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Relationships of Floras (& Faunas) Vicariance Vs. Dispersal How Do 4/1/19 Relationships of Floras (& Faunas) Vicariance vs. Dispersal how do you decide? Knowledge of earth and organism histories now permit closer examination of relationships of disjunct floras and faunas Vicariance • Southern Hemisphere temperate Biogeography has relied on • Southern Hemisphere tropics two sources of information • the Wallace Line • Eastern Asian - Eastern North American temperate 1. Phylogenetic trees - clades Disjunct (vicariad) species Disjunct continental areas 2. Knowledge of splitting events of areas - continents, mountain erection, etc. Dispersal What is missing? 3. Times for branching events of clades relative to geological event – clocks! Disjunct species Disjunct continental areas Relationships of Floras & Faunas Continents, Clades, and Clocks Bromeliaceae Rapateaceae Knowledge of earth and organism histories now permit closer examination of a2 b2 Taxon A Taxon A relationships of disjunct floras and faunas a1 b1 a3 b Taxon B 3 Taxon B • Southern Hemisphere temperate • Southern Hemisphere tropics a4 b Taxon C 4 Taxon C • the Wallace Line • Eastern Asian - E N American temperate Maximum likelihood tree DNA “rate smoothed” with different DNA rates tree along each branch 15 mya Taxon A * Taxon B Taxon C 2 dispersals! Bromeliaceae 30 20 10 Million yrs ago Rapateaceae * Biogeographical interpretation Fossil calibrated tree 1 4/1/19 Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Flora 35 species of trees and shrubs, evergreen and deciduous, restricted to South Interesting contrast between the floras of the southern hemisphere temperate America, New Zealand, Australia, Tasmania, New Caledonia, New Guinea, and tropical floras. and fossilized in Antarctica [The northern hemisphere continents are far more affected by recent glaciation events, so their distribution patterns are far more complex]. Absent from Africa! — “odd continent out” Nothofagaceae ✟✟✟✟ Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Flora Connections between South America and Australasia pronounced: Proteaceae comprise 1700 species of woody plants placed in 79 genera predominantly of the southern hemisphere. The family, unlike Nothofagaceae, • Subg. Nothofagus — South America occurs in south Africa and Madagascar, and extends into southern China. • Subg. Fuscospora — S. Am., N. Zeal., Tasmania • Subg. Lophozonia — S. Am., N. Zeal., Tasmania, Austr. The 16 genera from Africa are endemic and comprise only 3 lineages. In • Subg. Brassospora — New Caledonia, New Guinea comparison, South America and Australasia share roughly half of the genera in common. All tribes within the latter two areas are shared. Africa — “odd continent out”! Nothofagaceae Protea Roupala Banksia ✟✟✟✟ Proteaceae 2 4/1/19 Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Flora Restionaceae comprise 520 species of grass-like plants placed in 58 genera Restionaceae comprise 520 species of grass-like plants placed in 58 genera predominantly of the southern hemisphere. predominantly of the southern hemisphere. The 350 species from Africa are unique and belong only to 11 genera of the Restio group. In contrast, South America and Australasia share many genera including some species. Africa — “odd continent out”! Askidiosperma — Restionaceae Cape Region Fynbos, S. Africa — Restionaceae Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Flora Why is Africa the “odd continent out” when it comes to the temperate southern 1. All three continents separated from Gondwana at about 100-110 mya in the hemisphere flora? early Cretaceous, but South America and Australia linked with temperate Antarctica until about 50 mya (and via small water passages until 27 mya) Three reasons: broken Restionaceae made Proteaceae ✟✟✟✟ Nothofagaceae Estimates in millions of years BP when migration routes between land masses were broken or made. 3 4/1/19 Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Flora 2. Africa drifted further north and experienced greater climatic change through 3. Africa made secondary contact with temperate Eurasia around 17 mya; long this latitudinal journey. Greater extinction of temperate biota; which is now contact further differentiated the temperate flora of Africa relative to South restricted to small area of south Africa. America and Australia Temperate flora Triassic broken Temperate flora made Present Positions of Labrador, Africa, and Australia in the Triassic (200 mya) Estimates in millions of years BP when migration routes between land masses were broken or made. and at the present Vicariance vs. Dispersal? Vicariance vs. Dispersal? Temperate Gondwanan disjuncts – vicariance and/or dispersal ? Temperate Gondwanan disjuncts – vicariance and/or dispersal Timing of organism divergence vs. timing of geological divergence critical Timing of organism divergence vs. timing of geological divergence critical 120 mya 50 mya Nothofagus marsupials southern beeches ratite birds 90 mya 20 mya Gondwana Jurassic – Cretaceous border ~ 150 million years ago 4 4/1/19 Southern Hemisphere Temperate Flora Southern Hemisphere Temperate Fauna southern beeches? – ratites – flightless birds? vicariance and a lot of dispersals vicariance and a lot of dispersals A. Geological connections Australia <—> New Zealand dispersals B. Geological story C. Mitochondrial story (different story) Science 2014 Mitchell et al. Southern Hemisphere Temperate Fauna Southern Hemisphere Temperate Fauna ratites – flightless birds ratites – flightless birds Yonezawa et al. (2017) Phylogenomics and morphology of extinct paleognaths reveal the origin and evolution of the ratites. Current Biology intercontinental dispersals independent loss of flight 5 4/1/19 Southern Hemisphere Temperate Fauna Vicariance vs. Dispersal? Relationship of the Australian Marsupials GBE Tracking marsupial evolution using genomic data - Dromiciops Fossils of both marsupials and placental mammals found in marsupial and placental mammalsResolving diverged Marsupial Phylogeny Using Retroposonsin the Late Cretaceous Antarctica ~85 Ma Tasmanian devil Western quoll Gallus et al. 2015 Dasyuromorphia Fat-tailed dunnart Numbat Notoryctemorphia Southern marsupial little bush monkey mole Eastern barred Peramelidae bandicoot Peramelemorphia Southern brown bandicoot Fossil marsupial (Eocene 40 Ma) Greater bilby Australidelphia x13 Diprotodontia Tammar wallaby Microbiotheria Monito del monte Nilsson et al. 2010 Paucituberculata Marsupialia Silky shrew Why did marsupials but NOT placental opossum x10 Didelphimorphoa Grey short-tailed mammals migrate on to Australia? 8 0 7 0 6 0 5 0 4 0 3 0 2 0 1 0 opossum mya CRETACEOUS PALEOGENE NEOGENE Q* Figure 2. Phylogenetic tree of marsupials derived from retroposon data. The tree topology is based on aF presence/absenceIG.1.—Phylogenetic retroposon tree of the seven marsupial orders based on 16 of the 20 phylogenetically informative retrotransposon insertions and sequence matrix (Table 1) implemented in a heuristic parsimony analysis (Figure S3). The names of the seven marsupial ordersdata. are Markers shown infor red, different and the nodes icons in the marsupial tree are shown asorangecircles.BranchesshowingretrotransposonsupportbasedonNilsson et al. (2010) marsupialare representative disjunction of each of the orders: Didelphimorphia, between Virginia opossum; South Paucituberculata, America shrew opossum; Microbiotheria, monitomarsupial del monte; migration via Antarctica by KT event, Notoryctemorphia, marsupial mole; Dasyuromorphia, Tasmanian devil; Peramelemorphia, bilby; Diprotodontia,are indicated kangaroo. as gray Phylogenetically circles. The tree includes representatives of all seven living marsupial orders, and has been scaled to divergence time (Mitchell et al. informative retroposon insertions are shown as circles. Gray lines denote South American species distribution,2014). and The black phylogenetic lines Australasian tree is based on a Bayesian analysis of 32,253 nt from 28 nuclear gene fragments (supplementary fig. S1, Supplementary Material and Australiamarsupials. The cohort Australidelphiavicariance is indicated asor well asdispersal? the new name proposed for the four ‘‘true’’ Australasian orders (Euaustralidelphia)adaptive. radiation, and then later vicariance doi:10.1371/journal.pbio.1000436.g002 online). The branches leading to Peramelemorphia, Dasyuromorphia, and Notoryctemorphia have been collapsed. Additional markers supporting the relationship between Peramelemorphia (red shaded area), Dasyuromorphia (green shaded area), and Notoryctemorphia (brown shaded area) are shown separately in figure 2.AustralidelphiareferstothegroupingoftheSouthAmerican Microbiotheria and the four Australian orders (Szalay 1982). Q*, reach a topology consistent with the retroposon markers. among mammals [24], and canquaternary even mislead period. phylogenetic However, both studies gave low support for the position of reconstruction when mixed with nuclear data. Microbiotheria, illustrating the difficulties in resolving a short The position of Microbiotheria has been intensely debated since branch using sequence data under difficult conditions, such as the cohort Australidelphia was first suggested based on tarsal possible nucleotide composition bias problems and randomization evidence [1]. After decades of uncertaintyMiocene derived (23.5–11 from molecular Ma) represent mainland Australian fossil record from Dasyuromorphia, phylogenetic conclusions, of fast evolving sites. The support from two independent sources of and
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