BIO 4202

Evolutionary Ecology

(Vasanth 2019)

Ullasa Kodandaramaiah

MODULE: PHYLOGENETICS AND

BIOGEOGRAPHY

Part B: Historical Biogeography

Alfred Wallace (Zoogeographic regions)

Neartic

Neotropical

Ethiopian (Afrotropical) Oriental

(Asian/Indo-Malayan)

Palearctic

Australasian + Oriental + Afrotropical = Old World TropicsPalearctic + Nearctic = HolarcticNearctic + Neotropical = New World

Australasian

SpeciationSpeciation

Allopatric Allopatric - newly formed species occupy different geographic areas

SympatricSympatric – newly formed species occupy the same geographic areas

Historical biogeography –

how did species come to be distributed the way they are?

disjunct distributionsdisjunct distributions

~ place of origin

Parapatric: Speciation with limited geneflow

Peripatric: Small peripheral,isolated population.

founder effect, genetic drift, etc (e.g island colonization)

Species 1 Species 2

Ancestor

Ancestor

Vicariant speciation

Speciation

by dispersal

Historical biogeography perspective -allopatric, peripatric & parapatric: disjunct distributions

ColonizationColonization before before v/sv/s after after

Vicariant speciation between areas A+BC and B+C

A B C

A BC

Speciation by dispersal from area X to Y

XY

Y(sp3)

X YX

(sp2)X

(sp1)

X

BC

A(sp1)

ABC

Ancestral areas

B(sp2)

C(sp3)

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

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

Alfred Wegener – 'Theory of Continental Drift' (accepted in 1960s)

Evidence – shape, fossils, mountain belts, etc

Illustration by Osvaldocangaspadilla, Wikimedia

Soil & rock samples from mountain ranges

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.

Continental Drift - movement over geological time of the earths large land masses as a result of plate tectonics

Image:Rick Cheel

Pangea

Wikimedia

200 MYA – Laurasia begins to separate from Gondwanaland

225 MYA - Pangea

200

Pangea

Breakup of Pangaea/Gondwana

130

India+Madagascar+Seychelles beginning rifting

120

105

Africa breaks off

95

90

88

India-Seychelles separated from Madagascar

Nasikabatrachus sahyadrensis (Pig-nosed frog/Purple frog) Biju and Bossuyt 2003, Nature 425, 711-714

Sooglossidae (Seychelles)Nasikabatrachidae (Western Ghats)

Divergence – ca 130 mya

Photo: Karthick Bala, Wikimedia

85

(New Zealand+New Caledonia) from Australia+Antarctica

Zanis M J et al. PNAS 2002;99:6848-6853©2002 by National Academy of Sciences

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).

Image: Wikimedia

Photo: Thomas J. Lemieux

Amborella trichopoda: New Caledonia

Sister to otherflowering plants?

80

70

India from Seychelles ca. 65 mya

60

53

45

Australia starts drifting northwards, split completely by 40

37

SA starts drifting northwards, split completely by 35

30

20

12

10

05

Breakup of Gondwana: summary

India+Madagascar+Seychelles break off 130-80 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)

Laurasia

Eastern and Western Palaearctic,Eastern and Western Nearctic

Connected, disconnected and reconnected in different combinations

- Transatlantic bridges - Beringian bridges

South and North America

● isolated since separation of Laurasia from Gondwana

● Reconnected 3 mya

● Isthmus of Panama

Modified from slide by John McCall

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

Great American Interchange

● Fauna exchanged adapted to savannas

● Isthmus >> savanna?

Modified from slide by John McCall

Panamanian Isthmus : Vicariance

Image http://www.shmoop.com/speciation/speciation-geography.html

Altheus (snapping shrimp)Hurt et al 2008 Evolution 63-2: 514–530

Sp 1

Sp 2

Sp 3

Sp 4

Africa

New Zealand

South America

Australia

! simplified

Africa

New Zealand

South America

Australia

taxon-area cladogram

1966 - Lars Brundin - chironomid midges and break-up of Gondwana.

'Phylogeny reflects vicariance' - phylogenetic biogeography

Africa

New Zealand

South America

Australia

India 1

Africa 2South

America 4

New

Zealand 3

Australia 5

Antarctica

PalaeognathaeLiving ratites and tinamous

(large, mainly flightless birds)

7 68

25

1

1. Tinamous: SAm2. Kiwis: NZ3. (Moas: NZ)4. (Elephant birds: Africa)5. Cassowaries: Aus-NG6. Emus: Aus7. Ostrich: Afr-EurA8. Rheas: SAm

(extinct groups)

Slide by Ian Hutchinson

Taxon: 1 2 3 4 5 6 7 8Area: S.Am NZ NZ Ma Au,NG Au Af S.Am

Slide by Ian Hutchinson

Gondwanan vicariancesee also Cooper et al 2001. Nature 409 (6821): 704–70

1

7

4

2,35,68

Gondwana

Illustration by Ian HutchinsonFig: Pearson Prentice Hall Inc

Dispersal

31 spp.

pansies,

buckeyes.

Biogeography of the genus Junonia

(Nymphalidae: Nymphalinae):

Testing hypotheses of vicariance and dispersal

Kodandaramaiah & Wahlberg 2007 Journal of Evolutionary Biology 20: 2181-2191

Extant distribution

17 (15)

10 (8)

3

4 (4)

17 (15) 10 (8)

8 (1)

4 (4)

simple VICARIANCE scenario

simple DISPERSAL Scenario

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

3 genes

1 mitochondrial - COI 2 nuclear - EF1-alpha & wingless

> 3000 bp : 22 spp.

Molecular phylogeny

Kodandaramaiah & Wahlberg 2007. Journal of Evolutionary Biology 20(6) 2181-2191DOI: 10.1111/j.1420-9101.2007.01425.x

Strict Consensus of 791 equally MPT

Bayesian tree

Africa

Oriental (Asia) Asia + Australia

AustraliaAfrica

Africa + Asia + Aust.

New World

Oriental (Asia)

Outgroups (Africa)

Junonia

Predictions:

● simple VICARIANCE scenario:

Species within different regions form monophyletic

groups.

● simple DISPERSAL scenario ?

?

The beginning of things

Outgroups

New World

Molecular Dating estimate (minimum)

20 myCollision of Africa+Arabia with Eurasia

Madagascar

● ca. 80% endemic (95% of the reptiles, 99% of amphibians, and 100% of land mammals excluding bats).

● Radiations: Lemurs, tenrecs, chameleons, dung beetles, etc

Photo: Reuters / www.dailymail.co.uk

Lemur catta (Ring-tailed lemur) Calumma parsonii(Parson's Chameleon)

Photo: Wikimedia / Frank Wassen

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

Sister groups of Malagasy clades

Most divergences in the Cenozoic (65 mya to present)

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 45–26 Mya [53] is shaded. Poux et al. BMC Evolutionary Biology 2008 8:102 doi:10.1186/1471-2148-8-102

Radiation of tenrecs

Photo: Wikimedia/Wilfried Berns

Echinops telfairi(Lesser Hedgehog-tenrec)

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) 687-697

2) SE Asia: e.g. Skinks (Datta-Roy et al 2012 Mol Phylogenet Evol 63(3):817–824)

Majority of life forms have affinities to SE Asia (Cenozoic dispersal)

3) Africa: e.g. Junonia

4) Eurasia/Holarctic: E.g Argynnis hyperbius

Photo: Wikimedia/Alpsdake

Argynnis hyperbius(Indian Fritillary)

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): 351-356)

4) Neotropical: Gondwanan, dispersal from NA (Great American Interchange)

5) Holarctic: Gondwanan, dispersal from Oriental, African (see Sanmartín et al 2001. BJLS 73: 345-390)

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

Speciation by 'Dispersal'

Geodispersal

● Dispersal when a barrier is lost

e.g Great American Interchange

Dispersal between Africa & Asia across the Arabian Peninsula