Posted on Authorea 12 Oct 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.160253833.37656353/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 7 Spain Catalonia, Barcelona, Barcelona, 6 5 4 USA DC, ton, 3 2 Spain Catalonia, celona, 1 GONZ JACOB pelagic mobile highly FERRER-OBIOL events. JOAN of paleoceanographic dispersal of and role alpha speciation important potentially to of the trait revisions drivers on phenotypic light suggesting the key divergence, new on conditions. genomic shed a understanding local of and our be and patterns seabirds extend founder to strategies and findings that life taxonomy appears our current found with Globally, mass the associations we taxonomy. between Body significant and incongruences shows currents, found it diversification. also ocean and of We diversification, surface processes shearwater by during main speciation. favoured rapid selection the and be under dispersal are to high events of appears We vicariant period (ILS). dispersal a and by sorting that followed lineage molecular extinction, showed incomplete performed megafauna of analyses marine We levels Pliocene of high Biogeographic the seabirds. distributed role the of for the pelagic globally effect account explored mobile major to are we a (MSC) highly identified that data, approach these coalescent sequencing phylogeny multispecies in representation well-resolved a reduced speciation applying a Using and mobile dating, with diversification. diversification highly rapid seabirds on in and changes even pelagic slow paleoceanographic speciation, of both and of group diversification periods a of show are drivers and Shearwaters as act organisms. can changes marine paleoceanographic environments, marine In Abstract 2020 12, October 4 3 2 1 Gonzales-Solis Obiol Ferrer Joan seabirds rapid promoted pelagic in Pliocene diversification late the in changes Paleoceanographic eateto isine,Dra nvriy uhm UK Durham, de University, Durham Universitat Biosciences, Biologia, of de Department Facultat Ci`encies Ambientals, i Ecologia Evolutiva, Biologia de Departament USA MD, Laurel, d’ Center, Centre Research Wildlife Patuxent Survey, Geological U.S. Washing- Institution, Smithsonian History, Natural of Museum Spain National Zoology, Catalonia, Vertebrate Barcelona, of (IRBio), Department Biodiversitat la de Recerca de Institut Bar- Barcelona, de Universitat Biologia, de Estad´ıstica, Facultat Gen`etica, i Microbiologia de Departament uhmUiest eateto Biosciences of Department University Durham d’ Centre Biologia Institution de Smithsonian Facultat Barcelona de Universitat tdsBooiusd hze NS&L ohleUieste 96,Vlir nBi,France Bois, en Villiers Universit´e, 79360, Rochelle La & Chiz´e, CNRS de Biologiques Etudes ´ tdsBooiusd hze NS&L ohleUniversit´e Rochelle La & Chiz´e, CNRS de Biologiques Etudes ´ ALEZ-SOL ´ 1 ui Rozas Julio , 1 ee James Helen , 1,2,* ´ IS EE .JAMES F. HELEN , 6,2 UI ROZAS JULIO , 1 nran Welch Andreanna , 2 .Chesser R. , 1,2 3 .TRYCHESSER TERRY R. , NRAN .WELCH J. ANDREANNA , 1 2 icn Bretagnolle Vincent , 4 n at Riutort Marta and , 3,4 ICN BRETAGNOLLE VINCENT , 7 3 AT RIUTORT MARTA , Jacob , 1 1,2,* 5 , Posted on Authorea 12 Oct 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.160253833.37656353/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. pce iiswe sdtgte ihohrdt ye uha hntpcadeooia information. ecological on and phenotypic perspective as additional such 2018; provide types Yang, can data & data other Rannala, with genomic (Leach´e, Zhu, disagreement together delimitation that of used exists species extent when the agreement to limits Despite 2017), applied species taxa. Knowles, be morphological sympatric limits & should or of species changes parapatric Sukumaran data cases on in when genomic hypotheses In conclusive informed how especially be provide 2018). can about stasis, can Malard, and occurs, data morphological taxa & this genomic allopatric by (Fiˇser, When Robinson, traits information, of minimal 2013). hampered behavioural at al., are limited relatives further et and allopatry close be Seddon stasis in their 2008; may niche from (Price, species ecological differ selection delimit to in social Pelletier, tend to of (Carstens, isolation forms first ability radiations reproductive other allospecies our young and of Many in sexual 2013). evolution to especially Cutter, gradual subjected challenging, 2013; the the delimitation Satler, of widespread, & species paradox” evaluations Reid, make geographically species explicit can “marine are systems. for allopatry marine the that need in in as complexes a differentiation described of is species patterns there been In driving Thus, has in 1994). taxa processes among Palumbi, selective marine differentiation 2003; of of fine-scale David, role barriers number of highly & physical evidence in a Bonhomme, obvious counterintuitive especially in of (Bierne, However, prevail, lack species 2013). will the and isolation-by-distance, al., where or et populations 2012), panmixia, (Moura envi- Nosil, of marine species promote 2004; processes in Orr, mobile that neutral than & mechanisms that terrestrial Coyne and the Life, in suggest 2012; space for would of understood over al., Evidence Tree better operate et 2014). currently that the (Butlin Zardoya, are processes ronments across speciation & abiotic biodiversity Vargas distributed and and species-level differentiation 1953; unevenly biotic population of Simpson, from is generation result 2009; richness distribution po- the (Benton, Species of of time for patterns adaptation 2017). responsible current and Pennell, driver its evolution & and major independent Schluter a the 2010; from as results (Kopp, acts that ultimately process and evolutionary pulations key a is Speciation INTRODUCTION events. paleoceanographic seabirds of pelagic mobile role highly important shows of Keywords: the dispersal the it on and between our light speciation and Globally, incongruences of new taxonomy. diversification, drivers alpha shed found the to and shearwater also on revisions understanding We suggesting during our divergence, conditions. extend selection genomic findings local of appears under patterns and mass and potentially strategies Body taxonomy diversification. current life trait of currents, with phenotypic processes ocean rapid main associations key and surface the significant by are a dispersal events favoured high vicariant be of be and identified period to to founder We a appears that (ILS). by found dispersal followed sorting we that extinction, and lineage showed megafauna multispecies incomplete analyses marine a of Biogeographic Pliocene and levels applying the speciation. high dating, diversification of the molecular effect on for performed major changes account We well-resolveda paleoceanographic to seabirds. a (MSC) pelagic of with approach mobile role seabirds coalescent highly reduced speciation, the Using pelagic these diversification. explored and of in rapid and we speciation group diversification slow both data, a of of sequencing are periods drivers show representation Shearwaters and as distributed organisms. globally act are marine that phylogeny mobile can highly changes in paleoceanographic even environments, marine In ABSTRACT Biologia, de [email protected] Estad´ıstica, Facultat *E-mail: i Spain; Gen`etica, Microbiologia Catalonia, de Barcelona, Barcelona, Departament de to: Universitat sent be to Correspondence ebrs iesfiain pcain oeua aig igorpy taxonomy biogeography; dating; molecular speciation; diversification; seabirds; and [email protected] 2 Posted on Authorea 12 Oct 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.160253833.37656353/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. ftegopb nlsn eoi atrso eetsae netyaddffrnito mn shearwater taxonomy among current differentiation the and of ancestry validity shared the recent assess of we for patterns Finally, of responsible genomic size. surface history forces analysing taxa. body and by ecological biogeographic trait, vicariance group the the phenotypic events, the explore key founder infer of we a of Furthermore, then roles in diversification. We variability the phylogeny. their evaluating the driving shearwater and in account the ranges currents to affecting ancestral MSC ocean estimating ILS the pelagic by using of of group tree group levels the species this shearwater in high speciation time-calibrated al- the and first diversification for for the of (PE-ddRAD-Seq) produce drivers the we sequencing Specifically, explore DNA to seabirds. site-associated taxa shearwater the restriction pelagic extant all double-digest during endangered most paired-end events these use of vicariant we conservation and Here, the founder for 2016; potential of Velando, implications great importance important & their have relative Paiva, seabirds. despite can Genovart, the shearwaters species (Munilla, Understanding of seabird reported 1985). species, evolution most shearwater Lien, been several in The have & in event shearwaters. However, events Storey rare of 2008). foundation Bernatchez, evolution a colony & the Weimerskirch, be contemporary during (Milot, to events dispersal believed founder long-range of is for importance colonies our the of to oceanographic is and critical foundation environmental interest are of particular light models in Of biogeographic world 2015). in events. alternative the of throughout Ellegren, of incongruence diversification evaluation & effects shearwater phylogenetic and of Smeds, the of ranges understanding Suh, ancestral source incorporating 1997; of common Maddison, by reconstruction events, Reis, 2016; most The solution such al., Dos the For accurate et likely & (Edwards 2016). more is events (Angelis Hahn, which a diversification timescales & rapid (ILS), biases offers Mendes recent sorting to (MSC) 2011; lineage lead at model Knowles, incomplete can particularly coalescent & concatenation estimates, multispecies Peterson, on Delaney, age the based Heled, Analyses misleading events, McCormack, times. and speciation divergence 2015; of lengths accurate timing branch the estimate to to in changes necessary oceanographic is and closely vocalisation environmental it speciation. between historical and within as differ relate delimitation diversification such accurately species may promote To resolving traits, that size, in factors assist phenotypic body the will few data of particular, genomic context a in using the study and only (Austin, comprehensive stasis A differences indeed, morphological species. colour related 1996); high to plumage Austin, due slight mostly 2004; controversial, characteristics, and 2018; Pasquet, are strategies Austin, limits & foraging & Lavers, different species Wenner, as Lombal, Bretagnolle, Sixth, 2007; such 2011). al., factors al., et biotic et Smith, (G´omez-D´ıaz, intrinsic (Friesen, Rayner and speciation speciation 2015) promote Gonz´alez-Sol´ıs, of also al., drivers as can promote et allochrony historical such Silva founder to changes as 2006; of known Page, act paleoceanographic roles are & instance, can the Peinado, factors for oscillations evaluate biotic Procellariiformes; climatic to and related Pleistocene model abiotic and diversification the ideal shearwaters Fifth, rapid degrees an the analyses. and of in them biogeographic ecologies speciation makes periods using different mobility exhibit vicariance clear high genera and showing their recognised events Fourth, phylogeny three richness. well-resolved the species Third, a recognises of review). First, taxonomy with study under current environments. case al. species the marine excellent et Second, 30 in allopatry. (Ferrer-Obiol an in and speciation offer mostly genera and breed they differentiation three and and distributed population Procellariidae, globally of are family 2007). McCoy, mechanisms shearwaters the is & the in which Burg, 2006; examining (Friesen, group 2002), al., differentiation for monophyletic (Coulson, et genetic a 2015). grounds reinforce (Shaffer are therefore breeding Pinet, year) Shearwaters and their a & flow to gene km Phillips, philopatry limit 120,000 Guitteaud, to high than expected show Delord, more also Weimerskirch, some (covering perform Procellariiformes 2007; Earth and speciation However, ability Ruiz, on marine dispersal the migrations & high of animal a Gonz´alez-Sol´ıs, Oro, examples with longest extreme Croxall, seabirds most the pelagic the mobile of of highly some are present Procellariiformes Procellariiformes paradox. order the of Seabirds 3 Posted on Authorea 12 Oct 2020 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.160253833.37656353/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. eoye fteprsml eddphwsblw5o bv 5,o ftegntp ult a eo 30. below mask was to quality genotype data, the missing if or without 150, SNPs above with biallelic or file 5 VCF only below the include was two processed depth to of further read 2011) We format per-sample maximum rate. al., the call including a evolutionary variant if analyses et selected genotypes the to the (Danecek variants we in performed v.0.1.15 we called heterogeneity method, rate, exported reduce VCFtools this substitution we to nucleotide for and single transitions a dataset total) only (R. assumes in suitable v.2.5.0 SNAPP individuals a 2 Because of (51 (VCF). (Bryant, BEAST prepare approach subspecies program v.1.3. To MSC per the SNAPP 2019). 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Data may be preliminary. uek,&Hde,21) ugop eeicroae notetm-airtdserae reuigthe R. using in Stecher, tree 2019) shearwater Schliep, (Kumar, & time-calibrated database (Paradis the TimeTree outgroups package into the the ape incorporated the between from were from times retrieved no Outgroups function Divergence were bind.tree 2017). which clades. outgroups Hedges, their (for the & within lineages among Suleski, ranges outgroup and of shearwaters related totality and the closely the represent most to they parameter two comparable because j statistically the common the recent being rostrata genus of most without available): not Pseudobulweria shearwaters’ ranges and are the for the of with data criticised range used models genetic ancestral been most we the the has when (MRCA), infer for To model Sanmart´ın, present, ancestor and 2018). scenario & j the (AICc) (Ree best-fit + Criterion to model the DEC DEC Information Pliocene select Akaike the the Corrected to because dispersal from used separately, 1). limited currents were that (Figure weights ocean another occurred and AICc diversification surface areas shearwater major all compared the under by between We of (j) dispersal parameter connected 1). unrestricted speciation areas founder-event (Figure allowed the between that tree without one shearwater and with time-calibrated (Matzke, scenarios: models two the 1.1.2 these dispersal-extinction- of (Landis, and BayArea v. estimates models: 2013), and range three BioGeoBEARS 1997), Huelsenbeck, ancestral package of Ronquist, & (DIVA; versions R dispersal-vicariance Moore, likelihood The 2008), using Matzke, Smith, between 2001). ranges & limit Ree (Sutton, ancestral the (DEC; Front estimate set cladogenesis Subtropical We to breeding (E). the used shearwater Sea was at Mediterranean contemporary 2013) B and (C), on Ocean and Ocean Pacific Atlantic based Tropical A and North chosen North areas and (B), were (D), Ocean shearwater areas Southern Ocean of (A), Indian five Zealand patterns Tropical New The examine and Australia to Southern areas. and ranges: broad ranges five ancestral estimate across to dispersal performed were analyses Biogeographic estimation range Ancestral and (times) lengths branch both of estimate estimation the of allows value Densitree SNAPP in ( in sizes visualised implemented population were model trees mean finite-sites run SNAPP to The of 2013). set distributions Bouckaert, heights & posterior & node and Baele, 2010). with (Heled (Bouckaert, burn-in, Xie, trees TreeAnnotator v.2.2.7 as Drummond, (MCC) with discarded maximum-clade-credibility (Rambaut, each was estimates generate v.1.7.1 runs, MCMC age to Tracer each replicate stationarity merged of using three and were 10% 300) Convergence conducted replicates first iterations. > we The (MCMC) sizes analysis, 2018). 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. npro f1000gnrtosada xr 0,0 CCieain ape vr ,0 generations. 1,000 every sampled burn- iterations a the MCMC with and 100,000 clusters, matrix extra into coancestry PE- an a individuals 8,049 and infer assign generations for the to to 100,000 haplotypes of used used of was 75% of was RADpainter least period set SNPs. at algorithm in a 63,492 in clustering of in present MCMC total loci resulting fineSTRUCTURE a for POPULATIONS, containing haplotypes using matrix loci exported co-ancestry format ddRAD We a ancestry events. RADpainter derive shared coalescent to to recent the individuals information most infer linkage species, to the haplotype among 2018) on uses structure Falush, based fineRADstructure & genetic individuals. 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. iainei nipratmd fseito nseraes h lgtylwrAC o IAIEmodels DIVALIKE for AICc lower slightly that The than suggesting AICc shearwaters. included, in lower not speciation had was of models than (j) mode DEC important AICc parameter dis- and an event lower DIVALIKE is founder had restricting vicariance the currents, 1). matrix (Table when ocean dispersal matrix especially surface dispersal present models, a BAYAREALIKE unrestricted and including historical an analyses, main with estimation by models connected range areas ancestral between models, persal tested all Under Analysis shearwaters. of Biogeographic history assuming evolutionary age the when without in analysis 34,477–52,790) size SNAPP HPD: population However, in in (95% species variation 2020). changes individuals shearwater International notable frequent individuals Birdlife a N=43,485 a N=63,555 years; for showed and was (19 estimated constraints 2016), value SNAPP time estimated al., by generation highest et estimated lowest the Genovart average). the species on years; assuming shearwater older when S2, (13 (0.4% all (Figure 50,390–77,155) estimates across root using age HPD: the size on review) (95% on effect population constraint estimates (under little age older mean very single slightly had al. 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Figure S2; (Table a nerdt e45 Mya 4.52 be to inferred was Ardenna ). Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. oyms hne a eipratdrn pcain h neta tt eosrcino migratory of reconstruction state ancestral that The showing of shearwaters, MRCAs speciation. in the during common are important that clades showed be sister may behaviour between changes (Fig- mass latitude mass body breeding in good body maximum differences a for also striking recovered was 2b, correlation between latitude Figure strongest correlation Breeding R the positive year-round). adjusted with the species S5c; shows mass, a ure S5b body by (Figure mean occupied range latitudinal range of (Figure mass latitudinal and predictor mass and body lightest) with body range in the associated variance mass mean species the is body in sedentary/dispersive of shearwaters variance pre- and strategy 67% in the heaviest all explained migratory size of the range with Overall, were 75% body correlations migrants explained that factors. long-distance significant strategy suggesting ecological S5a; showed migratory predictors, by Indeed, mass best influenced capacity. the body strongly movement were Mean is range trait latitudinal S3). this and Table that (FDR 2a; suggesting correlations (Figure dictors, significant predictors several the recovered and analyses size PGLS body The of squares the least whereas generalized 0.45), (0.16). Phylogenetic = both or Pacific (0.27) Tropical area and Zealand North New 0.45, - of Australia = MRCA South Pacific the Tropical of and area ancestral North Tropical hand, and support and of other Atlantic highest North origin the the the (North The On by showed followed S4). area (0.54). 0.44), Figure area Zealand = and Zealand New probability 1 - ML 10 (Figure (marginal Australia x (0.33) shearwaters 1.9 South Pacific of = the best region (P model, the range ancestral favoured between j the strongly of (LRT) + as was of DIVALIKE test rate j probability ratio best + the higher DIVALIKE likelihood the a that the and Under showed have Indeed, 0.1733 events models expansion. j founder to range + that DIVALIKE 0.0874 than showing and data models from DIVALIKE smaller, the in magnitude ranged of However, of most parameter order vicariance. explaining j an widespread was the of (d) importance speciation, expansion the event supported founder further including models DEC to compared suiu lsesb h nSrcuecutrn loih Lwo,Hleta,Mes auh 2012), Falush, recovered were & study Myers, the Hellenthal, in (Lawson, included algorithm subspecies clustering and fineStructure phylogeny species the shearwater the recent by all most clusters and the unique review), supported genera as under largely shearwater group three al. each the et within to (Ferrer-Obiol subdivisions corresponding Further clusters 3). major (Figure three identified (Weber, analysis sizes fineRADStructure The population high taxonomy with and species divergence in Genomic strongest is agreement gBGC in 2014). are of Ellegren, results & of impact These number Jarvis, the the 0.94). Romiguier, with that = Boussau, correlation probability hypothesis significant posterior G positive the 0.684, of a = had with fixation coefficient GC* preferential GC-biased that (correlation the with found been pairs consistent to we have However, breeding leads heterozygotes, may AT/GC dataset. that This in our mechanism correlations C S4). of recombination-associated and significant (Table rates a any LHT substitution (gBGC): for find the the conversion slower not in and gene variance were did dataset high evolution we PE-ddRAD-Seq the yet (mtDNA) our by 1998), DNA of influenced Stanley, rate mitochondrial & substitution of (Nunn the rates Procellariiformes between that the in found taxa study larger rate previous a substitution from the Results on traits life-history 0.37, of and Effects 0.47 = probability ML (marginal of species MRCA sedentary the Conversely, a or respectively). migrant 0.86, respectively). short-distance and a 0.94 either = likely probability most ML marginal S6; (Figure 2 022.A hw ntepeormo neta tt eosrcin o oyms in mass body for reconstructions state ancestral of phenogram the in shown As =0.212). Puffinus Calonectris a siae sete h ot n rpclPcfi 03) the (0.37), Pacific Tropical and North the either as estimated was Calonectris and a nueuvcloii nteNrhr Hemisphere Northern the in origin unequivocal an had 8 Ardenna Ardenna a lotae oteSuhAsrla-New - Australia South the to traced also was eems ieyln-itnemigrants long-distance likely most were < .5 ewe oysz measures size body between 0.05) Puffinus was -5 ). Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. eea tde aesonta id r ao eemnn ffrgn agsadmgaoyrue of routes migratory and ranges 1). foraging (Table of currents determinant ocean major surface a by are connected winds areas conclusions that to firm marginal shown dispersal draw have to restricted studies cannot limited that Several we are matrix nevertheless, and dispersal ( currents; a wild that log-likelihood without suggest ocean in the differences analyses surface in reduced biogeographic by exist the The favoured given island 2009). is same al., et dispersal the Indeed, Forero, shearwater inhabiting (Navarro, shearwaters. species Procel- in species parapatric other described sister between Unlike been overlaps of not mobility. has high mechanisms records speciation and main few sympatric distribution the 2007), very global al., probably their et are by Smith, events expected (Friesen, lariiformes founder as shearwaters, and in vicariance speciation that of Reis, indicate Dos analyses biogeographic & Our (Angelis ILS shearwaters of of role 2016). history Drummond, the error & Biogeographic consider age Xie, fully node Heled, of to Ogilvie, degree failing 2017; higher Drummond, by accuracy. a times & caused by reduce Bouckaert, divergence caused potentially significantly Ogilvie, older likely analyses, 2015; the to most given latter that are limitation and argue the S2) study, this we (Table this in phylogeny, analyses expect in shearwater concatenation estimated not the (Ferrer by size by do likelihoods estimated population encompassed overall marginal therefore timescales high best We shallow relatively the for the strict the model given obtained review). the clock limitations, model under of a these of clock instead select Despite use strict to 2007) al. analyses the Yang, the et previous that & from rates Nonetheless, Obiol showed Rannala substitution benefit 2018). dataset 2006; likely al., in PE-ddRAD Rambaut, et would heterogeneity this (Stange & which the by Phillips, implemented review), is Ho, model under than limitation clock (Drummond, sizes As additional clock al. is population S3). An relaxed et assumption larger (Figure a (Ferrer-Obiol with This constraint versa. lineages lineages age vice for shearwater 2018). any equal overestimated and without among al., slightly of results estimation et be assumption SNAPP overall might (Stange the the times phylogeny the in is divergence the shown our estimation as of result, time analysis, branches a our divergence all in absolute on violated for sizes clearly approach population SNAPP constant the and of SNAPP limitation using A estimation divergence. time genomic divergence of of patterns the Limitations and uncovered potential taxonomy evidence consider current the we used the using we and between Lastly, diversification, range incongruences evolution. shearwater ancestral explain its in their shaped to shearwater size have infer here may in body events that to of forces founder as evolutionary role and for and well vicariance the ecological analysis as dispersal, discuss of group, also biogeographical paleoceanographic role We the the the and diversification. of evaluate explore and diversification tree explicitly analyses to group the biogeographic species us driven major formal allowed have time-calibrated a This may in phylogeny. MSC that resolved speciation first events fully and a diversification the on of based constructing drivers shearwaters the by of seabirds, analysis fundamental of a presents study This DISCUSSION ease to 4b Figure 4b). in Figure delimited in interval interval (second (first between distances subspecies genetic Atlantic same the distances different the addition, the genetic In However, The within comparison). distances 4). taxonomy. qualitative (Figure genetic current overlap of the some showed distribution with species consistent among were and between distances within distances genetic of of distributions distributions the Overall, and for except .carneipes A. .creatopus A. .biln nicolae bailloni P. Calonectris htwr nec ae lsee noasnl population. single a into clustered case, each in were that , and .carneipes A. and pce eewti h nevlo eei itne mn ieetsubspecies different among distances genetic of interval the within were species .biln dichrous bailloni P. n between and , < nt)btenacsrlrneetmto oeswt or with models estimation range ancestral between units) 3 9 , .mauretanicus P. .mauretanicus P. .boydi P. and and and .yelkouan P. .yelkouan P. .baroli P. n ewe the between and , and , eewti the within were , .creatopus A. Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. optto o etstspasa motn oei iiigppltoso ml rclaifre,such Procellariiformes, small of population populations of limiting cause in principal role a important is an and species 2012) plays threatened al., sites alien as those invasive et nest identifies by (Croxall for that Predation seabirds competition history. for metric evolutionary threat a gene among unique current is reduced declines main their which have the of 2), may is (e.g. because of (Table species genera attention year sizes scores shearwater particular the population EDGE deserve other throughout The highest that to sites richness. the compared breeding species had dispersal higher their many lower promoted to Their and close 2020). flow remain of al., origin that et Pacific dispersers Tropical Ramos a or supports migrants further short-distance samples which for obtain 2004), not advantages other could al., providing we most et Although 2016), to 1961). (Miller, al., (inaccessible record Pacific et fossil the for the (Shoji and of analyses m of waters estimation MRCA 55 range subtropical the ancestral and of where 2001), tropical depths Burger, poor to seabirds; nutrient tropical dive the routinely in to prey reaching them allow birds represent these likely 2001); migrants Atlantic North long-distance of the also ancestors in 2014; were The period. found taxa Shaffer, non-breeding fossils extinct the the & of that during that MRCA Silk, and died suggest the Phillips, Hemisphere, that We reconstruction, Southern (Carey, S6). state the coasts ancestral (Figure in our breeding European migrant on long-distance and that based a migrants American and trans-equatorial likely North 2006) long-distance al., on are et species common Extant Shaffer of locally record. origin fossil Atlantic be the has North can on the as with based Panama, disagrees 1996) of which (Austin, 1), isthmus by (Figure the Ocean Southern of the closure colonised gradual that the 2008). indicate G´omez-D´ıaz of analyses (Lessios, North Our Mya; al., result organisms the (˜3.44 et marine the and rates other as mtDNA Pacific in event on North observed vicariant based the been a breeding estimates between the suggests previous Mya) not to and (˜3.54 were found similar time 2006) were very divergence Olson, the fossils is estimated these considering 2009; clades where The However, Atlantic Olson, regions species. genus. the 2008; the the that possibility for (Olson, for the areas of Mya hotspot eliminate MRCA not ˜14 the speciation can that we a to given S6), as and back genus area dating the this of Atlantic mobility supporting North 2001), the Rasmussen, from & record. ancestral fossil described the the been as in Atlantic have uncertainties North these of the by MRCA of misled The suggestions been Florida) earlier have Formation, Thus, may Hawthorne review). shearwaters Miocene, in (lower of species al., area fossil et shearwater shearwater (Ferrer-Obiol Atlantic oldest uncertain North the is oldest be record the to fossil of believed shearwater was position which rich which phylogenetic relatively hypotheses, The the ( 1954). previous on species Kuroda, than based fossil 1996; shearwaters area (Austin, ancestral of area the area this was ancestral in Atlantic analyses the North biogeographic current the of On the that species hypothesis that 2013). extinct suggested suggest probable Remsen, from observations and fossils These more & shearwater 1961). 2012) (Dickinson a of (Miller, al., diversity area represent ones highest et oldest single the the (Croxall any harbours of biodiversity California in some seabird of and breeding coast global species the of hand, shearwater other South hotspot of the The a number region S4). currently greatest (Figure ancestral is Pacific the the Tropical area as and has Zealand Northern area the New Zealand by that New - followed suggests - support Australia Procellariiformes. highest Australia study the the South our with in the shearwaters hence, dispersal inferred of currents; analyses species ocean estimation of surface range determinant of Ancestral important driver an primary be a Grissac, also also de could are Louzao, winds Weimerskirch, Winds 2009; 2012). (Gonz´alez-Sol´ıs Delord, al., Procellariiformes & et the in especially seabirds, Puffinus .subalaris P. hawtr Mner,Rms&Fres19;Rms otio oa oi,19) tsea, At 1997). Moniz, & Sola, Monteiro, Ramos, 1996; Furness & Ramos (Monteiro, shearwaters .raemdonckii P. rmteGlpgs napeiu td hsseisfre ld with clade a formed species this study previous a in Galapagos; the from Calonectris Puffinus Puffinus pce Rd´ge ta. 09.Ehne ypeain nr-adinter-specific and intra- predation, by Enhanced 2019). al., (Rodr´ıguez et species curdtesrnetdvn dpain ftetregnr Osn Rasmussen, & (Olson, genera three the of adaptations diving strongest the acquired Ardenna a ot aicadNrhAlni itiuin osl fa es species 5 least at of Fossils distribution. Atlantic North and Pacific North a had and .arvernensis P. a ot utai e eln rgnad hratr oelineages some thereafter, and, origin Zealand New - Australia South a had ssilucer(lo,18)adteaeof age the and 1985) (Olson, unclear still is ) Calonectris 10 Puffinus otpoal rgntdbsdo h current the on based originated probably most a rbbyaln-itnemgat(Figure migrant long-distance a probably was Puffinus Puffinus otextant Most . pce edt esaland small be to tend species Ardenna .nativitatis P. Puffinus Ardenna a lomost also was .micraulax, P. pce are species proposed (Austin Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ewae o irtr n nlsdnsigseis(anaig&Sre,21;Mii&Dyn 2003). Dayan, & to Meiri tends 2019; Rule Street, 2 Bergmann’s & (Figure to (Mainwaring mass conformity species body that nesting variation mean shown enclosed size have and and that body latitude migratory studies breeding intraspecific for weaker previous between to despite be correlation apply Meiri, significant S5C), to 2002; positive Figure shown (Ashton, and a debate been found much also also ( to has we Shearwater subject pattern Streaked are the this the trend although at shearwaters, in 1848) this world the (Bergmann, of the level In throughout generation interspecific inhabiting birds 2011). the and lower for 2003) individuals for and holds Dayan, environments responsible species, & size warmer mechanisms Meiri body endothermic inhabiting 2002; in those related (Ashton, pattern than intraspecific closely geographical larger the This be of 1848). to group tend (Bergmann, latitudes latitudes a higher or and habitats correlation. species surface colder this endothermic body weaker of by The an effect corrected deposits. additive Within when fat and mass behaviour twofold of body migratory the accumulation mean hand, higher supports and other a S3) strategy migratory the (Table migratory to in On between all leading of wings 2a). correlation resources correlations significant (Figure longer additional significant but strategy (i.e. the of migratory by larger exploitation shown with be the as measures allows 2015) to size al., tend likely et body is species Minias other correlation 1995; migratory the This Richman, hand, 2). & (Figure Price, one mass Marchetti, the species; body On mean additive. for mass predictor and evaluated body twofold best affecting the merely was be are strategy may that Migratory findings that our interpreting drivers at distributional taken be and must results poorly correlational. caution behavioural Our are although variation potential 1998). Procellariiformes, Stanley, in mass the on & in body success (Nunn variation light of Procellariiformes breeding drivers new the the affects by some level, exhibited interspecific progenitors shed variation the high the at Aanes, the hand, Sether, of despite instance, other Tveraa, For understood size) 1995; the level. Jouventin, On & intraspecific body 1998). Weimerskirch, the Chastel, Erikstad, overall at 1999; & by Chastel, fitness & to corrected (Barbraud related species mass closely several trait (body a condition is mass as body body such 2015). trait Procellariiformes, groups Malaquias, the phenotypic & In seabird key (Eilertsen other species a sea in as deep detected in mass even been Body and also 2020) al., has et Vianna period 2006). 2019; group. G´omez-D´ıaz this al., al., that the 1998; et during Walker, et shifts affected (Cole climatic & diversification penguins severely the Pleistocene (Avise in by from extinction bottlenecks increase promoted are and Pliocene three probably An species splitting was late the geographic fossil in dispersal the stems and promoted extinct that long the speciation probably known (O’Dea the suggests of be the with habitats burst this to together of subsequent coastal 1), 2001); hypothesised The ˜36% with Rasmussen, (Figure been shearwaters, & associated O’Dea genera has In Olson, megafauna 2006; 1985; shearwater reduction 2017). of al., Olson, This al., et rate 1971; (Fedorov et reduction. (Howard, extinction cap Pliocene Pimiento such the Ni˜no, ice 2007; of in el Arctic al., cause increase permanent the et the three-fold of of a been formation end have of the significant the the and may cause over as a Panama reduction 2016) such suffered of gradual al., changes, Isthmus and period, et oceanographic fluctuation the Global extreme breeding of by 2017). foraging closure al., the followed the main al., et Pliocene during the et the Miller sea-level especially represent (Pimiento of 2007; which end global shearwaters, Pleistocene Raymo, the waters, largest large at & Neritic the reduction and (Lisiecki 2001). sudden Cenozoic, Billups, Pleistocene medium the & and for During Thomas, Pliocene grounds Sloan, radiation. the Pagani, rapid in Zachos, a occurred 2005; during oscillations Ocean speciation and main rapid Indian changes the and the high of of and instance, some period For a Ocean, likely 1). as are appeared (Figure boundary dispersal These Pliocene-Pleistocene and 2016). the genera, al., three the et Across extinction (Genovart to place species in some put why drive reasons are could that measures one for conservation Gonz´alez-Sol´ıs, and & threat unless Cort´es, 2017) Arcos, main 2008; a Neves, & also is bycatch fisheries Puffinus hawtr aetehgetnme fednee pce mn h shearwaters. the among species endangered of number highest the have shearwaters aoeti leucomelas Calonectris Puffinus Puffinus pedfo h aict h ot tatc h Southern the Atlantic, North the to Pacific the from spread 11 hawtr Bgn,Mnii otio Nascimento, Monteiro, Mancini, (Bugoni, shearwaters aaooe l,21) mn hawtrspecies, shearwater Among 2016). al., et Yamamoto ; Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ape uigti td,priual aafo h rpclPcfi htbedi eoeilnsadhave and islands be remote not in could breed species that sizes. that taxa use population Pacific the a low should tropical include and the including studies to distributions from taxonomic Future revaluation required taxa limited ongoing also morphological very particularly with study, status. is thorough this together research species during a Further data, granted sampled with vocalisations. genomic be of data our Parkin, evaluation not genomic consideration, & detailed should Knox, combining final taxa Helbig, approaches a Collinson, these As delimitation Sangster, that 2010; 2020). suggest Olson, al., debate, 2009; et al., et Ramos Gomez-Diaz 2005; population 2013; of al., subspecies: et patterns different (Genovart weak among showed interval Atlantic species the three In shearwater within the distances other pressures. Navarro, genetic selection pairs, 2000; strong and species species be under (Barbraud, structure single aforementioned rapidly selection to evolve a the under known could within to thus be are found addition and can which 2009), be section), Gonzalez-Solis, size, can previous & phases (see body Kaliontzopoulou, pale behaviour and of and migratory lack colouration Dark with (Penhallurick the plumage covaries shearwaters. differentiation on in of mtDNA based differ low species conspecific (i.e., pairs on within considered species even and et be traits These 1962) Gill labile should (Bourne, 2019; 2004). they Bonan, size Wink, that & and & argued (Carboneras, colour have checklists in taxonomic authors differentiation in correspon- some uniform species nuclear but of different & analyses, 2020), lack 2014). two Contreras-Diaz, al., isotopes a Gonzalez-Solis, as Juste, (Genovart, recently, & considered stable species widely Kaliontzopoulou, more characters, two are Gomez-Diaz, the However, phenotypic between Militao, between admixture 2012; to 2002). found Oro, attributed Parkin, was was and level & mtDNA, Amengual, individual Helbig, and (Heidrich, the Knox, of sequences at individuals Sangster, b dence between cytochrome 1998; observed using Wink, divergences monophyly genetic & reciprocal lowest the and the 4). with osteological were (Figure overlapped morphological, species subspecies and diverged recently same species these the of between pair divergence any genetic amongst the pairs, divergence, species fineRADStruc- species-level dataset, two for PE-ddRAD between our structure of divergence resolution no creatopus genomic high showed and the analysis structure Despite taxonomy. population ture 1996). inform fine-scale (Bretagnolle, provide can limited a to that are potential estimates vocalizations being shearwater the despite have of analyses datasets addition, (Austin, analyses delimitation, Genome-wide datasets In and G´omez-D´ıaz, species genetic 2009). studies, different for 2013; between Gonz´alez-Sol´ıs, trait morphological Peinado, also remarkably al., promising amongst and & their et microsatellites), arisen to and Genovart has due mtDNA 1996; (i.e., Conflict partly data 2004). topic, genetic controversial al., on and based et challenging (Austin a morphology is similar shearwaters in delimitation Species taxonomy shearwater of 1995). (Barbraud al., Considerations Procellariiformes et body of Higher Chastel mass. species 2005; several Weimerskirch, body on in & act success size Barbraud to breeding 1999; those of selection higher to Chastel, sexual cause with compared & for likely resources associated substrate available most the been the provide has the exploiting might condition dimorphism at This be efficient. efficient to size less more shown are are and that that been hand, sexes individuals differences has one between mass between segregation the body arise larger segregation may On ecological hand, (Gonz´alez-Sol´ıs, twofold. other ecological Indeed, Procellariiformes the likely other On 2019). for in in 2004). is dimorphism al., allow differences range et by may latitudinal Felipe and/or and areas (De range migratory foraging mass be larger body to exploiting study. between latitudes this association higher in strong explored in The be living indirectly not species be could also of which could correlations tendency behaviour, these diving higher However, body latitudes. a colder by by and corrected higher driven mass in thrive body to mean adaptation and better latitude breeding (R maximum between surface strongest was correlation The .pacifica A. and 2 .8;TbeS) ugsigta eve ois needn fbd ie ih rvd a provide might size, body of independent bodies, heavier that suggesting S3), Table 0.387; = .carneipes A. n oeseisehbtacniumfo aet ak(i.e., dark to pale from continuum a exhibit species some and ) Calonectris Fgr ) utemr,atog ed o osdrteet eagntccutoff genetic a be to there consider not do we although Furthermore, 3). (Figure pce.Teeseiscmlxsaetesbeto non aooi debate taxonomic ongoing of subject the are complexes species These species. .mauretanicus P. and 12 .yelkouan P. .mauretanicus P. eegatdseissau ae on based status species granted were .creatopus A. .mauretanicus P. .boydi P. and .yelkouan P. and and .baroli P. .carneipes A. .Bd size Body ). and , and , A. Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. idieItrainl(00 UNRdLs o id.Dwlae rmht:/w.idieogon http://www.birdlife.org from Downloaded birds. for List Cornell Red World. the IUCN of Birds 17/08/2020. (2020) (2020). S. International T. paradox. https://birdsoftheworld.org/bow/home Schulenberg, BirdLife USA. & marine-speciation NY, G., the P. 1399–1406. Ithaca, Ornithology, Rodewald, 270(1522), and of K., Sciences, preference B. 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Bayesian Reis, on Dos & K., Angelis, REFERENCES Investigaci´on Cient´ıfica de Equipos a “Ayudas Research (program illustrations. 062 comments Fundaci´on shearwater BBVA her code wonderful the for project his Paris by 2017”, R. Mart´ı for Franch Josephine provided and thank manuscript was to this like funding of would for we version Finally, Kansas authorities early pertinent of work. an research: the this University on thank this for We the needed for History, Naturelle. Science, permits loans d’Histoire Natural the Natural National issuing tissue of of Museum for provided Museum the Museum skins and that American University Institute, museum for institutions Museum, State Biodiversity from Louzao Burke many Louisiana Washington samples History, Maite the of tissue Natural thank and University providing of to Martinez-Gomez, and/or Museum like E. 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Data may be preliminary. ui oa http://orcid.org/0000-0002-6839-9148 Rozas Julio Gonz´alez-Sol´ısJacob http://orcid.org/0000-0002-8691-9397 Bretagnollehttp://orcid.org/0000-0002-2320-7755 Vincent Chesserhttp://orcid.org/0000-0003-4389-7092 Terry R. http://orcid.org/0000-0002-2495-6133 James F. Helen Ferrer-Obiolhttp://orcid.org/0000-0002-1184-5434 Joan read ORCID authors to All contributed authors. authors all all from input and with collection manuscript manuscript. the data final wrote to the and contributed approved data and the J.F. analysed and J.F. A.J.W. design. study V.B., J.G., H.F.J., R.T.C., CONTRIBUTIONS AUTHOR for files ( XML Repository and Digital SNAPP Dryad MCC format and the VCF files in input including: fineRADStructure deposited analysis, stages are number COEVOL several trees, for accession file the at format under PHYLIP data (ENA) analyses, Archive SNAPP processed Nucleotide containing European the Files on PRJEB38458. archived are reads PE-ddRAD Raw in aberrations and ACCESSIBILITY rhythms, DATA Trends, (2001). K. 686–693. 292(5517), Billups, visualization Science, & for present. E., package to Ecology Thomas, R Ma in L., 65 an Methods climate Sloan, ggtree: global data. M., (2017). associated Pagani, Y. other J., T. and Zachos, T. covariates Lam, their 28–36. & with 8(1), Y., Evolution, trees Guan, leucomelas. and phylogenetic H., Calonectris of Zhu, shearwater annotation K., (2016). streaked and D. A. the Takahashi, Smith, seabird, G., . . Yu, pelagic . R., a 801–808. 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States Noll, penguin Weber, United R., of the L. drivers Pertierra, of reveal V., Sciences Figueir´o, analyses H. of J., Academy Genome-wide M. Frugone, (2020). N., MA. K. A. Sunderland, C. F. life. Fernandes, of A., J. tree Vianna, The (2014). R. Zardoya, & P., Vargas, 10.1046/j.1365-2656.1998.00234.x doi: 699–704. 21 https://doi.org/1XXXXX ). Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. r umrzdi al 2 lutain yMr´ rnhaerpeettv hawtrseisdepicted species shearwater representative times divergence are are of Mart´ı Franch estimates and by Pleistocene. Posterior BioGeoBEARS Illustrations early (Inset). lineages. in map - their S2. the currents by to Table Pliocene ocean according in late coded surface summarized tips present the and are and nodes during dispersal restricting at historical peak matrix boxes times main dispersal as speciation a divergence shown by using the of connected model j densities Note areas + Posterior DIVALIKE between the tree. tree. under estimated the species were above ranges the Ancestral labeled below are shown epochs are and periods Geological topology. 1 Figure TABLES AND FIGURES http://orcid.org/0000-0002-2134-7674 Riutort Marta Welchhttp://orcid.org/0000-0002-5947-4695 J. Andreanna ieclbae pce reo h hawtr sn osrito h otaeadafixed a and age root the on constraint a using shearwaters the of tree species Time-calibrated 22 Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. irtr taeyadtelttdnlrnefrec species. each genera: for three range the latitudinal indicate the colours and Edge strategy clades. migratory sister R between mass *** adjusted (FDR: body the Calonectris testing mean multiple show in for differences tile correction abrupt FDR each showing by within p-values adjusting Numbers after < significant when test. green each in for coloured recovered were coefficients R adjusted 2 Figure 0.001 > irtr taeyadlttdnlrneaetebs rdcoso oysz.()Hampshowing Heatmap (a) size. body of predictors best the are range latitudinal and strategy Migratory ** 2 (purple), ausfrPL nlsso oysz esrsaanttepeitr.Pstv correlations Positive predictors. the against measures size body of analyses PGLS for values < 0.01 > Ardenna * < .5.()Peormo enbd ascntutdi htos(eel 2012) (Revell, Phytools in constructed mass body mean of Phenogram (b) 0.05). gen and (green) Puffinus 23 oag) etasnx otepeormso the show phenogram the to next Heatmaps (orange). 2 ausadare and values Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. iue4 Figure most that and genera three clusters. the unique discernible represent as clusters a recovered major are represents probability study three name posterior the the Every a in that by included Note defined (black). subspecies coefficients, and high tree. coancestry species finestructure to of the matrix (yellow) pairwise in low the 0.9 from on based coded cluster colour discrete are coefficients try 3 Figure lsee nRDtutr onetymti ae n809P-dA oi ariecoances- Pairwise loci. PE-ddRAD 8,049 on based matrix coancestry fineRADStructure Clustered h itiuin fgntcdsacswti n mn pce vra.()Dsrbto of Distribution (a) overlap. species among and within distances genetic of distributions The 24 > Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. hawtrseisadsbpce ntestudy. the in subspecies and species shearwater 2 Table AICc highest the with model the case each for and separately bold. shown in are shown (j) is weight parameter event founder the 1 Table and within distances Atlantic genetic of three distributions the the of Zoom-in among between (b) creatopus subspecies, Comparisons right. among proposed to subspecies, species. show left within bars among from values Vertical genera for taxonomy. among panels panel: current and across middle to species visualisation species, according assist within genera) to among panel: limits (upper orientative panel: shearwaters lower the and in species levels among taxonomic different at distances genetic oprsno oeso neta ag siainfrteseraes oeswt n without and with Models shearwaters. the for estimation range ancestral of models of Comparison ubro reigpis osrainsau,eouinr itntesadEG crsfor scores EDGE and distinctness evolutionary status, conservation pairs, breeding of Number ali h ihnsbpce ag,adcmaiosbetween comparisons and range, subspecies within the in fall Calonectris pce ali h mn useisrange. subspecies among the in fall species .mauretanicus P. 25 and .yelkouan P. .baroli P. n between and and .boydi P. .carneipes A. n between and and A. Posted on Authorea 12 Oct 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.160253833.37656353/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. 26