MODULE: PHYLOGENETICS AND BIOGEOGRAPHY Part B: Historical Biogeography

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1 BIO 4202 Evolutionary Ecology (Vasanth 2018) Ullasa Kodandaramaiah MODULE: PHYLOGENETICS AND BIOGEOGRAPHY Part B: Historical Biogeography

2 Alfred Wallace (Zoogeographic regions) Neartic Palearctic Neotropical Ethiopian (Afrotropical) Oriental (Asian/Indo-Malayan) Australasian Australasian + Oriental + Afrotropical = Old World Tropics Palearctic + Nearctic = Holarctic Nearctic + Neotropical = New World

3 Historical biogeography how did species come to be distributed the way they are? restricted distribution ~ place of origin Speciation Allopatric - newly formed species occupy different geographic areas Sympatric newly formed species occupy the same geographic areas disjunct distributions

4 Peripatric: Small peripheral,isolated population. founder effect, genetic drift, etc (e.g island colonization) Parapatric: Speciation with limited geneflow

5 Historical biogeography perspective - allopatric, peripatric & parapatric: disjunct distributions Ancestor Ancestor Species 1 Species 2 Vicariant speciation Speciation by dispersal Colonization before v/s after

6 Ancestral areas A BC A B C Vicariant speciation between areas A+BC and B+C X Y A (sp1) X (sp1) B (sp2) X (sp2) Speciation by dispersal from area X to Y X ABC BC C (sp3) XY Y (sp3)

7 Allopatric speciation - disjunct distributions pre-1960 s era - belief in a stable earth - maximum-dispersal model 'dispersalism', identification of centres-of-origin 1960 s - Acceptance of plate tectonic theory (continental drift) - phylogenetics Willi Hennig

8 Plate tectonics Was noted for several centuries that the outlines of some continents appeared as though they could fit together like a jigsaw best when the outlines are edges continental shelves. Adapted from Rick Reel's slides

9 Alfred Wegener 'Theory of Continental Drift' (accepted in 1960s) Evidence shape, fossils, mountain belts, etc Illustration by Osvaldocangaspadilla, Wikimedia

10 Soil & rock samples from mountain ranges

11 Plate Tectonics - theory that the earth s crust, including the continents and ocean floors, is made up of a series of plates, as plates collide volcanoes occur, and may result in oceanic islands, mountain ranges etc. Image: Rick Cheel Continental Drift - movement over geological time of the earths large land masses as a result of plate tectonics

12 Pangea Wikimedia

13 225 MYA - Pangea 200 MYA Laurasia begins to separate from Gondwanaland

14 2 0 0 Breakup of Pangaea/Gondwana Pangea

15 1 3 0 India+Madagascar+Seychelles beginning rifting

16 1 2 0

17 1 0 5 Africa breaks off

18 9 5

19 9 0

20 8 8 India-Seychelles separated from Madascar

21 Nasikabatrachus sahyadrensis (Pig-nosed frog/purple frog) Biju and Bossuyt 2003, Nature 425, Photo: Karthick Bala, Wikimedia Sooglossidae (Seychelles) Nasikabatrachidae (Western Ghats) Divergence ca 130 mya

22 8 5 (New Zealand+New Caledonia) from Australia+Antarctica

23 Sister to other flowering plants? Amborella trichopoda: New Caledonia Image: Wikimedia Photo: Thomas J. Lemieux 50% majority rule tree derived from those trees sampled after burn in. (Bayesian Analysis). Posterior probabilities are indicated above the branches (note: not an ultrametric tree) by National Academy of Sciences Zanis M J et al. PNAS 2002;99:

24 8 0

25 7 0 India from Seychelles ca. 65 mya

26 6 0

27 5 3

28 4 5 Australia starts drifting northwards, split completely by 40

29 3 7 SA starts drifting northwards, split completely by 35

30 3 0

31 2 0

32 1 2

33 1 0

34 0 5

35 Breakup of Gondwana: summary India+Madagascar+Seychelles break off mya Africa splits off 105 mya, collision with Eurasia ca 20 mya New Zealand isolated ca. 85 mya India separated from Madagascar ca 88 mya (collision with Asia ca 40 mya, Himalayas) India from Seychelles ca 65 Australia from Antarctica ca.40 mya SA from Antarctica ca 35 mya (Antarctica circumpolar current & glaciation)

36 Laurasia Eastern and Western Palaearctic, Eastern and Western Nearctic Connected, disconnected and reconnected in different combinations - Transatlantic bridges - Beringian bridges

37 South and North America isolated since separation of Laurasia from Gondwana Reconnected 3 mya Isthmus of Panama Modified from slide by John McCall

38 Great American Interchange 50% of extant land mammal genera in South America descend from North American immigrants. Corresponding figure for North America: 29% Great American Interchange migrants. Those with a cross by their name became extinct in their new continent. Modified from slide by John McCall

39 Fauna exchanged adapted to savannas Isthmus >> savanna? Modified from slide by John McCall

40 Panamanian Isthmus : Vicariance Altheus (snapping shrimp) Hurt et al 2008 Evolution 63-2: Image

41 Lars Brundin - chironomid midges and break-up of Gondwana. 'Phylogeny reflects vicariance' - phylogenetic biogeography Sp 4 Sp 3 Sp 2 Sp 1 Australia South America New Zealand Africa Australia South America New Zealand Africa! simplified taxon-area cladogram

42 Australia South America South America 4 Africa 2 India 1 New Zealand Antarctica Africa New Zealand 3 Australia 5

43 Palaeognathae Living ratites and tinamous (large, mainly flightless birds) Tinamous: SAm 2. Kiwis: NZ 3. (Moas: NZ) 4. (Elephant birds: Africa) 5. Cassowaries: Aus-NG 6. Emus: Aus 7. Ostrich: Afr-EurA 8. Rheas: SAm (extinct groups) Slide by Ian Hutchinson

44 Taxon: Area: S.Am NZ NZ Ma Au,NG Au Af S.Am Slide by Ian Hutchinson

45 Gondwanan vicariance see also Cooper et al Nature 409 (6821): , ,6 Gondwana Illustration by Ian Hutchinson Fig: Pearson Prentice Hall Inc

46 Dispersal

47 Biogeography of the genus Junonia (Nymphalidae: Nymphalinae): Testing hypotheses of vicariance and dispersal 31 spp. pansies, buckeyes. Kodandaramaiah & Wahlberg 2007 Journal of Evolutionary Biology 20:

48 Extant distribution 4 (4) 4 (4) 10 (8) 17 (15) 10 (8) 17 (15) 8 (1) 3

49 simple VICARIANCE scenario

50 simple DISPERSAL Scenario

51 Phylogeny: predictions simple VICARIANCE scenario: species within different regions form monophyletic groups. simple DISPERSAL scenario: Asian spp. nested within African Australian spp nested within Asian New World spp nested within African

52 Molecular phylogeny 3 genes 1 mitochondrial - COI 2 nuclear - EF1-alpha & wingless > 3000 bp : 22 spp.

53 Bayesian tree Strict Consensus of 791 equally MPT Kodandaramaiah & Wahlberg Journal of Evolutionary Biology 20(6) DOI: /j x

54 Outgroups (Africa) Africa Junonia Oriental (Asia) Asia + Australia Oriental (Asia) Australia Africa Africa + Asia + Aust New World

55 Predictions: simple VICARIANCE scenario: Species within different regions form monophyletic groups. simple DISPERSAL scenario?

56 Outgroups? The beginning of things New World

57 Molecular Dating estimate (minimum) 20 my Collision of Africa+Arabia with Eurasia

58 Madagascar

59 ca. 80% endemic (95% of the reptiles, 99% of amphibians, and 100% of land mammals excluding bats). Radiations: Lemurs, tenrecs, chameleons, dung beetles, etc Lemur catta (Ring-tailed lemur) Calumma parsonii (Parson's Chameleon) Photo: Reuters / Photo: Wikimedia / Frank Wassen

60 Testing vicariance versus dispersal Gondwanan vicariance: - older divergence times - sister clades in Asia or other Gondwanan fragments Post-Gondwanan dispersal - recent divergence times - sister clades in Africa

61 Sister groups of Malagasy clades Most divergences in the Cenozoic (65 mya to present)

62 Radiation of tenrecs Echinops telfairi (Lesser Hedgehog-tenrec) Photo: Wikimedia/Wilfried Berns Timing of tenrec speciation events and Madagascar colonization. Divergence times by a Bayesian relaxed molecular clock method, with six time constraints from fossil calibrations. One of them, the paenungulate radiation is represented on the chronogram. Black circles indicate the divergence from the non-malagasy sister group (node 2) and the initial divergence of Malagasy tenrecs (node 3). Standard deviations are indicated by grey bars, and 95% credibility intervals by open bars. The period of a putative land bridge between Madagascar and Africa at Mya [53] is shaded. Poux et al. BMC Evolutionary Biology :102 doi: /

63 Historical biogeography of India 1) Gondwanan (Out-of-India hypothesis) e.g. Nasikabatrachus sahyadrensis reviewed in Datta-Roy & Karanth 2009 J of Biosciences 34 (5) ) SE Asia: e.g. Skinks (Datta-Roy et al 2012 Mol Phylogenet Evol 63(3): ) Majority of life forms have affinities to SE Asia (Cenozoic dispersal) 3) Africa: e.g. Junonia 4) Eurasia/Holarctic: E.g Argynnis hyperbius

64 Argynnis hyperbius (Indian Fritillary) Photo: Wikimedia/Alpsdake

65 1) African: Gondwanan, Dispersal from Holarctic, Oriental 2) Australasian: Gondwanan, Dispersal from SE Asia 3) New Zealand: predominantly Dispersal from Oriental/Australia (see Waters & Craw 2006 Syst Biol 2006, 55 (2): ) 4) Neotropical: Gondwanan, dispersal from NA (Great American Interchange) 5) Holarctic: Gondwanan, dispersal from Oriental, African (see Sanmartín et al BJLS 73: )

66 Speciation by 'Dispersal' pre 1960 s: dominant view 1960 s: the science of the improbable, the rare, the mysterious and the miraculous Now??: dispersal is fundamental to allopatric speciation

67 Geodispersal Dispersal when a barrier is lost e.g Great American Interchange Dispersal between Africa & Asia across the Arabian Peninsula

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