5 1) Allopatric Speciation - populations become reproductively isolatedduring periods of geographic separation (e.g. bymountains, land bridge disappearance, sea strait).- this mode is recognized as important forspeciation of many plants and animals;- the best evidence comes from birds (Darwin'sfinches, Hawaiian honeycreepers), mammals,butterflies and Drosophila (e.g. Hawaii)
6 One of the best examples is the accepted history behind Darwin’s Galapagos finches. The initial colonist is believed to be a finch from Cocos Island, off the mainland of Costa Rica.The Cocos finch is the sole finch species on Cocos Island, whereas many species occupy the Galapagos islands owing to emigration followed by allopatric speciation on ‘new’ islands in the Galapagos.Habitat and food specialization and geographic isolation facilitated speciesCocos Island
8 Hawaiian Honeycreepers same process as per Darwin's finches except the scale of radiation and morphological variation in beak size was far more extensive. Process occurred in <10 million yearsS. Olson (2004) Evolution in Hawaii
9 Hawaiian DrosophilaKevin Edwards, Illinois State Univ.Hawaiian Drosophila: ~1000 diverse species from a single introduction of an Asian Drosophila species that spread before the current Hawaiian Islands were formed, and colonized the chain by 'island hopping' about 26 million years ago. New islands were colonized as they became suitable ecologically. Lava flows and geographic separation fragmented species' ranges, causing bottlenecks that facilitated evolution of species
10 Wrasse SpeciationA classic example of allopatric speciation as a result of separation of populations by the uplifting of the Isthmus of Panama (<3 million years ago) separating some species into Atlantic and Pacific populations
11 2) Sympatric Speciation - more controversial and in most cases lesscommon than allopatric speciation- best examples are from plants withpolyploidy (multiplication of chromosomenumber resulting in instantaneous speciation)- this mode is successful only if new individualscan interbreed through assortative mating,inbreed (self-mating), or become polyploid- believed responsible for ~40% of plant speciesdiversity, far less in animals, but is found inparasitic Chalcidoidea, and may occur in parasiticand hyperparasitic species through shifts in hostselection.
12 Hawaiian Silverswords Hawaiian silversword alliance consists of about 30 species in 3 genera (Wilkesia, Argyroxiphium, Dubautia).The species exhibit an extraordinary range of anatomical, morphological, and ecological adaptationsgenetically very closeevolved from a single ancestor that colonized Hawaii by way of long-distance dispersal from North America
14 3-Spine Sticklebacks in British Columbia Lakes Limnetic MaleBenthic MaleThe similarity of their mtDNA (<0.5% sequence divergence) suggests that each pair arose independently by sympatric speciation, following a single invasion of each lake by ancestral marine sticklebacksOther possibility is an initial invasion from the sea, with individuals developing benthic feeding mode, followed by a later 2nd invasions, with individuals focusing on planktonic foodsIncipient speciation is driven by positive assortative matingVamosi, & Schluter Evolution
15 Cichlid Fishes in African Great Lakes hundreds of species in each of Lakes Victoria, Malawi and Tanganyika, that have both habitat and feeding specialization, the latter driven largely by mouth morphology
16 3) Parapatric model- speciation occurs among adjacent populations ofthe same species as a result of strong and differingselective pressures in the local environments- there is only limited evidence, perhaps bestexemplified by local 'races' of plants likeDeschampsia caespitosa that establish on minewastes rich in heavy metals like copper and nickel;these populations are different ecologically andgenetically from adjacent individuals on non-toxicsoils.
17 Deschampsia caespitosa is an extremely variable grass with a wide distribution, though mainly in the northern hemisphere40 subspecies or varietiesin North America there exist 5 morphologically different formstaxonomy of the group, particularly in Central Europe, is still puzzling, because of the extreme morphological variability and ambiguous boundaries between the taxaAdaptation to particular environments (metal vs. nonmetal) may result in the formation of these ecotypes and represent the beginning of speciationChiapella (2000) Biol. J. Linn. Soc.
18 Parapatric speciation in Orioles? May occur in zones where distinct species have range overlap and hybridizeSibley & Short (1964) Condor
19 How does speciation occur? Gradual divergence in the face of differing selective pressures for different, isolated populations; and/orgenetic revolution in which more than 1 genetic constitution is favoured by selection for populations with the same environmental conditions.Speciation in the latter model occurs if a population switches from one adaptive peak to another, most likely occurring because of genetic drift or by polyploidization, which causes instantaneous speciation.
20 Diversification has occurred both continuously and sporadically, with dramatic increases in diversity following each of the mass extinctions. Without repeating the history, we can review the times of major diversification of particular phyla…Cambrian: sea invertebrates: archaeocyathans, inarticulate branchiopods, trilobites
21 Family Level Gains and Losses 1900Family Level Gains and Losses
22 Rest of Paleozoic: corals, articulate branchiopods, cephalopods (e. g Rest of Paleozoic: corals, articulate branchiopods, cephalopods (e.g. squids), ostracods, crinoids, starfish, graptolites (extinct colonial group), first land plants; insects in late Paleozoic
23 Mesozoic: bivalves, gastropods, malacostracans, echinoids, bony fishes, marine reptiles, first gymnosperms and (later) angiosperms; dinosaurs appear.Cenozoic: social insects; modern and placental mammals; hoofed mammals and apes; man.Most geologic records are based on marine taxa, specifically invertebrates.
24 Family-level Diversification: Benton (1995) used data collated from 90 experts for taxa including microbes, algae, fungi, protists, plants and animals for 7186 different families.Benton selected family level distinctions for a number of reasons, the main one being that he could avoid uncertainty regarding species designations by working at a higher level of resolution.Family level diversification shows the following history:
25 Family Diversification through time based on fossil record Benton (1995) Science
26 Speciation through Time log (diversity) thru timenew families in period
27 Extinction through Time Benton (1995) Science# families lost thru time% families lost thru time% families lost in period
28 Loss of families as a function of the number that started the period and the length of the period
29 Benton (1995), Science1) family-level diversity increased in a burst during the Vendian (Precambrian);2) diversity fell to 120 families during late Cambrian;3) diversity increased to 450 during Ordovician;4) diversity rose to ~600 during remaining Paleozoic;5) fell to 420 at beginning of Triassic;6) rose to 2600 during Cretaceous (144 myr) and during Pleistocene and Holocene (<2 myr);7) most continental families arose following Silurian – land plants, insects and vertebrates;
30 8) by contrast, many marine families arose during the Vendian and early Cambrian period followed by massive losses during the late Cambrian and late-Permian; then exploded in diversity until the late Cretaceous; further expansion occurred at the beginning of the Tertiary (65 myr);9) family diversity increases were clearly logarithmic for all taxa and for continental taxa, though a more complex pattern of radiation and extinction occurred with marine families.
31 Overall Patterns:many of the radiations occurred following massive extinctions;most of the early radiations were marine, most of the later ones terrestrial;huge increases during the Eocene (55 myr) correspond with radiation of the insectsinsect and flowering plants appeared to undergo radiation together suggesting coevolutionary radiations
32 mass extinctions occurred during the early Cambrian, late Ordovician, middle and late Devonian, late Carboniferous, late Permian, end Triassic, end-Jurassic, mid-Cretaceous, late Eocene4 of extinction periods were protracted (not fast)some extinctions (and radiations) were not well recorded in the fossil record (referred to as slippage)different measures of extinction (per family, total rate etc.) each have their own weakness (e.g. dividing by total extant diversity where the small number of families early on skews that statistic)
33 For the 5 accepted mass extinctions: Time Period % of families ExtinctLate Ordovician Late Devonian Late Permian Late Triassic end-Cretaceous Remember, the clock ticks faster for genera or for species than for families, so large family losses would correspond with huge species losses
34 General patterns in extinctions have been identified: terrestrial vertebrate taxa that went extinct were large-bodied (e.g. dinosaurs; end-Pleistocene megafauna). That would be consistent with the expected population biology of these species: low population numbers, long-lived, slow maturing, low reproductive rates, large home ranges and high food demands.Widespread genera fared extinction periods much better than those with narrow geographic limits.
35 Factors (e.g. species richness) that protected genera from background extinctions were ineffective protection during the mass extinction that occurred at the end of the Cretaceous.Tropical marine taxa (corals) were harder hit during mass extinctions than taxa from higher latitudes.Species diversity losses of marine bivalves took, on average, 10 million years to recoup through new radiations.Species responses to changing environments (particularly following the Pliocene-onset [<5.2 myr]) were highly individualistic.
36 Thus, current reserve designs should reflect the broad requirements and environmental responses of different taxa if large-scale biodiversity is to be successfully preserved.In many cases, a rich diversity of protected microhabitats may prove more important in reserve design than maximizing reserve area.
37 References:Axelrod, D.I A theory of angiosperm evolution. Evolution 6:29-60.Axelrod, D.I Mesozoic paleogeography and early angiosperm history. Bot. Rev. 36:Benton, M.J Diversification and extinction in the history of life. Science 268:52-58.Futuyma, D.J Evolutionary Biology, 2nd Edition. Sinauer, Sunderland, MA.Jablonski, D Extinctions: a palaeontological perspective. Science 253:Groom, M.J., Meffe, G.K. and C.R. Carroll Principles of Conservation Biology. Sinauer. 3rd ed.Primack, R.B Essentials of Conservation Biology. Sinauer, Sunderland, MA.Sisk, T.D., A.E. Launer, K.R. Switky and P.R. Ehrlich Identifying extinction risks. Bioscience 44:Sun, G., D.L. Dilcher, S. Zheng, Z. Zhou In Search of the First Flower: A Jurassic angiosperm, Archaefructus, from Northeast China. Science 282: