BIRTH, SEX, AND DEATH: THE DEMOGRAPHY OF SINGLE POPULATIONS

Populations
Organisms are all to some extent sociable beings, interacting with one another to survive. Their interactions create
populations and communities. Populations are loose collections of individuals belonging the same species. Red deer (Cervus
elaphas) in Britain constitute a population. All of them could interbreed, should the opportunity arise. In practice, most
populations, including the red deer population in Britain, exist as sets of local populations or demes. A local red deer
population lives in the grounds of Lyme Park, Cheshire. Its members form a tightly linked, interbreeding group and display
features typical of many populations (Box 4.1).
Population growth
A population changes due to natality and immigration, mortality and emigration. Natality covers a variety of ways of
'coming into the world' —live birth, hatching, germinating, and fission. In mammals, natality is the same thing as the birth
rate. Mortality

4
POPULATIONS
Most species exist as groups of interbreeding individuals—populations. This chapter covers:
• the form and function of single populations
• ways in which populations survive
• human exploitation and control of populations
covers the only way of 'leaving the world' —dying. Death may occur through old age (senescence). It is more likely to result
from disease, starvation, or predation. Two aspects of reproduction are distinguished. Fertility measures the actual number of
new arrivals in a population. In humans, the fertility rate averages about one birth every eight years per female of
childbearing age. Fecundity measures the potential number of new arrivals. In humans, the fecundity rate is about one birth
per nine to eleven months per female of childbearing age. Not many women fulfil that particular potential!
Most classic population studies focused on natality and mortality, and assumed that immigration and emigration balance
another or are too small to contribute greatly to population change. Obviously, emigration and immigration are important
components of long-term biogeographical change. They are also important in more recent population studies that emphasize
small-scale comings and goings within a species range. This idea, in the guise of metapopulation theory, will be discussed
later.
Exponential growth
Populations grow exponentially when the population increase per unit time is proportional to population
Box 4.1
THE RED DEER POPULATION IN LYME PARK, CHESHIRE, ENGLAND
Lyme Park lies on the western flanks of the Pennines. Its 535 ha consist of park grassland, moorland, and woodland. Red
deer (Cervus elaphas) have lived in Lyme Park for over 400 years. They are a remnant of a larger population once present in
Macclesfield Forest, which used to extend many kilometres along the west side of the Pennines. Up to 1946, while the park
was privately owned, the population numbered 170 to 242. Some deer were shot for sport and some for the dinner table.
After ownership was passed to the National Trust and Stockport Metropolitan Borough Council, the population rose to over
500. Culling was introduced in 1975 and the population is now maintained at 250 to 300.
A study of the red deer population was carried out from 1975 to 1983 (Goldspink 1987). The population was dominated
by hinds. There were over three times more hinds than stags. Hinds were recorded in three main areas—the park, the
moorland, and Cluse Hay (which is a subdivision of the moorland) (Figure 4.1). All tagged hinds remained close to their
areas of capture. Hinds tended to form large groups of 30 or more throughout the study area. Most calves were born in the
first two weeks of June. Maximum densities of hinds and calves per km2 varied from 30 in the park to 77 on the moorland.
Three bachelor herds of stags were present. Moor stags spent much of the summer on a small area (30 ha) of ground above
Cluse Hay. They dispersed more during winter, probably to avoid exposure and windchill. The Knott group, which lives in
the park, consisted of prime stags, more than 6 years old, during the winter. Younger stags were recorded on Cage Hill.
Stags in the park area were more tolerant of disturbance than those living on the moorland. The rut started in mid-September
and went on well into November. The median date of observed copulations was 16 October. The apparent gestation
POPULATIONS 85
period (to the median date of calving) was about 237 days. Stags from the Knott rutted on Cluse Hay, while those from Cage
Hill preferred the moor or the park. Moor stags appeared to play but a small part in the rut, despite their numerical
abundance. Dominant stags, which commonly had 40-hind harems for two to three weeks during the rut, were usually
replaced within two years.
Figure 4.1 The main areas occupied by the red deer (Cervus elaphas) population, Lyme Park, Cheshire, England. Source:
After Goldspink (1987)
Growth rates of stags and hinds were low on the moorland. Calf-to-hind ratios were generally low and within the range
0.20–0.34; maxima of 0.55– 0.60 were recorded in woodland sites. Conditions on
POPULATIONS 86
the moorland were severe during the winter due to the poor quality of herbage, mainly purple moorgrass (Molinia caerulea),
and a lack of shelter. Annual calf mortality varied from 10 to 30 per cent. It was particularly high during the cold winters of
1978–9 and 1981–2. Population density and winter food were probably principal factors limiting the
size and is unconstrained. This is readily appreciated by a simple example. Take two individuals. After a year they have
produced four offspring. Reproducing at the same rate, those four offspring will themselves have produced eight offspring
after a further year. The growth of the population follows a geometric progression—2, 4, 8, 16, 32, 64, and so on. This
exponential growth rapidly produces huge numbers. A female common housefly (Musca domestica) lays about 120 eggs at a
time (Leland Ossian Howard, cited in Kormondy 1996:194). Around half of these develop into females, each of which
potentially lays 120 eggs. The number of offspring in the second generation would therefore be 7,200. There are seven
generations per year. If all individuals of the reproducing generation should die off after producing, the first fertile female
will be a great, great, great, great, great grandmother to over 5.5 trillion flies.
Growth in populations with overlapping generations and prolonged or continuous breeding seasons may be described by
the equation:
dN/dt=rN
Notice that the growth rate, dN/dt, is directly proportional to population size, N, and to the intrinsic rate of increase (per
capita rate of population growth), r. The intrinsic rate of increase, sometimes called the Malthusian parameter, is defined as
the specific birth rate, b, minus the specific death rate, d. In symbols, r=bd. (Specific rates are rates per individual per unit
time.) Exponential growth, then, simply depends upon how many more births there are than deaths, and on how large the
population is. No other factors restrict population growth. Unhindered growth of this kind is the biotic potential of a
population
(RNChapman 1928). It is described by the solution to the population growth equation:
N=N
0
ert
This equation shows that the starting population, N
0
, increases exponentially (e is the base of the natural logarithms) at
rate determined by the intrinsic rate of natural increase, r. It describes a J-shaped curve (Figure 4.2). The maximum intrinsic
rate of natural increase varies enormously between species. In units of per capita per day, it is about 60 for the bacterium
Escherichia coli; 1.24 for the protozoan Paramecium aurelia; 0.015 for the Norway rat (Rattus norvegicus); 0.009 for the
domestic dog (Canis domesticus); and 0.0003 for humans. For the red deer population in Lyme Park, from 1969 to 1975, it
was 0.07696.
The doubling time is the time taken for a population to double its size. It may be calculated from values of r. The
doubling times for the species mentioned above, assuming geometric increase prevails, are: Escherichia coli—0.01 days;
Paramecium aurelia— 0.56 days; Rattus norvegicus—46 days; Canis domesticus —77 days; and humans—6.3 years. For
Lyme Park red deer, it is 9.0 years.
Logistic growth
The English house sparrow (Passer domesticus) was introduced to the United States around 1899. Concern was expressed
that in a decade, a single pair could lead to 275,716,983,698 descendants, and by 1916 to 1920 there would be about 575
birds per 40 ha (Kormondy 1996:194). In the event, there were only 18 to 26 birds per 40 ha, less than 5 per cent of the
performance of deer on the moor. Growth rates of stags and hinds in the park were high, but calf-to- hind ratios were low at
0.15–0.26. Culling has reduced levels of natural mortality but further improvements in performance are unlikely to be
achieved without a reduction in sheep stocks, some improvement in habitat, and the provision of shelter.
POPULATIONS 87
Figure 4.2 Exponential growth of a hypothetical population. Notice that, with a bit of artistic licence, the population growth
describes a J-shaped curve
expected density. Patently, something had prevented the unfettered geometric increase in the immigrant house sparrow
population.
A growing population cannot increase indefinitely at a geometric rate. There is a ceiling or carrying capacity, a limit to
the number of individuals that a given habitat can support. The biotic potential of a population is curtailed by an
environmental resistance (RNChapman 1928). After a period of rapid growth, the house sparrow population reached its
population ceiling, partly because native hawks and owls took to the new English dish on the menu.
The population ceiling or carrying capacity is usually denoted by the letter K. It alters the population growth equation in
this way:
dN/dt=rN(1-N/K)
This equation for growth rate is called the logistic equation. It describes a sigmoid or S-shaped curve (with the top of the 'S'
being pushed to the right) where population grows fast initially but tapers off towards the carrying capacity (Figure 4.3).
When the population size is small, compared with the carrying capacity, the environmental resistance to growth is minimal.
The population responds by virtually unbounded exponential growth. This is the bottom part of the 'S'. When the population
size is large and
approaching the carrying capacity, environmental resistance is great. The population barely grows. This is the distorted top
part of the 'S'. If the population should exceed the carrying capacity, negative growth occurs; in other words, the population
falls until it drops below the carrying capacity.
Population growth (or decline) rates are affected by density-dependent factors and by density- independent factors. The
effects of density- independent factors do not vary with population density and they will affect the same proportion of
organisms at any density. Weather, pests, pathogens, and humans commonly affect populations in that way. The effects of
density-independent factors do vary with population density, so that the portion of organisms affected increases or decreases
with population density. Birth rates and death rates both depend on population density: birth rates tend to decrease with
increasing density, while death rates tend to increase with increasing density.
Population irruptions
Irruptions are common features of many populations. They involve a population surging to a high density and then sharply
declining. This boom- and-bust pattern occurs in some introduced ungulates. It happened to the Himalayan tahr (Hemitragus
Figure 4.3 Logistic growth of a hypothetical population. Notice that the population growth curve is a stretched S-shape
POPULATIONS 88
jemlahicus) in New Zealand (Caughley 1970). The tahr is a goat-like ungulate from Asia. Introduced into New Zealand in
1904, it spread through a large part of the Southern Alps. After its introduction, the population steadily rose. The birth rate
fell slightly but the death rate increased, largely due to greater juvenile mortality. After a period when the population was
large, a decline set in. This decline was caused by reduced adult fecundity and further juvenile losses. Changes in the tahr
population may have resulted from changes in food supply. As the tahr grazed, they altered the character of the vegetation
that they fed on. In particular, snow tussocks (Chionochloa spp.), which are evergreen perennial grasses, were an important
food source late in winter. They cannot abide even moderate grazing pressure and disappeared from land occupied by the
tahr. With no snow tussocks to eat, the tahr browsed on shrubs in winter and managed to kill some of these.
Another irruption occurred on Southampton Island, Northwest Territories, Canada (DCHeard and Ouellet 1994). By
1953, Caribou (Rangifer tarandus groenlandicus) had been hunted to extinction on Southampton Island. In 1967, 48 caribou
were captured on neighbouring Coat's Island and released on Southampton Island. The population of caribou older than 1
year grew from 38 in 1967 to 13,700 in 1991. The annual growth rate during this period was 27.6 per cent and did not
decrease with increasing population density. On Southampton Island, caribou did not suffer high winter mortality in some
years, but they did on Coat's Island. Caribou density was higher on Coat's, which suggests that adverse weather has a
minimal effect when animal density is low.
Irruptions also occur because of unusually high immigration. In the winter of 1986–7, many more rough-legged buzzards
(Buteo lagopus) moved into Baden-Württemberg, south-west Germany, than in any other winter during at least the preceding
century (Dobler et al. 1991). From 14 January to 7 April, rough-legged buzzards were observed daily. The highest numbers
observed were 109 individuals on 1 February and 110 on 8 February. The buzzard influx was caused by high snow cover and
cold spells in the eastern parts of Central Europe. Depressions over
Southern Europe may have blocked the route south with cloud and snowfall.
Population crashes
Some populations often crash. In early 1989, two- thirds of the Soay sheep (Ovis aries) population on St Kilda, in the Outer
Hebrides, Scotland, died within a twelve-week period. The cause of the crash was investigated by post-mortem examination
and laboratory experiments (Gulland 1992). Post-mortem examination showed emaciated carcasses with a large number of
nematode (Ostertagia circumcincta) parasites, and that the probable cause of death was protein-energy malnutrition.
However, well-nourished Soay sheep artificially infected with the parasite in the laboratory showed no clinical signs or
mortality, even when their parasite burdens were the same as those recorded in the dead St Kilda sheep. Thus, parasites
probably contributed to mortality only in mal-nourished hosts, exacerbating the effects of food shortage.
Marine iguanas (Amblyrhynchus cristatus) living on Sante Fe, in the Galápagos Islands, suffered a 60 to 70 per cent
mortality due to starvation during the 1982–3 El Niño—Southern Oscillation event (Laurie and Brown 1990a, b). Adult
males suffered a higher mortality than adult females while food was short, but size explained most of the mortality
differences between the sexes. Almost no females bred after the event. In the next year, the frequency of reproduction
doubled, the age of first breeding decreased, and mean clutch size increased from two to three. These demographic changes
are examples of density- dependent adjustments of population parameters that help to compensate for the crash.
 Population crashes may have repercussions within communities. In Africa, savannahs are maintained as grassland partly
because of browsing pressures by large and medium herbivores. In Lake Manyara National Park, northern Tanzania, bush
encroached on the grassland between 1985 and 1991, shrub cover increasing by around 20 per cent (Prins and Van der Jeugd
1993). Since 1987, poaching has caused a steep
POPULATIONS 89
decline in the African elephant (Loxodonta africana) population in the Park. However, shrub establishment preceded the
elephant population decline, and is not attributable to a reduction in browsing. In two areas of the Park, shrub establishment
coincided with anthrax epidemics that decimated the impala (Aepyceros melampus) population. In the northern section of
the Park the epidemic was in 1984; in the southern section it was in 1977. An even-aged stand of umbrella thorn (Acacia
tortilis) was established within the grassland in 1961, which date coincided with another anthrax outbreak among impala.
Similarly, another even-aged stand of umbrella thorn was established at the end of the 1880s, probably following a
rinderpest pandemic. The evidence suggests that umbrella-thorn seedling establishment is a rare event, largely owing to high
browsing pressures by such ungulates as impala. Punctuated disturbances by epidemics, which cause population crashes in
ungulate populations, create narrow windows for seedling establishment. This process may explain the occurrence of even-
aged umbrella thorn stands.
Chaotic growth
Irregular fluctuations observed in natural populations, including explosions and crashes, were traditionally attributed to
external random influences such as climate and disease. In the 1970s, Robert M.May recognized that these wild fluctuations
could result from intrinsic non-linearities in population dynamics (May 1976). For instance, a relatively simple population
growth equation of the form
N
t+1
and Wedin 1991). Monocultures of ticklegrass were planted at two different densities on ten different soil mixtures.
Populations growing in unproductive soils, low in nitrogen, maintained fairly stable aboveground biomass. Populations
growing on richer soil showed biomass oscillations. Populations growing on the richest soils displayed as 6,000-fold crash
(Figure 4.5). No other species growing in the garden where the experiment was conducted crashed in 1988, which suggests
that environmental agencies were not responsible. The dynamics for the crashing populations were shown to be chaotic. The
chaotic regime resulted from the growth inhibition by litter accumulation. The litter causes a one-year delay between growth
and the inhibition of future growth. Litter production is greater in more productive plots. The magnitude of the time- delayed
inhibitory effect therefore increases with productivity. This may lead to an oscillation between a year of low litter and high
living biomass, and a year of high litter and low living biomass. The implications of this study are profound—litter feedback
may cause population oscillations and possibly chaos in productive habitats. However, litter accumulates where it falls, and
litter-driven chaotic dynamics occurs at small spatial scales—it may avoid detection if looked for at medium and large
scales.
Since the late 1980s, part of theoretical ecology has focused on the spatio-temporal dynamics generated by simple
ecological models. To a large extent, the results obtained have changed views of complexity (Bascompte and Sole 1995).
Specifically, simple rules are able to produce complex spatio- temporal patterns. Consequently, some of the
=N
te[r(1-N
t /K)]
complexity underlying Nature does not necessarily have complex causes. The emerging framework has which is applicable
to population that suffer
far-reaching implications in ecology and evolution. It epidemics
at high densities, possesses an amazing
is improving the comprehension of such topics as the range of
dynamic behaviour—stable points, stable
scale problem, the response to habitat fragmentation, cycles, and
chaos—depending on the growth rate, r
the relationships between chaos and extinction, and (Figure 4.4).
how higher diversity levels are supported in Nature. Much of the
work on chaotic population dynamics is theoretical. Some studies have shown that chaos does occur in natural populations.
The
Age and sex structure
ticklegrass or hairgrass (Agrostis scabra) displays
The aggregate population size, N, obscures the fact oscillations
and chaos in experimental plots (Tilman
that individuals differ in age and in ability to reproduce.
POPULATIONS 90
Figure 4.4 Internally driven population dynamics: stable points, stable cycles, and chaos. NB The growth rates, r, in (d) and
(e) are the same, but the ratios of the initial in(e), Source: N
After 0
/k=1.5.
May (1981)
population to the carrying capacity, N
0
/k, are different: in (d), N
0
/k= 0.075;
The population age-structure is the proportion of individuals in each age class. A stable age distribution with a zero growth
rate has about the same number of individuals in each age class, except for fewer in post-reproductive classes. In contrast, a
growing population has a pyramid-shaped structure, with large numbers of pre-reproductive individuals.
Life tables
A life table, like an actuarial table, shows the number of individuals ex pected in each age class based upon fecundity and
survival rates. Survival rates, s, are related to death rates, d, in the following way: s=1-d. Life-table statistics are normally