HOW POPULATIONS COHERE: FIVE RULES FOR COOPERATION

- Cooperation means that a donor pays a cost, c, for a recipient to get a benefit b
- Multicellular organisms exist because of cooperation between their cells
1. Kin selection
2. Direct reciprocity
3. Indirect reciprocity
4. graph selection
5. group selection

SINGLE-SPECIES DYNAMICS

- species whose population ecology can be studied without also explicitly including the dynamics of
other species in the community
- if birth and death rate remain constant then the consequent increase or decrease in population
numbers occurs exponentially population dynamics occurs on a geometric rather than an arithmetic scale

The rate of population growth

- unstructured population: birth and death rates are identical across individuals
- males have no effect on population growth rate (true for most, but not all organisms)
- = 0
- a disease will spread only if r=b-d>0 : every initial infection must leave at least one secondary
infection
- a population has a discrete breeding season
- a population is age-structured: demographic rates vary with age but are constant within an age
class
- the members in the youngest age class are given by the numbers in each age class the season
before, multiplied by their fecundity
- damping ratio: ratio of the largest eigenvalue of the projection matric to the second largest
eigenvalue (provides information whether there is a smooth or oscillatory approach to the long-term
growth rate
- in America adult Magicicada are abundant only once in 17 years, and this pattern could be
explained by initial conditions (for example all year classes are wiped except for one) plus lack of cohort
mixing; in fact, this is highly unlikely, the power of even minor cohort mixing to destroy the imprint of
starting values is so strong that ecologists have universally rejected this hypothesis and sought active
processes to maintain the synchronized cohorts; because the length of the life cycle is a prime number
(and 11- and 13- year cicada populations are found) the classical explanation is that it is a means of
escaping predation, as it is hard for predator populations with life cycle that are not exact divisors of the
cicada life cycle to increase in density

(, + 1) = [(, ) + (, )](, )
=0

(0, ) = ()(, )
0

- the numbers of individuals at age 0 are simply the numbers of current individuals in the population
multiplied by their age-specific birth rates
Density dependence

- as populations exceed their resources, there will be either a

decline in birth rates or an increase in death rates
- globally stable equilibrium: a single equilibrium point
- the Allee effect = presence of a minimum population density
below which the population goes extinct
- budworm moth (Choristoneura fumiferana) is hypothesised
normally to be controlled at a relatively low population density by a
suite of specialist invertebrate predators; if, however, the population
is perturbed such that control breaks down, the moth increases in
density to a second equilibrium which is set by bird predation (these
predators ignore spruce budworm unless they are very common); a
further perturbation is needed to switch it back to the low-density
equilibrium
- if the angle is more than 45 (growth curve is below the 45 line
to the left of the equilibrium) we have an unstable equilibrium
- cannibalism is much more common in animal kingdom than often realised, and when it occurs it is
nearly always size-related, with older larger individuals consuming their smaller conspecifics

Chaos
- Ricker equation as it is commonly used in applied population biology, particularly for fisheries (nt is
population density, r=fecundity factor)
(1 )
+1 =
- chaos: the trajectory never converges on a simple cycles,
but fluctuates aperiodically around very many values of n, never
repeating itself
- this has the structure of a fractal
- Ricker model predicts chaos for all r>2.69
- all chaotic systems are governed by strange attractors
and, determining that a systems attractor is fractal is one way
of identifying chaos in nature
- Tribolium castaneum: adults and larvae cannibalise eggs, while adults also eat pupae: by varying a
single parameter (pupal mortality) the dynamics of the system moved from stability to chaos and then to a
3-point cycle)

Randomness
- poor years have a greater negative effect on population growth than the positive effects of good
years
- a single year with 0 net production renders the long-term growth rate 0
- demographic stochasticity can be ignored with more than 50 or so female breeders
ANALOGY: PINBALL MACHINE (buffers)
 METAPOPULATIONS AND THEIR SPATIAL DYNAMICS

- Levins metapopulation idea: patches of suitable habitat for a species are distributed across a
landscape over time, there is a dynamical process of colonization and extinction
o empty habitat
o eradication threshold: habitat destruction has the effect of lowering the colonization curve and
making it less steep towards the origin;
if the slope of the colonization curve at the origin becomes smaller than the slope of the
extinction curve, extinction is inevitable and this does not require all or even necessarily
a substantial fraction of patches to be destroyed
extinction debt
Allee effect: either birth rates decrease or death rates increase at low densities (=inverse density
dependence)
bimodal distribution of patch occupancy
o source-sink metapopulations: source =net outflow of individuals(surplus production of
individuals); sink=population only persists because of immigration into it from source habitats
Arctic ground squirrels: prime land for colonies is scarce and squirrels that are forced
out burrow in the banks of creeks that are prone to flooding, which drowns them
Temperate plants have the option of a seed bank, while tropical ones don't
humans are a sink habitat for
Ebola virus
entirely conceivable that
most members of a species
live in a sink habitat
(especially if it is not too bad)
inferences about a
species preferred habitat
based on where we typically
find it may be completely
wrong
CONSERVATION: better to
save a small amount of
source
habitat
than a big
amount of
sink habitat
o two-species Levins metapopulations:
patch extinction can be a function of patch age: the
colonization rate be greater than 1/A (A is the average age of the
patches)
habitat destruction has no effect on equilibrium prey
abundance
(extinction: wood-rotting fungi will find that their patch ultimately rots
completely way and epiphytic mosses will ultimately find that their
tree falls over)
- local populations connected by dispersal, but without the extinction
of the local populations
- populations in patches of good habitat sustain populations in poor
habitats
 PREDATOR-PREY INTERACTIONS