Week 12: Altruism

Darwin’s solution to fatal problem: Darwin suggested that selection operated on parents, and those parents who produced more effective worker offspring in
the first instance, were likely to have, later on, greater numbers of reproductive offspring

For Cooperative Brood Care, Reproductive Castes, Overlap of Generations

Solitary ( - - - )
Quasisocial ( + - - ) Example: individuals same generation, they cooperate in rearing brood, but all lay eggs
Semisocial (+ + - )
Eusocial ( + + + ) Example: : individuals from > 1 generation, with queen and worker castes
Kin Selection: Hamilton’s Rule - b*r - c > 0
Mathematical Definition of Relatedness: the probability that two individuals share the same genes by common descent from a recent ancestor (like parents,
grandparents, etc). Conceptually, relatedness is a measure of the genetic similarity between individuals: as the value moves from 0 to 1 relatedness is higher.

Degree of relatedness in diploid and haplodiploid

Relationship Diploid Haplodiploid

Sister and sister 1/2 3/4

Mother daughter 1/2 1/2

Sister brother 1/2 1/4

In a haplodiploid species - the inclusive fitness of female workers is highest if they help produce more sisters - genetic return for rearing sisters is greater than
for rearing their own offspring

Cooperative breeding in Seychelles Warbler to make sure there was a territory to live in

Reciprocal altruism in Ficedula hypoleuca

In Ficedula hypoleuca, neighbours can jointly ‘mob’ potential predators, or else they can let their neighbour do the mobbing and avoid the risk of harm. If
reciprocal altruism occurs, then birds should join in mobbing if their neighbour helped out previously, but not join mobbing if their neighbour did not help in
the past
 Repeated interactions
 Ability to recognize individuals from previous encounters
 Ability to remember what those individuals did in previous encounters - Well developed cognitive and memory abilities
 Generally, this will require animals that are longlived

We can now say that Kin Selection will tend to maximize the Inclusive Fitness of altruistic individuals, and… ….that in some populations Darwinian selection
(direct fitness) is a subset of Kin Selection (which considers both direct and indirect fitness)

Group Selection Theory

Put forward more formally by V.C. Wynn-Edwards in 1960’s, who worked on bird ecology.
Wynn-Edwards was struck by some island nesting birds that seemed to altruistically hold back on reproduction.
Although there may be enough physical space on islands for all birds to reproduce, surrounding fish stocks are not sufficient for all pairs to lay eggs •If all
birds reproduce, the population will grow beyond the carrying capacity. •Then the entire population will collapse. •But populations that ‘constrained’ their
reproduction would survive •Eventually, the surviving populations (who constrained their reproduction) would be able to colonize the now-empty islands of
the failed populations
Flawed because no account of time scales and migration (what if selfish individual enters?)

Inbreeding facilitates eusociality… Australian gall forming thrips

single foundress (queen) who produces most of the brood. • gall is defended by soldiers (male and female) who are either sterile or have low reproductive
output. • New foundresses mate with their brothers before dispersing from the gall, leading to high inbreeding. (highest known for any insect) • Some
soldiers reproduce (brother and sisters) and lay offspring in the maternal gall as they are unable to disperse. • Why would the foundress allow soldiers to
reproduce?

Why do birds spend 1-3 years helping? - gaining experience, access to resources
Week 10: Community Ecology
Disturbances can eliminate organisms and alter resource availability -
moderate to high species richness may improve stability and have higher
biomass and are better at recovering from environmental stress and more
resistance to introduced species
Competitive exclusion principle as two species with exactly the same niche
are not able to coexist
Resource partitioning - realised niche vs fundamental niche (tree length
example)
Character displacement - Darwin’s finches in sympatric have exaggerated
extremes in beak depth but allopatric areas don’t
Competition -/- Exploitation (predation, parasitism, herbivory) +/- Positive
(commensalism, mutualism) +/0 and +/+
Exploitation: Social parasitism - brood parasites laying eggs in nests of
other species, hymenopteran families in ants, bee social parasites with
females inhabiting nest of other host bees and relying on the resources
from host to the nest to survive and rear.
Emery’s rule - social parasites among insects tend to be closely related to
their host species
Exploitative (speed and ability to acquire resource) and interference
competition (tall tree shading neighbours or one species battling another
or preventing access to resource)
Energetic Hypothesis: Length of food chain is limited by inefficiency of
energy transfer between each tropic level (~10% transferred between
levels
Bottom up vs top down control
Species of impact: foundation species (removal will impact community
structure, often provide habitat or food supply), examples: (a) corals, (b)
mangroves, (c) giant kelp, and (d) freshwater macroalgae. Keystone species
(exerting influence on community that isn’t reflected in its abundance).
Ecosystem Engineer (magnetic termites in the NT construct mounds to
transform nutrient-poor soils into nutrient rich islands)
Equilibrium Model - Climax Community vs Nonequilibrium Model
Intermediate Disturbance Hypothesis
Ecological Succession ( Process of change in the species structure of an
ecological community over time following major or minor
disturbance/environmental change) - Primary or Secondary
Geographical impacts on community - evapotranspiration, area effect:
larger area = more species (habitat diversity)
Island equilibrium Model (MacArthur and Wilson) Equilibrium number of nor helps the other.
species (Q), rates of immigration and extinction influenced by • distance Conjunctive symbiosis – relationship where the two symbionts form a single
from mainland • size of the island
Week 11: Coevolution
Moth and orchid - either longer spurs selected for optimal contact with
pollinator body or pollinator shift as alternate hypothesis for long nectary in
other species
Coevolution - reciprocally induced evolutionary change - the mutual
evolutionary influence between two species, each species causes a change in
the frequency of alleles (genetic makeup) of the other species
Principles of coevolution - Frequent interactions • Impact on Reproductive
Success • Relative Evolutionary Potential
examples of interspecies interactions that might provide the selection
pressure for coevolution: Plant- Herbivore • Predator-Prey • Parasite-Host •
Mutualism
Interspecies interactions as evidence for coevolution: Cospeciation – two
species lineages diverge in a similar phylogenetic pattern (not proof alone but
supports)
Preliminary evidence: individual adaptations of the interacting species such as
Some flowers have evolved to
actually resemble bees and Insect Herbivores
PLANT DEFENSES - examples • Physiology -Toxins – Secondary compounds –
i.e. tannins or toxic alkaloids • Morphology – seed shells, spines, thorns and
stiff hairs made of tough material, Brambles
HERBIVORE RESPONSES - examples • Diet Shift – become generalised feeder –
eat a variety of plants • Specialised physiology or morphology – combat toxins
or breakdown toughened cuticle (gut microflora), changes to teeth/jaw
structure
Example: Figs and Fig wasps: 500 species of figs (Ficus) pollinated by
Agaonidae (Chalcidoidae) wasp • Figs have Unique entry for wasps •
Phenology of female flower receptivity and pollen production synchronise
with wasp life cycle • Seeds at different heights (limited availability). • Wasp
adaptations include Morphological Adaptation in female • Load and release
pollen behaviours • Specific between fig species and wasp species
Coevolution is more likely for predators that concentrate mainly on one or a
few species of prey and prey that have only a few predators
Encounter avoidance AND Reduce Detectability
Thwart Predator Approach AND Subjugation – prevent injury AND Aposematic
Colouration (Warning Colouration)
Batesian vs Mullerian mimicry
Darwinian medicine - manipulating the evolution of pathogens • altering
selection for host resistance • using pathogens as biological control agents.
Intraspecies coevolution - sexual selection driving changes in secondary sexual
characteristics – As Male peacock feathers – Female preferences for mates
change
Arms race not purposeful or predetermined (Common among many
plant/herbivore systems and host/parasite)
Geographic mosaic theory of coevolution
Guild (diffuse) multiple to multiple - Specific (Pairwise) – Polygenic AND
Escape and Radiate – example: plant-herbivore
Commensalism – relationship that benefits one symbiont but neither harms
entity.
Disjunctive symbiosis – relationship where there is no physical union between
the two symbionts.
Endoparasite – a parasite that lives inside the body of its host.
Ectoparasite – a parasite that lives on the outer surface of the host.
Facultative symbiosis – relationship where the symbionts are able to survive
independently.
Mutualism – relationship where the interspecific interaction benefits both
symbionts.
Obligatory symbiosis – relationship where one or both symbionts are entirely
dependent on each other for survival.
Parasitism – relationship where one symbiont (parasite) derives resources
from the other (host).
1. A alga–fungus symbiosis (lichen) Mutualism
2. An epiphyte (Platycerium sp. stag horn) commensalism
3. A mistle-toe (parasite)
4. Rhizobium root nodules of a legume (mutualism)
5. Rust fungus (Melamsora lini) on a flax plant (parasite)
Week 9: Life History Strategies
Hermaphroditism - individuals produce both egg and sperm but don’t usually self fertilise but reproduce by exchanging sperm (snails, flatworms,
earthworms)
Sequential hermaphrodites - First one sex then the other (clownfish)
External Fertilisation
Often synchronised by temperature, day length and hormones. Fertilisation is ensured by spawning next to mate, whereas in internal fertilisation it is
copulation and transferring sperm (copulation requires the right timing, environmental cues, pheromones, courtship behaviour)
Some animals use BOTH sexual and asexual reproduction like aphids (depending on summer, fall, winter and spring)
Maximising Survival and Reproduction
time, energy and nutrients are limited so balanced strategy to maximise chances of successful reproduction needed (trade offs)
what affects strategies of maximising successful reproduction chances? longevity and age of first reproduction, number of breeding attempts, number of
offspring produced, level of parental investment, selection favours traits that improve survival and reproduction BUT→maximising one can decrease the
other
Timing of Reproduction
Semelparity - meaning→one reproductive event in lifetime (favoured by highly variable and unpredictable environment) examples (Antechinus stuartii and
salmon)
Iteroparity – many reproductive events in lifetime example→Bristlecone pines surviving thousands of years and reproducing year after year
issues with strategy: total lifespan, age at which reproduction first begins, effort in each reproductive episode, effort needed to bring an offspring to maturity,
effect of reproduction on subsequent survival, quality of offspring in any one episode
r-selected strategy components↓small, short lived, semelparity, many small offspring, little parental investment, high offspring mortality
k-selected strategy components: large, long lived, iteroparity, few large offspring, high parental investment, high offspring survival, based off of what→logistic
growth
r-selected strategies description: density-independent selection, maximises r, the per capita rate of increase, suited to life in an unstable environment,
reproduce early and abundantly and allow for a low survival rate of offspring, example→locusts
k-selected strategies description: density-dependent selection due to high competition for limited resources, few offspring but high investment in each
(examples: mammals with stable population sizes, forest trees, many birds), additional element→parental protection (in plants→usually none but
occasionally sheltering; in animals none = egg scattering / spawning, moderate = selective egg deposition; and high = extended egg incubation)
Unusual Life-History Strategies
sex reversal - examples↓wrasse (large males have access to females and if largest male in the group dies→next largest female changes to male) and oyster
(small male adult can produce many sperm but only large adult female can produce many eggs and they reproduce early as males but then change to female
when they become bigger)
sex role reversal: sea horses (males invest more than females and become pregnant)
paternal care in giant water bug as female lays eggs on top of male and males aerate and protect the eggs
maternal sacrifice as hump earwig (anechura harmandi) eggs are laid during the early spring when food is scarce and mothers are then consumed by their
first instar nymphs (phenomenon is called→matriphagy)
Evolution of Clutch Size
'Lack clutch' after biologist David Lack prediction→parents should produce clutch sizes that maximise the total number of offspring produced from that clutch
food limitation hypothesis→if too many chicks then parents not able to feed all and some will die
testing hypothesis showed after VanderWerf (1992) meta-analysis that there is no real evidence for it
number of fledglings may not be the only important parameter, what else is important? - survival rates of adults to next breeding opportunity, fecundity of
adults at their next breeding attempt, survival of fledglings to reproductive maturity, fecundity of the offspring when they come mature,
these factors studied in→Ficedula albicollis - results from enlarged clutch sizes showed ↓
(i) juveniles were less likely to survive to following season (ii) fecundities of mothers were reduced in subsequent year (iii) daughters had lower fecundities,
BUT (iv) survival rates of parents weren’t affected, All of these mean that the food limitation hypothesis is not sufficient to predict optimal clutch sizes
Ecology
Scale of Ecology - Organismal, Population, Community, Ecosystem, landscape (biome), global (biosphere)
Where do organisms live?
depends on two kinds of factors↓biotic (competition (intra and interspecific), predation, beneficial interactions with other species) abiotic (sunlight,
[determined by solar radiation intensity: efficiency of photosynthesis, availability of liquid water] water , organisms in tidal zone can dry out quickly,
some organisms are adapted to tolerate limited water, example of plant→desert plants that have high desiccation tolerance and water storage and reduced
leaf surface area, example of animal→Spinifex hopping mouse, doesn't need liquid water; acquires it all from food
plants and herbivore case study results→more large mammal herbivores when moisture is intermediate and plant available nutrients are high

Temperature - cells may freeze and rupture below 0° C while most proteins denature above 45° C
two temperature strategies in animals (ectotherm - they require less energy to maintain body temperature and they use behavioural methods to maintain
correct body temperature - what could potentially kill them?↓ short periods of low or high temperature or long periods of moderately low temperature and
endotherm Comparatively better able to maintain a constant body temperature than ectotherms. Thermoneutral zone: consumes energy at a basal rate,
Environmental temperature outside thermoneutral zone→more energy consumed, Consumes energy faster than ectotherm of similar body size
what characteristics limit plant distribution? physical structure, pH, mineral composition (example→salt licks or clays and dirt or sodium)
Determining an Organism's Distribution Flowchart
Why is species X absent? limited dispersal yes?→area inaccessible or insufficient time no? limited behaviour yes?→habitat selection no? biotic factors yes?
→predation, parasitism, competition, disease no? abiotic factors
Population Ecology definition→The study of the dynamics of populations & how populations interact with the environment. Environmental influences on
density and distribution, age structure, and population size
what is a population?→Group of individuals of the same species living and interbreeding in the same area. A species consists of one, a few, or many
populations.
Types of populations - Panmictic description→all individuals are potential partners as there is no isolation of any aspect of the population (how much gene
flow?→maximum)
Fragmented - description→populations break into fragments with reduced gene flow, due to?→habitat fragmentation (humans !! but not always), can lead
to→extinctions if fragments are too small
Relictual - description→relicts from a previous broad distribution. Change happens due to geological and climate reasons, example→Spanish Juniper - broad
distribution during glacial period Pleistocene but now only found on mountain tops
Meta-populations
definition→Groups of semi-independent populations linked by immigration and emigration that occupy discrete patches of suitable habitats among a sea of
unsuitable ones - imposes genetic structure on population
what happens if a population goes extinct?→patch is recolonised
dynamics influenced by habitat size, quality and heterogeneity, spatial arrangement of individuals, asynchronous population dynamics, individuals from a
source population spill out into sink populations
Populations on the move
Migration - definition→cyclical movement of a population, from one place to another then back again — target destination!
Whole population moves, impact→important ecological consequences! Population takes biological activities with it!
patterns of migration, Long temporal cycles: seasonal movements OR Short temporal cycles such as diurnal and tidal
OR long distance such as→yearly migration between hemispheres
Probability of return - Multiple return: some birds, One return: some fish, One way only: some insects
Dispersal - definition→the movement of individuals away from each other (ex. high density to low density / away from parents / from birth location to
reproducing area)
ways of dispersing - major mechanisms: Passive versus active dispersal, Directional versus haphazard dispersal, Combined strategy, example→clam larvae
drift in current (haphazard) but seasonal release of larvae to prevailing current direction (directional)
Week 7: Genetics and Selection Week 7: Genetics and Selection
Genes made up of DNA, many genes code for proteins while others will have Mutation:
other functions (controlling expression of other genes. Some genes have no Mutations are changes in the nucleotide sequence of DNA. They can cause
function that we know of and some are never expressed. new genes and alleles to arise.
Humans have from hundreds to over 2M bases. And between 20,000 and
25,000 genes. Point mutations • Change in one base pair in a gene (substitution,
insertion or deletion) Can have a significant impact on phenotype &
Natural selection: fitness.
If all individuals reproduced successfully, populations would increase Large scale mutations: chromosomal mutations affect many loci.
exponentially
But population size usually stable Mutation rates • Tend to be low in animals and plants • Average about
• Resources are limited -> this will lead to competition over resources one mutation in every 100,000 genes per generation • Are most rapid in
• Natural enemies viruses
• Not all individuals will survive or reproduce
• Only SOME will! Sexual Recombination:
Individuals vary in their heritable traits. Sexually reproducing organisms have great scope to produce genetic
Due to this differential survival and reproductive potential populations will variation in their offspring
change over time
More individuals with favourable traits. • Reshuffling of genes during the production of eggs & sperm during
MEIOSIS • Combining genes of two individuals (mother & father)
Selected for and against: Must reduce number of chromosomes
• A trait is referred to as being ‘selected for’ if it is likely to increase in • From diploid to haploid (reduction by 50%) • Fusion of male and female
frequency in the population over time gamete will return the zygote to diploid state.
• A trait is referred to as being ‘selected against’ if it is likely to decrease in
frequency in the population over time Selection maintaining diversity: heterozygote advantage (anemia and
malaria), frequency-dependent selection (flies and bee), alternate
Adaptation and Fitness: selection pressures (green, red pea aphid eaten by wasp and visual
• An adaptation is a heritable trait that increases an individual’s survival and ladybird)
ability to reproduce
• Fitness is the contribution everyone makes to the gene pool (= number of ** Neutral allele theory - redundancy as 64 codon sequences but only 23
offspring) relative to other individuals amino acids so 3rd position in codon often no effect on amino acid
• Those with more offspring have higher fitness outcome. Thus, some nucleotide substitutions don’t affect protein
• Individuals are not altruistic and are not concerned about species function - invisible to selection.
• Individuals are concerned about own survival and reproduction – they are
‘selfish’
INDIVIDUALS DO NOT EVLOVE ONLY POPULATION Week 8: Concept of Species and Speciation
Mendel’s Genetic Idea:
Law of segregation of genes (“First law”): Morphological species concept
• Every individual organism contains two alleles for each trait, and that these
alleles segregate (separate) during meiosis such that each gamete contains
only one of the alleles. based on looking similar in structure and appearance. However, problem
• Gamete = sperm or egg is intraspecific genetic polymorphisms like Batesian mimicry in African
• Each pea in the F1 has inherited one ‘purple’ allele and one ‘white’ allele
from its parents swallowtail (papilio dardanus). Another problem is plasticity (genotype
Law of dominance (“Third law”): producing different phenotypes) such as social insect castes and seasonal
If the two alleles at a locus differ:
• Then one, the dominant allele, determines the organism’s appearance in Bicyclus wing patterns relative to risk of predation. Another issue is
(phenotype)
• The other allele, the recessive allele, has no noticeable effect on the sexual dimorphism such as in anglerfish and Australian quacking frog
organism’s phenotype Crinia georgiana. Another issue is developmental differences such as larva
to adult. Another issue is cryptic species such as trapdoor spiders where
females are indistinguishable and male (short-lived, seasonal, rare)
genitalia is the only way to tell species apart; also giraffes

Biological Species Concept

reproductive isolation with independent genetic system - all individuals

Incomplete dominance Phenotype of the F1 generation is somewhat a mixture
between the phenotype of each parent
Codominance - If two dominant alleles produce a separate phenotype.
Frequency of dominant genes • Dominant genes are not always more frequent
in the population than recessive genes
Law of independent assortment (“Second law”): For two or more traits
Each pair of alleles segregates independently during gamete formation.
drawn from a common gene pool with species evolving as a unit.
Prezygotic and postzygotic barriers do not AIM to create new species but
speciation can be as a result of these mechanisms. Prezygotic barriers:
habitat isolation, temporal isolation, behavioural isolation, mechanical
isolation (coupling role reversal), gametic isolation. Postzygotic barriers:
reduced hybrid viability, reduced hybrid fertility, hybrid breakdown
(second generation sterile)

** conceptual problems: sibling species, ring species, asexual species

(rotifers)

** functional/operational definition: morphological SC, paleontological SC,

Calculating genotype frequencies Hardy Weinberg Law ecological SC useful for polymorphic species, phylogenetic SC.
2 2
AA= p , Aa=2 pq and aa=q ** Speciation Processes - two basic patterns of evolutionary change -
If two alleles ‘A’ and ‘a’ occur with frequencies p and q, THEN genotypes
will occur in the frequencies of and allele frequencies will not change over anagenesis and cladogenesis. Cladogenesis can occur in 3 ways: allopatric
time speciation, sympatric speciation (three mechanisms: polyploidy, new
Main assumptions that we’ll keep in mind:
1. No selection on genotypes – Selection reduces genetic diversity ecological niches [lake Malawi] and sexual selection [assortive mating])
2. Random mating
3. Populations are very large
** evidence for sympatric speciation: social parasites in ants suggested to
Genes:
Gene pool: Is the total aggregate of genes in a population at any one time. be sister groups to their hosts by phylogenetic studies (temporal barrier to
Consists of all gene loci in all individuals of the population. gene flow between cheats and non-cheats). Rhagoletis mating on host
Genetic Variation: Genetic variation in populations contribute to evolution.
Two major processes produce variation in gene pools that contributes to plants but mating arenas changed when apples were introduced as hosts.
differences among individuals.
Mutation and Sexual recombination. Conflicting evidence for social parasites and a single judicious extinction
event could have made these allodapine bees look like evidence for
sympatric speciation. Parapatric speciation