Outline 1. The What: Defining species 2. The How: Mechanisms for speciation Speciation Part One: What is a Species, and How Does a Species Form? Sections 1.1 and 9.2 Three different definitions 1. Morphological species 2. Biological species 3. Phylogenetic species Brainstorm: What is a species? (1) The Morphological Species definition A species is a group of organisms that shares certain structural features. based on comparing structural of organisms Advantage: Simple! features e.g., body shape, size, colouration Limitations: Morphological variation within species Morphological similarities between species (2) Biological species definition A species is a group of interbreeding organisms that can produce viable, fertile offspring under natural conditions. Requirements of a biological species: • Viable offspring • Offspring are able to develop to birth, and live. • Fertile offspring • Offspring are capable of producing children of their own. Based on reproductive compatibility between organisms. • Natural conditions • In nature, the organisms are likely to encounter each other and mate. All of these organisms are hybrids. Why are the parents still considered separate species? Advantages widely-used Lion + Tiger = Liger (sterile) Limitations spatially-isolated populations of the same species • can’t interbreed in nature can’t apply to fossil species asexually-reproducing organisms? Jackal + Dog = Hybrid (fertile) Donkey + Horse = Mule (sterile) (3) Phylogenetic species definition A species is a group of organisms that shares a common ancestor and evolutionary history. When does a species begin? When reproductive isolation occurs between related groups. Examines the evolutionary line of descent of organisms, through DNA analysis When does a species end? When lineages become extinct. Learning Checkpoint Advantage can be applied to extinct species Limitation evolutionary histories are not known for all species • The definition of species has evolved over time. • The phylogenetic definition of a species is the ultimate goal of modern classification. • However, the biological definition is still the most commonly-used among biologists and students. Speciation is... Speciation happens through the • the formation of new species from existing ones ▫ an ancestral species branches off into two or more new lineages reproductive isolation of populations. eventual result: two populations cannot interbreed requires a stop of gene flow between populations • a type of macroevolution gene flow Population A gene flow between populations maintains reproductive compatibility Population B Pop’ns A and B remain part of the same species. Reproductive isolation in Lizards (2:38) Speciation happens through the reproductive isolation of populations. eventual result: two populations cannot interbreed requires a stop of gene flow between populations Reproductive isolation requires barriers to gene flow: Reproductive Isolating Mechanisms gene flow SPECIES A genetic differences accumulate in each population; prevents reproduction The two populations diverge into two separate species. SPECIES B Pre-zygotic Post-zygotic aka “pre-fertilization” aka “post-fertilization” prevention of: • mating • fertilization affect the survival or viability of hybrid offspring Pre-zygotic mechanisms act before fertilization. Prevention of mating: • behavioural isolation • ecological/habitat isolation • temporal isolation • mechanical isolation Prevention of fertilization: • gametic isolation Behavioural isolation Pre-zygotic mechanisms • behavioural isolation •ecological/habitat isolation •temporal isolation •mechanical isolation •gametic isolation Sexual cues/mating rituals are not recognized by individuals of different species. Example: Songs of different species of birds or insects; mating “dances” Temporal isolation Pre-zygotic mechanisms • behavioural isolation •ecological/habitat isolation •temporal isolation •mechanical isolation •gametic isolation Different species are fertile at different times, preventing mating. different seasons, different times of day Example: B. americanus and B. fowleri can produce fertile hybrids in a lab; in nature they are fertile during different seasons. Habitat isolation Pre-zygotic mechanisms • behavioural isolation •habitat isolation •temporal isolation •mechanical isolation •gametic isolation Geographical isolation prevents individuals from meeting and mating. Can occur even within the same area, if the populations occupy different habitats. Example: Two species of garter snakes (genus Thamnophis) occur in the same geographic area. One lives mainly in water, and the other is terrestrial. Mechanical isolation Pre-zygotic mechanisms • behavioural isolation •ecological/habitat isolation •temporal isolation •mechanical isolation •gametic isolation Genitals are mechanically incompatible (though copulation may be attempted). Example: Insects Plants (mode of pollination) Pre-zygotic mechanisms • behavioural isolation •ecological/habitat isolation •temporal isolation •mechanical isolation •gametic isolation Gametic isolation Mating and sperm transfer may take place, but gametes are incompatible. sperm may not survive in female reproductive tract fusion is prevented by some biochemical mechanism Post-zygotic mechanisms act after fertilization. • zygotic mortality/hybrid inviability • hybrid sterility • hybrid breakdown Example: Common in marine invertebrates, which utilize external fertilization. Zygotic mortality or Hybrid inviability Post-zygotic mechanisms • hybrid inviability • hybrid sterility • hybrid breakdown Zygote does not survive to birth, or surviving hybrid is frail. Example: Goat/sheep hybrid zygote mortality Hybrid sterility Post-zygotic mechanisms • hybrid inviability • hybrid sterility • hybrid breakdown The hybrid cannot produce normal gametes, so it cannot reproduce. When parents differ in chromosome number, meiosis cannot occur normally. Example: Mules (donkey/horse hybrid). Hybrid breakdown Post-zygotic mechanisms • hybrid inviability • hybrid sterility • hybrid breakdown F1 hybrids are viable and fertile, but subsequent generations are not. Example: Some rice cultivars produce robust hybrids, but the next generation is small and sterile. Case study: Salamanders of California (3:22) Four species of leopard frog: What keeps these species isolated? • mating calls differ • defective hybrid embryos 1) Rana pipiens 2) R. blairi 3) R. utricularia 4) R. berlandieri • Reproductive isolating mechanisms maintain species by preventing gene flow and hybridization. Which specific mechanism? ▫ Can be categorized as either pre-zygotic or post-zygotic. • Post-zygotic mechanisms act after fertilization. Hybrids are produced, but with reduced viability or fertility. • • • • Read 9.2 (pg. 360-363) Pg. 363 #13, 15, 17 Pg. 373 #1-4 worksheet: Speciation • Speciation involves reproductive isolation of populations. Pre-zygotic or post-zygotic? • Pre-zygotic mechanisms prevent the production of hybrids, by preventing either mating or fertilization. Homework Summary