a desert; plants near a black walnut, which releases the toxin juglone
RandomResources are evenly spread and individuals neither attract nor repel each other. Rare in nature.Dandelions grown from wind-blown seeds in a uniform lawn; some forest trees and invertebrates
Dispersion describes how individuals are spaced within the area, which is a separate question from how many there are.

Estimating population size

For plants and slow-moving animals, ecologists use quadrats: square frames of known area (often 1 m² or 0.25 m²) placed at random points in the habitat. Every individual inside each quadrat is counted. Random placement matters, because choosing spots that look interesting biases the result. The sample density is the total count divided by the total area sampled (number of quadrats times the area of one quadrat), and the population estimate is that density multiplied by the total area of the habitat: . When conditions change across a habitat, such as from a pond edge up a slope or from a forest edge into a field, a transect is used instead: a line along which organisms are recorded at set intervals (a line transect) or within a strip of fixed width (a belt transect). Transects reveal how density changes along a gradient, which random quadrats would blur.

Animals that move, hide or live in water need a different method. In mark–recapture, researchers capture a sample, mark each animal harmlessly (ear tags, leg bands, a clipped fin, a dab of paint) and release them. After the marked animals have had time to mix back in, a second sample is captured. The key idea is that the fraction of marked animals in the second sample should equal the fraction of marked animals in the whole population: . Rearranging gives the Lincoln–Petersen index.

M = number marked in the first capture; n = total caught in the second capture; m = marked animals recaptured in the second capture.
Mark–recapture only works if its assumptions hold
The estimate assumes: (1) the population is closed during the study, with no births, deaths, immigration or emigration between samples; (2) marks are not lost and do not harm the animal or change its behaviour; (3) marked animals mix randomly back into the population; (4) every individual, marked or not, has the same chance of being caught. If marked animals learn to avoid traps (trap-shy), too few are recaptured, is too small and is overestimated. If they learn traps mean free food (trap-happy), is too large and is underestimated. Lost tags also shrink and inflate .

Births, deaths and survivorship

Natality is the birth rate: the number of offspring produced per unit time, often expressed per 1000 individuals. Fecundity is the potential reproductive capacity of an individual under ideal conditions (a female cod may release millions of eggs), which is usually far higher than fertility, the number of offspring actually produced. Mortality is the death rate: the number of individuals dying per unit time. To see how mortality changes with age, ecologists build a life table. They follow a cohort (a group born at the same time) and record how many are still alive at the start of each age interval. The mortality rate for an interval is the number that died during it divided by the number alive at its start. Plotting the number of survivors against age gives a survivorship curve. The survivor axis is usually logarithmic, so a straight line means a constant proportion dies in each interval.

10001001010255075100Percentage of maximum life spanNumber of survivors (log scale)Type IType IIType IIIType I — humans, large mammals: most survive to old age, then die over a short spanType II — songbirds, some lizards: about the same chance of dying at every ageType III — fish, oysters, many plants: most die young; the few survivors live long
Fig 2Survivorship curves on a log scale: Type I survivors fall late in life, Type II fall at a constant rate, Type III fall steeply early.
CurvePattern of mortalityTypical life historyExamples
Type ILow mortality through youth and middle age; most deaths in old age.Few offspring, much parental careHumans in countries with good health care, Dall sheep, elephants
Type IIRoughly constant proportion dies at every age (a straight line on a log scale).Moderate offspring numbers; risk from predators or accidents at every ageMany songbirds such as the American robin, grey squirrels, some lizards
Type IIIVery high mortality among the young; the few that survive early life live relatively long.Huge numbers of offspring, little or no parental careOysters, Atlantic cod, frogs, sugar maple seedlings
Real populations often fall between types or change type across the life span.

Age structure is the proportion of a population in each age class, usually grouped as pre-reproductive, reproductive and post-reproductive. A population with many young individuals has built-in potential to grow as they reach breeding age; one dominated by older individuals will tend to decline. Small populations face a special danger. The minimum viable population (MVP) is the smallest population size that can persist for a set time (for example a 95% chance of surviving 100 years) despite chance events. Below it, a population becomes vulnerable to three kinds of randomness: demographic (by chance, one year's births are mostly male), environmental (one harsh winter or fire) and genetic (inbreeding and genetic drift erode variation). Individuals may also struggle to find mates. These effects feed on each other, a spiral sometimes called an extinction vortex. The Vancouver Island marmot, which fell to fewer than 30 animals in the wild in the early 2000s before captive breeding and releases, is a Canadian example of a population pushed below a safe size.

Worked examples

Reveal one step at a time and try to beat it to the next line.

Example 1
To estimate the snowshoe hares in a boreal forest plot, researchers live-trap and ear-tag 120 hares, then release them. Two weeks later they trap 150 hares, of which 30 carry tags. Estimate the population and state one assumption the estimate depends on.
Example 2
A student places five 1.0 m² quadrats at random in a 2.0 ha woodland (1 ha = 10 000 m²) and counts white trilliums: 4, 7, 3, 6 and 5. Estimate the trillium density and population size.
Example 3
A cohort of 1000 frog eggs is followed. Survivors at the start of each year: age 0: 1000; age 1: 100; age 2: 60; age 3: 40; age 4: 25. Calculate the mortality rate for the first year and for age 3 to 4, and identify the survivorship type.

Where marks get lost

  • Mixing up the letters in Lincoln–Petersen. and go on top and the recaptured marked count goes on the bottom; writing gives a number smaller than the number you marked, which is impossible.
  • Dividing the quadrat total by the number of quadrats instead of the total area sampled. Five 0.25 m² quadrats cover 1.25 m², not 5 m², so the density is the count divided by 1.25 m².
  • Confusing density with dispersion. Two populations can have the same density (individuals per m²) but very different spacing (clumped versus uniform).
  • Treating fecundity and natality as the same thing. Fecundity is the maximum possible output; natality is the actual birth rate, which is usually far lower.
  • Reading a straight survivorship line as a constant number dying each year. On the usual logarithmic axis, a straight line (Type II) means a constant proportion dies each interval.

Flashcards 10

Population
Population density
Clumped, uniform, random dispersion
Lincoln–Petersen index
+ 6 more

Practice

1
Northern gannets nest on cliff ledges with each nest just beyond pecking distance of its neighbours. What dispersion pattern is this, and what causes it?
2
Biologists band 45 ducks at a marsh. A week later they capture 60 ducks, 9 of which are banded. Use the Lincoln–Petersen index to estimate the population.
ducks
3
Four 0.25 m² quadrats placed at random in a 500 m² field contain 5, 3, 6 and 4 dandelions. Estimate the number of dandelions in the field.
dandelions
4
An aerial survey counts 240 white-tailed deer in a 12 km² conservation area. What is the population density?
deer/km²
5
In a life table, 400 individuals of a cohort are alive at the start of age 5 and 300 are alive at the start of age 6. What is the mortality rate for that year, as a percentage?
%
6
Which organism is most likely to show a Type III survivorship curve?
7
During a mark–recapture study of chipmunks, marked animals learn that traps contain bait and return to them more often than unmarked animals. How does this affect the population estimate?
8
What term describes the maximum potential number of offspring an individual can produce under ideal conditions?
9
Put the steps of a mark–recapture study in order.
  1. 1Calculate N = Mn/m
  2. 2Capture a second sample and count the total (n) and the marked animals (m)
  3. 3Release the marked animals and allow them to mix with the population
  4. 4Capture a first sample of animals
  5. 5Mark each animal harmlessly and record the number marked (M)
10
Match each survivorship type or dispersion pattern with the best example.
Type I survivorship
Type II survivorship
Type III survivorship
Clumped dispersion
Random dispersion
11
What is the term for the smallest population size that is likely to persist for a set period despite random genetic, demographic and environmental events? (Give the full term or its abbreviation.)

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