Lecture Note:

Animal Ecology (Biol. 423)

Chapter 2. Population Ecology
2.3. Life History Patterns and
Reproduction Strategies
2.3. Life History Patterns and reproductive strategies
2.3.1. Life history evolution
Study of life history patterns helps in find answers to the
following and other similar questions
1. Why is it that swifts usually produce clutches of three eggs,
when other birds produce larger clutches, and the swifts
themselves are physiologically capable of doing so?
2. Why is it that the ratio between age at maturity and average
lifespan is often roughly constant within a group of
organisms but markedly different between groups (e.g.
mammals 1.3, fish 0.45)?
3. Why does orchids produce vast numbers of tiny seeds while
tropical Mora trees produce just a few enormous ones?
4. Why does mammals produce very few off springs while
fishes release thousands of eggs?
• The study of the evolution of life histories is a search for
patterns – and for explanations for those patterns.
• We must remember, however, that every life history, and
every habitat, is unique.
2.3.2. Components of life histories
• What are the most important components of any organism’s
life history?
A. Individual Size
B. Rapid Development
C. Size of offspring
A. Large Size
Advantages of individual size
• increase an organism’s competitive ability
• increase individual’s success as a predator
• decrease individual’s vulnerability to predation
• Has higher stored energy (can survive better in times of
reduced availability of food/nutrients), and
• Produce more off springs
• Disadvantages of larger size:
– Vulnerable to some physical forces (e.g. a larger tree is
more likely to be felled in a gale)
– Preferred by predators (e.g. many predators exhibit a
preference for larger prey), and
– larger individuals typically require more resources and may
therefore be more prone to a shortage of them.
• Hence, in many organisms intermediate sized, not maximum
size, are optimal (e.g. Damselfish Figure 4.16).
Figure 4.16 For adult male damselflies, Coenagrion puella, the predicted optimum size
(weight) is intermediate (upper graph), and corresponds closely to the modal size class in
the population (histogram below). The upper graph takes this form because mating rate
decreases with weight, whereas lifespan increases with weight
B. Rapid Development
• Development is the progressive differentiation of parts,
enabling an organism to do different things at different stages
in its life history.
• Rapid development can increase fitness because it leads to
the rapid initiation of reproduction.
• Rapid development leads to early onset of reproduction.
c. Size of offspring
• Large sized of off springs are often better competitors
• They obtain better nutrients
• They are better at surviving in extreme environments.
• Hence, they often have a better chance of surviving to
reproduce themselves.
• Combining all of this detail, life histories are often described
in terms of a composite measure of reproductive activity
known as ‘reproductive allocation’ (also often called
‘reproductive effort’).
• This is best defined as the proportion of the available resource
input that is allocated to reproduction over a defined period
of time;
• Figure 4.17 shows an example involving the allocation of
nitrogen, a crucial resource in this case.
Figure 4.17 Percentage allocation of the crucial resource nitrogen to different
structures throughout the annual cycle of the perennial plant Sparaxis grandiflora
in South Africa. The plant sets fruit in the southern hemisphere spring
(September–December). The plant grows each year from a corm.
4.3.3. Reproductive value
• Natural selection favors those individuals that make the
greatest proportionate contribution to the future of the
population
• Contribution of individuals to future generation is ultimately
realized through their effects on fecundity and survival
• One of the measures of fitness of organism is through the
reproductive value
Reproductive value
I. Is the sum of the current reproductive output and the
future reproductive outputs (Residual Reproductive Value,
RRV)
II. RRV combines expected future survival and expected future
fecundity. Thus, RRV takes into account the contribution of
an individual to future generations, relative to the
contributions of others
III. Natural selection favours the ones for which the sum of
current reproductive output and RRV is highest
• Reproductive value (RV) changes with age (Fig. 4.18)
• RV is low for young individuals when each of them has only a
low probability of surviving to reproductive maturity
• It increases steadily as the age of first reproduction is
approached
• Reproductive value is then low again for old individuals
Figure 4.18 Reproductive value
generally rises and then falls with
age, as explained in the text. (a)
The annual plant Phlox
drummondii (b) The sparrow-
hawk, Accipiter nisus, in southern
Scotland. Solid symbols refer to
breeders only; open symbols
include non-breeders).
2.3.5. Trade-offs
• Any organism’s life history must be a compromise allocation
of the resources that are available to it
• Usually resources devoted to one trait are unavailable to
others
• A ‘trade-off ’ is a negative relationship between two life
history characteristics in which increases in one are associated
with decreases in the other as a result of such compromises
• For instance, Douglas fir trees (Pseudotsuga menziesii) benefit
both from reproducing and from Growing but the more cones
they produce the less they grow (Figure 4.19a)
• Male fruit-flies benefit both from a long period of
reproductive activity and from a high frequency of matings,
but the higher their level of reproductive activity earlier in life
the sooner they die (Figure 4.19b).
• The longevity of male fruit-flies (Drosophila melanogaster)
generally increases with size (thorax length).
• However, longevity was reduced in males provided with one
virgin and seven mated females per day (1) compared with
those provided with eight mated famales (o), because of the
increase in courtship activity, and reduced still further in
males provided with eight virgins per day (8)
Figure 4.19 Life history trade-offs.
(a) The negative correlation between
cone crop size and annual growth
increment for a population of Douglas (b) The longevity of male fruit-flies
fir trees, Pseudotsuga menziesii. (Drosophila melanogaster) generally
increases with size (thorax length).
• If there is variation between individuals in the amount of
resource they have at their disposal, then there is likely to be
a positivecorrelation between two apparently alternative
processes
• some individuals will be good at everything, others
consistently awful.
• In Figure 4.20, the aspic vipers (Vipera aspis) in the best
condition produced larger litters but also recovered from
breeding more rapidly, ready to breed again.
Figure 4.20 Female aspic vipers (Vipera aspis) that produced larger litters also
recovered more rapidly from reproduction
2.3.4. The cost of reproduction
• Here, ‘cost’ is used in a particular way to indicate that an
individual, by increasing its current allocation to reproduction,
is likely to decrease its survival and/or its rate of growth, and
therefore decrease its potential for reproduction in future
• Thus, individuals that delay reproduction, or restrain their
reproduction to a level less than the maximum, may grow
faster, grow larger or have an increased quantity of resources
available for maintenance, storage and, ultimately, future
reproduction
• Figure 4.22 The cost of reproduction in ragwort . The line divides plants
that survive (0), from those that have died by the end of the season (+).
For a given size (rootstock volume), only those that have made the
smallest reproductive allocation (number of capitula) survive, although
larger plants are able to make a larger allocation and still survive
• Any ‘cost’ incurred by contemporary reproduction is likely to
contribute to a decrease in residual reproductive value (RRV).
• Natural selection favors the life history with the highest
available total reproductive value: the sum of two quantities,
one of which (contemporary reproductive output) tends to go
up as the other (RRV) goes down
2.3.7. The number and fitness of offspring
• A second key trade-off is that between the number of
offspring and their individual fitness which is at its simplest,
this is a trade-off between the size and number of offspring
• That is, a reproductive allocation can be divided into fewer,
larger offspring or more, smaller offspring
• Negative correlations have also been observed between egg
size and number of different species (e.g. Figure 4.23a, b)

(a) A negative correlation between
the number of propagules per stem
produced by six species of goldenrod
plants and the weight of single
propagules.
Figure 4.23 Evidence for a trade-off between
the number of offspring produced in a clutch
by a parent and the individual fitness of those
offspring.
(b) A negative correlation between clutch size
and egg volume amongst Hawaiian species of
Drosophila, cultured in different conditions
X = relatively poor and restricted resource
□ = on bacteria within decaying leaves
= = productive resources – yeasts
2.3.7. The number and fitness of offspring
• Sinervo (1990) manipulated the size of the eggs of an iguanid
lizard by removing yolk from them after they have been
produced, giving rise to healthy but smaller offspring than un-
manipulated eggs.
• These smaller hatchlings had a slower sprint speed probably an
indication of a reduced ability to avoid predators, and hence of a
lower fitness (Figure 4.23c).
• Within natural populations, this species produces smaller
clutches of larger eggs in California than in Washington
• the comparison between the two populations does indeed
appear to reflect a trade-off between the number of offspring
and their individual fitness.
Figure 4.23 (c) The mass and sprint speed of California hatchlings of the lizard,
are lower from eggs with some yolk removed (open circle) than from un-
manipulated control eggs (solid circle). Also shown are the means for the
control California hatchlings (CA) (fewer, larger eggs) and those for two
samples from Washington (WA).
2.4 Options sets, fitness contours and a classification of
habitats

• An options set describes the whole range of combinations of

two life history traits that an organism is capable of exhibiting
• The options set describes, for any given level of present
reproduction, the range of growth increments that the
organism can achieve, and for any given growth increment,
the range of levels of present reproduction that the organism
can achieve.
• The outer boundary of the options set represents the trade-
off curve. For any point on that boundary, the organism can
only increase mx by making a compensatory reduction in
growth and vice versa.
• An options set may be convex-outwards, implying in the
present reproduction only slightly affects growth (Figure
4.24a).
• Alternatively, the set may be concave-outwards implying that
substantial growth can only be achieved when the level of
reproduction is considerably less than the maximum (Figure
4.24b).
Figure 4.24 (a, b) Options sets – the combinations available to an organism of,
in this case, present reproduction and present growth. As explained in the
text, the outer boundary of the options set is a trade-off curve: (a) is convex-
outwards, (b) concave-outwards.
• A fitness contour is a line joining combinations of mx and
growth for which fitness (reproductive value) is constant
(Figure 4.24c).
• Contours further away from the origin represent
combinations with greater fitness.
• The shapes of fitness contours reflect not the organism’s
intrinsic properties but the habitat in which it lives.
(c) Fitness contours linking combinations of present reproduction
and present growth that have equal fitness in a given habitat.
Hence, contours further from the origin have greater fitness.
2.4.1. Habitats: a classification (118)
• In the case of reproductive fitness habitat is described and
classified from the point of view of the organism concerned,
rather than whether we feel the habitat is patchy or
homogeneous, harsh or benign.
• The shapes of fitness contours reflect an organism’s habitat.
The graphs reflect the effect of the habitat on that particular
organism or the response of that organism to the habitat.
• For established individuals (i.e. not newly emerged or newly
born offspring) two contrasting habitat types can be
recognized.
1. High CR (cost of reproduction) habitats
• High CR are habitats in which any reduced growth that results
from present reproduction has a significant negative effect on
RRV, and hence on fitness.
• Thus, similar fitness can be achieved by combining high
reproduction with low growth or low reproduction with high
growth. Fitness contours therefore run diagonally with a
negative slope (Figure 4.25a)
2. Low CR habitats,
• In Low CR, RRV is little affected by the level of present growth.
• Fitness is thus essentially determined by the level of present
reproduction alone and will be much the same whatever the
level of present growth.
• The fitness contours therefore run approximately vertically
(parallel to the ‘growth’ axis; Figure 4.25a)
Figure 4.25 A demographic classification of habitats. (a) Habitats of established
individuals can be either: (A) relatively high CR or (B) relatively low CR
• Habitats can be relatively high CR for at least two reasons.
1. When there is intense competition amongst established
individuals with only the best competitors surviving and
reproducing .
– Reproduction may be costly because it reduces growth
and hence substantially reduces competitive ability in the
future, and thus reduces RRV
2. Whenever diminutive adults are particularly susceptible to an
important source of mortality from a predator or some abiotic
factor.
– Reproduction may be costly because it maintains adults in
these vulnerable size classes.
• On the other hand, habitats can be relatively low CR for at least
three different reasons.
1. Much mortality may be indiscriminate and unavoidable, so
that any increase in size caused by reproductive restraint is
likely to be worthless in future.
2. The habitat may be so benign and competition-free for
established individuals that all of them have a high probability
of surviving, and a large future reproductive output,
irrespective of any present lack of reproductive restraint.
3. A habitat may be low CR simply because there are important
sources of mortality to which the largest individuals are
especially prone.
• A related classification of habitats for newly born offspring
can also be constructed.
• There are two contrasting types, assuming that larger
offspring can be produced only if there are fewer of them.
(Figure 4.25b)
1. ‘Offspring size-sensitive’ habitats, in which the
reproductive value of individual offspring rises
significantly with size
2. ‘Offspring size-insensitive’ habitats, in which the
reproductive value of individual offspring is little affected
by their size
Figure 4.25: (b) Habitats of recently produced offspring can be
either: (C) relatively offspring size-sensitive or (D) relatively
offspring size-insensitive.
Figure 4.26 (a) Options sets and fitness contours (see Figure
4.25) suggest that relatively high CR habitats should favor
relatively small reproductive allocations.
(b) The distribution of four biotypes (A–D) of the
dandelion Taraxacum officinale, amongst three
populations subject to low, medium and high levels of
disturbance (i.e. habitats ranging from relatively high CR to
relatively low CR).
(c) The reproductive allocations (RAs) of the different
biotypes from the different sites of origin, showing that
biotype A, which predominates in the relatively low CR
habitat, has a relatively large RA, and so on.
4.11 The size and number of offspring
• the division of a given reproductive allocation into a smaller
number of larger offspring is expected in relatively offspring
size-sensitive habitats
• In guppies offspring size was larger where predation was most
concentrated on the smaller juveniles (table 4.6)
• In Drosophila offspring size was larger in habitats where
competition was likely to be most intense (Fig. 4.23)

(a) A negative correlation between
the number of propagules per stem
produced by six species of goldenrod
plants and the weight of single
propagules.
Figure 4.23 Evidence for a trade-off between
the number of offspring produced in a clutch
by a parent and the individual fitness of those
offspring.
(b) A negative correlation between clutch size
and egg volume amongst Hawaiian species of
Drosophila, cultured in different conditions
X = relatively poor and restricted resource
□ = on bacteria within decaying leaves
= = productive resources – yeasts
4.11.1 The number of offspring: clutch size
• In this section we will try to answer the following question.
“How is it that particular clutch sizes, or particular sizes of
seed crop, have been favoured”
• Lack (1947) proposed that:
‘natural selection will favour not the largest clutch size but a
compromise’
• According to the proposal clutch size by balancing the number
produced against their subsequent survival, leads to the
maximum number surviving to maturity.
• Results of a number of studies suggested that Lack’s proposal
is wrong.
• Experimental increases in clutch size often lead to apparent
increases in productivity
• Nevertheless Lack’s has been important in directing ecologists
towards an understanding of clutch size
• A number of reasons for the lack of fit are now apparent and
two are particularly important
1. First, many of the studies are likely to have made an
inadequate assessment of the fitness of individual offspring
2. Cost of reproduction was not considered in the studies
1. inadequate assessment of the fitness of individual offspring
• It is important to look beyond the number of eggs. We should
consider:
How well do they survive the following winter?
How many chicks do they have themselves?
• For example, in Oxford (UK) great tits (Parus major)
– Addition of eggs to a normal clutch produced more cheeks
(10.96)
– The normal clutch size was 8.68
– Removal of eggs resulted in less number of cheeks (5.68)
• However recruitment was high from un-manuplated clutches
(Fig 4.29)
Figure 4.29 (a) The ‘Lack clutch size’. If the fitness of each individual
offspring decreases as total clutch size increases, then the total fitness
of a clutch (the product of number and individual fitness) must be
maximized at some intermediate (‘Lack’) clutch size.
(b) The mean observed number of
young recruited per nest ± SE
relative to experimental
manipulations (additions to, or
removals from, clutches) in great
tits.
(c) However, if there is also a cost
of reproduction, then the
‘optimum’ clutch size is that where
the net fitness is greatest,
2. Considering cost of reproduction
• Natural selection will favor a lifetime pattern of reproduction that gives rise to the
greatest fitness overall
• A large and apparently productive clutch may extract too high a price in terms of
RRV. A large and apparently productive clutch may extract too high a price in terms
of RRV.
• The favored clutch size will then be less than what appears to be the most
productive in the short term (Figure 4.29c).
– Females (Clethrionomys glareolus) were treated with gonadotropin hormones
– Treated females were considerably more productive at the time the litters
were born
– However, the treated females also paid a significant cost for their increased
reproductive efforts
– higher mortality during nursing, decreased body mass gain and a
decreased probability of producing a subsequent litter.
4.12 r and K selection
• This is the concept of r and K selection, originally propounded
by MacArthur and Wilson (1967; MacArthur, 1962) and
elaborated by Pianka (1970)
• The letter r refers to the intrinsic rate of natural increase
(above) and indicates that r-selected individuals have been
favored for their ability to reproduce rapidly (i.e. have a high r
value)
• The letter K will not be introduced properly until intraspecific
competition is discussed fully in the next chapter, but for now
we need note only that it refers to the size (‘carrying
capacity’) of a crowded population, limited by competition.
• The concept is therefore based on there being two contrasting
types of habitat: r-selecting and K-selecting.
• It originally emerged the contrast between species that were
good at rapidly and species that were good at maintaining
themselves on islands once many colonizers had reached
there (K species). colonizing relatively ‘empty’ islands (r
species).
• A K-selected population lives in a habitat that imposes few
random environmental fluctuations on it. A K-selected
population lives in a habitat that imposes few random
environmental fluctuations on it.
• By contrast, an r-selected population lives in a habitat that is
either unpredictable in time or short lived.