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Speciation Walter Salzburger: Charles R. Darwin (1809-1882) What is a species? Bombina bombina Bombina variegata What is a species? What is a species? ‣ There is no agreement on how species should be defined... ‣ Species are fundamental (real) natural units ‣ Species are independent evolutionary units. Members of the same species share a gene pool. Mutation, drift and selection operate within species. frequency ‣ Taxonomists define species by means of morphological or phenetic characters Variation within/between Species: ‣ Individuals within a species are variable. There is no “ideal” or “typical” individual ? trait time What is a species? What is a species? ‣ There is no agreement on how species should be defined... For example, the Blue Tit (Parus caeruleus) is a paraphyletic species. The North African subspecies P. c. degener and P. c. ultramarinus are the sister group to the European Blue Tit (P. c. caeruleus) plus the Eurasian Azul Tit (P. cyanus) with four subspecies (P. c. cyanus, flavipectus, tianshanicus, and yaniseensis) Salzburger et al. (2002) The evolution of part of the original species into a new one renders the remaining populations paraphyletic. P. ca Eu erul rop eus e Two species appear to be present at one place, but those two “species” are connected by a series of forms that are geographically arranged in a ring. No phenetic character could be used, except arbitrarily, to divide the ring into two species. A division would be meaningless, as there really is a continuum, not a number of clear-cut, separate species. P. c Eu yanu ras s ia Paraphyletic Species: Stebbins (1994) Ring Species: P. ca N- erul Af eus ric a ‣ There is no agreement on how species should be defined... Biological Species Concept (BSC) Biological Species Concept (BSC) ‣ “Species are groups of interbreeding natural populations that are reproductively isolated from other such groups” (Mayr 1963) ‣ “Species are groups of interbreeding natural populations that are reproductively isolated from other such groups” (Mayr 1963) ‣ reproductive isolation: no interbreeding with members of other species ‣ The BSC places the taxonomy of a species within the framework of population genetics ♥ ♥ Biological Species Concept (BSC) ‣ problems: ‣ intermediate stages (“young species”, ring species) ‣ BSC fails to deal with low levels of gene flow between taxa (e.g., introgression) ‣ asexual organisms or organisms with partially uniparental reproduction ‣ using interbreeding and/or reproductive isolation alone may distort evolutionary history ‣ The BSC explains why members of a species resemble one other, and differ from other species. Ecological Species Concept (ESC) ‣ Species are defined ecologically by a shared ecological niche ‣ Members of a species occupy an adaptive zone that is different from that of any other species (in its range) ‣ ESC is based on evolutionary cohesion ‣ main problem: how to a priori define the adaptive zone characteristic for a species? Phylogenetic Species Concept (PSC) Reproductive Isolation “speciation continuum” ‣ A “phylogenetic species” is a basal cluster of organisms that is diagnosably distinct from other such clusters (Cracraft 1989) species A ‣ A species is the smallest monophyletic group of common ancestry (de Queiroz & Donoghue 1988) ‣ PSC is based on evolutionary history ‣ problems: the phylogeny of a gene might not reflect the entire evolutionary history; paraphyly of species poses a problem, too. no ancestral species partial imperfect reproductive isolation complete reproductive isolation species B time Reproductive Isolation ‣ premating isolating barriers: mechanisms that prevent the formation of hybrid zygotes by impeding gene flow before the transfer of sperm or pollen ‣ postmating, prezygotic isolating barriers: mechanisms that prevent the formation of hybrid zygotes after sperm or pollen transfer but before fertilization ‣ postzygotic isolating barriers: mechanisms that reduce the viability or fertility of hybrid zygotes Premating isolating barriers Behavioral Mutual attraction between the sexes of different species is weak or absent, preventing them from initiating courtship Habitat isolation. The adaptation to different habitats in the same general area prevents or limits gene exchange Ecological Temporal isolation. Gene flow between two taxa is impeded on the basis of different breeding/flowering times Pollinator isolation. Gene flow between angiosperm species is reduced by differential interactions with pollinators. Mechanical Inhibition of normal copulation or pollination between two species due to incompatibility of reproductive structures Mating system Evolution of partial or complete self-fertilization or asexual reproduction can result in the creation of a new taxon Premating isolating barriers Premating isolating barriers ‣ Cichlid fishes of Lake Victoria, East Africa ‣ Orchids of the genus Ophrys: pollinator isolation 0 - P. nyererei P. pundamilia + P. nyererei x pundamilia 0 - P. nyererei *no color vision P. pundamilia Seehausen et al. (1997) Seehausen & van Alphen (1998) Postmating, prezygotic isolating barriers Copulatory Behavior Gametic Isolation www.grandsitesaintev ictoire.com + Preference of P. nyererei female monochromatic light* Preference of P. nyererei female normal (white) light Behavior of an individual during copulation is insufficient to allow normal fertilization Postzygotic isolating barriers Extrinsic Noncompetitive gametic isolation. Intrinsic problems with transfer, storage, or fertilization of heterospecific gametes in single fertilizations between members of different species Competitive gametic isolation. Heterospecific gametes are not properly transferred, stored, or used in fertilization when competing with conspecific gametes (“conspecific sperm or pollen preference”) Ecological inviability. Hybrids develop normally but suffer lower viability because they cannot find an appropriate ecological niche Behavioral sterility. Hybrids have normal gametogenesis but are less fertile than parental species because they cannot obtain mates Hybrid inviability. Hybrids suffer developmental difficulties causing full or partial lethality Intrinsic Hybrid sterility. Hybrids suffer problems in the development of the reproductive system or gametes or suffer neurological or physiological lesions that render them incapable of successful courtship Postzygotic isolating barriers Reinforcement fitness ‣ Reproductive isolation may be reinforced by natural selection species A aabb species B AABB ‣ evidence: prezygotic isolation between sister-species occurring in sympatry is greater than between pairs of species that live in allopatry AA bb ‣ Reinforcement is the evolution of divergent mate preferences in response to selection against hybrids (if hybrids have low fitness there is a high cost of mating with a wrong species) Bb B loc us us A Aa loc BB aa Reinforcement Biogeography of speciation parapatric speciation isolation index sister taxa (in sympatry) allopatric speciation sympatric speciation sister taxa (in allopatry) 0 m 1/2 migration rate genetic distance Coyne & Orr (1997) Gavrilets (2004) Biogeography of speciation Biogeography of speciation parapatric speciation sympatric speciation time allopatric speciation ‣ allopatric speciation is the origin of new species from geographically isolated populations. There is no gene exchange between the diverging populations ‣ parapatric speciation is the origin of new species when gene flow between diverging populations is neither zero nor the maximum possible ‣ sympatric speciation is the origin of new species without geographic isolation. It is the emergence of new species from a population where mating is random with respect to the birthplace of the mating partners Allopatric speciation Allopatric speciation Vicariance* speciation ‣ reproductive isolation evolves as byproduct of independent evolution: ...in Alpheus snapping shrimp from both sides of the Isthmus of Panama snapping shrimp (Alpheus) ‣ Selection. Adaptation to different environments ‣ Mutation. Different mutations will arise in the populations ‣ Drift. Random change in gene frequencies NASA TW Doeppner *is the splitting in the range of a taxon, e.g., through continental drift, mountain formation, glaciation Knowlton & Weight (1998) ‣ In allopatric speciation, a gene pool is initially divided by an external barrier to dispersal 26 Kauai Parapatric speciation Peripatric* speciation 12 Oahu 29 ‣ Parapatric speciation involves the evolution of reproductive isolation between two populations that exchange genes 40 Maui complex www.pacificislands.com ‣ Clinal models require environmental gradients. Subpopulations become adapted to their local habitats and speciate ‣ In ”stepping-stone” models only adjacent populations exchange genes ‣ an intermediate step in parapatric speciation is a hybrid zone, i.e. an area where hybridization occurs 26 Drosophila *a new species emerges from a small, isolated subpopulation on the periphery e.g., colonization of an island (“founder effect speciation”) NASA Hawaii Parapatric speciation Parapatric speciation “Stepping-stone” model: most realistic scenario? “Stepping-stone” models Clinal models colonization dry environmental gradient dry local adaptation wet river hybrid zone hybrid reproductive isolation colonization 0 time population expansion local adaptation population expansion gene flow local extinction speciation! wet re-colonization Sympatric speciation ‣ “No aspect of speciation is as controversial as the view that new species can begin to form sympatrically” (Coyne & Orr 2004) ‣ Mathematical models and computer simulations show that sympatric speciation is theoretically plausible (under certain conditions) ‣ However, evidence for sympatric speciation in nature is scarce... ‣ ...since - in many putative cases - alternative scenarios (allopatric, parapatric) cannot be ruled out Sympatric speciation 1. antagonism between selection and recombination 2. development of sufficient ecological differences to allow coexistence reproductive isolation and ability to coexist must evolve simultaneously trait selection interbreeding “ecological speciation” Sympatric speciation ‣ apple maggot fly: sympatric host races palm trees on Lord Howe Island Savolainen et al. (2006) crater lake cichlids www.organicgardenpests.com apple maggot fly larvae of the original population of apple maggot fly (Rhagoletis pomonella) feeds on hawthorn www.gartenliebe.de www.wdr.de host shift of one population to apple in the USA in the mid-nineteenth century (Bush 1993) within ca. 150 years, two ecologically distinct species evolved Feder et al. (2000) ‣ evidence Feder et al. (2000) ‣ two “fundamental” problems (Coyne & Orr 2004) frequency Sympatric speciation Sympatric speciation Arrow cichlid (A. zaliosus) Laguna de Apoyo Sympatric speciation Midas cichlid (Amphilophus citrinellus) Laguna de Apoyo Ecological speciation Nicaraguan crater lake cichlids ‣ ... is the evolution of barriers to gene flow resulting from ecologically-based divergent selection ✓sympatric occurrence ✓ecological difference ‣ (as opposed to ‘mutation-order speciation’, i.e. the evolution of reproductive isolation by the chance occurrence and fixation of different alleles between populations adapting to similar selection pressures ✓reproductive isolation Barluenga et al. (2006) ✓monophyly ‣ e.g. ‘ecomorphs’ of stickleback fish along environmental gradients Polyploidy and Hybrid Speciation Polyploidy and Hybrid Speciation Hybrid speciation in Louisiana irises ‣ Polyploids arise naturally and are interfertile among themselves, but reproductively isolated from parental species ‣ Hybrids may have a fitness advantage under certain conditions ‣ Polyploidy and hybrid speciation is common in plants, but also animals appear to have originated in this way swamp freshwater marsh I. fulva hybrid Polyploidy and Hybrid Speciation Polyploidy and Hybrid Speciation Introgressive hybridization “hybrid species” I. hexagona Heliconius heurippa back-crosses predominantly with one parental species (Rieseberg & Wendel 1993) Mavarez et al. (2006) Xiphophorus clemenciae species 1 stabilized introgressant species 2 Meyer et al. (2006) Arnold & Bennett (1993); images: M. Arnold ‣ New species may originate by polyploidization or as a consequence of hybridization Natural Selection... ...Sexual Selection ‣ ...“is the selection on mating behavior, either through competition among members of one sex (usually males) for access to members of the other sex or through choice by members of one sex (usually females) for certain members of the other sex” (Ridley 1996) ...Sexual Selection Natural Selection... images: www.crbs.umd.edu, www.smh.com.au images: www.idscaro.net, www.wikipedia.com ‣ ... “is the process by which the forms of organisms in a population that are best adapted to the environment increase in frequency relative to less well-adapted forms over a number of generations” (Ridley 1996) ...Sexual Selection Natural Selection... ‣ ...operates if the following conditions are met: competitors reproduction sexual selection individual fitness other members of the same sex heredity natural selection fitness of the genotype other individuals in the same population trait variation variation in fitness organisms must reproduce to form new generations offspring resemble parents (“like must produce like”) individuals in natural populations vary in (adaptive) traits individuals in natural populations vary in the number of their offspring that survive to reproduce (‘lifetime reproductive success’) (Ridley 2004; Stearns & Hoekstra 2005) fitness ...Sexual Selection ...Sexual Selection ‣ ...seems to play an important role in speciation (in allopatry, parapatry, and sympatry) Sex trait favored by sexual selection traits that females find attractive ‣ Traits involved in mate choice, courtship and mating evolve faster than most other traits. Male traits that persuade females to mate traits that discourage females from mating again (copulation plugs, mating injuries, transmitted chemicals, etc.) image: M. Schartl ‣ Sexual selection involves co-evolution of female preference and male signals Xiphophorus hellerii Female choice of the best males (best genes, largest territory, least parasites, etc.) resistance to male persuasion ...Sexual Selection ‣ large divergence in male ornaments ‣ low morphological divergence of females ‣ large divergence of female preference ‣ large divergence in male nuptial coloration ‣ low morphological divergence of females ‣ large divergence of female preference image: E. Schraml Cichlid fishes of Lakes Malawi & Victoria images: E. Schraml ‣ small ecological divergence www.smh.com.au Birds of paradise