Palestra Swansea Abril 2017

Biogeography and macroevolution of Atlantic Forest anurans Diogo Borges Provete Federal University of São Carlos at Sorocaba Brazil [email protected] | diogoprovete.weebly.com 1

Hello everyone, thanks for comming. First of all, I’d like to thank Laura for the invitation and for the introduction as well. Well, first when Laura invited me to give this talk I first thought I could maybe talk about one of my papers that came out recently. But then I thought, nah, that’s too easy… and I decided to show you a working in progress, which are results I got from my previous post doc in Gothenburg (Göteborg), Sweden. For the 2nd half I’ll show the questions I’m trying to answer right now with the current post doc project, and hopefully receive some feedback. So as I said I did a post post in Gothenburg in 2015, spent 6 months there (short project), and then I moved back to Brazil and started four months ago with this new project.

Background | Atlantic Forest | Partial Results | Current project | Conclusion 1. Background – The Atlantic Forest (Hotspot) • Models for explaining biodiversity in AF • Anuran amphibians diversity in AF 2. Results from previous post doc at Göteborg, Sweden • GeoSSE – rates of speciation and extinction between refugia/nonrefugia • Phylogenetic community structure • Partitioning PhyloBetaDiversity – nestedness-resultant and turnover 3. Quick overview of the questions of my current project • Linking macroevolutionary and ecological mechanisms to understand current distribution of vertebrate diversity • Phenotypic hotspots • Equilibrium/non-equilibrium mechanisms • Adaptive landscape of body size and clutch size evolution • Role of climatic niche on the rate of evolution of body size in a continental fauna 2

Background | Atlantic Forest | Partial Results | Current project | Conclusion

Almost a third (26.6%) of the Brazilian population is concentrated along the East cost. 11.4 - 16 % of the original coverage remains Many endemic species ~ 92% of AF amphibians are endemic to this biome

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Ribeiro et al. 2009 Biol. Conserv.


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MODELS TO EXPLAIN AF BIODIVERSITY

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Climatic stability throughout the Pleistocene Imprint on present species composition and intraspecific lineage diversification Climatic refugia may have contributed to species diversification Phylogeographical studies found expanding populations in southern AF

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Carnaval et al. 2009 Science, Carnaval & Moritz 2008 J. Biogeog


odelled ranges of the Atlantic forest under narrow definitions, 6 ka bp, and 21 ka bp climate using d MAXENT modelling approaches. Scale m.

ps obtained under current, 6 ka bp and 21 ka bp yielded by both BIOCLIM and MAXENT models. ned area map depicts areas which were potentially forests during the climatic oscillations of the late and Holocene. Refugia or historically stable areas d as those grid cells for which forest presence was ll models and time projections.

atic model validation and comparisons rns of endemism and with raphical data

ast ranges were qualitatively evaluated against ypes inferred from pollen records at sites located urrent or predicted range of the Atlantic forest, as areas currently occupied by the neighbouring

Historically stable areas Atlantic forest broader definition

Pernambuco refuge S. Francisco river Bahia refuge Doce river

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Figure 3 Summary maps of historically stable areas for the Carnaval & Moritz 2008 J. Biogeog Atlantic forest under narrow and broader definitions, obtained by summing across BIOCLIM and MAXENT output grids for forest absence/presence under current, 6 ka bp and 21 ka bp climatic scenarios. White lines depict the Sa˜o Francisco (top) and Doce (bottom) rivers. The arrow indicates the small refuge predicted at the boundaries of the states of Espı´rito Santo and Rio de Janeiro.

21 ka bp periods between latitude 3! S and 33.7! S and longitude 58! W and 34.8! W, excluding sites located in the Amazonian forest biome. For the 6 ka bp period, records were available from 11 localities, representing 10 spatially unique points at 30¢¢ resolution; for the 21 ka bp period, data existed for four sites (Tables 1 & 2, Fig. 4). Hypothesized refuge areas were compared with published patterns of species diversity and endemism for birds (da Silva et al., 2004), mammals (Costa et al., 2000), butterflies (Tyler et al., 1994), bamboo species (Soderstrom et al., 1988) and woody plants (Prance, 1982). They were also contrasted with molecular data available for the local fauna (see below). In both instances, we would expect to find evidence of high


Geographic barriers (e.g., Mountain ranges of southeast) Allopatric and parapatric speciation (most) Anurans are highly dispersal limited, depend on humidity Many small-ranged species in the AF Strong environmental gradients that could foster ecological specialization

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Villalobos, Provete et al. 2011 PLoS One, Thome et al. 2008 Mol. Ecol


”Atlantis" Forest Hypothesis Forest expansion during LGM into the [now submerged] continental shelf Against the ”refugia hypothesis" DNA coalescence model for small mammals Increase of suitable areas all most species

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Regionalization based on taxonomic species composition Marked differences in species composition

Figure 1. Major Atlantic Forest eco-regions, modified from the World Wildlife Fund designations (A): AMF = Araucaria Moist Forests; APF = Alto Parana´ (semideciduous/deciduous) Forests; BCF = Bahia Coastal (moist) Forests; BIF = Bahia Interior (semidecidual/decidual) Forests; PCF = Pernambuco Coastal (moist) Forests; PIF = Pernambuco Interior (semideciduous) Forests; and SMCF = Serra do Mar Coastal (moist) Forests. Biogeographic regions based on the anuran fauna generated through k-means clustering with v-fold cross-validation (B). doi:10.1371/journal.pone.0104130.g001

Vasconcelos et al. 2014

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and what we up [26], with,range at least for amphibians, these species along the atlantic forest that we try to explain levelend of error maps represent the areas where a marked perspective,diﬀerences however, they in may function composition very well at grains particular species can be expected to occur, and it will be expected to be found only in suitable habitats within these areas [6]. Thus, overprediction is an inherent methodological limitation of these kinds of range maps [27]. Within a macroecological PLOS ONE | www.plosone.org

greater than 50650 km [28]. That is to say, although the IUCN amphibian maps might include either over- or underpredictions [26], using range maps is presumed to be as reliable as more refined information regarding a given species distribution (e.g., 3

August 2014 | Volume 9 | Issue 8 | e104130


[WORK IN PROGRESS]

WHAT WE DID SO FAR

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Testing the role of pleistocene climatic refugia on anuran amphibian species distribution in the AF Partitioning phylobetadiversity (turnover and richness difference components) Phylogenetic community structure Differential diversification rates - GeoSSE

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Atlantic Forest Refugia (Carnaval and Moritz model)

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Background | Atlantic Forest | Partial Results | Current project | Conclusion Rhinella fernandezae Rhinella hoogmoedi Rhinella margaritifera Rhinella crucifer Rhinella ornata Rhinella abei Rhinella jimi Rhinella cerrade nsis Rhinella schne ideri Rhinella icteric Rhinella acha a vali Rhinella Rhinella rubescens Rhinella arenarum henseli Rhinell Rhinell a pygmae Rhinel a granulo a Pseudola bergi sa Pseudo paludicol paludi a my Pseud Pseu opalud cola falcstacalis Pseu dopalud icola min ipes Ple dopalud icola sal eira icola tica Ple urodema Physurodema bib ternetzi ala dip roni Phys ala emus aglolister Phys uirrei ala emus an Eupe gren mphemus alb Phys Phys alae ix natte ifronssis alae mus Phys re lis m Ph alae us ei ri mus jord Ph ysalae mus evananensis Phyysalae er ge P sa m P hysa laem us m ikae listai P hysa laem us si axim P hysa laem us cr gnife us P hysa laem us om r P hysa laem us eryt biei P hysa laem us atla hros P hysa laem us deimntic P hysa laem us mar aticus P hysa la us sp m us Phhysa laeemus cicainigeroratus m ys la P ca da P hys ala em us h m P hys ala em us en aca P hy ala em us kro selii n P hy sala em us na yeri P hy sala em us ma nus P hy sala em u m cu P hy sa e u s o ore live P hy sa lae mu s in lfers ira ntr P hy sa lae mu s o sp ii e is P hy sa lae m s bte era P hy sa lae m us rio c tu Ph hys salalae mu us b cae grantus s m s ili te de P y a P h sa la em us so g ns is P h ys la em u b a on P h ys ala em u s g arr res ige S hy ys ala em u s a ra io i rus P c s ala e u s lb cil i P a yth ala e m s cu on is Pa ararate rop emmu us ir bok vieri ota tu ra te lm hry us s ru ro erm s te lm a s c p ratu a lm a to s en es s n ni ato tob biu aw tra tris s biu ius c aya lis s ga ard e po ig os ec ea oi ilo e ga ste r

Rhinella dorbignyi Dendrophryniscus brevipollicatus Dendrophryniscus berthalutzae Dendrophryniscus krausae Dendrophryniscus leucomystax stawiarskyi Dendrophryniscus s proboscideus Dendrophryniscu cus carvalhoi is Dendrophrynis bucens Frostius pernam ophthalmuss peritu Frostius erythr ryniscus moreirae Melanoph ryniscusadmirabilis Melanoph ryniscusfulvoguttatus ilis Melanoph ryniscus spectab s Melanophphryniscuscus dorsalius Melanoanophrynis atrolute sis cus Mel phrynis cambaraen osus Melano iscus granul ioi ophryn macroiscus alip ns Melan ryniscus ifro x ryn oph Melanoph cus tum ple i nis Melan s sim ophry niscu rvalhos Melanlanophry nus ca canu Me ontophry s ameri ltripesi cu pio nu Od tophry hrynus s aliceps sti Odon ontop nioglottu ta Od ge phrys cribguttatris su Macro hi erato ys ys palus hirc Proceratophr ophr ys sc hringi Proc oceratatophr moecururus Pr ocer hrys ys ep Pr atop ophr laticboiei a ys er at ProcProceratophrophrysiculaton nd og us er at ProcProcers appeanop om ii ry mel lost viot toph rys phylrys pabraunia os s cera toph rys Pro cera toph ratophphrybigibbelinois Pro cera roce rato rys s av losu ix Pro P Procetoph phry venu itatr us im e cera ratoalus s pha tlas ProProce eph phalueso lus alatusoi e m a chyc yc s ph acu ard us Tra Trachphalu yce romibern pidingi i le ig yce Trachs n lus d lus reen annoi ch a n i lu ha ha ph s g erm ru fi Tra ep ep ce nti ok b orf s yc yc hy a b on sd e s ch ch c om on od ng aeth su s Tra Tra Traoryth nod henla la yrin culo olukyi C pherasp tihy s g ma luteuts reri x s a o te s s a e ara Ap Itap dy yte dyte s k uch ystatus Ap yllo od llo yte w m ta tris s Ph yll y d s no nc os u s Ph Phhyllodyte ela pu vir ulosatu oi P yllo s m tes bre rc in lm la ii e m e o y Ph dytellod ytes tub cu ed nic sch is o y d s a s la a tr s yll Ph llo te s te p m lus u Ph Phy ody dyteody hus ara pa surd yll llo yll nc s c us us Ph Phy Ph rhy hu nch ch o c y n en yn rh rhy ha orh no no Sp aen hae ae h p ph Sp S S

Sp Sp S p Sp Sp hae hae ha ha ha no no en en en rh rh orh orh orh yn yn yn yn yn chu chu ch ch ch s s us S u u pa pra m Sc Scin cin s bros o ulo sin irim ina Sc ax ax m rop alv u x o ina pe faiv eli hil in s Sc btr x p rpu ov colaus i in ia e s ic Sc Sc Scin ax ngurereillushi ina ina ax be la ca S S x x b rt tu Sc cin cina cathalbicellohae s ina ax x a a ni x c lit ri n S lo arn to na s Sci S ScinScin cin ngili evareuse na cin ax ax ax ne llii x fu ax p a me us sc ac ach riad lloi om um yc ne S ScicinaxScin argininaturus a n Sci ax trcuspx simatuss a id n S ax pic atuilis Sci nax S cinaxcald heiro s ca cina lit aru i Sci Sci mpo x dutoralism nax na ssea ar tei fla x hi brai SciScina voguem alis nax x cu ttatu Scinluizot ricicas Scin Sc Scin ax agavioi ax ar inax ax ha ilis Scingyreoratratuyii Scina ax na natu s x x− sic s ScinaScina signatuus x fus x hum s ilis co Scina x rizvarius Sc Scinax ibilis er Sc inax pinalt ima Scinainax cre Scina x angretatus x ns Scina granulatuis s Sci x car oi Scinaxnax aurdos atu ins Scinax peratu s Scinax kautskysi Scinaxmachadoi eury Scinax dice brie Scinax jureini a Scinax heye ri x ranki Scinax Scina squalirostri s Scinax v−sign atus Scinax crospe dospilu Scinax arduouss Scinax uruguayu Pseudis platensiss Pseudis bolbodactyla

Green = occur in both regions Blue = refugia endemic Red = endemic to non-refugia

Megaelosia lutzae Megaelosia jordanensis Megaelosia apuana Hylodes nasus

Hy ps H yp ib Hy oa H sib s y

Hy psib alb ps oas ib Hy Hy psib oa om oa pa Hy H psibps oa s p arg s lurda ps yp o ibo s p ulc ina n lis d h ib s a a H Hy o ib s s ra e tu ii Hy H yps psibas oas sec becsinullus s ps yps ib o ma ca ed ke s ibo ib oa as rg in en ri H Hy yp Hyp as oas s gu bis ina gua s sib s se a e ch tu p Hy sib oa ibo m tla nth off s p o s a ig n i Ap Hyp sibo as le fre s jo uttaticueri Ap las sib as p ica aq tu s Ap Apla lastotodis oas polytolin nec uini s la st d la ta e a Ap stod odis iscucus p tistr enatus e iu la is cu s e Ap stod cus s coflumrviriatus s Ap Ap lasto iscu albo ch ine idis last last dis s ca sig ran us A odis odis cus llip natuae cus cus mu ygiu s Apl plasto as leu sib sicu s AplAplas todi discus ila copy tu s asto todi scus ca scus eh vi giuss Apl discus co rh Apl Apl as ib wey ar la Phry asto astoditodisc irapi golddti no disc sc us ta ti Ph med us us eu arildnga geni ae Phry ryno usa albofre m bo Phry Phrynomededusa kermnatusoi nom nom usa fim anni m br ed ed Phyll usa apusa vaarginaiata nz ta om PhyllPhyllom eduspendicuolinii omed edus a ba lata Ph us a ihehiana Phyll yllomeda tetra ringii omed us ploide Ph usa a distin a burm Phylloyllomedu cta me sa eister i Phyllo Phyllodusa no sauvag medu ii rde me sa hyp dusa stina ure PhyllomPhyllome ochonaz drialisa edusa dusa roh megac Phyllom dei edusa ephala Phasma ayeaye Phasma hyla guttata hyla jand Phasma hyla aia Phasmacochranae Phasmah hyla yla spec exilis tabilis Phasmahy la timbo Hylomantis Hylom granulosa Limnomedusa antis aspera macroglossa Megael goeldii Megaelosia osia boticariana Megaelosia massarti Megaelosia bocainensis

531 species

sis us s en ss lu rin lo n ta g ata ni cty ca olito ng oh da ni us s b ca cks ero ofe ph u a e uth sh is m ph us iz le g ns ra m h us e irin lo ra mp ph hus s d edreueli c yc lo ra C yc clo rammp phu us migalae ius nsis C y clo ra m ph us v b ne C y clo ra m ph us du a ni C rg e lo is Cy yc clorarammph hus s o dus s C y clo ra mp hu s ulu oi iens C y lo ra mp hu arv alh ce m tus a C yc lo ra p p rv ora ru C yc lo ram us ca b tzo alm sis iroi C yc lo en us us lu mip en be C yc ha en ph us se sili ari C ac ha m ph us bra and oi Z ac lora m ph s ir alh Z c ra hu m rv i Cy yclolorammp hus s ca ustoegeri s C yc lora mp hu s fa ejn C yc lora mp hu st er irensi C yc ra mp hus asp de sus C yclo ra mp hus ban ino C yclo ra mp us fulig irim C yclo ra ph us im si stes C yclo ram ph us ju au na oh ko osus C yclo ram ph ul C yclo ram phus rhya C yclo ram phus gran C lo m us loraramphiliaris is Cyc m lo a m xatil Cyc panu Cyc op a sa i Thororop a lutzegatymlitanaus Th orop a m tropo aene tomusi Th orop a pe lus chys audi Th orop acty s tra udichm Th ssoddactylu s gatzoru Croosso dactylu s lu idti Cr osso dactylu schm r Cr osso ctylus dispamaschii Cr ossoda ctylus cara is Cr soda ctylus grand inus Cros soda ctylus cyclosp Cros soda lus dantei Cros sodacty lus Cros sodactyi Cros nsi es ua Hylod es merte eri Hylod es heyber Hylod es gla i Hylod s fred es Hylode s phyllod Hylode asper s Hylode ornatus esi Hylodes magalha Hylodes pipilans Hylodes sazimai Hylodes amnicola Hylodes vanzolinii Hylodes atus Hylodes perplic cinus Hylodes dactylo Hylodes babax naetes Hylodes charadra Hylodes regius Hylodes lateristrigatus Hylodes meridionalis Hylodes otavioi Hylodes cardosoi

Pseudis paradoxa Pseudis fusca Pseudis minuta Pseudis cardosoi Lysapsus limellum Xenohyla truncata Xenohyla eugenioi Dendropsophus cruzi seniculus Dendropsophus jimi Dendropsophus s oliveiraii Dendropsophu hus ruschi ns Dendropsop phus elega Dendropso phus dutrai nus Dendropso domeridia ophus pseu us elianeae Dendrops Dendropsoph us werneri soph utus Dendropsophus min neri bran drop s Den psophu dulus Dendrophus rubicun borni pso ophus san anus dro Den ndrops us meridi ae lutz De opsoph ber tha tuss Dendropsophus s bipuncta nanu Dendr ropsophusophussoaresi Dend Dendropsophus ancepsri us hdere rop ai Dend ropsoph us na lim Dend soph ophuss rheas ps ophu op rop icr ri Dend Dendro ropshus mgie sle nd ns De psop hus cipie ae ro op Dendndrops hus destuder adi i De ropsop hus haddvais op us ai Dendndrops sophphus noreng si De dropopso alva ar tinchii Denendr hyla la mmas ea D anno nohy ra tart ae rm man la cala as cian ta hy hy lu da x er Boke Bok annoanno hyla rcum hyla s erm erm no la ci hyla eudi oi hy no ps alh ei Bok Bok erman no do rv g an Bok man erm pseula ca lancolae la xi er ok ohy sa nia a Bok Bnohylannohy nn la lca ne an rma a ohy vu ahe zaea erm oke okermann hyla yla anu os ta Bok B B rm nno nohyla n luctu na eai ig ke a n Bo rm rma nnoh hyla res ouvydra a ke la g o c Bo Boke rma ann la c yla leps ipo lum ke hy oh c it a sis Bo okermnno ann hylayla ibymb ctatu s B rma rm nno oh s c un oenatus s e ke ok a ann oa s p cip e icu li Bo B kermerm psib iboa as milin ph mba ps Bo Bok Hy yps sibos se ogra po nice llae s H yp oa ge as ra ste tatu s H ib as ibo as as c alu us ps ibo yps ibo o puneph ian ans s y ib s s H p H p ps o oc go pit sti er lb Hy Hy Hy s a ten as cre exa fab oa s o s s s ib as sib oa oa oa ps ibo yp sib sib ib Hy yps H yp yp yps H H H H

i nn i ii i a n tz to rm h a lu in ke so ir s s p bo eck que biulode es s iz nti ato ty lod de ma a lm ac ty lo s un a te od dac cty biu jura cari la ra ss o a to a u Pa ro ss sodlma ra araana ei acty C ro s te e ra n om ed i C ro ra om e ra th la x ann C a en om e ra hy ty P d n om e ra dip ermorata tus s ic u k A de n om e A de en om era bo armavoptacin A d en om era m fl ys us sis A d en m era lus m sc n tes A d no m ty s fu tale dy A de no ac tylu s na lo s A de tod dac tylu s trog de A ep to ac tylu s ylo ae L ep tod ac tylu s h len L ep tod ac lu s e ixi L ep tod acty lu sp i ktitess L ep tod acty lus joly toa u L ep d cty us no tace L epto da ctyl us mys lis o L epto da ctyl us raci ej ni L epto da ctyl us g rtan an L epto da ctyl s se aum ius L epto da tylu s pl rnar L epto ac tylu s fu stus thicus L ptod ac tylu s va rin ra Le ptod ac tylu laby aqua Le ptod ac tylus cam eus s Le ptod ac lus cupr lariuis Le ptod acty lus nicu ns Le ptod acty lus cu aque Le ptod acty lus ch trans e Le ptod acty lus la atinga iae Le ptod acty s ca amba Le ptod actylu s mar is Le ptod actylu s virid ipinus Le ptod tylu podic sus Le ptodac tylus latina ac us Le ptod tyl phax Le ptodacctylus sy natha Le ptoda a euryg la Le reoran a parvu scopa Vit reoran urano sticta Vit reorana is leuco ueira ntiq Vit mocle a is ma Chias mocle is atlantic ias Ch mocle is gnoma nctata opu Chias mocle Chias mocleis alb ubarti Chias ocleis schcis Chiasm ocleis cru eiroi Chiasm ocleis cord oanus Chiasm cleis alag i Chiasmo cleis carvalho aba Chiasmo cleis capix Chiasmo leis sapiranga Chiasmoc ops histrio Stereocycl s incrassatus Stereocyclop s parkeri Stereocyclop llii Arcovomer passare muelleri notus Dermato leis erythrogaster Elachistoc Elachistocleis bicolor Elachistocleis ovalis Myersiella microps Dasypops schirchi Rana palmipes Pipa carvalhoi Ameerega flavopicta Ameerega picta Allobates olfersioides Fritziana fissilis Fritziana ohausi Fritziana goeldii Gastrotheca Gastrotheca fissipes Gastrothec flamma Gastrothe a albolineata Gastrothe ca fulvorufa ca microdisc Gas us Bractroth erne hyceeca phalus stoi Brachy Brachy cephalu didactylus Brachy cephaluss ephippium cephal nodote Brachy cephal us ver teb rga Brachy Brachy cephal us hermo ralis cepha us alipioi genesi Brachy lus cep Bra ize Bracchycephhalus brucksohni Brac hyce alus po nneus mbali Isc hyce phalus Isc hnocnephalus pernix Isc hnocne ma sa ferrug Isc hnocne ma ge mbaquiinus hr ti Ischhnocne ma Isch nocn ma spanios Isch nocn ema penaxa nocn ema lactea vant Isch inho Isch nocn em rand Isch nocn emaa holti orum nocn ema epip Isch Is no em ju ed Is chno cnem a ve ipoca a rruc Is chno cnem a oc Is chno cnem a m tavi osa anezoi Is chno cnem a bo Is chno cnem a pa lbod inho Is chno cnem a ni rana acty Ischchno cnem gr la a en Is n cn na iven sis tr Is chn ocn em a iz su Is ch ocn em a p ecksta is Is ch nocn em a usi oh Is chnnoc e a ghen lla ni Is ch oc ne ma ue selii Is ch no ne ma oe nthe ri A ch no cne ma ery a H de no cne ma pa th H ad lop cn m g rv rome P ad da hryem a h ua a ra Pri risti dad dus ne a ve oeh lteri P s m u p p n n B ris tim an s lic ac an ei N ary tim a tis bin ife hy cio H o c a nti ra ota r dactyi H o ble ho nti s p m tu la H o lo lla lo s a ag s E o loa ad b s vin ulo ii E u lo d en ilin tern h du E u pa ad en lu e e ai trai E u pa rk e p e ata tz C up parkrke eren braholederw i era a e re lla d te a to rkererell lla braei r ldti ph ll a co s ry a trid ch ilie s ro a ra n s au bu cty n is rita sta la ae

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AF species that have polygons in the IUCN. prunned the phylogeny Pyron. We also coded the species regarding their endemism level

Expectations

More nestedness between refugia and non-refugia Turnover between refugia

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Background | Atlantic Forest | Partial Results | Current project | Conclusion Mean of richness difference

Mean turnover

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No clear pattern between refugia and non-refugia regions, as expected. No richness differences between refugia and non-refugia Instead, mountaneous regions, like the Serra do Mar in the southeast, border betweeen AF and other biomes, and the Bahia refugium, have high values for turnover 18


The predictions would only make sense if the lineages that are endemic to refugia were closely related…

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… instead, they are overdispersed in the phylogeny

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The distribution of endemic species to refugia vs non-refugia along the phylogeny is not random, compared either to the null model or to the species pool of the Atlantic Forest Conversely, refugia endemic lineages are more phylogenetically distant than the endemic to non-refugia, compared to the species pool of the AF This suggest that many lineages (e.g., different families) are endemic to refugia Traits that may be associated with endemism in refugia may have have evolved several times 21


x = extinction rate s = speciation rate d = range expansion

A = endemic to refugia B = endemic to non-refugia AB = both 22


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WHAT THE RESULTS MEAN Species endemic to refugia have lower rates of range extension than other species Contrary to our expectations, but… Makes sense if endemic species were ”specialists” Speciation is higher in less stable regions Areas with higher perturbation frequency, and then varying selective pressures promote higher speciation in a broad scale Higher speciation rates in non-refugia suggest diversification in AF as a whole is not dependent on refugia alone 24


Conclusion

The refugia from the Plio-Pleistocene seem not to have generated the species diversity of anurans in the Atlantic Forest, since most species are older than the refugia itself… but how do I known that?

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500 200 100 50 20

N

1

2

5

10

Most of the lineages were already there!

−200

−150

−100

−50

Time (in Millons of years)

This is a lineage through time plot of the phylogeny for the AF species

0 26


Conclusion

Probably, refugia were important to maintain genetic diversity at the intraspecific level, as phylogeographical studies have demonstrated (e.g., Carnaval et al. 2009; Tonini et al. 2013), but are not strongly involved in species diversification per se.

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Next steps

Test for an effect of refugia, current, and past climate on the metacommunity structure of anurans For example, is the distribution of species endemic to refugia more correlated with past climatic conditions due to Phylogenetic Niche Conservatism? Are communities in refugia older than in non-refugia?

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Leibold et al. (2010) Ecol Lett


Patterns and processes in the origin and diversification of terrestrial vertebrates in the Brazilian Atlantic Forest

Supported by FAPESP #2016/13949-7

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Multi-taxa evaluation of distribution patterns Non-volant mammals Anurans Birds

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Causes of the spatial distribution of functional diversity of terrestrial vertebrates in the Brazilian Atlantic Forest

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QUESTION How equilibrium and non-equilibrium mechanisms interact to shape the species richness and functional diversity of vertebrates in the AF.

To distinguish the contribution of equilibrium and nonequilibrium mechanisms, I will test how speices richness and functional diversity are related with evolutionary time, diversification rate, productivity, climatic variables, and topographic heterogeneity. 32


Traits

FDis Map FDis

Topographic heterogeneity Climatic variables Diversification rate Productivity Evol time (MPD)

Structural Equation Modelling using GLS

For each group separately 33


HYPOTHESIS If equilibrium mechanisms were more important, I expect that current climatic variables and gradients of productivity are strongly and positively related to species richness and functional diversity. Climatic seasonality would be a strong environmental filter, constraining the niche space available in regions with more seasonal climate, decreasing its functional diversity. This effect should be more marked for poikilotherms than homeotherms 34

Oliveira et al. 2016 Glob. Ecol. Biogeog | Wiens et al. 2006 Am Nat


Adaptive dynamics of functional traits along the phylogeny

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QUESTION Is there phenotypic convergence for body size in distantly related lineages that occur in the same region (e.g., distant species that would expand their populations towards the south of the AF recently) ?

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Clavel & Morlon 2017 PNAS

R package bayou multi-regime OU model

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HYPOTHESIS Patterns of phenotypic convergence can indicate that distant lineages are under the same selection regime, suggesting that environmental factors are important to determine the variation in traits. Example, environments with harsh climate (highly seasonal) strongly constrain the variation in traits at the community level 39

Mahler et al. 2013 | Webb et al. 2002


Influence of the climatic niche on the rate of phenotypic evolution

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QUESTION

Assuming that diversification rates are correlated with rates of phenotypic evolution… Did climatic niche evolution influence phenotypic variation of species at broad spatial scales?

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Rabosky & Adams 2012 Evolution | Rabosky et al. 2013 Nat. Comm. | Lawson & Weir 2014 Ecol. Lett

Climatic variables CHELSA

OMI

Extract eigenvectors to describe the climatic niche

Estimate evolutionary rate 42

Body size

R package EvoRAG

Climatic niche

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WHAT DO I EXPECT? Gradients of temperature and rainfall seasonality are important to differentiate the climatic niche of species. On its turn, the niche will influence the rate of trait evolution (e.g., body size) differentially for the three groups, due to different thermal tolerances More strong in poikilotherms (amphibians) than in homeotherms (birds and mammals) Differences in the north (warmer and more stable) and south portions of AF (cooler and unstable) => groups with life history traits adapted to each region 44

Carnaval et al. 2014 Proc. Royal Soc.


Acknowledgements

Prof. Fernando R. da Silva UFSCar

João Tonini

José Hidasi Neto

Daniel Caetano

Alexander Zizka

Prof. Alexandre Antonelli GU

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[email protected] dbprovete.weebly.com http://fernandoecologia.wixsite.com/fernandorodrigues/ 46