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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
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
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
© 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!"#$%&'(%)* %+,,%-#.+/)+0%1+02-+,3%45#62#4%6)-5$24#%.7#%820"#$.#9$+.#4:;%9<.%"#$=%*#>%?#0)-#4%*$)-%.7#4#%)$?+024-4%7+"#%9##0%4#@<#06#AB%C#%7+"#:%.7#$#*)$#:%*)$-#A%+%8D,)9+,%E0"#$.#9$+.#%D#0)-264%F,,2+06#;%1DEDF3B%!<$%20.#0.%24%.)%9<2,A%+%6),,+9)$+.2"#%0#.>)$G%)* %A2"#$4#%462#0.24.4%.)%.+6G,#%-+H)$%67+,,#0?#4%1#B?B:%45#62#4%4#,#6.2)0:%4+-5,#%6),,#6.2)0%+0A%4.)$+?#:%4#@<#06#%+44#-9,=:%+00).+.2)0:%+0+,=.26+,%.)),43%+44)62+.#A%>2.7%?#0)-#I.$+046$25.)-#%4#@<#0620?%+6$)44%+%,+$?#%.+J)0)-26%45#6.$<-B%C#%+2-%.)%5$)-).#%4.+0A+$A4%.7+.%>2,,%*+62,2.+.#%6)-5+$+.2"#%+55$)+67#4%.)%20"#$.#9$+.#%?#0)-264%+0A%6),,+9)$+KWLRQV�DFURVV�WKH�LQWHUQDWLRQDO�VFLHQWLÀF�FRPPXQLW\��&DQGLGDWH�VWXG\�WD[D�LQFOXGH�VSHFLHV�IURP�3RULIHUD��&WHQRSKRUD��&QLGDULD��3ODFR]RD��0ROOXVFD��$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.)$4%+0A%42?0+,20?%5+.7>+=4:%2A#0.2*2#A%
20%-)4.%-#.+/)+0%?#0)-#4%4.<A2#A%4)% *+$% 1Q$2"+4.+"+%#.%+,B%����:%����3BE0"#$.#9$+.#4:%2B#B%+02-+,4%>2.7)<.%9+6G9)0#4:%#06)-5+44%
+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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
#6)4=4.#-4:%206,<A20?%-+0=%.7+.%+**#6.%5#)5,#B%Q)-#%+$#%5+$K+42.#4%+0A%A24#+4#%"#6.)$4%.7+.%+**#6.%.7#%7#+,.7%)* %7<-+04:%,2"#4.)6G:%+0A%5,+0.%6$)54B%F4%20"+42"#%45#62#4:%.7#=%6+0%7+"#%
Journal of Heredity 2014:105(1):1–18doi:10.1093/jhered/est084
by guest on Decem
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Developmental Biology
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Species Major group Subgroup Size (MB)
URL Citation
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Ecology
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Fisheries and Aquaculture
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+,,% 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,#%
by guest on Decem
ber 15, 2013http://jhered.oxfordjournals.org/
Dow
nloaded from
7
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Threatened, Endangered, and Remarkable Species
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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%)$%92).#670),)?26+,%
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$#6#0.%?#0)-#%462#06#%202.2+.2"#4%+0A%6))$A20+.2)0%0#.>)$G4%+0A%>2,,%4=067$)02/#%#**)$.4%>2.7%4<67%?$)<54%.7$)<?7%*<.<$#%SURMHFWV��%HFDXVH�*,*$� LV� DQ� LQWHUQDWLRQDO� FRQVRUWLXP�RI �462#0.24.4:% +?#062#4:% +0A% 204.2.<.2)04:% >#% >2,,% +,4)% +92A#% 9=%.7#% $<,#4% )* % ?,)9+,% *<0A20?% +?#062#4% *)$% A+.+% $#,#+4#% 1#B?B:%WKRVH� SURSRVHG� E\� WKH� *OREDO� 5HVHDUFK� &RXQFLO�� 7..5OII>>>B?,)9+,$#4#+$676)<062,B)$?3B% C#% +$#% +>+$#% .7+.% A2**#$K#0.%0+.2)04%7+"#%A2**#$#0.%6)04.$+20.4%+0A%$#?<,+.2)04%)0%.7#%<4#%)* %92),)?26+,%4+-5,#4B%D2"#0%.7#%20.#$0+.2)0+,%0+.<$#%)* %DEDF:%>#%>2,,%>)$G% .)%#04<$#% .7+.%0+.2)0+,%?#0)-26% ,#?+K62#4% +$#%5$).#6.#A% +0A%>2,,% 6)04<,.%>2.7% .7#%5#$.20#0.% ?)"K#$0-#0.+,% +?#062#4% 20% .7#% 6)<0.$2#4% *$)-% >7267% 4+-5,#4%)$2?20+.#B%C#%>2,,%A#5)42.%4#@<#06#%A+.+%20%5<9,26%A+.+9+4#4%�H�J���*HQ%DQN���DV�ZHOO�DV�GHSRVLW�'1$�YRXFKHUV�LQ�SXEOLKFDOO\�DFFHVVLEOH�UHSRVLWRULHV��H�J��*OREDO�*HQRPH�%LRGLYHUVLW\�1HWZRUN��6PLWKVRQLDQ��DEDF%24%+0%206,<42"#%#0.#$5$24#%.7+.%20"2.#4%+,,%20.#$#4.#A%
5+$.2#4% .)% H)20% .7#%#**)$.%)* % 20"#$.#9$+.#%?#0)-264B%C#%>2,,%+..#-5.%.)%6+5.<$#%.7#%2-5+6.%)* %.7#%#**)$.%20%.7#%>2A#$%462K#0.2*26%+0A%5<9,26%+$#0+4%9=%*),,)>20?%$#,#"+0.%5<9,26+.2)04%+0A%).7#$%5$)A<6.4%.7+.%$#4<,.%*$)-%DEDF%202.2+.2"#4B
by guest on Decem
ber 15, 2013http://jhered.oxfordjournals.org/
Dow
nloaded from
Journal of Heredity
8
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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)
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Pycnogonida (1323)
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Remipedia (18)
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Mystacocarida (13)
Cephalocarida (13)
Copepoda (15976)
Arachnida (110615)
Ostracoda (7882)
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Insecta (1020007)
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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.$+.#%)0%#J5+0A20?%.7#%DEDF%
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�IRU�ODWH�������)XQGLQJ�WKH�VHTXHQFLQJ�RI �WKRXVDQGV�)* %20"#$.#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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Received July 15, 2013; First decision September 3, 2013; Accepted October 18, 2013
Corresponding Editor: Stephen Karl
by guest on Decem
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