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Gallirallus_pendicule Gallirallus_philippen Gallirallus_philippen Gallirallus_philippen Gallirallus_ripleyi Gallirallus_wakensis Tricholimnas_sylves Gallirallus_rovianae Gallirallus_rovianae Nesoclopeus_wood Gallirallus_owstoni Habropteryx_insigni Habropteryx_torqua Habropteryx_torqua Habropteryx_torqua Habropteryx_okinaw Cabalus_modestus Eulabeornis_castan Gallirallus_australis_ Gallirallus_australis_ Tricholimnas_lafresn Habroptila_wallacii Gallirallus_calayane Gallirallus_striatus Gallirallus_pectoralis Aramidopsis_platen 0.6 0.4 0.2 0 H DE AB AB L O N J J J H I B A A G M DE F F K C A AB AB B Gallirallus_pendicule Gallirallus_philippen Gallirallus_philippen Gallirallus_philippen Gallirallus_ripleyi Gallirallus_wakensis Tricholimnas_sylves Gallirallus_rovianae Gallirallus_rovianae Nesoclopeus_wood Gallirallus_owstoni Habropteryx_insigni Habropteryx_torqua Habropteryx_torqua Habropteryx_torqua Habropteryx_okinaw Cabalus_modestus Eulabeornis_castan Gallirallus_australis_ Gallirallus_australis_ Tricholimnas_lafresn Habroptila_wallacii Gallirallus_calayane Gallirallus_striatus Gallirallus_pectoralis Aramidopsis_platen 1 0.8 0.6 0.4 0.2 0 Not Fly Fly Fly Not Not Not Not Not Not Not Not Fly Fly Fly Not Not Fly Not Not Not Not Not Fly Fly Not Linked trait+dispersal model allows statistical inferences not possible with either individually. Although this case is “obvious,” in many datasets, the influence of traits on dispersal may be unknown. Flightlessness is more frequent at the tips, so models that allow reversal are favored. Regardless, the trait inference is improved (not perfect, see root). Inferring m2 is difficult, but fixing to 0 is feasible and nearly as good (~1 LnL). Founderevent speciation is a key process, as is founder+flight loss. The base model maOers a lot. This must be part of the model choice! These models ignore lineage extinction, but ClaSSE models infeasible. Models with more areas will be slower, but BioGeoBEARS is parallelized. Check out BioGeoBEARS, updated code at PhyloWiki: hOp://phylo.wikidot.com/ Trait-dependent dispersal models for phylogenetic biogeography, in the R package BioGeoBEARS Abstract Acknowledgements: BioGeoBEARS was developed during support by: NSF Award 0919124, and by the National Institute for Mathematical and Biological Synthesis (NIMBioS), an Institute sponsored by the National Science Foundation through NSF Award #DBI-1300426 with additional support from The University of Tennessee, Knoxville. NJM was also supported by the Australian Research Council’s Discovery Early Career Researcher Award #DE150101773, and by The Australian National University. Nicholas J. Matzke *,1,2 QR code: CV, website, R code! Organism traits must be important in historical biogeography. In particular, rates of dispersal (both range expansion dispersal, and jump dispersal leading to founderevent speciation) must depend to some degree on traits such as flight and its loss, and seed dispersal mechanisms and the dispersal abilities of animals that transport seeds. However, to date no probabilistic historical biogeographical models have been available that allow geographic range and traits to coevolve on the phylogeny, with traits influencing dispersal ability. In purely continuoustime Markov models, adding a trait is just a maOer of doubling the size of the rate matrix; however, biogeographical models also include a much more complex discretetime model describing how geographic range can change during cladogenesis. Traits might also influence this process. I present an addition to the R package BioGeoBEARS that enables an evolving discrete trait to influence dispersal ability for both anagenetic and cladogenetic range change. This model can be freely combined with models adding jump dispersal (e.g., DEC+J), distance as a predictor of dispersal (+x models, with dispersal rate multiplied by distance^x), and other variants. I test the model against simulations and datasets where large evolutionary changes in dispersal ability are highly likely (e.g., Pacific rails, which have repeatedly lost flight). BioGeoBEARS trait + dispersal model Conclusions 1 Discovery Early Career Researcher Award (DECRA) Fellow; Division of Ecology, Evolution, and Genetics; Research School of Biology; The Australian National University, Canberra *nick.ma)[email protected] References & more info: Use QR code or go to: h4p://phylo.wikidot.com/nicholasjmaAke 2 Work began at: National Institute of Mathematical and Biological Science (NIMBioS) and Department of Ecology & Evolutionary Biology, University of Tennessee, Knoxville Results: Ancestral geography TwiOer and photos are OK! @NickJMakke Study system: “typical” rails of the Pacific G. pacificus, from Tahiti, “known only from a painting made by Georg Forster on James Cook’s second voyage.” Adolf Meyer (1898), The Birds of Celebes and the neighbouring islands. https://en.wikipedia.org/wiki/Snoring_rail Data from: Kirchman (2012) Collaborator: Jeremy J. Kirchman, Curator of Birds, New York State Museum Source: Kirchman, Jeremy J. (2012). Speciation of Flightless Rails on Islands: A DNA-Based Phylogeny of the Typical Rails of the Pacific, The Auk, 129(1):56-69. (Data provided by collaborator Jeremy J. Kirchman) Statistical Model Choice Base model rate of range expansion rate of range loss weight of founder5event speciation rate of flight 5> flightlessness rate of flightlessness 5> flight dispersal multiplier when trait=1 (flight) dispersal multiplier when trait=2 (flightless) rate of switch to flightlessness during founder event rate of switch to flight during founder event number of free parameters log5likelihood delta AICc model weight d e j t 12 t 21 m 1 m 2 jt 12 jt 21 np LnL dAICc mw DEC 3.84 1.94 0 3.31 0.94 1 0 0 0 4 +119.6 37.6 0.0% DEC 0.18 0.00 0.28 5.18 1.65 1 0 0 0 5 +103.9 9.4 0.9% DEC 3.81 2.01 0 2.58 0 1 0 0 0 3 +120.3 36.2 0.0% DEC 0.15 0.00 0.16 2.84 0 1 0 0 0 4 +107.5 13.5 0.1% DEC 2.50 1.69 0 3.60 1.28 1 0 1.00 0 5 +119.4 40.3 0.0% DEC 0.18 0.00 0.82 4.37 1.12 1 0 0.75 0 6 +104.2 13.3 0.1% DEC 3.31 1.98 0 2.57 0 1 0 1.00 0 4 +120.2 38.9 0.0% DEC 0.15 1E+12 0.16 2.84 0 1 0 0.00 0 5 +107.5 16.6 0.0% DIVALIKE 1.99 0.82 0 5.16 1.48 1 0 0 0 4 +115.2 28.9 0.0% DIVALIKE 0.24 0.00 0.22 5.73 1.71 1 0 0 0 5 +104.1 9.8 0.7% DIVALIKE 2.68 1.50 0 2.60 0 1 0 0 0 3 +117.7 31.0 0.0% DIVALIKE 0.16 0.00 0.10 2.61 0 1 0 0 0 4 +109.3 17.1 0.0% DIVALIKE 2.35 0.90 0 4.38 1.11 1 0 0.91 0 5 +115.3 32.2 0.0% DIVALIKE 0.24 0.00 0.49 5.49 1.49 1 0 0.62 0 6 +104.1 13.2 0.1% DIVALIKE 2.56 1.45 0 2.70 0 1 0 0.58 0 4 +117.7 33.8 0.0% DIVALIKE 0.17 1E+12 0.64 2.81 0 1 0 0.87 0 5 +109.3 20.1 0.0% BAYAREALIKE 6.40 3.70 0 4.13 2.37 1 0 0 0 4 +129.7 57.8 0.0% BAYAREALIKE 5.34 3.70 0.95 4.14 2.38 1 0 0 0 5 +122.3 46.2 0.0% BAYAREALIKE 3.32 3.91 0 2.67 0 1 0 0 0 3 +130.8 57.2 0.0% BAYAREALIKE 3.34 3.91 0.78 2.67 0 1 0 0 0 4 +123.0 44.4 0.0% BAYAREALIKE 4.36 3.46 0 5.66 2.45 1 0 0.98 0 5 +129.9 61.4 0.0% BAYAREALIKE 0.09 0.16 0.35 9.24 3.01 1 0 0.42 0 6 +97.51 0.0 96.5% BAYAREALIKE 3.36 2.66 0 2.61 0 1 0 0.00 0 4 +132.5 63.5 0.0% BAYAREALIKE 0.07 1E+12 0.14 2.59 0 1 0 0.14 0 5 +103.4 8.4 1.5% Black: free parameter that is estimated. Gray: fixed parameter. Best Model under these constraints DEC: “Dispersal- Extinction- Cladogenesis”(of Lagrange, Ree & Smith 2008) Likelihood version of Ronquist (1997)’s Dispersal-Vicariance Analysis (no vicariance) Gallirallus_pendicule Gallirallus_philippen Gallirallus_philippen Gallirallus_philippen Gallirallus_ripleyi Gallirallus_wakensis Tricholimnas_sylves Gallirallus_rovianae Gallirallus_rovianae Nesoclopeus_wood Gallirallus_owstoni Habropteryx_insigni Habropteryx_torqua Habropteryx_torqua Habropteryx_torqua Habropteryx_okinaw Cabalus_modestus Eulabeornis_castan Gallirallus_australis_ Gallirallus_australis_ Tricholimnas_lafresn Habroptila_wallacii Gallirallus_calayane Gallirallus_striatus Gallirallus_pectoralis Aramidopsis_platen 1 0.8 0.6 0.4 0.2 0 Not Not Not Not Not Not Not Not Not Not Fly Not Not Not Not Not Not Fly Fly Not Not Not Fly Not Fly Fly Fly Not Not Not Not Not Not Not Not Fly Fly Fly Not Not Fly Not Not Not Not Not Fly Fly Not Not Not Not Not Not Not Not Not Not Not Not Not Fly Not Not Not Not Not Not Fly Fly Not Not Not Fly Not Not Not Not Not Not Not Not Not Not Not Not Fly Not Not Not Not Not Not Fly Fly Not Not Not Fly Gallirallus_pendicule Gallirallus_philippen Gallirallus_philippen Gallirallus_philippen Gallirallus_ripleyi Gallirallus_wakensis Tricholimnas_sylves Gallirallus_rovianae Gallirallus_rovianae Nesoclopeus_wood Gallirallus_owstoni Habropteryx_insigni Habropteryx_torqua Habropteryx_torqua Habropteryx_torqua Habropteryx_okinaw Cabalus_modestus Eulabeornis_castan Gallirallus_australis_ Gallirallus_australis_ Tricholimnas_lafresn Habroptila_wallacii Gallirallus_calayane Gallirallus_striatus Gallirallus_pectoralis Aramidopsis_platen 1 0.8 0.6 0.4 0.2 0 Not Fly Fly Fly Not Not Not Not Not Not Not Not Fly Fly Fly Not Not Fly Not Not Not Not Not Fly Fly Not Standard trait model Gallirallus_pendicule Gallirallus_philippen Gallirallus_philippen Gallirallus_philippen Gallirallus_ripleyi Gallirallus_wakensis Tricholimnas_sylves Gallirallus_rovianae Gallirallus_rovianae Nesoclopeus_wood Gallirallus_owstoni Habropteryx_insigni Habropteryx_torqua Habropteryx_torqua Habropteryx_torqua Habropteryx_okinaw Cabalus_modestus Eulabeornis_castan Gallirallus_australis_ Gallirallus_australis_ Tricholimnas_lafresn Habroptila_wallacii Gallirallus_calayane Gallirallus_striatus Gallirallus_pectoralis Aramidopsis_platen 1 0.8 0.6 0.4 0.2 0 Not Not Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Not Fly Fly Fly Fly Fly Not Not Not Not Fly Fly Fly Not Not Not Not Not Not Not Not Fly Fly Fly Not Not Fly Not Not Not Not Not Fly Fly Not Not Not Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Not Fly Fly Fly Fly Fly Not Not Not Not Not Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Fly Not Fly Fly Fly Fly Fly Not Not Not Trait+dispersal model Flying: Flightless: Millions of years ago (Ma) Millions of years ago (Ma) Millions of years ago (Ma) Ancestral area estimate: Phillipines & nearby (the 2-area species are actually more widespread) Founder-events colonizing remote islands are new areas are associated with rapid (evolutionarily) loss of flight ANU Research School of Biology. Poster at Earlyand MidCareer Conference. R. N. Robertson Lecture Theatre, Bldg. 46. Feb. 11th, 2016, 4:45 pm.
Transcript
Page 1: Trait-dependent dispersal models for phylogenetic ...phylo.wdfiles.com/.../2016-02-11_ANU_trait-based_dispersal_v2.pdf · 2016-02-11  · Cook’s second voyage.” s. l Data from:

Gallirallus_pendiculentusGallirallus_philippensis_melloGallirallus_philippensis_19403Gallirallus_philippensis_42507Gallirallus_ripleyiGallirallus_wakensisTricholimnas_sylvestris_1896Gallirallus_rovianae_28014Gallirallus_rovianae_826433Nesoclopeus_woodfordi_immaculaGallirallus_owstoniHabropteryx_insignisHabropteryx_torquatus_celebensHabropteryx_torquatus_sulcirosHabropteryx_torquatus_torquatuHabropteryx_okinawaeCabalus_modestusEulabeornis_castaneoventrisGallirallus_australis_australiGallirallus_australis_greyiTricholimnas_lafresnayanusHabroptila_wallaciiGallirallus_calayanensisGallirallus_striatusGallirallus_pectoralisAramidopsis_plateni

1 0.8 0.6 0.4 0.2 0Millions of years ago

HDEABABLONJJJHIBAAGM

DEFFKCA

ABABB

BioGeoBEARS DEC+J+t12+t21+jt12+m1 on Pacific Railsancstates: global optim, 2 areas max. d=0.0935; e=0.1565; j=0.3493; t12=9.2439; t21=3.0112; jt12=0.4168; LnL=−97.51

Gallirallus_pendiculentusGallirallus_philippensis_melloGallirallus_philippensis_19403Gallirallus_philippensis_42507Gallirallus_ripleyiGallirallus_wakensisTricholimnas_sylvestris_1896Gallirallus_rovianae_28014Gallirallus_rovianae_826433Nesoclopeus_woodfordi_immaculaGallirallus_owstoniHabropteryx_insignisHabropteryx_torquatus_celebensHabropteryx_torquatus_sulcirosHabropteryx_torquatus_torquatuHabropteryx_okinawaeCabalus_modestusEulabeornis_castaneoventrisGallirallus_australis_australiGallirallus_australis_greyiTricholimnas_lafresnayanusHabroptila_wallaciiGallirallus_calayanensisGallirallus_striatusGallirallus_pectoralisAramidopsis_plateni

1 0.8 0.6 0.4 0.2 0Millions of years ago

NotFlyFlyFlyNotNotNotNotNotNotNotNotFlyFlyFlyNotNotFlyNotNotNotNotNotFlyFlyNot

BioGeoBEARS DEC+J+t12+t21+jt12+m1 on Pacific Rail flightancstates: global optim, 1 areas max. d=0.0935; e=0.1565; j=0.3493; t12=9.2439; t21=3.0112; jt12=0.4168; LnL=−97.51

Linked  trait+dispersal  model  allows  statistical  inferences  not  possible  with  either  individually.  Although  this  case  is  “obvious,”  in  many  datasets,  the  influence  of  traits  on  dispersal  may  be  unknown. •  Flightlessness  is  more  frequent  at  the  

tips,  so  models  that  allow  reversal  are  favored.  Regardless,  the  trait  inference  is  improved  (not  perfect,  see  root).

•  Inferring  m2  is  difficult,  but  fixing  to  0  is  feasible  and  nearly  as  good  (~1  LnL).

•  Founder-­‐‑event  speciation  is  a  key  process,  as  is  founder+flight  loss.

•  The  base  model  maOers  a  lot.  This  must  be  part  of  the  model  choice!

•  These  models  ignore  lineage  extinction,  but  ClaSSE  models  infeasible.

•  Models  with  more  areas  will  be  slower,  but  BioGeoBEARS  is  parallelized.  

Check  out  BioGeoBEARS,  updated  code  at  PhyloWiki:  hOp://phylo.wikidot.com/  

Trait-dependent dispersal models for phylogenetic biogeography, in the R package BioGeoBEARS

Abstract

Acknowledgements: BioGeoBEARS was developed during support by: NSF Award 0919124, and by the National Institute for Mathematical and Biological Synthesis (NIMBioS), an Institute sponsored by the National Science Foundation through NSF Award #DBI-1300426 with additional support from The University of Tennessee, Knoxville. NJM was also supported by the Australian Research Council’s Discovery Early Career Researcher Award #DE150101773, and by The Australian National University.

Nicholas J. Matzke*,1,2 QR  code:  CV,    website,  R  code!

Organism  traits  must  be  important  in  historical  biogeography.  In  particular,  rates  of  dispersal  (both  range-­‐‑expansion  dispersal,  and  jump  dispersal  leading  to  founder-­‐‑event  speciation)  must  depend  to  some  degree  on  traits  such  as  flight  and  its  loss,  and  seed  dispersal  mechanisms  and  the  dispersal  abilities  of  animals  that  transport  seeds.

However,  to  date  no  probabilistic  historical  biogeographical  models  have  been  available  that  allow  geographic  range  and  traits  to  co-­‐‑evolve  on  the  phylogeny,  with  traits  influencing  dispersal  ability.    In  purely  continuous-­‐‑time  Markov  models,  adding  a  trait  is  just  a  maOer  of  doubling  the  size  of  the  rate  matrix;  however,  biogeographical  models  also  include  a  much  more  complex  discrete-­‐‑time  model  describing  how  geographic  range  can  change  during  cladogenesis.  Traits  might  also  influence  this  process.

I  present  an  addition  to  the  R  package  BioGeoBEARS  that  enables  an  evolving  discrete  trait  to  influence  dispersal  ability  for  both  anagenetic  and  cladogenetic  range  change.    This  model  can  be  freely  combined  with  models  adding  jump  dispersal  (e.g.,  DEC+J),  distance  as  a  predictor  of  dispersal  (+x  models,  with  dispersal  rate  multiplied  by  distance^x),  and  other  variants.  I  test  the  model  against  simulations  and  datasets  where  large  evolutionary  changes  in  dispersal  ability  are  highly  likely  (e.g.,  Pacific  rails,  which  have  repeatedly  lost  flight).

BioGeoBEARS trait + dispersal model

Conclusions

1  Discovery  Early  Career  Researcher  Award  (DECRA)  Fellow;  Division  of  Ecology,  Evolution,  and  Genetics;  Research  School  of  Biology;  The  Australian  National  University,  Canberra *nick.ma)[email protected]

References & more info: Use  QR  code  or  go  to:    h4p://phylo.wikidot.com/nicholas-­‐‑j-­‐‑maAke

2  Work  began  at:  National  Institute  of  Mathematical  and  Biological  Science  (NIMBioS)  and  Department  of  Ecology  &  Evolutionary  Biology,  University  of  Tennessee,  Knoxville

Results: Ancestral geography

TwiOer  and  photos  are  OK! @NickJMakke

Study system: “typical” rails of the Pacific

G. pacificus, from Tahiti, “known only from a painting made by Georg Forster on James

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Source: Kirchman, Jeremy J. (2012). Speciation of Flightless Rails on Islands: A DNA-Based Phylogeny of the Typical Rails of the Pacific, The Auk, 129(1):56-69.

Areas&(simplified&from&Kirchman)(Data  provided  by  collaborator  

Jeremy  J.  Kirchman)

Statistical Model Choice

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DEC 3.84 1.94 0 3.31 0.94 1 0 0 0 4 +119.6 37.6 0.0%DEC 0.18 0.00 0.28 5.18 1.65 1 0 0 0 5 +103.9 9.4 0.9%DEC 3.81 2.01 0 2.58 0 1 0 0 0 3 +120.3 36.2 0.0%DEC 0.15 0.00 0.16 2.84 0 1 0 0 0 4 +107.5 13.5 0.1%DEC 2.50 1.69 0 3.60 1.28 1 0 1.00 0 5 +119.4 40.3 0.0%DEC 0.18 0.00 0.82 4.37 1.12 1 0 0.75 0 6 +104.2 13.3 0.1%DEC 3.31 1.98 0 2.57 0 1 0 1.00 0 4 +120.2 38.9 0.0%DEC 0.15 1E+12 0.16 2.84 0 1 0 0.00 0 5 +107.5 16.6 0.0%

DIVALIKE 1.99 0.82 0 5.16 1.48 1 0 0 0 4 +115.2 28.9 0.0%DIVALIKE 0.24 0.00 0.22 5.73 1.71 1 0 0 0 5 +104.1 9.8 0.7%DIVALIKE 2.68 1.50 0 2.60 0 1 0 0 0 3 +117.7 31.0 0.0%DIVALIKE 0.16 0.00 0.10 2.61 0 1 0 0 0 4 +109.3 17.1 0.0%DIVALIKE 2.35 0.90 0 4.38 1.11 1 0 0.91 0 5 +115.3 32.2 0.0%DIVALIKE 0.24 0.00 0.49 5.49 1.49 1 0 0.62 0 6 +104.1 13.2 0.1%DIVALIKE 2.56 1.45 0 2.70 0 1 0 0.58 0 4 +117.7 33.8 0.0%DIVALIKE 0.17 1E+12 0.64 2.81 0 1 0 0.87 0 5 +109.3 20.1 0.0%

BAYAREALIKE 6.40 3.70 0 4.13 2.37 1 0 0 0 4 +129.7 57.8 0.0%BAYAREALIKE 5.34 3.70 0.95 4.14 2.38 1 0 0 0 5 +122.3 46.2 0.0%BAYAREALIKE 3.32 3.91 0 2.67 0 1 0 0 0 3 +130.8 57.2 0.0%BAYAREALIKE 3.34 3.91 0.78 2.67 0 1 0 0 0 4 +123.0 44.4 0.0%BAYAREALIKE 4.36 3.46 0 5.66 2.45 1 0 0.98 0 5 +129.9 61.4 0.0%BAYAREALIKE 0.09 0.16 0.35 9.24 3.01 1 0 0.42 0 6 +97.51 0.0 96.5%BAYAREALIKE 3.36 2.66 0 2.61 0 1 0 0.00 0 4 +132.5 63.5 0.0%BAYAREALIKE 0.07 1E+12 0.14 2.59 0 1 0 0.14 0 5 +103.4 8.4 1.5%

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Gallirallus_pendiculentusGallirallus_philippensis_melloGallirallus_philippensis_19403Gallirallus_philippensis_42507Gallirallus_ripleyiGallirallus_wakensisTricholimnas_sylvestris_1896Gallirallus_rovianae_28014Gallirallus_rovianae_826433Nesoclopeus_woodfordi_immaculaGallirallus_owstoniHabropteryx_insignisHabropteryx_torquatus_celebensHabropteryx_torquatus_sulcirosHabropteryx_torquatus_torquatuHabropteryx_okinawaeCabalus_modestusEulabeornis_castaneoventrisGallirallus_australis_australiGallirallus_australis_greyiTricholimnas_lafresnayanusHabroptila_wallaciiGallirallus_calayanensisGallirallus_striatusGallirallus_pectoralisAramidopsis_plateni

1 0.8 0.6 0.4 0.2 0Millions of years ago

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BioGeoBEARS BAYAREALIKE on traitonly M0_unconstrainedancstates: global optim, 1 areas max. a=1.1954; j=0; LnL=−14.83

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Gallirallus_pendiculentusGallirallus_philippensis_melloGallirallus_philippensis_19403Gallirallus_philippensis_42507Gallirallus_ripleyiGallirallus_wakensisTricholimnas_sylvestris_1896Gallirallus_rovianae_28014Gallirallus_rovianae_826433Nesoclopeus_woodfordi_immaculaGallirallus_owstoniHabropteryx_insignisHabropteryx_torquatus_celebensHabropteryx_torquatus_sulcirosHabropteryx_torquatus_torquatuHabropteryx_okinawaeCabalus_modestusEulabeornis_castaneoventrisGallirallus_australis_australiGallirallus_australis_greyiTricholimnas_lafresnayanusHabroptila_wallaciiGallirallus_calayanensisGallirallus_striatusGallirallus_pectoralisAramidopsis_plateni

1 0.8 0.6 0.4 0.2 0Millions of years ago

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BioGeoBEARS BAYAREALIKE on traitonly M0_unconstrainedancstates: global optim, 1 areas max. a=1.1954; j=0; LnL=−14.83

Standard trait model

Gallirallus_pendiculentusGallirallus_philippensis_melloGallirallus_philippensis_19403Gallirallus_philippensis_42507Gallirallus_ripleyiGallirallus_wakensisTricholimnas_sylvestris_1896Gallirallus_rovianae_28014Gallirallus_rovianae_826433Nesoclopeus_woodfordi_immaculaGallirallus_owstoniHabropteryx_insignisHabropteryx_torquatus_celebensHabropteryx_torquatus_sulcirosHabropteryx_torquatus_torquatuHabropteryx_okinawaeCabalus_modestusEulabeornis_castaneoventrisGallirallus_australis_australiGallirallus_australis_greyiTricholimnas_lafresnayanusHabroptila_wallaciiGallirallus_calayanensisGallirallus_striatusGallirallus_pectoralisAramidopsis_plateni

1 0.8 0.6 0.4 0.2 0Millions of years ago

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BioGeoBEARS DEC+J+t12+t21+jt12+m1 on Pacific Rail flightancstates: global optim, 1 areas max. d=0.0935; e=0.1565; j=0.3493; t12=9.2439; t21=3.0112; jt12=0.4168; LnL=−97.51

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Trait+dispersal model

Flying:

Flightless:

Millions of years ago (Ma) Millions of years ago (Ma)

Millions of years ago (Ma)

Ancestral area estimate: Phillipines & nearby (the 2-area species are actually more widespread)

Founder-events colonizing remote islands are new

areas are associated with rapid

(evolutionarily) loss of flight

ANU  Research  School  of  Biology.  Poster  at  Early-­‐‑and  Mid-­‐‑Career  Conference.  R.  N.  Robertson  Lecture  

Theatre,  Bldg.  46.    Feb.  11th,  2016,  4:45  pm.

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