BIOL B242 Evolutionary Genetics: Coevolution Escape_and_radiate coevolution evolutionary innovation enables adaptive radiation, i.e. speciation due to availability of ecological opportunity. What is coevolution? Coevolution is: Evolution in two or more evolutionary entities brought Concordant and non_concordant phylogenies about by reciprocal selective effects between the entities. If the phylogenies are concordant (see overheads), this may from Ehrlich and Raven (1964): imply: "Butterflies and plants: a study in coevolution" Ø That cospeciation has occurred, or Ø That one of the groups (often the parasite) has Examples we have already encountered: "colonized" the other (the host). Here, the host shifts may well correspond to phylogeny because closely related Sex and recombination may have evolved because of: hosts are more similar. a coevolutionary arms race between organisms and In other cases, phylogenies may not be concordant, because their parasites the parasite may be able to switch between host lineages fairly frequently (see examples on overheads). sexual selection: potentially a coevolutionary phenomenon between Host/parasite and predator/prey coevolution female choice and male secondary sexual traits. Concordant phylogenies does not prove coevolution i.e. coevolution within a single species. We must look at individual adaptations of the exploiter and Here we deal with interspecific coevolution only. the exploited

Coevolution might occur in any interspecific interaction. Examples: Defences of plants against herbivores For example:Interspecific competition for food or space "Secondary chemistry" Ø Parasite/host interactions e.g. tannins and other phenolic compounds, alkaloids Ø Predator/prey interactions like nicotine and THC, or cyanogenic glycosides Ø Symbiosis Often toxic Ø Mutualisms Animals, such as insects, have obviously adapted to Mimicry, for example potentially coevolutionary, can be: feeding on plants parasite/host interaction (in Batesian mimicry) If plants have evolved defensive chemistry, ? or mutualism (Müllerian mimicry). plant/insect coevolution. Argument of Ehrlich & Raven Batesian mimicry however, model finds it hard to escape its parasite. Critics argue that: So coevolutionary chase unlikely. Ø phytophagous insects are usually rare, and therefore do not pose a threat to their host plants Müllerian mimicry Ø secondary chemistry may be a byproduct of normal the more abundant and noxious species trapped by its metabolic processes, rather than necessarily defensive own pattern; the rarer or less noxious species gains by mimicking Evidence for insect/plant coevolution the more common or noxious species, Thus: mutual convergence is unlikely. Central American plant "bullshorn Acacia" Ø Large spines against mammalian herbivores Types of coevolution Ø Lacks cyanogenic glycosides. "How likely is coevolution?" Ø Thorns are particularly large, hollow, provide shelter to a depends what you mean by “coevolution”! species of Pseudomyrmex ant Ø Plant also provides proteinaceous food bodies (Müllerian Various types: bodies) on the tips of the leaflets; ants eat Specific coevolution = coevolution (narrow sense) Ø Ants are nasty! Defend against caterpillars, even One species interacts closely with another mammals and plant (vines) competitors. Changes in one species induce changes in the other Ø Plants not occupied by ants are heavily attacked. Either polygenic or gene_for_gene coevolution.

Concordant speciation or cospeciation Speciation in one form causes speciation in another Cospeciation doesn't necessarily require coevolution

Diffuse coevolution = guild coevolution Groups of species interact in non-pairwise fashion e.g. Group of plant species may be fed on by a particular family of insects c.f. Ehrlich & Raven’s original idea

1 Related Acacia species Ecological release (the reverse of Gause’s principle) Ø Lack hollow thorns, food bodies Ø Have no specific associations with ants Ø A species colonizes area where no competitors Ø Many cyanogenic glycosides in their leaves Ø May experience ecological release Ø Grows to very large population sizes v bullshorn Acacia has evolved a close, mutualistic Ø Disruptive selection to evolve apart association with the ants to protect from herbivores (and plant competitors). v Adaptive radiation v cyanogenic glycosides that are found in other species have a defensive role; a role which has been taken Often on islands: over by Pseudomyrmex. e.g. Darwin's finches of the Galapagos islands e.g. Hawaiian honeycreepers Egg_mimicry in Passiflora Defenses of cyanogenic glycosides, alkaloids Sometimes on “ecological islands” breached by Heliconius e.g. lakes in the North temperate zone in last 10,000 years

Predator_prey coevolution Sticklebacks (Gasterosteus) in Canada benthic (deep water) and limnetic (shallow water) forms Predator offensive evolution keep to their own habitat, mate assortatively e.g. Mammalian predators must be fast, strong, cunning Trout family Prey defence Ø Atlantic char (Salvelinus), Thingvallavatn, Iceland Ø Large size and strength Ø FOUR different trophic forms Ø Protective coverings such as shells or hard bony plates Ø Defensive weapons, such as stings or horns Cichlids in African Lakes Ø Defensive coloration (see mimicry lecture) Ø 300 spp. in the last 12,400 years in Lake Victoria Ø Unpalatability and nastiness Ø Partly sexual selection, partly ecological divergence

v examples of coevolution Adaptations leading to ecological release; "escape and radiate" coevolution As we have seen, mimics may not coevolve with their models, but do coevolve with their predators Possession of a unique adaptation Highly coevolved pollination systems may also allow adaptive radiation

Yucca and Yucca moths (Tegeticula) Ø Brian Farrell: herbivory on flowering plants Figs and figwasps v massive amounts of speciation (overhead). · Larvae are seed/flower eaters · Plant is dependent on herbivore for pollination Ø Resin_ or latex_bearing canals in plants Ø Latex and resin is a physical defence against herbivorous v Tightly coevolved mutualism insects.

Yucca and Yucca moths v more rapid speciation rate

· Sometimes the mutualism has breaks down Conclusions · Moth reverts to a parasitism; does not pollinate. Coevolution v In fig wasps, and most Yucca moths, these mutualisms have become very specific, and essential An area where genetics, ecology, phylogeny interact to both species. (A general theme we have stressed in this course!)

Similar to ancient prokaryotic mutualisms: Majority of diversity of life and life forms Mitochondria & chloroplasts with archaebacterial cells not just due to adaptation to static environments producing eukaryotes instead, due to biotic interactions

Coevolutionary competitive interactions and adaptive Biotic environment itself constantly evolving radiation v orders of magnitude more diversity than by simple, static “Gause’s principle” adaptations If two species have identical resources Ø competitive exclusion

v less well adapted species will go extinct

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