Chapter 10

Populatio
n
Dynamic
contrast,s hemlock did

| In
not
reach its present range limits
until just 2,000 years ago.

6,000 yr

----------------------0------\ V----

000 yr
8.0
10.0

yr
yr
^ 10,000 yr
12,0

yr

Hemlock spread
north and west
from
the
southeast.

Hemlock
(Tsuga canadensis)

Figure 10.6 The northward expansion of two tree species in North America following glacial retreat (data from Davis 1981).
ago. In contrast, hemlock didn't reach the northwestern
limit
of its present distribution until 2,000 years ago.
The pollen preserved in lake sediments indicates that
forest trees in eastern North America spread northward following the retreat of the glaciers at the rate of 100 to 400
m
(0.1-0.4 km) per year. This rate of dispersal is similar to
that
of some large mammals such as the North American elk.
However, it is 1/100 the rate of dispersal shown by
Eurasian
collared doves in Europe and 1/1,000 the dispersal rate of
Africanized honeybees across South, Central, and North
America.
The previous examples concern dispersal by populations in the process of expanding their ranges. Significant
dispersal also takes place within established populations
whose
ranges are not changing. Movements, within established
ranges, can be an important aspect of local population
dynamics. We will consider two examples.

Dispersal in Response
to Changing Food Supply
Predators show several kinds of responses to variation in
prey density. In addition to the functional response we discussed in chapter 6, C. S. Holling (1959) also observed
numerical responses to increased prey availability. Numerical responses are changes in the density of predator
populations in response to increased prey density. Holling studied
populations of mice and shrews preying on insect cocoons

and attributed the numerical responses he observed to

increased reproductive rates. He commented that because
the reproductive rate of small mammals is so high, there
was
an almost immediate increase in density with increase in
food. However, some other predators, with much lower
reproductive rates, also show strong numerical responses.
These numerical responses to prey density are almost
entirely due to dispersal.
In some years, northern landscapes are alive with small
rodents called voles, Microtus spp. Go to the same place
during other years and it may be difficult to find any voles. In
northern latitudes, vole populations usually reach high densities every 3 to 4 years. Between these peak times, population densities crash. Population cycles in different areas are
not synchronized, however. In other words, while vole population density is very low in one area, it is high elsewhere.
Erkki Korpimiiki and Kai Norrdahl (1991) conducted a
10-year study of voles and their predators. The study began
in 1977 during a peak in vole densities of about 1,800 per
square kilometer and continued through two more peaks in
1982 (960/km 2 ) and 1985-86 (1,980 and 1,710/km 2 ). The
researchers estimated that between these population peaks
vole densities per square kilometer fell to as low as 70 in
1980
and 40 in 1984. During this period, the densities of the
European kestrel, Falco tinnunculus, short-eared owls, Asioflammeus, and long-eared owls, Asio orus, closely tracked vole
densities (fig. 10.7). How do kestrel and owl populations
track these variations in vole densities?
What mechanisms produce the numerical responses
by kestrels and owls to changing vole densities? Look at

236
Population Ecology

figure 10.7 for a clue. The peaks in raptor densities in

1977,
1982, and 1986 match the peaks in vole densities almost
perfectly. If reproduction was the source of numerical
response
by kestrels and owls, there would have been more of a
delay,
or time lag, in kestrel and owl numerical response. From
this
close match in numbers, Korpimaki and Norrdahl proposed
that kestrels and owls must move from place to place in
response to local increases in vole populations.
Is there any supporting evidence for high rates of
movement by kestrels and owls? Korpimaki (1988) marked and

Kestrel and owl densities

closely follow variation
in vole densities in

Voles
@ Kestrels and owls

2.0
00

In the colonization
cycle, upstream and
downstream
dispersal

1.500-

1.000

Many organisms engage

in
upstream
movements
that appear to compensate
for downstream drift.

Figure

10.7

Dispersal

and

numerical

response

by

predators

(data from Korpimaki and Norrdahl 1991).

Drift moves organisms

downstream, sometimes
; actively as behavioral
drift, sometimes passively
with floods.

and
reproduction
have
major influences on
stream populations.

Figure 10.8 The colonization cycle of stream invertebrates.

recaptured 217 kestrels, a large proportion of their study

population. Because European kestrel populations have an
annual survival rate of 48%' to 66%, he predicted a high
rate
of recapture of the marked birds. However, only 3% of the
female and 13% of the male kestrels were recaptured. These
very low rates of recapture indicated that kestrels were moving out of the study area. From their data, Korpimaki and
Norrdahl concluded that the kestrels and owls in western
Finland are nomadic, moving from place to place in
response to changing vole densities.
These studies documented the contribution of dispersal
to local populations of kestrels and owls. Earlier in this section, we saw how studies of expanding populations have
shed light on the contribution of dispersal to local
population
density and dynamics. Many other local populations are
strongly influenced by dispersal. One of the environments
in
which dispersal has a major influence on local populations
is
in streams and rivers.

Dispersal in Rivers and Streams

One of the most distinctive features of the stream and river
environment is current, the downstream flow of water. What
effect does current have on the lives of stream organisms?
As
you may recall from chapter 3, the effects of current are
substantial and influence everything from the amount of
oxygen

in the water to the size, shape, and behavior of stream

organisms. In this section, we stop and consider how stream populations are affected by current.
Lets begin with a question. Why doesnt the flowing
water of streams eventually wash all stream organisms,
including fish, insects, snails, bacteria, algae, and fungi, out
to
sea? All stream dwellers have a variety of characteristics
that
help them maintain their position in streams. Some fish such
as trout are streamlined and can easily swim against swift
currents, while other fish like sculpins and loaches are well
designed for avoiding Ihe full strength of currents by living
on
the bottom and seeking shelter among or under stones.
Microorganisms resist being washed away by adhering to
the
surfaces of stones, wood, and other substrates. Many stream
insects are flattened and so stay out of the main force of the
current, while others are streamlined and fast-swimming.
Despite these means of staying in place, stream organisms do get washed downstream in large numbers, particularly during flash floods, or spates. To observe this
downstream movement of organisms, put a fine or medium
mesh net in a stream or river and you will soon capture
large
numbers of stream insects and algae along with fragments
of
leaves and wood. Stream ecologists refer to this downstream
movement of stream organisms as drift. Some drift is due to

displacement of organisms during flash floods. However,

some is due to the active movement of organisms downstream.
Whatever its cause, stream organisms drift downstream
in large numbers. Why doesnt drift eventually eliminate
organisms from the upstream sections of streams? Karl
Muller (1954, 1974) hypothesized that drift would
eventually
wash entire populations out of streams unless organisms
actively moved upstream to compensate for drift. He proposed that stream populations arc maintained through a
dynamic interplay between downstream and upstream dispersal that he called the colonization cycle. The colonization
cycle is a dynamic view of stream populations in which
upstream and downstream dispersal, as well as
reproduction,
have major influences on stream populations (fig. 10.8).
Many studies support Mullers hypothesized colonization cycle, especially among aquatic insects. As larvae,
aquatic insects disperse upstream as well as downstream by
swimming, crawling, and drifting. Because of continuous

dispersal, which reshuffles stream populations, new

substrates put into streams are quickly colonized by a wide
variety of stream invertebrates, algae, and bacteria. Most of
these dynamics are difficult to observe because they occur
too
quickly, within the substratum, or at night, or they involve
microorganisms impossible to observe directly without the
aid of a microscope. However, a snail that lives in a tropical
stream in Costa Rica provides a well-documented example
of
the colonization cycle.
The Rio Claro flows approximately 30 km through
tropical forest on the Osa Peninsula of Costa Rica before
flowing
into the Pacific Ocean. One of the most easily observed
inhabitants of the Rio Claro is the snail Neritina latissima. which
occupies the lower 5 km of the river. The eggs of Neritina
hatch
to produce free-living planktonic larvae that drift down to
the
Pacific Ocean. After the larvae metamorphose into small
snails they reenter the Rio Claro and begin moving
upstream
in huge migratory aggregations of up to 500.000
individuals
(fig. 10.9). These aggregations move slowly and may take
up
to I year to reach the upstream limit of the population.
The population of Neritina in the Rio Claro consists of
a mixture of migrating and stationary subpopulations, with

exchange between them. Individual snails migrate

upstream
for some distance and then leave the migrating wave and
enter a local subpopulation. At the same time individuals
from the local subpopulation enter the migratory wave and
move upstream. Thus, individuals move upstream in steps
and
immigration
continuously
adds
to
local
subpopulations,
while emigration removes individuals. Because an
organism
that is visible to the naked eye does all this in a clear
stream,
and does it at a snails pace, we are provided with a unique
opportunity to observe that dispersal can
strongly
influence
local population density. Dispersal dynamics,
though
difficult
to study, deserve greater attention.

Concept 10.1 Review

L. Why might a species such as the Eurasian collared
dove be less threatened by rapid climate change than
hemlock or maple trees?
2. Some ecologists who have hung clear plastic sheets
coated with adhesive capable of trapping flying insects
from river bridges have found that the side of the
sheets facing downstream trap more adult aquatic
insects than the upstream-facing side. Explain.

Ongoing dispersal can join numerous subpopulations to

form a metapopulation. Populations of many species
occur not as a single continuously distributed population
but
in spatially isolated patches, with significant exchange of
individuals among patches. A group of subpopulations
living
on such patches connected by exchange of individuals
among
patches make up a metapopulation. The population of
Glanville fritiIlary butterflies, Melitaea cinxia, which lives
in
dry meadows scattered through the landscape of southern
Finland (see chapter 8. p. 204). is a metapopulation. In the
discussion of this butterfly metapopulation, we reviewed
how the exchange of individuals among subpopulations has
been well documented (Saccheri et al. 1998). However, the
metapopulation of Melitaea in southern Finland is only one
of many that are well known. Here is another example of a
butterfly metapopulation.

A Metapopulation
of an Alpine Butterfly
Once population biologists began to include the concept
(b)

I ijmre 10.9

Tlte colonization cycle

in

action, (o) A wave of

migrating snails, Neritina latissima, in the Rio Claro, Costa Rica;

(/>) a close-up of the migrating snails.

of metapopulations in their thinking, they found them

everywhere. Butterflies have been well represented in
studies
of metapopulations. One ol these butterflies is the Rocky
Mountain Parnassian butterfly, Pcirnassins smintliens (figure 10.10). The range of P. smintliens extends from northern
New Mexico along the Rocky Mountains to southwestern.
Alaska. Along this range P. smintheus caterpillars leed
mainly on the leaves and flowers of stonecrop. Sedum sp..
in