Introduction to Comparative Methods
BM CC EB What can we learn from a tree?
Net diversification (r) Relative extinction (ε)
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yllaceae yllaceae
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M Ma a lv lv a ac c ea e ae e 6 6 Myrtales Gunnerales Myrtales Gunnerales Bixaceae Bi xa ceae
Cist C ist a 9 ac ce 9 e ae 15 ae 15 Dipt Saxifragales Commelinales Dipt Saxifragales Commelinales er B e o ro B ca ca rpace rp a ce a a r e Poales r e Poales Ca C e r ceae assicaceae ar icacea a ic acea id a id assicaceae ce h rch a rc e O e O
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e eae acea ac Eudicotyledoneae ac Eudicotyledoneae c il ila Sm Sm
Kr Kr ame am e ria ria ce ce ae Rosidae ae Rosidae
Zy Z g ygo op p hy hy llac llac eae e ae ae ceae ia liace Lil Li Q Qu uil laj ill aj ac ac eae Huerteales eae Huerteales
Pol Pol yg yg a 14 a ae laceae la ceae ce 14 e a thi ae thiac an n el ela 26 M 26 M e ea aceae ac on on em em St St Su Su ria ria na Malvales nac Malvales ce e ae ae
e e ea a c ace tha th lan an Commelinidae4 Cyc Commelinidae4 Cycl
ae ceae reace o corea osc os Gunneridae Di Gunneridae Di Fabaceae Brassicales Fabaceae Brassicales
e ea ae tac itace ari r ch cha dro Hy Hydro
e e ea ea ac ac smat mat Ali Alis
e cea a naceae eton eto og g tam Po Potamo
e cea ceae ina ina ag nc ncag Asparagales Ju Asparagales Ju e e ea acea iac eldi eld 21 Tofi 21 Tofi Zygophyllales Petrosaviidae Zygophyllales Petrosaviidae
e eae Aracea Arac Fabales raceae Fabales raceae Pandanales Aco Pandanales Aco
e ceae ea ndia diac rna rnan He He
ae eae iace im nimiac Liliales 3Mon Liliales 3Mo aceae raceae Ros Rosa ace r ce ae Lau ae Lau
E 13 El lae aeagnacea 13 ag nac eae e
e ceae ea rmata rmatac rospe ospe Athe Ather eae e gac gacea rte orte Gomo Gom
eae ae nac nace paru paru Rhamn Si Rhamn Si a ace ceae ae
ae eae hace nt canthac Calyca Caly ae ae ce iace matia omat Eupo Eup
U U lma lmac ceae Nartheciidae eae Nartheciidae
Cann C ann Annonaceae aba abac ceae 17 Annonaceae eae 17
U ceae U ceae rticac dra rticac ndra ea an ea anta e Himant ceae e Him ceae neria neria Alismatales Dege Alismatales Dege
e ae acea ticace Rosales Myristic Rosales Myris ae ae oliace oliace Mesangiospe1rmae Monocotyledoneae Magn Mesangiospe1rmae Monocotyledoneae Magn
ae ae ridace ridace Lacto Lacto
Mor Mor aceae Laurales aceae Laurales
hiaceae hiaceae Aristoloc Aristoloc
An An isophyll isophylle eaceae aceae
raceae raceae Sauru Sauru
Coriar Coriar iaceae iaceae aceae Cucurbi Cucurbi r tac Magnoliales tac Magnoliales eae raceae eae Pipe 7 20 Pipe 20 ae 7 ae Winterace Winterace
ae Cucurbitales Canellace Cucurbitales Canellaceae Bego Bego niaceae niaceae
eae ceae Magnoliidae Chloranthac Magnoliidae Chlorantha
eae ae Piperales Trimeniac Piperales Trimeniace
eae ceae Schisandrac Schisandra
yaceae yaceae Austrobaile Austrobaile Dat Dat iscaceae iscaceae e Cabombaceae Cabombacea
Notho Notho fagaceae Fabidae fagaceae Fabidae e e Nymphaeacea Nymphaeacea
e e Hydatellacea Hydatellacea
e Amborellaceae Amborellacea
Gnetaceae Gnetaceae
e ae Canellales Welwitschiacea Canellales Welwitschiace
Fagaceae Fagaceae
Pinaceae Pinaceae
Myricaceae Fagales Gnetophyta Cupressaceae Myricaceae Fagales Gnetophyta Cupressaceae
Juglandaceae Juglandaceae Angiospermae Nymphaeales Podocarpaceae Angiospermae Nymphaeales Podocarpaceae
Betulaceae Betulaceae
Zamiaceae Zamiaceae
Casuarinaceae Casuarinaceae
Coniferae Cupressophyta Cycadaceae Coniferae Cupressophyta Cycadaceae Cycadales Ginkgoaceae Cycadales Ginkgoaceae
Phrymaceae Phrymaceae Celastraceae Spermatophyta Acrogymnospermae Celastraceae Spermatophyta Acrogymnospermae Lepidobotryaceae Celastrales Lepidobotryaceae Celastrales
Orobanchaceae Orobanchaceae Huaceae Connaraceae Oxalidales Huaceae Connaraceae Oxalidales
Oxalidaceae Oxalidaceae Paulowniaceae Paulowniaceae Brunelliaceae 300 Brunelliaceae 300 ae Elaeocarpace Elaeocarpaceae
Lamiaceae Lamiaceae Cunoniaceae Cephalotaceae Cunoniaceae Cephalotaceae eae e Humiriac Humiriacea
eae eae Achariac Achariac
Lacistemataceae Lacistemataceae
Salicaceae Salicaceae
T Thoma homandersiac ndersia Malpighiales eae Malpighiales ceae
Verbenaceae Verbenaceae
raceae raceae Passiflo Passiflo
Lenti Lenti bularia bulariacea ceae e
Violaceae Violaceae
e ceae Goupiacea Goupia 200 eae 200 ceae thac Ixonan Ixonantha
ceae Irvingiaceae Irvingia
Acanthace Acanth ae aceae
e iacea Clusiaceae Clus
ae ae ce etiace Bonnetia Bonn
By By blida blidacea ceae e
ae ae lace phyllace hyl Calo Calop
Bigno Bigno niace nia ae ceae
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Pedaliac Pedaliac eae eae
M M artyniaceae artynia eae ceae ricaceae ricac Hype Hype
ae e race racea dend dend Picro Picro Scrophula Scro ph r ular iaceae iaceae
e ae cea tha anthace llan Phyll Phy
P 100 Pla lant nta aginacea gin 100 ac e eae
Stil St b ilb Lamiales aceae Lamiales aceae e ceae ea ia hiac pig Malpigh Mal
Ges Gesn ne riace eria ae ceae ae eae nace tinac Elati Ela
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Cal C c alc e eola e eola r a r acea iace ace iace ae a Ochn 22 Ochn 22 e Te Tet Solanales trachond Solanales rac hond raceae rac 19 19 eae e eae racea rac Pe Pe
O O leacea leac e Superasteridae 24e r ε Superasteridae 24ae Plocospe B Plocospe B or or agi ag nac ina e cea Berberidopsidales10 ae Berberidopsidales10 e
Hydrol H ydrol S eacea S eac phenoclea phenoclea ea e e Mo Mo nti n n ceae tin cea iaceae iacea e r Santalales Pentapetalae rmataceae 0.00 0.00 Santalales Pentapetalae e mataceae Co Co n n volvu volvu la lac ceae eae ae aceae ace or or ph zoph zo Rhi Rhi
eae laceae xy xylac ro o yth ythr Er Lamiidae Er Lamiidae
e e ea acea ac nd Pand Caryophyllales Pa Caryophyllales e e acea Sola Sola Lin n Euphorbiaceae Linacea na a ce cea Euphorbiaceae ae e
ae eae ce aca placac opl eae tr eae entro en C C pac opac no la Balan Ba
e eae a ace al et petalac ap ich Dicha 16 D 16 ae ae iace iace on igon ig Tr ae Tr ae ce iace ia Gentianales on Gentianales phron phr Eu Eu Garryales 0.05 Garryales A A po poc cy yna na e ceae ea ce ac ae ni le Ctenolophonaceae Dilleniaceae 0.20 Ctenolophonaceae Dil
e cea laca Olacaceae Dipsacales8 O Dipsacales8
laceae ta Santalaceae Bruniales San Bruniales
ae ceae ce a Cornales ili Cornales p Opilia Apiales18 O Apiales18 e a eae e Genti c Ge pfia ntian epfiac a e nac o ac Scho Asteridae ea Sch Asteridae e ae e
e ae e ac hacea th n rant ra Lo Campanulidae G Lo Campanulidae G el el s s emiac emi aceae eae
23 L L o 23 o gan g e e a a n ia ia ce ce c a ae e a id e a e e Ericales ps ac Ericales ic opsidaceaxicaceae o x rid o 0.10 rid to xt x e e e e rb a A e A e Berbe B c aceae ia n eni e k k ran ysobalanaceae ran ysobalanaceae F F ae r ace r ricaceae ric ama ama T Aquifoliales T Aquifoliales Ch Ch e ea ac on g ly olygonaceae o P Asterales 0.40 P Asterales
R R e ub e ubi a iaceae e ac ac e nacea in ae g bagi ba lum Plum P
e a eae e c rac ra e s o Drose Dr e a eae e e c a a e c th a e n ll e yllaceae e y a h p h e e p c Nepenthac acea N o a s d e ae la a e c e Drosop oclad c Dro o c e a r a a ll t ll e e y is y c a h aceae c h a e e p r n p r a Ancistr aceaeeae A o d ac nd c n i ce e n e a a d n e io o nds e Dionco dode ondsi D d o ysenac b ys h a m h eiac peiaceae h m p P i P o Rhab Simm R S r ste Astero 0.15 A V V ahliacea ahl ia c eae Ga Ga e rr rr y e y Eucom aceae E a ea u c e ac co ll a yllaceae y mm e h mia h op op ia y y r Icacinacea cea r Ic ce a ac a Ca C e inac e
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A A piac p ia Oncothecaceae c e e a Oncothecaceae a e e
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c e c a a a eae r e r pa 0.60 c e pa ta a ce e c a c ea matac aceae a m a r e r e e e p im spe Lime s L o n gno g e Ste St
A A r r al al i i ace e a a cea e ceae ae c e oa oa z z Ai Ai
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ce ia tiace Ad t Ad n o o o x x Mon a M acea ce e ae a ae e P P e a a r r ac ac alinace alinac T Co r T C r yphia y o p lu l u h me m i B B a r c r e c u lli e A B u ll e n n ia a a a ia i cea e a c e cea ce e a e a e e e
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n n g g e ce e e g e g t i t i C C C C o a o a e i e i ac a a a m a m a a
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S S r r aceae aceae Cactaceae Cactaceae
Aquifoliaceae Aquifoliaceae
Ericaceae tisiaceae Ericaceae tisiaceae r r
Cu 0.30 1.00 Cu
Balsaminaceae Balsaminaceae making trees is hard.... because trees are information-rich the coming age of the megaphylogeny* the coming age of the megaphylogeny*
* trees that are too big for your brain ensembl
Why do you want to make or use trees? What do you hope to learn? What can you do?
• Diversification (speciation and extinction) • Character evolution • Characters and diversification • Biogeography • Testing complex evolutionary models Diversification (speciation and extinction) Sharks Polypteriformes Chondrostei Amiiformes Elopomorpha Osteoglossomorpha Ostariophysi Clupeomorpha Esociformes Salmoniformes Galaxiiformes Stomiiformes Osmeriformes Argentiniformes Aulopiformes Myctophiformes Beryciformes+Stephanoberyciformes Ophidiiformes Percomorpha Scombridae Polymixiiformes Lampriformes Percopsiformes Zeiforms Latimeridae Dipnoi Gymnophiona Anura Caudata Monotremes Marsupials Boreoeutheria Afrotheria+Xenarthra Sphenodon NongekkoSquamates Taxa Gekkos 1 Pleurodira 5 Cryptodira Crocodylinae 10 Gavialidae 50 Camininae 100 Alligatorinae 500 Tinamiformes 1000 Struthioniformes 5000 Neoaves 10000 Galliformes 20000 Anseriformes 500 400 300 200 100 0 MYA
Character evolution ● ● ● ● ● ● ● ● ● ● ● ● ● ● ● ●
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● ● ● ● Siren intermedia
Siren lacertina
Pseudobranchus striatus
Pseudobranchus axanthus
Onychodactylus fischeri
Ranodon sibiricus
Paradactylodon mustersi
Paradactylodon gorganensis shift direction
Pachyhynobius shangchengensis
Hynobius formosanus
Hynobius sonani
Hynobius retardatus
Hynobius boulengeri
Hynobius kimurae
Hynobius abei
Hynobius lichenatus Hynobius tokyoensis 1.00 Hynobius nigrescens
Hynobius takedai
Hynobius stejnegeri
Hynobius katoi
Hynobius chinensis
Hynobius yiwuensis
Hynobius amjiensis
Hynobius guabangshanensis
Hynobius leechii 0.80
Hynobius okiensis
Hynobius tsuensis
Hynobius dunni
Hynobius nebulosus
Hynobius hidamontanus
Salamandrella keyserlingii
Liua tsinpaensis Liua shihi 0.50 Batrachuperus yenyuanensis
Batrachuperus tibetanus
Batrachuperus pinchonii
Batrachuperus londongensis
Cryptobranchus alleganiensis
Andrias japonicus
Andrias davidianus
Necturus punctatus
Necturus maculosus 0.20
Necturus lewisi
Necturus beyeri
Necturus alabamensis
Proteus anguinus
Rhyacotriton variegatus
Rhyacotriton cascadae
Rhyacotriton kezeri Rhyacotriton olympicus 0.00 Amphiuma tridactylum
Amphiuma means
Amphiuma pholeter
Desmognathus auriculatus
Desmognathus brimleyorum
Desmognathus monticola
Desmognathus fuscus
Desmognathus welteri
Desmognathus ochrophaeus point.seq -0.20
Desmognathus imitator
Desmognathus marmoratus
Desmognathus quadramaculatus
Desmognathus aeneus
Desmognathus wrighti
Phaeognathus hubrichti
Aneides flavipunctatus Aneides ferreus -0.50 Aneides lugubris
Aneides hardii
Aneides aeneus
Ensatina eschscholtzii
Hydromantes platycephalus
Hydromantes italicus
Karsenia koreana Plethodon albagula -0.80 Plethodon glutinosus
Plethodon cylindraceus
Plethodon teyahalee
Plethodon cheoah
Plethodon shermani
Plethodon meridianus
Plethodon amplus
Plethodon montanus Plethodon ouachitae -1.00 Plethodon caddoensis
Plethodon jordani
Plethodon metcalfi
Plethodon kentucki
Plethodon yonahlossee
Plethodon dorsalis
Plethodon ventralis
Plethodon angusticlavius
Plethodon welleri
Plethodon punctatus
Plethodon wehrlei
Plethodon hubrichti
Plethodon nettingi
Plethodon richmondi
Plethodon shenandoah
Plethodon cinereus
Plethodon hoffmani
Plethodon serratus
Plethodon idahoensis
Plethodon vandykei
Plethodon neomexicanus
Plethodon dunni
Plethodon vehiculum
Plethodon elongatus
Eurycea tridentifera
Eurycea neotenes
Eurycea sosorum
Eurycea nana
Eurycea rathbuni
Eurycea wilderae Bolitoglossinae
Eurycea junaluska
Eurycea bislineata
Eurycea cirrigera
Eurycea longicauda
Eurycea lucifuga
Eurycea quadridigitata
Eurycea spelaea
Eurycea tynerensis
Eurycea multiplicata
Pseudotriton ruber
Pseudotriton montanus
Gyrinophilus porphyriticus
Stereochilus marginatus
Batrachoseps major
Batrachoseps attenuatus posterior rates
Batrachoseps wrightorum
Cryptotriton nasalis
Cryptotriton alvarezdeltoroi
Nyctanolis pernix
Dendrotriton rabbi Dendrotriton cuchumatanus rep(-0.5, length(point.seq)) Nototriton barbouri
Nototriton richardi Nototriton guanacaste 3.011 Nototriton picadoi
Nototriton abscondens
Oedipina uniformis
Oedipina pacificensis
Oedipina poelzi
Oedipina pseudouniformis
Oedipina cyclocauda
Oedipina gephyra
Oedipina elongata 0.519
Oedipina parvipes
Oedipina complex
Oedipina savagei
Bradytriton silus
Thorius troglodytes
Thorius dubitus
Chiropterotriton magnipes Chiropterotriton chondrostega 0.090 Chiropterotriton multidentatus
Chiropterotriton priscus
Chiropterotriton lavae
Chiropterotriton dimidiatus
Chiropterotriton arboreus
Pseudoeurycea scandens
Pseudoeurycea galeanae
Pseudoeurycea cephalica
Pseudoeurycea naucampatepetl 0.015
Pseudoeurycea gigantea
Pseudoeurycea boneti
Pseudoeurycea bellii
Pseudoeurycea goebeli
Pseudoeurycea brunnata
Pseudoeurycea exspectata
Pseudoeurycea rex Pseudoeurycea smithi 0.003 Pseudoeurycea anitae
Pseudoeurycea cochranae
Pseudoeurycea gadovii
Pseudoeurycea melanomolga
Pseudoeurycea longicauda
Pseudoeurycea robertsi
Pseudoeurycea altamontana
Pseudoeurycea unguidentis
Pseudoeurycea saltator point.seq 0.000
Pseudoeurycea juarezi
Pseudoeurycea werleri
Pseudoeurycea mystax
Pseudoeurycea nigromaculata
Pseudoeurycea lynchi
Pseudoeurycea leprosa
Pseudoeurycea firscheini Lineatriton lineolus 0.000 Parvimolge townsendi
Ixalotriton parvus
Ixalotriton niger
Bolitoglossa hartwegi
Bolitoglossa rufescens
Bolitoglossa occidentalis
Bolitoglossa odonnelli
Bolitoglossa lignicolor
Bolitoglossa yucatana 0.000
Bolitoglossa striatula
Bolitoglossa mexicana
Bolitoglossa platydactyla
Bolitoglossa flaviventris
Bolitoglossa macrinii
Bolitoglossa riletti
Bolitoglossa hermosa Bolitoglossa rostrata 0.000 Bolitoglossa helmrichi
Bolitoglossa engelhardti
Bolitoglossa lincolni
Bolitoglossa franklini
Bolitoglossa porrasorum
Bolitoglossa longissima
Bolitoglossa decora
Bolitoglossa synoria
Bolitoglossa celaque
Bolitoglossa morio
Bolitoglossa dunni
Bolitoglossa diaphora
Bolitoglossa conanti
Bolitoglossa carri
Bolitoglossa dofleini
Bolitoglossa subpalmata
Bolitoglossa pesrubra
Bolitoglossa gracilis
Bolitoglossa marmorea
Bolitoglossa minutula
Bolitoglossa cerroensis
Bolitoglossa schizodactyla
Bolitoglossa colonnea
Bolitoglossa sima
Bolitoglossa biseriata
Bolitoglossa palmata
Bolitoglossa peruviana
Bolitoglossa altamazonica
Bolitoglossa medemi
Bolitoglossa adspersa
Hemidactylium scutatum
Pleurodeles waltl
Pleurodeles poireti
Tylototriton vietnamensis
Tylototriton hainanensis
Tylototriton asperrimus
Tylototriton wenxianensis
Tylototriton kweichowensis
Tylototriton verrucosus
Tylototriton taliangensis
Echinotriton andersoni
Echinotriton chinhaiensis
Pachytriton brevipes
Pachytriton labiatus
Paramesotriton deloustali shift probability
Paramesotriton fuzhongensis
Paramesotriton guangxiensis
Paramesotriton hongkongensis
Paramesotriton chinensis
Paramesotriton caudopunctatus Paramesotriton laoensis rep(-0.5, length(point.seq)) Cynops ensicauda
Cynops pyrrhogaster Cynops cyanurus 1.000 Cynops chenggongensis
Cynops orphicus
Cynops orientalis
Neurergus strauchii
Neurergus microspilotus
Neurergus kaiseri
Neurergus crocatus
Ommatotriton vittatus
Ommatotriton ophryticus 0.875
Calotriton asper
Calotriton arnoldi
Triturus carnifex
Triturus dobrogicus
Triturus karelinii
Triturus cristatus
Triturus marmoratus Triturus pygmaeus 0.750 Lissotriton boscai
Lissotriton italicus
Lissotriton helveticus
Lissotriton montandoni
Lissotriton vulgaris
Mesotriton alpestris
Euproctus platycephalus
Euproctus montanus
Notophthalmus viridescens 0.625
Notophthalmus perstriatus
Notophthalmus meridionalis
Taricha rivularis
Taricha sierrae
Taricha torosa
Taricha granulosa
Mertensiella caucasica Chioglossa lusitanica 0.500 Lyciasalamandra atifi
Lyciasalamandra helverseni
Lyciasalamandra antalyana
Lyciasalamandra billae
Lyciasalamandra fazilae
Lyciasalamandra luschani
Lyciasalamandra flavimembris
Salamandra salamandra
Salamandra atra
point.seq 0.375
Salamandra corsica
Salamandra lanzai
Salamandra infraimmaculata
Salamandra algira
Salamandrina perspicillata
Salamandrina terdigitata
Ambystoma gracile Ambystoma maculatum 0.250 Ambystoma macrodactylum
Ambystoma californiense
Ambystoma rosaceum
Ambystoma dumerilii
Ambystoma rivulare
Ambystoma lermaense
Ambystoma granulosum
Ambystoma altamirani
Ambystoma velasci 0.125
Ambystoma taylori
Ambystoma tigrinum
Ambystoma ordinarium
Ambystoma andersoni
Ambystoma flavipiperatum
Ambystoma mexicanum
Ambystoma amblycephalum Ambystoma jeffersonianum 0.000 Ambystoma laterale
Ambystoma annulatum
Ambystoma cingulatum
Ambystoma bishopi
Ambystoma opacum
Ambystoma barbouri
Ambystoma texanum
Dicamptodon aterrimus
Dicamptodon ensatus
Dicamptodon tenebrosus Dicamptodon copei photos from wiki commons Characters and diversification Silvestro et al. 2013 Evolution Biogeography Ree and Smith 2008 Sys Biol Testing Complex Evolutionary Models Price et al. 2014 Nature