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American Genetic Association ISSN 0022-1503 (Print) ISSN 1465-7333 (Online) journal of HEREDITY Volume 105 | Number 1 | January/February 2014 www.jhered.oxfordjournals.org An international journal of genomic and evolutionary diversity

American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

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Page 1: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

journal of V

olume 105 | N

umber 1 | January/February 2014

HE

RE

DIT

Y

journal ofHEREDITYAn international journal of genomic and evolutionary diversity

Perspective

The Global Invertebrate Genomics Alliance (GIGA): Developing Community Resources to Study Diverse Invertebrate GenomesGIGA Community of Scientists 1

Original Articles

Conservation Genetics and the Implication for Recovery of the Endangered Mitchell’s Satyr Butterfly, Neonympha mitchellii mitchelliiChristopher A. Hamm, Victoria Rademacher, Douglas A. Landis, and Barry L. Williams 19

Out of the Alps: The Biogeography of a Disjunctly Distributed Mountain Butterfly, the Almond-Eyed Ringlet Erebia alberganus (Lepidoptera, Satyrinae)Dirk Louy, Jan Christian Habel, Werner Ulrich, and Thomas Schmitt 28

Impact of Population Expansion on Genetic Diversity and Structure of River Otters (Lontra canadensis) in Central North AmericaJessica R. Brandt, Adam L. Brandt, Frank K. Ammer, Alfred L. Roca, and Thomas L. Serfass 39

Phylogeography of the Dark Fruit-Eating Bat Artibeus obscurus in the Brazilian AmazonWallax Augusto Silva Ferreira, Bárbara do Nascimento Borges, Symara Rodrigues-Antunes, Fernanda Atanaena Gonçalves de Andrade, Gilberto Ferreira de Souza Aguiar, José de Sousa E Silva-Junior, Suely Aparecida Marques-Aguiar, and Maria Lúcia Harada 48

Range Expansion of House Sparrows (Passer domesticus) in Kenya: Evidence of Genetic Admixture and Human-Mediated DispersalAaron W. Schrey, Andrea L. Liebl, Christina L. Richards, and Lynn B. Martin 60

Genetic Variation and Population Structure in the Endangered Hermann’s Tortoise: The Roles of Geography and Human-Mediated ProcessesMelanie Perez, Barbara Livoreil, Sara Mantovani, Marie-Catherine Boisselier, Barbara Crestanello, Jawad Abdelkrim, Céline Bonillo, Vassilis Goutner, Josie Lambourdière, Massimo Pierpaoli, Bogoljub Sterijovski, Ljiljana Tomovic, Sibelle T. Vilaça, Stefano Mazzotti, and Giorgio Bertorelle 70

The Elephants of Gash-Barka, Eritrea: Nuclear and Mitochondrial Genetic PatternsAdam L. Brandt, Yohannes Hagos, Yohannes Yacob, Victor A. David, Nicholas J. Georgiadis, Jeheskel Shoshani, and Alfred L. Roca 82

Strong Population Structure and Shallow Mitochondrial Phylogeny in the Banded Guitarfish, Zapteryx exasperata (Jordan y Gilbert, 1880), from the Northern Mexican PacificAna Castillo-Páez, Oscar Sosa-Nishizaki, Jonathan Sandoval-Castillo, Felipe Galván-Magaña, María-del-Pilar Blanco-Parra, and Axayácatl Rocha-Olivares 91

Phylogeographical Patterns of an Alpine Plant, Rhodiola dumulosa (Crassulaceae), Inferred from Chloroplast DNA SequencesYan Hou and Anru Lou 101

How Much Does Inbreeding Contribute to the Reduced Fitness of Hatchery-Born Steelhead (Oncorhynchus mykiss) in the Wild?Mark R. Christie, Rod A. French, Melanie L. Marine, and Michael S. Blouin 111At Risk of Population Decline? An Ecological and Genetic Approach to the Threatened Palm Species Butia eriospatha (Arecaceae) of Southern BrazilAlison G. Nazareno and Maurício S. dos Reis 120

Brief CommunicationsGenetic Background Alters Dominance Relationships between mat Alleles in the Ciliate Tetrahymena thermophilaSujal S. Phadke, Tiago Paixão, Tuan Pham, Stephanie Pham, and Rebecca A. Zufall 130

Characterization of Microsatellite Loci and Repeat Density in the Gooseneck Barnacle, Pollicipes elegans, Using Next Generation SequencingLouis V. Plough and Peter B. Marko 136

Letter to EditorVolvox, Rolling out from under the Shadow of Chlamydomonas with Support from the AGADavid Roy Smith 143

Announcements 145

The cover.Invertebrate organisms for potential genome/transcriptome projects through the GIGA consortium (see pp. 1–18). See inside cover for key to organisms.1. Gorgonocepahlus chilensis (basketstar; Echinodermata), 2. Vallicula multiformis (platyctenid; Ctenophora), 3. Paraseison kisfaludyi (Rotifera), 4. Bryozoa, 5. Spadella sp. (arrow worm; Chaetognatha), 6. Glass sponge (Porifera), 7. Millnesium sp. (water bear; Tardigrada), 8. Collosondeis sp. (sea spider; Pycnogonida), 9. Phoronopsis californica (horseshoe worm; Phoronida),10. Pherecarida sp. (fireworm; Annelida), 11. Enchiridium cf. evelinae (flatworm; Platyhelminthes, photo G. Giribet), 12. Priapulus sp. (penis worm; Priapulida), 13. Epimeria robusta (amphipod; Crustacea),14. Xenoturbella sp. (Xenoturbellida), 15. Peripatopsis alba (albino velvet worm; Onychophora, photo G. Giribet), 16. Micromelo undatus (miniature melo; Mollusca, photo G. Giribet), 17. Flabellum sp. (solitary coral; Cnidaria), 18. Vieitezia luzmurubeae (ribbon worm; Nemertea) inside the atrium of Ciona intestinalis (sea squirt; Tunicata, photo Jacinto Pérez, Hydronauta). Image credits, Greg Rouse, except where indicated otherwise.

American Genetic AssociationISSN 0022-1503 (Print)ISSN 1465-7333 (Online)

journal ofHEREDITYVolume 105 | Number 1 | January/February 2014

www.jhered.oxfordjournals.org

An international journal of genomic and evolutionary diversity

Page 2: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

journal of V

olume 105 | N

umber 1 | January/February 2014

HE

RE

DIT

Y

journal ofHEREDITYAn international journal of genomic and evolutionary diversity

Perspective

The Global Invertebrate Genomics Alliance (GIGA): Developing Community Resources to Study Diverse Invertebrate GenomesGIGA Community of Scientists 1

Original Articles

Conservation Genetics and the Implication for Recovery of the Endangered Mitchell’s Satyr Butterfly, Neonympha mitchellii mitchelliiChristopher A. Hamm, Victoria Rademacher, Douglas A. Landis, and Barry L. Williams 19

Out of the Alps: The Biogeography of a Disjunctly Distributed Mountain Butterfly, the Almond-Eyed Ringlet Erebia alberganus (Lepidoptera, Satyrinae)Dirk Louy, Jan Christian Habel, Werner Ulrich, and Thomas Schmitt 28

Impact of Population Expansion on Genetic Diversity and Structure of River Otters (Lontra canadensis) in Central North AmericaJessica R. Brandt, Adam L. Brandt, Frank K. Ammer, Alfred L. Roca, and Thomas L. Serfass 39

Phylogeography of the Dark Fruit-Eating Bat Artibeus obscurus in the Brazilian AmazonWallax Augusto Silva Ferreira, Bárbara do Nascimento Borges, Symara Rodrigues-Antunes, Fernanda Atanaena Gonçalves de Andrade, Gilberto Ferreira de Souza Aguiar, José de Sousa E Silva-Junior, Suely Aparecida Marques-Aguiar, and Maria Lúcia Harada 48

Range Expansion of House Sparrows (Passer domesticus) in Kenya: Evidence of Genetic Admixture and Human-Mediated DispersalAaron W. Schrey, Andrea L. Liebl, Christina L. Richards, and Lynn B. Martin 60

Genetic Variation and Population Structure in the Endangered Hermann’s Tortoise: The Roles of Geography and Human-Mediated ProcessesMelanie Perez, Barbara Livoreil, Sara Mantovani, Marie-Catherine Boisselier, Barbara Crestanello, Jawad Abdelkrim, Céline Bonillo, Vassilis Goutner, Josie Lambourdière, Massimo Pierpaoli, Bogoljub Sterijovski, Ljiljana Tomovic, Sibelle T. Vilaça, Stefano Mazzotti, and Giorgio Bertorelle 70

The Elephants of Gash-Barka, Eritrea: Nuclear and Mitochondrial Genetic PatternsAdam L. Brandt, Yohannes Hagos, Yohannes Yacob, Victor A. David, Nicholas J. Georgiadis, Jeheskel Shoshani, and Alfred L. Roca 82

Strong Population Structure and Shallow Mitochondrial Phylogeny in the Banded Guitarfish, Zapteryx exasperata (Jordan y Gilbert, 1880), from the Northern Mexican PacificAna Castillo-Páez, Oscar Sosa-Nishizaki, Jonathan Sandoval-Castillo, Felipe Galván-Magaña, María-del-Pilar Blanco-Parra, and Axayácatl Rocha-Olivares 91

Phylogeographical Patterns of an Alpine Plant, Rhodiola dumulosa (Crassulaceae), Inferred from Chloroplast DNA SequencesYan Hou and Anru Lou 101

How Much Does Inbreeding Contribute to the Reduced Fitness of Hatchery-Born Steelhead (Oncorhynchus mykiss) in the Wild?Mark R. Christie, Rod A. French, Melanie L. Marine, and Michael S. Blouin 111At Risk of Population Decline? An Ecological and Genetic Approach to the Threatened Palm Species Butia eriospatha (Arecaceae) of Southern BrazilAlison G. Nazareno and Maurício S. dos Reis 120

Brief CommunicationsGenetic Background Alters Dominance Relationships between mat Alleles in the Ciliate Tetrahymena thermophilaSujal S. Phadke, Tiago Paixão, Tuan Pham, Stephanie Pham, and Rebecca A. Zufall 130

Characterization of Microsatellite Loci and Repeat Density in the Gooseneck Barnacle, Pollicipes elegans, Using Next Generation SequencingLouis V. Plough and Peter B. Marko 136

Letter to EditorVolvox, Rolling out from under the Shadow of Chlamydomonas with Support from the AGADavid Roy Smith 143

Announcements 145

The cover.Invertebrate organisms for potential genome/transcriptome projects through the GIGA consortium (see pp. 1–18). See inside cover for key to organisms.1. Gorgonocepahlus chilensis (basketstar; Echinodermata), 2. Vallicula multiformis (platyctenid; Ctenophora), 3. Paraseison kisfaludyi (Rotifera), 4. Bryozoa, 5. Spadella sp. (arrow worm; Chaetognatha), 6. Glass sponge (Porifera), 7. Millnesium sp. (water bear; Tardigrada), 8. Collosondeis sp. (sea spider; Pycnogonida), 9. Phoronopsis californica (horseshoe worm; Phoronida),10. Pherecarida sp. (fireworm; Annelida), 11. Enchiridium cf. evelinae (flatworm; Platyhelminthes, photo G. Giribet), 12. Priapulus sp. (penis worm; Priapulida), 13. Epimeria robusta (amphipod; Crustacea),14. Xenoturbella sp. (Xenoturbellida), 15. Peripatopsis alba (albino velvet worm; Onychophora, photo G. Giribet), 16. Micromelo undatus (miniature melo; Mollusca, photo G. Giribet), 17. Flabellum sp. (solitary coral; Cnidaria), 18. Vieitezia luzmurubeae (ribbon worm; Nemertea) inside the atrium of Ciona intestinalis (sea squirt; Tunicata, photo Jacinto Pérez, Hydronauta). Image credits, Greg Rouse, except where indicated otherwise.

American Genetic AssociationISSN 0022-1503 (Print)ISSN 1465-7333 (Online)

journal ofHEREDITYVolume 105 | Number 1 | January/February 2014

www.jhered.oxfordjournals.org

An international journal of genomic and evolutionary diversity

Page 3: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

Editor-in-ChiefC. Scott Baker, Oregon State University

Managing EditorAnjanette Baker, American Genetic Association

Editorial BoardErnest Bailey, University of KentuckyAdalgisa Caccone, Yale UniversityAndrew J. Crawford, Universidad de los Andes, ColombiaJerry Dodgson, Michigan State UniversityRobert C. Fleischer, Smithsonian InstitutionSusan Gabay-Laughnan, University of IllinoisStewart Grant, Alaska Dept of Fish & GameJ. Perry Gustafson, University of MissouriFangpu Han, Chinese Academy, ChinaScott Hodges, University of California, Santa BarbaraJennifer Jackson, British Antarctic Survey, United KingdomPrem P. Jauhar, USDA-ARS, North Dakota State UniversityWarren E. Johnson, Smithsonian Conservation Biology InstituteF. Andrew Jones, Oregon State UniversityStephen A. Karl, Hawaii Institute of Marine BiologySusan J. Lamont, Iowa State UniversityJohn Logsdon, University of IowaJose V. Lopez, Nova Southeastern University Oceanographic CenterTherese Markow, University of California, San DiegoWilliam S. Modi, American Genetic AssociationWilliam Murphy, Texas A&M UniversityMohamed Noor, Duke UniversityTaras Oleksyk, University of Puerto RicoKenneth Olsen, Washington University, St. LouisElaine Ostrander, American Genetic AssociationReid G. Palmer, USDA, Iowa State UniversityJill Pecon-Slattery, Smithsonian Conservation Biology InstituteRoger H. Reeves, Johns Hopkins UniversityHoward Ross, University of Auckland, New ZealandOliver Ryder, San Diego ZooHoracio Schneider, Universidade Federal do Para, BrazilHector Seuánez, Instituto Nacional de Cancer, BrazilBrad Shaffer, University of California, Los AngelesWilliam Sherwin, University of New South Wales, AustraliaChristopher I. Smith, Willamette UniversityTomoko Y. Steen, The Library of CongressJohn R. Stommel, USDA-ARS, Beltsville Agricultural Research CenterWilliam Tracy, University of WisconsinDavid B. Wagner, University of KentuckyRobin Waples, Northwest Fisheries Science Center, NMFSRobert Wayne, University of California, Los Angeles

The American Genetic Associationwww.theaga.org

PresidentRobin Waples

Past PresidentKerry Shaw

President ElectKatie Peichel

Executive Vice PresidentBrad Shaffer

SecretaryOliver Ryder

TreasurerKim Hughes

CouncilDavid BaumAnne BronikowskiLila FishmanSteve KarlCarlos MachadoMaria OriveElaine OstranderRobert WayneKelly Zamudio

Journal of Heredity

JHERED10501M

Page 4: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

© The American Genetic Association 2013. All rights reserved. For permissions, please e-mail: [email protected]

1

The Global Invertebrate Genomics Alliance (GIGA): Developing Community Resources to Study Diverse Invertebrate GenomesGIGA COMMUNITY OF SCIENTISTS*

Address correspondence to Dr. Jose V. Lopez, Oceanographic Center, Nova Southeastern University, 8000 North Ocean Drive, Dania Beach, FL 33004, or e-mail: [email protected].

*Authors are listed in the Appendix

Abstract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�DFURVV�WKH�LQWHUQDWLRQDO�VFLHQWLÀF�FRPPXQLW\��&DQGLGDWH�VWXG\�WD[D�LQFOXGH�VSHFLHV�IURP�3RULIHUD��&WHQRSKRUD��&QLGDULD��3ODFR]RD��0ROOXVFD��$UWKURSRGD��(FKLQRGHUPDWD��$QQHOLGD��%U\R]RD��DQG�3ODW\KHOPLQWKHV��DPRQJ�RWKHUV��*,*$�ZLOO�WDUJHW�LMMM%0)0204#6.I0)00#-+.)A#%45#62#4:%>2.7%+0%#-57+424%)0%-+$20#%.+J+%9#6+<4#%)* %.7#%<0$2"+,#A%57=,#.26%A2"#$42.=%20%.7#%RFHDQV��3ULRULWLHV�IRU�VHOHFWLQJ�LQYHUWHEUDWHV�IRU�VHTXHQFLQJ�ZLOO�LQFOXGH��EXW�DUH�QRW�UHVWULFWHG�WR��WKHLU�SK\ORJHQHWLF�SODFHPHQW��UHOHYDQFH�WR�RUJDQLVPDO��HFRORJLFDO��DQG�FRQVHUYDWLRQ�UHVHDUFK��DQG�WKHLU�LPSRUWDQFH�WR�ÀVKHULHV�DQG�KXPDQ�KHDOWK��:H�KLJKKOLJKW�EHQHÀWV�RI �VHTXHQFLQJ�ERWK�ZKROH�JHQRPHV��'1$��DQG�WUDQVFULSWRPHV�DQG�DOVR�VXJJHVW�SROLFLHV�IRU�JHQRPLF�OHYHO�GDWD�+66#44%+0A%47+$20?%9+4#A%)0%.$+045+$#06=%+0A%206,<42"#0#44B%N7#%DEDF%C#9%42.#%17..5OII?2?+B0)"+B#A<3%7+4%9##0%,+<067#A%.)%*+62,2.+.#%.724%6),,+9)$+.2"#%"#0.<$#BKey words: biodiversity, comparative genomics, consortium, evolution, GIGA, invertebrates, metazoa

N7#%,+4.%PMM%-2,,2)0%=#+$4%)* %#"),<.2)0%7+"#%9##0%-+$G#A%9=%WKH�GLYHUVLILFDWLRQ�RI �DQLPDO�OLIH��'HVSLWH�WKH�UDQJH�RI �ERG\�W\SHV� DQG� RUJDQLVPV� REVHUYHG� DPRQJ� WKH� 0HWD]RD�� VSDQK020?%4+,54:%45)0?#4:%47$2-54:%4@<2A4:%+0A%4#+%4.+$4:%+,,%+02-+,4%DURVH�IURP�D�FRPPRQ�DQFHVWRU��0HWD]RDQV�VKDUH�D�QXPEHU�)* %*#+.<$#4%.7+.%A24.20?<247%.7#-%*$)-%).7#$%)$?+024-4O%.7#=%+$#%+,,%-)4.,=%-<,.26#,,<,+$:%7#.#$).$)5726:%+0A%672#*,=%-).2,#%#<G+$=).#4% >2.7% 20.#$6#,,<,+$% H<06.2)04% +0A% +0% #J.$+6#,,<,+$%-+.$2J%)* %6),,+?#0%+0A%?,=6)5$).#204B%C2.7%.7#%#J6#5.2)0%)* %45#62#4% .7+.% +,4)% 5$)5+?+.#% +4#J<+,,=:% 4)-#% 4<55)4#A,=% *)$%+%,)0?%.2-#%1'DQFKLQ�HW�DO������3:%-#.+/)+04%A#"#,)5%*$)-%#-9$=)4%+$2420?%*$)-%+%A25,)2A%/=?).#%+0A%5+4420?%.7$)<?7%+%9,+4.<,+%4.+?#:%>7267%24%*),,)>#A%9=%6#,,%A2**#$#0.2+.2)0%+0A%-)$57)?#0#424% 16ODFN� HW� DO�� ������ 9DOHQWLQH� ������ 1LHOVHQ�������(UZLQ�DQG�9DOHQWLQH�����3B%N7#4#%5$)6#44#4%+$#%)$67#4K.$+.#A%9=%+%6)04#$"#A%A#"#,)5-#0.+,%.)),G2.:%206,<A20?%+%"+$2K#.=%)* %.$+046$25.2)0%*+6.)$4%+0A%42?0+,20?%5+.7>+=4:%2A#0.2*2#A%

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+9)<.%&'(%)* %-#.+/)+0%A2"#$42.=%1=KDQJ�����D3B%N7#%6)06#5.%)* %20"#$.#9$+.#%>+4%*2$4.%5$)5)4#A%9=%/DPDUFN�������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Journal of Heredity 2014:105(1):1–18doi:10.1093/jhered/est084

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Page 5: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

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Journal of Heredity

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Why Focus on Invertebrates?Phylogeny

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&HSKDORFKRUGDWD�� 7XQLFDWD�� &UDQLDWD�� (FKLQRGHUPDWD��+HPLFKRUGDWD��1HPDWRGD��$UWKURSRGD��$QQHOLGD��0ROOXVFD��3ODW\KHOPLQWKHV�� DQG� 5RWLIHUD��� EXW� WKHUH� DUH� FXUUHQWO\� QR�SXEOLVKHG�JHQRPHV�IRU�WKH�RWKHU���� LQYHUWHEUDWH�SK\OD��:H�#J+-20#A%6<$$#0.%57=,)?#0#.26%7=5).7#4#4%+0A%4#,#6.#A%G#=%20"#$.#9$+.#% 45#62#4% .7+.% 45+0% .7#% 57=,)?#0#.26% A2"#$42.=%+0A%-)$57),)?26+,%A245+$2.=%)0% .7#%+02-+,% .$##%)* % ,2*#% 14##%6XSSOHPHQWDU\�0DWHULDO���1HZ�LQYHUWHEUDWH�JHQRPH�GDWD�FDQ�$#"#+,%0)"#,%4#@<#06#4%>2.7%4<**262#0.%57=,)?#0#.26%42?0+,%.)%$#4),"#%,)0?4.+0A20?%@<#4.2)04B

Developmental Biology

!0#%)* %.7#%-)4.%#J62.20?%+55$)+67#4%.)%<0A#$4.+0A20?%.7#%)$2?204% +0A% #"),<.2)0% )* % +02-+,% A2"#$42.=% 24% A#.+2,#A% 6)-K5+$+.2"#%$#4#+$67%)0%#-9$=),)?26+,%A#"#,)5-#0.%12B#B:%#"),<K.2)0+$=% A#"#,)5-#0.+,% 92),)?=:% )$% ,4#52,4#3B%C7),#K?#0)-#%+0A%.$+046$25.)-#%20*)$-+.2)0%6+0%*+62,2.+.#%.7#%4<66#44%)* %4<67%$#4#+$67:%H<4.%+4%6#7%?#0#%67+$+6.#$2/+.2)04%A2A%A#6+A#4%9#*)$#%1,NXWD�����3B%D#0#%6)0.#0.:%4=0.#0=:%?#0#%6)5=%0<-K9#$:%+0A%624K$#?<,+.)$=% 20*)$-+.2)0%+$#%4)-#%)* %.7#%6$<62+,%A+.+%.7+.%-+=%#0+9,#%<4%.)%+AA$#44%.7#%?#0).=5#X57#0).=5#%A2,#--+B%C7),#K?#0)-#%4#@<#0620?%5$)"2A#4%.7#4#%A+.+%20%+%6)-5$#7#042"#% *+472)0B% E0% .7#% ,+4.%A#6+A#:%#"#$=% 20"#$.#K9$+.#%?#0)-#%4#@<#06#%5+5#$%7+4%.<$0#A%.7#4#%4#,#6.%45#62#4%20.)%#-#$?20?%+0A%2-5)$.+0.%A#"#,)5-#0.+,%-)A#,%4=4.#-4%1'HKDO� HW� DO�� ������ 3XWQDP� HW� DO�� ����:% ������ Q$2"+4.+"+%HW� DO�� ����3B% N7#% +AA2.2)0+,% ,4#52,4#% 5<9,26+.2)04% 7+"#% ?#0K#$+.#A%-+H)$% 2042?7.4% 20.)% .7#%-),#6<,+$%9+424% )* %-)$57)K,)?26+,% A2"#$42.=% 1'DYLGVRQ� DQG�(UZLQ� ������+HMQRO� ����3B%E0"#$.#9$+.#%.+J+%+,4)%206,<A#%4)-#%)* %.7#%,)0?#4.%,2"20?%+02K-+,4%)0%.7#%5,+0#.:%4<67%+4%.7#%)6#+0%Y<+7)?%6,+-%8"9/'9)*'$()-2'9)% 1-+J2-<-% $#5)$.#A% +?#% 'ML% =#+$43:% :)+,(('1")9&')%WXEH�ZRUPV��a����\HDUV���0XQUR�DQG�%OLHU�����3:%6)$+,,20#%A#-)45)0?#%8$/"#$9(,")*;'((,<)-)%1'P'%Z%LM%=#+$43%1C[$7#2A#%����3:% ?),A%6)$+,%=,")"2')� VS�� ����²�����\HDUV���EODFN�FRUDO�:,'#%)/&,$*.()1,""'+)��HVW��a�����\HDUV���5RDUN�HW�DO������3:%+0A%.7#%2--)$.+,%6<2")%1%RHKP�HW�DO������3

Speciation, Radiations, and Evolutionary Rates

E0"#$.#9$+.#4%7+"#% 6)042A#$+9,#%5).#0.2+,% .)% 20*)$-% *<0A+K-#0.+,% @<#4.2)04% 20% ).7#$% +45#6.4% )* % #"),<.2)0+$=% 92),)?=:%4<67% +4% 7)>% 0#>% 45#62#4% *)$-:%>7=% $+A2+.2)04% )66<$:% +0A%.7#% ?#0#.264% )* % #"),<.2)0+$=% 4.+424B% E0"#$.#9$+.#% 424.#$% 45#KFLHV�SDLUV�QRZ�UHVWULFWHG�WR�WKH�&DULEEHDQ�RU�HDVWHUQ�3DFLILF�E\� WKH� XSOLIW� RI � WKH� ,VWKPXV� RI � 3DQDPD� SURYLGH� D� VXSHUE�

1'XQQ�HW�DO��������+HMQRO�HW�DO��������3KLOLSSH�HW�DO��������6FKLHUZDWHU�HW�DO��������3LFN�HW�DO��������5\DQ�HW�DO��������1RVHQNR�HW�DO���������7KH�SRVLWLRQ�RI �;HQDFRHORPRUSKD�LV�XQFHUWDLQ��GDVKHG�OLQHV��DQG�LV�EDVHG�RQ�UHFHQW�VWXGLHV��LW�FRXOG�EH�WKH�VLVWHU�JURXS�WR�%LODWHULD��+HMQRO�HW�DO���������RU�DV�SDUW�RI �'HXWHURVWRPLD��3KLOLSSH�HW�DO���������7KHUH�LV�QR�SK\ORJHQRPLF�GDWD�IRU�0HVR]RD�RU�0LFURJQDWKR]RD��DQG�WKHLU�SRVLWLRQV�DUH�EDVHG�RQ�D�OLPLWHG�DPRXQW�RI �GLUHFW�VHTXHQFLQJ�GDWD��0HVR]RD��FRPSULVLQJ�5KRPER]RD�DQG�2UWKRQHFWLGD��LV�VKRZQ�DV�D�VLQJOH�WHUPLQDO�EXW�PD\�QRW�EH�D�FODGH��7KH�SODFHPHQW�RI �3KRURQLGD�LV�EDVHG�LQ�WKH�WUDGLWLRQDO�SODFHPHQW�RI �WKH�JURXS�DV�FORVH�WR�%UDFKLRSRGD��7KH�HVWLPDWHG�QXPEHUV�IRU�H[WDQW�GHVFULEHG�DQG�YDOLG�VSHFLHV��LQGLFDWHG�LQ�5+$#0.7#4#4%+*.#$%.7#%.#$-20+,%0+-#:%+$#%*$)-%=KDQJ������D�%+0A%*)$%-+$20#%?$)<54:%*$)-%@<#$2#4%)* %.7#%C)$,A%\#?24.#$%)* %0DULQH�6SHFLHV��-XQH��������7..5OII>>>B-+$20#45#62#4B)$?I+572+B575]5^4.+.4��1RWHV�IRU�WHUPLQDOV�LQGLFDWHG�E\� ��0HVR]RD�16)-5$2420?%\7)-9)/)+%+0A%!$.7)0#6.2A+3%24%47)>0%+4%+%420?,#%.#$-20+,%9<.%-+=%0).%9#%+%6,+A#B%N7#%45#62#4%6)<0.%*)$%\).2*#$+%LQFOXGHV�$FDQWKRFHSKDOD�DQG�IRU�&QLGDULD�LQFOXGHV�0\[R]RD�

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5

Table 1� 3XEOLVKHG�RU�LQ�SURJUHVV�ZKROH�QRQLQVHFW�QRQQHPDWRGH�LQYHUWHEUDWH�JHQRPH�SURMHFWV

Species Major group Subgroup Size (MB)

URL Citation

6,(#12,(()*"#1>$/) F00#,2A+ &OLWHOODWD ������ 7..5OII?#0)-#BH?2K54*B)$?I+HOUR��+HOUR��KRPH�KWPO

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Journal of Heredity

6

-)A#,% 4=4.#-% *)$% 20"#4.2?+.20?% $+.#4% +0A% 46+,#4% )* % 45#62+K.2)0% 20% .7#% 4#+% 1.QRZOWRQ� DQG�:HLJW� ������ /HVVLRV� ����3B%Q5#6.+6<,+$% 20"#$.#9$+.#% $+A2+.2)04:% 4<67% +4% .7#% +-5725)A%FUXVWDFHDQV�RI �/DNH�%DLNDO� �0DF'RQDOG� HW� DO�� ����3:% 6)-K5,#-#0.% "#$.#9$+.#% ,+G#% $+A2+.2)04% #,4#>7#$#B% N7#% 4#+% 4.+$:%0"<%/)$/,"'-)*&<$/,"):%7+4%6,)6G#A%)0#%)* %.7#%*+4.#4.%45#62+.2)0%$+.#4% 1aPMMM%=#+$43%)0% $#6)$A% 13XULW]� HW� DO�� �������%\�FRQK.$+4.:% 20"#$.#9$+.#4% +,4)% 206,<A#% *)$-4% .7+.% +55#+$% .)% 7+"#%$#-+20#A%+.% ,#+4.%4<5#$*262+,,=%<067+0?#A%)"#$% ,)0?%5#$2)A4%)* %?#),)?26+,% .2-#B%D#0)-26%+0+,=424%-+=%$#"#+,%$#?<,+.)$=%4#@<#06#4% 20"),"#A% 20%-)$57),)?26+,% 4.+424% 1#B?B:%7)$4#47)#%6$+94:%9,<#%6)$+,%6,('#%#"):%A##5K4#+% ,)94.#$%G,#.(<%&,)* '-#%'5-)/):%46,#$)45)0?#43:%+4%>#,,%+4%5$)"2A#%2042?7.%20.)%$+.#4%)* %#"),<.2)0+$=%67+0?#B

Ecology

E0"#$.#9$+.#4%+$#%-+H)$%6)-5)0#0.4%)* %-+$20#:%*$#47>+.#$:%+0A%.#$$#4.$2+,%#6)4=4.#-4B%Q)-#%20"#$.#9$+.#4%+$#%#6)4=4.#-%#0?20##$4:%4<67%+4%.7#%46,#$+6.202+0%6)$+,4%.7+.%6)04.$<6.%$##* %KDELWDWV� FRYHULQJ�RQO\������RI � DOO�RFHDQLF� VXUIDFH� DUHD� \HW�VXSSRUWLQJ�XS� WR�����RI � WRWDO�PDULQH�ELRGLYHUVLW\� �\#+G+K.XGOD�����3B%E0"#$.#9$+.#4%45+0%+,,%.$)5726%,#"#,4:%*$)-%5$2K-+$=%6)04<-#$4%.7$)<?7%>2A#,=%<.2,2/#A%5$#=%1#B?B:%5,+0G.)026%6)5#5)A43% .)%5+$+42.#4% +0A%+5#J%5$#A+.)$4% 1#B?B:%T<-9),A.%4@<2A3B%E0%.#$-4%)* %92)-+44:%4)-#%20"#$.#9$+.#4%4<67%+4%G$2,,%1#<57+<422A43% +0A% 6)5#5)A4% A)-20+.#% 5#,+?26% *))A% >#94%1%XLWHQKXLV� HW� DO�� ������$WNLQVRQ� HW� DO�� �������%HQWKLF� DQG�5,+0G.)026% 20"#$.#9$+.#4% 5,+=% 6$<62+,% $),#4% 20% 6+$9)0% +0A%0<.$2#0.%6=6,20?%20%.7#%)6#+0B%E0%A##5#$%>+.#$4%>2.7)<.%4<0K,2?7.:%>+.#$K6),<-0%20"#$.#9$+.#4%6=6,#%6+$9)0%.7$)<?7%.7#2$%.$)5726%0#.>)$G4:%+0A%9#0.726%+02-+,4%1#B?B:%.<9#>)$-4:%-<4K4#,4:% 6,+-4:% ?+4.$)5)A43% *)$-%5+$.0#$47254%>2.7% 67#-)4=0K.7#.26% 9+6.#$2+% .)% A)-20+.#% +0A% A#*20#% #6)4=4.#-4% 4<67% +4%7=A$).7#$-+,%"#0.%6)--<02.2#4%+0A%-#.7+0#%4##54%1D#$-+0%HW�DO������3B%D#0)-#4%)* %)$?+024-4%.7+.%,2"#%20%7+$47%#0"2K$)0-#0.4%12B#B:%#J.$#-)572,#43%)$%20%20.2-+.#%4=-92)424%>2.7%).7#$%)$?+024-4%1#B?B:%6)$+,X+,?+,%4=-92)4#4:%72?7,=%+A+5.#A%5+$+42.#43%7+"#% .7#%5).#0.2+,% .)% $+A26+,,=%67+0?#%)<$%<0A#$K4.+0A20?%)* %7)>%.7#4#%)$?+024-4%4<$"2"#%15XVVHOO�HW�DO��������)ORW�HW�DO������3B

Fisheries and Aquaculture

E0"#$.#9$+.#4% +$#% 9#6)-20?% 206$#+420?,=% 2-5)$.+0.% 4)<$6#4%RI �SURWHLQ�IRU�KXPDQ�QXWULWLRQ�ZRUOGZLGH��3DUWLFXODUO\�ZLWK�.7#%6),,+54#%)* %+%0<-9#$%)* %"#$.#9$+.#%*247#$2#4:%20"#$.#9$+.#%*247#$2#4%+0A%+@<+6<,.<$#%A)-20+.#A%9=%-),,<4G4%+0A%6$<4.+K6#+04%5$)"2A#%*))A%+0A%#-5,)=-#0.%*)$%-2,,2)04%)* %5#)5,#%1*ODVHU� DQG�'LHOH� ������� 3URPLQHQW� H[DPSOHV� DUH� ELYDOYHV�1)=4.#$4:%6,+-4:%-<44#,43:%?+4.$)5)A4%1+9+,)0#:%@<##0%6)0673:%+0A%6#57+,)5)A4%14@<2A:%)6.)5<4:%+0A%6<..,#*2473%+-)0?%.7#%-),,<4G4%16WRQHU�����3:%+0A%+%"+$2#.=%)* %A#6+5)A4%147$2-54:%6$+94:% ,)94.#$4:% +0A% 6$+=*2473% +-)0?% 6$<4.+6#+04% 11HLODQG�HW�DO������3B%D))4#0#6G%9+$0+6,#4%+$#%+-)0?%.7#%-)4.%#J5#0KVLYH�FUXVWDFHDQ�VHDIRRG�LQ�:HVWHUQ�(XURSH�RQ�D�SHU�NLORJUDP�EDVLV�� (FKLQRGHUPV� �VHD� XUFKLQV� DQG� VHD� FXFXPEHUV�� KDYH�LPSRUWDQW� FRPPHUFLDO� ILVKHULHV�� DQG� VSHFLHV� RI � &QLGDULD��$QQHOLGD�� 7XQLFDWD�� &HSKDORFKRUGDWD�� DQG� 6LSXQFXOD� DUH�

+,,% 7+$"#4.#A% *)$% 7<-+0% 6)04<-5.2)0% 20% 4)-#% 6)<0.$2#4B%%Q)-#% 20"#$.#9$+.#4:% 4<67% +4% G$2,,:% +$#% 7+$"#4.#A% *)$% 7<-+0%0<.$2.2)0%-)$#%20A2$#6.,=:%"2+%+02-+,%*##A%)$%A2#.+$=%4<55,#KPHQWV��)LQDOO\��QRW�DOO�ILVKHG�VSHFLHV�DUH�IRRG�VRXUFHV��SHDUO�)=4.#$4:% +9+,)0#:% -<44#,4:% 40+2,4:% 9,+6GI$#A:% +0A% 4)*.% 6)$K+,4%+$#%7+$"#4.#A%*)$% .7#2$%"+,<#% 20% .7#% H#>#,$=%)$%+@<+$2<-%.$+A#4B%F% 0<-9#$% )* % ,+$?#%-),,<4G4% +$#% 42-2,+$,=% *247#A% .)%4<55)$.%.7#%.)<$24-%-+$G#.B

Invasive and Pest Species

F%0<-9#$%)* %20"#$.#9$+.#4%6+<4#%#0"2$)0-#0.+,%5$)9,#-4%+$)<0A%.7#%>)$,AB%Q)-#%+$#%20"+42"#%45#62#4%.7+.%7+"#%6),)K02/#A%+0A%#J5+0A#A%.7#2$%0<-9#$4%20.)%0#>%7+92.+.4:%)*.#0%WR� WKH� GHWULPHQW� RI � QDWLYH� VSHFLHV�� ([DPSOHV� LQFOXGH� WKH�?$##0%6$+9%10)"9'->$*+),-)$3%+0A%.7#%/#9$+%-<44#,%1!",'$$,-)*%#(<+#"%&)�� LQ� 1RUWK� $PHULFD�� WKH� RUDQJH� FXS� FRUDO�1A>1)$/"),)* 9#99'-,)�� LQ� WKH�*XOI �RI �0H[LFR�DQG�&DULEEHDQ��+0A%.7#%?2+0.%F*$26+0%,+0A%40+2,%1:'$$)9&)/'-)*F>('9)3%20%-+0=%.$)526+,%+0A%4<9.$)526+,%+$#+4B%N7#%/#9$+%-<44#,%6+<4#4%-2,K,2)04% )* % A),,+$4% )* % #6)0)-26% A+-+?#% #+67% =#+$% 9=% 6,)?K?20?%525#4%6+$$=20?%*$#47>+.#$:%+0A% .7#2$%A#4.$<6.2"#% $+52A%?$)>.7% +0A% 92)*)<,20?% .7$#+.#0% 0+.2"#% 45#62#4% )* % +,$#+A=%#0A+0?#$#A% 5#+$,% -<44#,4B% !.7#$% 20"#$.#9$+.#% 45#62#4% +$#%QDWLYH� EXW� KDYH� UHFHQWO\� EHFRPH� SHVWV�� LQ� WKH� RFHDQ�� WKLV�.=526+,,=%)66<$4%9#6+<4#%)* %206$#+4#4%20%0<.$2#0.%,#"#,4%.7+.%4<55)$.%72?7#$%$#6$<2.-#0.%)$%?$)>.7%$+.#4%)* %.7#%5#4.%45#K62#4:% )$% 9#6+<4#% )"#$*24720?% )* % .7#% 5$#A+.)$4% )* % .7#% 5#4.%45#62#4% 7+4% +,,)>#A% .7#-% .)% -<,.25,=% <06)0.$),,#A% 1D+,2,%����3B% W+$?#% +??$#?+.2)04% )* % 4)-#%-#A<4+% 45#62#4% 0#?+K.2"#,=% 2-5+6.% *247#$2#4:% .)<$24-:% +0A% 5)>#$% 5,+0.% 6)),20?%4.+.2)04B%!0%6)$+,%$##*4:%.7#%-)4.%A$+-+.26%4<67%5#4.%24%.7#%6$)>0K)*K.7)$04% 4.+$*247:% 89)-/&)$/,"* %()-9':% +% -+H)$% *+6K.)$%6)0.$29<.20?%.)%.7#%A#6,20#%)* %6)$+,%6)"#$%20%.7#%E0A)K3DFLILF��'H·DWK�HW�DO������3B

Symbiosis

F% >2A#% $+0?#% )* % 20"#$.#9$+.#4% *)$-4% )9,2?+.#% +44)62+.2)04%>2.7%-26$))$?+024-4% .)% #4.+9,247% +%7),)92)0.% 17)4.%b% .).+,%4=-92)0.% 6)--<02.=3% .7+.% #06)-5+44#4%-<,.25,#% 4=-92).26%20.#$+6.2)04B%Q=-92).26%-26$)9#4%6+$$=%)<.%45#62*26%-#.+9),26%+0A%57=42),)?26+,% +6.2"2.2#4% .7+.% +$#%-<.<+,24.26:% 6)--#04+,:%)$%0#<.$+,%>2.720%.7#2$%7)4.B%S)$%#J+-5,#:% .7#%-+442"#%6)$+,%$##* %4.$<6.<$#4:%6$2.26+,%.)%.$)526+,%-+$20#%#6)4=4.#-4%+0A%+#4K.7#.26+,,=% 5,#+420?% .)% .)<$24.4%>)$,A>2A#:% 6)<,A% 0).% A#"#,)5%>2.7)<.% .7#% 57).)4=0.7#.26% +6.2"2.=% )* % 4=-92).26% +,?+#% ,2"K20?%>2.720%6)$+,%.244<#4%10XVFDWLQH�DQG�&HUQLFKLDUL�����3B%F4%-#0.2)0#A% +9)"#:% 7=A$).7#$-+,% "#0.% 6)--<02.2#4% $#,=% )0%67#-)4=0.7#.26% 4=-92)4#4B% Q+6)?,)44+0% 4#+% 4,<?4% 6)04<-#%+,?+#%+4%*))A:%+0A%+,4)%$#.+20%+0A%20.#?$+.#%.7#2$%67,)$)5,+4.4%20.)% .7#2$% .244<#4% .)% 6)0.20<#% 57).)4=0.7#424% *)$% .7#% 7)4.%4,<?U4%0<.$2.2)0%13LHUFH�DQG�&XUWLV�����3B%S)$%7<-+04:%$#6#0.%´+XPDQ�0LFURELRPH�SURMHFWµ�KDV�VLJQLILFDQWO\�LPSURYHG�WKH�462#0.2*26%+0A%5<9,26%$#6)?02.2)0%)* %9#0#*262+,%4=-92)0.%#6),K)?=%+0A%*<06.2)04%*)$%7)4.%7#+,.7%+0A%A#"#,)5-#0.%1T<-+0%0LFURELRPH� 3URMHFW� ����3B% N7<4:% .7#% $#,+.2"#,=% 4-+,,% 9)A=%42/#4%)* %-+0=% 20"#$.#9$+.#4% 6)<5,#A%>2.7% .7#2$% .#0A#06=% .)%6<,.2"+.#%-26$)92+,%4=-92)4#4%206$#+4#%.7#2$%"+,<#%+4%.$+6.+9,#%

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7

-)A#,4%*)$%#J5,)$20?%4=-92)424%4.<A2#4%20%?$#+.#$%A#5.7%1%RVFK�DQG�0F)DOO�1JDL�����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�6LOYD�HW�DO������3B

Threatened, Endangered, and Remarkable Species

\#,+.2"#,=% *#>% 20"#$.#9$+.#4%7+"#%9##0% ,24.#A%+4%#J.206.:%9<.%+%0<-9#$%)* %.7#-%+$#%.7$#+.#0#A%)$%#0A+0?#$#AB%F9+,)0#4%16)('#/'$% 455B3% >#$#% .7#% *2$4.% 20"#$.#9$+.#4% ,24.#A% <0A#$% .7#%(QGDQJHUHG� 6SHFLHV� $FW�� DQG� DOO� VWRQ\� FRUDO� VSHFLHV� DUH�OLVWHG� XQGHU�$SSHQGL[� ,,� RI � WKH�&RQYHQWLRQ�RQ� WKH�7UDGH�RI � (QGDQJHUHG� 6SHFLHV� �&,7(6��� 2WKHU� LQYHUWHEUDWHV� IDVK620+.#% .7#% 5<9,26% 9=% "2$.<#% )* % .7#2$% 42/#:% 4<67% +4% .7#% ?2+0.%4@<2A% 18"9&'/,>/&'$* 2>73:% .7#% ?2+0.% 6,+-% 1A"'2)9-)* .'.)$3:% +0A%.7#%0#-#$.#+0%9)).,+6#%>)$-%:'-,>$* (#-.'$$'+>$:%>7267%-+=%UHDFK� ���P� LQ� OHQJWK�� SHUKDSV� WKH� ORQJHVW� OLYLQJ� RUJDQLVP�15XSSHUW� HW� DO�� ������� 7KH� GHFDGH�ORQJ� &HQVXV� RI �0DULQH�W2*#% 47)>6+4#A% 7)>% 4-+,,:% 9#+<.2*<,:% +0A% <0<4<+,K,))G20?%20"#$.#9$+.#4%6+0%#0?+?#%.7#%5<9,26%1.QRZOWRQ�����3B%N7<4:%?#0)-#%4#@<#06#4%*$)-%.7#4#%+0A%).7#$%20"#$.#9$+.#%45#62#4%6+0%9#%"2#>#A%+4%+,.#$0+.2"#%)$%5+$+,,#,%-#+4<$#4%.)%/))4%+0A%*$)/#0% .244<#%6),,#6.2)04% *)$% .7#% 6)04#$"+.2)0%)* %92),)?26+,%+0A%?#0#.26%A2"#$42.=%15\GHU�����3B

Relevance to Human Health

0DULQH� LQYHUWHEUDWHV� DUH� WKH� VRXUFH� RI � WHQV� RI � WKRXVDQGV�)* % 92)67#-26+,% 6)-5)<0A4% >2.7% 5).#0.2+,% 7<-+0% 7#+,.7%+55,26+.2)04% 1)DXONQHU�������%OXQW� HW� DO�� ����3B% E0% .7#% ,+4.%WZR� GHFDGHV� DORQH� �����²������� QDWXUDO� SURGXFWV� UHVHDUFK�RQ�3RULIHUD� �'HPRVSRQJLDH�� DQG�&QLGDULD� �$QWKR]RD�� KDV�$#4<,.#A% 20%.7#% ,+$?#4.%0<-9#$%)* %67#-26+,%8,#+A;%-),#6<K,+$%4.$<6.<$#4%*)$%A246)"#$=%)* %0)"#,%57+$-+6#<.26+,4%1W#+,%HW� DO�� ����3B% N7#4#% 206,<A#% .7#% 4#442,#% -+0?$)"#% .<026+.#:%L9/,'-)$9'2')*/>"1'-)/)M%+0A%.7#%A#"#,)5-#0.%)* %.7#%+0.2K6+0KFHU�GUXJ��(7������<RQGHOLV����*DVWURSRG�VSHFLHV�RI �0#->$%SURGXFH�PRUH� WKDQ� ���� ELRDFWLYH� SHSWLGHV� LQ� WKHLU� SRWHQW�"#0)-:%>2.7%,2..,#%)"#$,+5%)* %.)J204%+-)0?%45#62#4%1!,2"#$+%DQG�7HLFKHUW�����3B%N7#%.)J204%#J7292.%+%>2A#%$+0?#%)* %0#<K$),)?26+,%#**#6.4:%4<67%+4%5+$+,=424%+0A%+0+,?#42+:%+0A%)0#%)* %.7#%5#5.2A#4% 24%6,2026+,,=%+"+2,+9,#%*)$% .$#+.-#0.%)* %67$)026%5+20B%C2.7%$+$#%#J6#5.2)04%12OLYHUD�DQG�7HLFKHUW�����3:%.7#%.+$?#.4%+0A%$),#4%)* %.7#4#%67#-26+,4%20%.7#%+02-+,4%5$)A<6K20?% .7#-% +$#% 0).% G0)>0B% C7),#K?#0)-#% 4#@<#06#4% *)$%20"#$.#9$+.#%45#62#4%.7+.%5$)A<6#%92)+6.2"#%6)-5)<0A4%>2,,%5$)"2A#%2042?7.4%20.)%.7#2$%67#-26+,%$),#4%20%0+.<$#:%.7#2$%#")K,<.2)0:% +0A% .7#%-#67+024-4% .7+.% 6)0.$),% .7#2$% 5$)A<6.2)0B%Q#@<#06#4%-+=% +,4)% *+62,2.+.#% .7#%A246)"#$=%)* %).7#$% 6)-K5)<0A4%)$%5$)6#44#4%>2.7%7<-+0%7#+,.7%)$%92).#670),)?26+,%

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PolicyF%-+20% ?)+,% )* % DEDF% 24% .)% 9<2,A% +0% 20.#$0+.2)0+,% -<,.2KA24625,20+$=% 6)--<02.=% .)%5<$4<#% 6)-5+$+.2"#% 20"#$.#9$+.#%?#0)-26%4.<A2#4B%C#%4##G%.)%A#"#,)5%.)),4%.)%#0+9,#%+0A%*+62,K2.+.#%?#0)-26%$#4#+$67%+0A%#06)<$+?#%6),,+9)$+.2)0B%C#%>2,,%A#"#,)5%4.+0A+$A4%.7+.%#04<$#%A+.+%@<+,2.=:%6)-5+$+92,2.=:%+0A%LQWHJUDWLRQ��%\�FRRUGLQDWLQJ�VDPSOH�FROOHFWLQJ�DQG�VHTXHQFK20?% #**)$.4% +-)0?% 20"#$.#9$+.#% 92),)?24.4:% >#% +2-% .)% +")2A%A<5,26+.2)0%)* %#**)$.%+0A%,#"#$+?#%$#4)<$6#4%-)$#%#**262#0.,=B%C#%#0"242)0% +% 462#0.2*26% 6)--)04%>7#$#% 47+$#A% $#4)<$6#4:%A+.+:% A+.+% 4.+0A+$A4:% +0A% 200)"+.2)04%-)"#% .7#% ?#0#$+.2)0%+0A%+0+,=424%)* %20"#$.#9$+.#%?#0)-26%A+.+%.)%+%,#"#,%.7+.%,2G#,=%6)<,A%0).%9#%+672#"#A%>2.7%.7#%.$+A2.2)0+,%52#6#-#+,%420?,#K20"#4.2?+.)$XA$2"#0%+55$)+67BDEDF%#-9$+6#4%+% .$+045+$#0.%5$)6#44%)* %5$)H#6.% 6))$K

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Journal of Heredity

8

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Journal of Heredity

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Polypodiozoa (1)

Ctenophora (192)

Cubozoa (40)

Trachylina (157)

Scyphozoa (186)

Malacosporea (4)

Hydroidolina (3395)

Hexacorallia (3578)

Staurozoa (48)

Placozoa (1)

Octocorallia (3549)

Myxosporea (2180)

Heteroscleromorpha (5341)

Keratosa (558)

Calcaronea (514)

Haploscleromorpha (1084)

Hexasterophora (513)

Calcinea (167)

Amphidiscophora (177)

Myxospongiae (138)

Oscarellidae (19)

Plakinidae (68)

Demospongiae

Hexactinellida

Calcarea

Homoscleromorpha

Anthozoa

Myxozoa

Hydrozoa

glass sponges

sponges

Acraspeda

Porifera

Cnidaria

Bilateria(other animals)

Medusozoa

soft corals, sea pens, gorgonians

corals, sea anemones

stalked jellies

box jellies

true jellies

comb jellies

calcareous sponges

bath sponges

Hydra, siphonophores etc.

flesh sponges

marine & freshwater jellies

Figure 2.� &RQVHQVXV�SK\ORJHQ\�RI �QRQELODWHULDQ�DQLPDOV��7KH�3RULIHUD�WUHH�DQG�FODVVLÀFDWLRQ�LV�EDVHG�RQ�&iUGHQDV�HW�DO��������:%1RVHQNR�HW�DO��������:%+0A%:|UKHLGH�HW�DO����������7KH�&QLGDULD�WUHH�DQG�FODVVLÀFDWLRQ�LV�EDVHG�RQ�&ROOLQV��������DQG�(YDQV�HW�DO��1����:%����3B%S),,)>20?%.7#%.#$-20+,%0+-#:%20%5+$#0.7#4#4:%24%.7#%#4.2-+.#A%0<-9#$%)* %#J.+0.%A#46$29#A%+0A%"+,2A%45#62#4%*)$%.7+.%JURXS�EDVHG�RQ�WKH�:RUOG�5HJLVWHU�RI �0DULQH�6SHFLHV�DV�GHVFULEHG�LQ�WKH�)LJXUH���OHJHQG��DQG�0\[R]RD�HVWLPDWHV�VWHP�IURP�W)-%DQG�'\NRYi�������B%C7#0%+"+2,+9,#:%6)--)0%0+-#4%+$#%>$2..#0%+9)"#%.7#%9$+067#4B

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-+,4%+0A%206,<A#4%.7#%"#$.#9$+.#4%1)LJXUHV��%+0A%'3B%F-)0?%LWV� QRQYHUWHEUDWH� PHPEHUV�� 'HXWHURVWRPLD� FRPSULVHV� D�>2A#%A2"#$42.=%)* %*)$-4%$+0?20?%*$)-%-26$)46)526%.)%7<?#:%*+-2,2+$% .)% 92/+$$#:% ?#,+.20)<4% .)% +$-)$K5,+.#A:% +0A%>)$-KOLNH�WR�SODQW�OLNH��$SDUW�IURP�&UDQLDWD��LQFOXGLQJ�9HUWHEUDWD���WKH� JURXS� LQFOXGHV� IRXU� PDMRU� FODGHV�� (FKLQRGHUPDWD��+HPLFKRUGDWD��7XQLFDWD��DQG�&HSKDORFKRUGDWD��DOO�H[FOXVLYHO\�PDULQH�� (FKLQRGHUPDWD� DQG� +HPLFKRUGDWD� IRUP� D� FODGH�G0)>0%+4%F-9<,+6$+$2+% 1S2?<$#%'3:%67+$+6.#$2/#A%9=%42-2,+$%A25,#<$<,+%,+$"+,%-)$57),)?=%+0A%4<55)$.#A%20%-)4.%57=,)KJHQHWLF�DQDO\VHV��7XQLFDWD�DUH�WKH�FORVHVW�UHODWLYHV�WR�&UDQLDWD�DQG�DORQJ�ZLWK�&HSKDORFKRUGDWD� IRUP�&KRUGDWD� �)LJXUHV��+0A%'3B%E.%+,4)%7+4%9##0%4<??#4.#A%.7+.%d#0+6)#,)-)$57+%+$#%.7#%424.#$%?$)<5%.)%F-9<,+6$+$2+%$+.7#$%.7+0%9#20?%5,+6#A%20%+%-)$#%9+4+,%5)42.2)0%+4%4##0%20%)LJXUH��B

Diplopoda (7753)

Pycnogonida (1323)

Symphyla (197)

Remipedia (18)

Pauropoda (835)

Branchiopoda (1184)

Thecostraca (1424)

Mystacocarida (13)

Cephalocarida (13)

Copepoda (15976)

Arachnida (110615)

Ostracoda (7882)

Malacostraca (40065)

Xiphosura (4)

Chilopoda (3100)

Ichthyostraca (339)

Chelicerata

Myriapoda

Pancrustacea

Xenocarida

Oligostraca

Vericrustacea

horsehoe crabs

sea spiders

centipedes

millipedes

seed shrimps

spiders, scorpions, harvestmen, ticks, mites.

horsehoe shrimp

Entognatha (9734)

Insecta (1020007)

Mandibulata

Hexapoda

garden centipedes

fairy shrimps, water fleas, clam shrimp etc.

barnacles

fish lice, tongue worms

crabs, lobsters, amphipods, isopods, krill

copepods

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Journal of Heredity

12

ConclusionsN7#%+04>#$4%.)%*<0A+-#0.+,%@<#4.2)04%+9)<.%.7#%)$2?204%)* %+02-+,% ,2*#%+0A%.7#%#"),<.2)0%)* %.7#2$%A2"#$4#%57#0)K.=5#4%-+=%9#%7#,A%20%.7#%?#0)-#4%)* %A24.206.%20"#$.#9$+.#%57=,+B% Q#,A)-K4.<A2#A% .+J+% -+=% 9#% G#=% .)% A246)"#$20?%7)>%0#<$)04:%-<46,#4:%"242)0:%7)$-)0#4:%2--<02.=:%,2-9%$#?#0#$+.2)0:% 6+-)<*,+?#:% 92),<-20#46#06#:% 6))5#$+K.2)0:%)$?+024-+,%,)0?#"2.=:%+0A%57=42),)?26+,%6)-5,#J2.2#4%#"),"#A%1'HKDO�HW�DO��������+RIPDQQ�HW�DO��������3XWQDP�#.% +,B% ������ ������ 6ULYDVWDYD� HW� DO�� ������� 'HILQLWLYH�+04>#$4% .)% .7#4#% @<#4.2)04% 7+"#% #,<A#A% 92),)?24.4% *)$%

6#0.<$2#4:% 9<.% .#670),)?26+,% 9$#+G.7$)<?74% 7+"#% 0)>%-+A#%2.%*#+429,#%.)%5#$*)$-%?#0)-#K46+,#%4.<A2#4%)* %0)0K-)A#,%)$?+024-4B*HQRPLF� UHVHDUFK� LQ� WKH� ��4.% 6#0.<$=% .)<67#4% -+0=%

+$#+4%)* %92),)?26+,% #0@<2$=B%N7#%92??#4.% 67+,,#0?#% *+620?%.7#% 462#0.2*26% 6)--<02.=% 7+4% 472*.#A% *$)-% .7#% +6@<242K.2)0% )* % 4#@<#06#% A+.+% .)% .7#% +0+,=424% )* % .7#% ,+$?#% A+.+%4#.4% ?#0#$+.#AB% DEDF% >2,,% .7<4% +..#-5.% .)% 5$)"2A#%?#0)-264%4<55)$.%+,,)>20?%.7#%20"#$.#9$+.#%92),)?=%6)-K-<02.=% .)% )"#$6)-#% .7#4#% 67+,,#0?#4%>72,#% +55,=20?% .7#%,+.#4.% +A"+06#4% *$)-% .7#% 6)--<02.=% )* % 6)-5<.+.2)0+,%92),)?24.4B

Clitellata (9485)

Amphinomida (230)

Cirratuliformia (539)

Sipuncula (148)

Spionida (792)

Eunicida (1444)

Phyllodocida (4615)

Capitellida (1757)

Terebelliformia (995)

Sabellida (1705)

Canalipalpata

AnnelidaAciculata

Chaetodermomorpha (130)

Gastropoda (62000)

Neomeniomorpha (280)

Bivalvia (9532)

Polyplacophora (989)

Monoplacophora (30)

Cephalopoda (835)

Scaphopoda (578)

Aplacophora

Brachiopoda (388)

Nemertea (1285)

Phoronida (18)

spaghetti worms etc.

featherworms, pogonophores etc.

earthworms, leeches etc.

fireworms

mud-blister worms etc.

peanut worms

paddleworms, bloodworms etc.

lugworms, spoonworms etc.

palolo worms, bobbit worms etc.

Conchifera

Aculifera

Mollusca

chitons

octopus, squids etc.

tusk shells

snails, slugs etc.

clams, oysters, etc.

lamp shells

ribbon worms

horseshoe worms

solenogasters

Myzostomida (?155)

Figure 4.� &RPELQHG�SK\ORJHQ\�RI �0ROOXVFD�DQG�$QQHOLGD��7KH�0ROOXVFD�WUHH�LV�GUDZQ�IURP�6PLWK�HW�DO��������:%>72,#%.7#%6)04#04<4%57=,)?#0=%)* %F00#,2A+%+0A%Q25<06<,+%24%9+4#A%)0%5RXVH�DQG�3OHLMHO�������%+0A%6WUXFN�HW�DO��������B%F4%20%)LJXUH��:%*),,)>20?%.7#%.#$-20+,%0+-#:%20%5+$#0.7#4#4:%24%.7#%#4.2-+.#A%0<-9#$%)* %#J.+0.%A#46$29#A%+0A%"+,2A%45#62#4%*)$%.7+.%?$)<5%9+4#A%RQ�WKH�:RUOG�5HJLVWHU�RI �0DULQH�6SHFLHV��$GGLWLRQDO�UHIHUHQFHV�IRU�JURXSV�ZLWK�QRQPDULQH�VSHFLHV�DUH�%LYDOYLD��ZKLFK�DOVR�206,<A#4%-+0=%*$#47>+.#$%45#62#4%1*UDI ��������*DVWURSRGD��ZKLFK�LQFOXGHV�QXPHURXV�IUHVKZDWHU�DQG�WHUUHVWULDO�VSHFLHV��&DPHURQ��������DQG�&OLWHOODWD��ZKLFK�LQFOXGHV�PDQ\�IUHVKZDWHU�DQG�WHUUHVWULDO�VSHFLHV��6NHW�DQG�7URQWHOM�������:HW]HO�DQG�5H\QROGV�����3B%7KH�VSHFLHV�FRXQW�IRU�WKH�WD[RQ�&DSLWHOOLGD�LQFOXGHV�WD[D�VXFK�DV�(FKLXUD�DQG�SRO\FKDHWHV�IRUPHUO\�LQFOXGHG�LQ�6FROHFLGD��D�WD[RQ�0)>%47)>0%.)%9#%5),=57=,#.26%16WUXFN�HW�DO������3B%C7#0%+"+2,+9,#:%6)--)0%0+-#4%+$#%>$2..#0%+9)"#%.7#%9$+067#4B

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�HW�DO��������%HOODQWXRQR�HW�DO��������S[$4.#$%#.%+,B%������0R\D�HW�DO��������%DUVKLV�HW�DO��������3pUH]�3RUUR�HW�DO��������9LGDO�'XSLRO�HW�DO������3BS<.<$#%#**)$.4%>2,,%6)06#0.$+.#%)0%#J5+0A20?%.7#%DEDF%

6)--<02.=:% $#*2020?% 2.4% ?)+,4:% A#"#,)520?% 7=5).7#4#4:% +0A%#4.+9,24720?% ?#0)-264% $#4#+$67% +0A% #A<6+.2)0+,% $#4)<$6#4B%DEDF% >#,6)-#4% +,,% -#-9#$4% )* % .7#% 462#0.2*26% 6)--<K02.=% >7)% >247% .)% 6)0.$29<.#% .)% 6)-5+$+.2"#% +55$)+67#4%

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�IRU�ODWH�������)XQGLQJ�WKH�VHTXHQFLQJ�RI �WKRXVDQGV�)* %20"#$.#9$+.#%?#0)-#4%>2,,%$#@<2$#%6$#+.2"#%+0A%6),,+9)$+K.2"#%+55$)+67#4%.7+.%?)%9#=)0A%.7#%.$+A2.2)0+,%*<0A20?%-#67K+024-4% )* % 20A2"2A<+,% 5$20625+,% 20"#4.2?+.)$% ?$+0.4% 1!,#G4=G%HW�DO������3B%C72,#%.7#%DEDF%.#+-%7+4%4.+$.#A%#J5,)$20?%5)4K429,#% *<0A20?%)55)$.<02.2#4:%>#% 20"2.#% *##A9+6G%+0A%5+$.262K5+.2)0%*$)-%.7#%462#0.2*26%6)--<02.=%*)$%H)20.%*<0AK4##G20?%4.$+.#?2#4%+0A%2A#+4B

Supplementary MaterialQ<55,#-#0.+$=%-+.#$2+,%6+0%9#%*)<0A%+.%7..5OII>>>BH7#$#AB)J*)$AH)<$0+,4B)$?IB

Cephalochordata (33)

Ambulacraria

ChordataCraniata

acorn worms

acorn worms

acorn worms

featherstars & sea lilies

sea stars and sea daisies

brittle stars etc.

sea cucumbers

urchins etc.

salps

larvaceans

sea squirts

sea squirts

sea squirts

pyrosomes

doliolids

lancelets

vertebrates

Ophiuroidea (2073)

Crinoidea (635)

Asteroidea (1858)

Echinoidea (1076)

Holothuroidea (1688)

Echinodermata

Eleutherozoa

Ptychoderidae (37)

Harrimanidae (41)

Spengelidae (20)

Torquaratoridae (6)

Pterobranchia (27)

Hemichordata

Appendicularia (67)

Pyrosomatida (8)

Doliolida (25)

Stolidobranchia (1024)

Phlebobranchia (330)

Aplousobranchia (1526)

TunicataSalpida (45)Thaliacea

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Journal of Heredity

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Funding$PHULFDQ�*HQHWLF�$VVRFLDWLRQ�ZLWK�D�6SHFLDO�(YHQW�$ZDUG�.7+.% 5$)"2A#A% .7#% 5$2-+$=% *<0A20?% *)$% .7#% -+2A#0% DEDF%ZRUNVKRS�� 7KHRGRVLXV� 'RE]KDQVN\� &HQWHU� IRU� *HQRPH�%LRLQIRUPDWLFV� �5XVVLD�0LQLVWU\�RI �6FLHQFH�0HJD�JUDQW����*����������� WR� 6�-�� 2·%ULHQ�� 3ULQFLSDO� ,QYHVWLJDWRU��� /LIH�7HFKQRORJLHV� DQG� %LR1DQR*HQRPLFV�� 1DWLRQDO� 6FLHQFH�)RXQGDWLRQ·V� $́VVHPEOLQJ� WKH�7UHH� RI � /LIHµ� �'(%� DZDUGV����������������������������������������������

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Journal of Heredity

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Page 21: American Genetic Association ISSN 1465-7333 (Online ...medinalab.org/new/wp-content/uploads/2013/10/GIGA14.pdfEditor-in-Chief C. Scott Baker,Oregon State University Managing Editor

Journal of Heredity

18

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Received July 15, 2013; First decision September 3, 2013; Accepted October 18, 2013

Corresponding Editor: Stephen Karl

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