Phylogenies
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Behavioral Ecology-1
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Phylogenies
Phylogenetic inference: the process of reconstructing the evolutionary history
of species, genes or other biological entities by grouping them based on shared
ancestry. The results of phylogenetic inference are phylogenetic trees, which
are diagrams that show the evolutionary relationships between organisms.
Most phylogenetic inferences have been depicted in the form of hierarchical
bifurcating trees, meaning trees that reflect a series of branching processes in
which one lineage splits into two descendant lineages.
These trees can be based on morphological traits but are much more often
inferred from genetic characters. The positioning of taxa on a tree is generally
based on their genetic similarity to one another.
There are many different ways in which phylogenies can be reconstructed from
genetic data, but most of them fall into one of four categories: distance,
parsimony, maximum likelihood and Bayesian methods. The first two are
becoming nearly obsolete.
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Phylogenies
Distance methods are based on measures of evolutionary differences between all pairs of taxa
(Figure 6.6). They can be calculated from the number of nucleotide differences (DNA data),
allele frequency data (microsatellites, SNPs) and amino acid differences (protein data). The
goal is to build a tree that accurately reflects how much genetic change has occurred – and
therefore roughly how much time has passed – since lineages split from one other.
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Behavioral Ecology
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Molecular genetics and the study of behavior
Behavioral ecology includes several areas of research:
1. Mate choice
2. Brood parasitism
3. Co-operative breeding
4. Foraging behavior
5. Dispersal
6. Territoriality
7. Manipulation of offspring sex ratios
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Molecular genetics and the study of behavior
Behavioral ecology is traditionally based on either laboratory-based research or
fieldwork.
a) Laboratory-based research: limited because many species cannot be kept in
captivity or captive conditions cannot exactly mimic those in the wild.
b) Observations and experiments in the field involving wild populations have
also limitations; for example, it may not be possible to identify individuals or
to follow and observe them for prolonged periods.
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Molecular genetics and the study of behavior
Behavioral ecology is traditionally based on either laboratory-based research or
fieldwork.
a) Laboratory-based research: limited because many species cannot be kept in
captivity or captive conditions cannot exactly mimic those in the wild.
b) Observations and experiments in the field involving wild populations have
also limitations; for example, it may not be possible to identify individuals or
to follow and observe them for prolonged periods.
Molecular data are increasingly used to supplement the more traditional
approaches, particularly when studying individuals in the wild.
A key advantage of molecular approaches is that small amounts of blood, hair,
feathers or feces are required for genotyping.
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Mating systems
Monogamy: very rare in most taxonomic groups with the notable exception of bird species, 90% of
which are monogamous. Biparental care may partially explain why social monogamy is so common in
birds.
Polygyny may evolve when offspring can survive without paternal care and males can attract multiple
mates. In many species this occurs when resources such as food are patchily distributed, because males
can then defend high-quality territories that will each attract multiple females.
Polyandry: rare, females compete for and defend territories to
which they attract multiple males. Females tend to be larger
and more colorful than males. Males will then perform most of
the parental care. The American jacana (Jacana spinosa) is a
well-studied example.
In the range of this species, suitable nest sites are scarce and
predation is high. There is an advantage for female jacanas to
simultaneously lay multiple clutches because it is more likely
that a proportion of these survive, increasing the female
fitness.
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Mating systems
Polygynandry: situation in which two or more males within a group are socially
bonded with two or more females.
Promiscuity: very common in mammals, documented in more than 130
mammalian species. Notice though that there are ~5,000 mammalian species..
Promiscuity can have high fitness benefits to males if they can fertilize multiple
females.
Females may also benefit from promiscuous mating due to the increased
offspring survival when females mated with multiple males.
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Mating systems and parentage analysis
Parentage analyses through genetic data allow to identify the genetic relationships
of offspring and their putative parents, which is key to understanding of mating
systems.
A major finding of these studies is that social mating system can be very different
from a genetic mating system.
Biological fathers may be harder to identify because they may offer no parental care
(most mammals) or may unknowingly care for young that are not their own (many
birds).
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Extra-pair fertilizations
Parentage studies can reveal patterns of promiscuity that differ from expectations
given the mating systems.
Sometimes, species considered promiscuous are less so. For example, in the
Arctic ground squirrels (Spermophilus parryii plesius) genetic data have shown
that more than 90% of the pups whose mothers mated with more than one male
were fathered by her first mate.
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Extra-pair fertilizations
Parentage studies can reveal patterns of promiscuity that differ from expectations
given the mating systems.
Sometimes, species considered promiscuous are less so. For example, in the
Arctic ground squirrels (Spermophilus parryii plesius) genetic data have shown
that more than 90% of the pups whose mothers mated with more than one male
were fathered by her first mate.
More often genetic data show that males and females are more promiscuous
than their social mating systems would suggest.
This is known as extra-pair fertilizations (EPFs), which occurs when individuals
choose mates that are not their social partners.
In fact, in the majority of bird species tested — data are available for more than
200 species — a substantial proportion (sometimes exceeding 50%) of offspring
are sired by a male other than that providing parental care.
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Evolutionary repercussions of EPFs
1. The male’s fitness may be unrelated to the number of offspring that he rears.
2. Males may hedge their bets and provide parental care in proportion to their
confidence in paternity.
3. Even in socially monogamous species, males do not have to form pair bonds in order
to achieve reproductive success.
4. EPFs increases variance in male reproductive success, which can have a substantial
influence on effective population size.
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Extra-pair fertilizations and population size (Ne)
We know that variance in reproductive success (VRS) can influence the effective size of a
population (Ne).
“Variation in reproductive success (VRS): variation in numbers of viable offspring
produced throughout an individual’s lifetime.
Higher VRS will decrease Ne relative to Nc.”
EPFs may either:
1. Decrease VRS by enabling unpaired males to reproduce
2. Increase VRS by allowing a handful of males to father a disproportionately high number
of offspring.
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Mate choice
Mate choice may be exercised by both males and females, but in general females are
choosier than males because they usually invest more in eggs than males do in
sperm.
Two hypotheses that may explain mate choice, and which have particularly benefited
from molecular data:
1. The good genes hypothesis.
2. The genetic compatibility hypothesis.
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Mate choice
The good genes hypothesis states that mates will be chosen on the basis of some
characteristic that will always confer high fitness values on offspring. Examples: Atlantic
salmon, lek-breeding tree frogs, and probably many others.
The genetic compatibility hypothesis is based on the idea that a particular paternal allele
will increase the fitness of offspring only when it is partnered with a particular maternal
allele. Individuals will choose their mate on the basis of their combined genotypes.
Female mice (Mus musculus) and female sand lizards (Lacerta agilis), for example, tend to
choose mates whose MHC loci are as dissimilar to theirs as possible, to either increase
heterozygosity at the MHC in particular, or to decrease inbreeding in general. Evidence on
the role that MHC compatibility may play in human mate choice is equivocal, although in
some human populations mate choice is more likely among MHC dissimilar partners.
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Mate choice
Both the good genes hypothesis and the genetic compatibility hypothesis require
genetically ‘appropriate’ mates within the population, which should be more likely in
populations with high genetic diversity.
This has led to the hypothesis that extra-pair paternity should be higher in populations
with high genetic diversity. There has been some support for this hypothesis, for example
extra-pair paternity was very low (only 2% of offspring were extra-pair) in a population of
white-throated dippers (Cinclus cinclus), which also had low genetic diversity.
However, the authors of that study also point out that without some consensus on how
genetic diversity should be measured and compared among populations and species, it
will be difficult to quantify a more general relationship between genetic diversity and
frequency of EPFs.
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