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

journal of HEREDITY Volume 105 | Number 1 | January/February 2014

91 101 143 130 136 111 120 145

. (fireworm;. Annelida), Zapteryx

Tetrahymena . (Xenoturbellida),. . (water bear; Tardigrada), Vallicula Vallicula 2. (basketstar; Echinodermata), (flatworm; Platyhelminthes, G. photo

. (solitary coral; Cnidaria), 18. Vieitezia Patterns of an Alpine Plant, Patterns of an Alpine . (arrow worm; Chaetognatha), 6. Glass . (penis worm; Priapulida), robusta Epimeria 13. of Microsatellite Loci and . (sea spider; Pycnogonida), Phoronopsis californica 9. cf. evelinae in the Banded Guitarfish, in the Banded Guitarfish, (ribbon worm; Nemertea) inside the atrium of Ciona mat Alleles in the Ciliate (platyctenid; Ctenophora), 3. kisfaludyi Paraseison (Rotifera), Butia eriospatha (Arecaceae) of Southern (sea squirt; Tunicata, photo Jacinto Pérez, Hydronauta). , 5. Spadella sp Much Does Inbreeding Contribute to the Much Does Inbreeding Contribute to Risk of Population Decline? An Ecological Risk of Population Decline? An Ecological (albino velvet worm; velvet Onychophora,Peripatopsis (albino alba G. photo Enchiridium Collosondeis sp Bryozoa Gorgonocepahlus chilensis 8. The cover. Invertebrate organisms for potential genome/transcriptomethrough projects the GIGAconsortium (see pp. See 1–18). inside cover for key to organisms. 1. multiformis 4. sponge (Porifera), sp Millnesium 7. (horseshoe worm; Phoronida),10. Pherecarida sp (horseshoe Phoronida),10. worm; 11. Giribet), 12. Priapulus sp sp Xenoturbella Crustacea),14. (amphipod; 15. Giribet), 16. Micromelo undatus (miniature melo; Mollusca, photo G. sp Giribet), Flabellum 17. luzmurubeae intestinalis credits, otherwise. whereImage indicated Greg except Rouse, Brief Communications Relationships Genetic Background Alters Dominance between Letter to Editor Volvox, Rolling out from under the Shadow of Chlamydomonas with Support from the AGA David Roy Smith Announcements Strong Population Structure and Shallow Mitochondrial Structure and Shallow Mitochondrial Strong Population Phylogeny thermophila Sujal S. Phadke, Tiago Paixão, Tuan Pham, Stephanie Pham, and Rebecca A. Zufall Characterization Repeat Density in the Gooseneck Barnacle, Pollicipes elegans, Using Next Generation Sequencing Louis V. Plough and Peter B. Marko (Jordan y Gilbert, 1880), from the Northern exasperata (Jordan y Gilbert, Mexican Pacific Sosa-Nishizaki, Jonathan Ana Castillo-Páez, Oscar Galván-Magaña, María-del-Pilar Sandoval-Castillo, Felipe Rocha-Olivares Blanco-Parra, and Axayácatl Phylogeographical Rhodiola dumulosa (Crassulaceae), Inferred from Chloroplast DNA Sequences Yan Hou and Anru Lou How Reduced Fitness of Hatchery-Born Steelhead (Oncorhynchus mykiss) in the Wild? Marine, Mark R. Christie, Rod A. French, Melanie L. and Michael S. Blouin At Palm and Genetic Approach to the Threatened Species Brazil Alison G. Nazareno and Maurício S. dos Reis 1 19 28 39 48 60 70 82

Erebia alberganus of the Dark Fruit-Eating Bat Neonympha mitchellii mitchellii Variation and Population Structure of Population Expansion on Genetic Expansion of House Sparrows (Passer of the Alps: The Biogeography of a Elephants of Gash-Barka, Eritrea: Nuclear and Lontra canadensis) in Central North America genomic and evolutionary diversity genomic and An international journal of journal of An international Perspective (GIGA): Developing Community Resources (GIGA): Developing Original Articles for Conservation Genetics and the Implication Satyr Recovery of the Endangered Mitchell’s The Global Invertebrate Genomics Alliance The Global Invertebrate to Study Diverse Invertebrate Genomes to Study Diverse Invertebrate Scientists GIGA Community of Butterfly, Christopher A. Hamm, Victoria Rademacher, Christopher A. Hamm, Victoria Rademacher, Douglas A. Landis, and Barry L. Williams Out Disjunctly Distributed Mountain Butterfly, the Almond-Eyed Ringlet Impact Diversity and Structure of River Otters (Lepidoptera, Satyrinae) and Dirk Louy, Jan Christian Habel, Werner Ulrich, Thomas Schmitt ( Ammer, Jessica R. Brandt, Adam L. Brandt, Frank K. Alfred L. Roca, and Thomas L. Serfass Phylogeography Artibeus obscurus in the Brazilian Amazon Nascimento Wallax Augusto Silva Ferreira, Bárbara do Atanaena Borges, Symara Rodrigues-Antunes, Fernanda Souza Gonçalves de Andrade, Gilberto Ferreira de Aparecida Aguiar, José de Sousa E Silva-Junior, Suely Marques-Aguiar, and Maria Lúcia Harada Range domesticus) in Kenya: Evidence of Genetic Admixture and Human-Mediated Dispersal Aaron W. Schrey, Andrea L. Liebl, Christina L. Richards, and Lynn B. Martin Genetic in the Endangered Hermann’s Tortoise: The Roles of Geography and Human-Mediated Processes Melanie 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 The Mitochondrial Genetic Patterns Adam L. Brandt, Yohannes Hagos, Yohannes Yacob, Victor A. David, Nicholas J. Georgiadis, Jeheskel Shoshani, and Alfred L. Roca journal of journal HEREDITY

journal of HEREDITY Volume 105 | Number 1 | January/February 2014

91 101 143 130 136 111 120 145

. (fireworm;. Annelida), Zapteryx

Tetrahymena . (Xenoturbellida),. . (water bear; Tardigrada), Vallicula Vallicula 2. (basketstar; Echinodermata), (flatworm; Platyhelminthes, G. photo

. (solitary coral; Cnidaria), 18. Vieitezia . (arrow worm; Chaetognatha), 6. Glass . (penis worm; Priapulida), robusta Epimeria 13. . (sea spider; Pycnogonida), Phoronopsis californica 9. cf. evelinae (ribbon worm; Nemertea) inside the atrium of Ciona mat Alleles in the Ciliate (platyctenid; Ctenophora), 3. kisfaludyi Paraseison (Rotifera), Butia eriospatha (Arecaceae) of Southern (sea squirt; Tunicata, photo Jacinto Pérez, Hydronauta). , 5. Spadella sp (albino velvet worm; velvet Onychophora,Peripatopsis (albino alba G. photo Enchiridium Collosondeis sp Bryozoa Gorgonocepahlus chilensis 8. The cover. Invertebrate organisms for potential genome/transcriptomethrough projects the GIGAconsortium (see pp. See 1–18). inside cover for key to organisms. 1. multiformis 4. sponge (Porifera), sp Millnesium 7. (horseshoe worm; Phoronida),10. Pherecarida sp (horseshoe Phoronida),10. worm; 11. Giribet), 12. Priapulus sp sp Xenoturbella Crustacea),14. (amphipod; 15. Giribet), 16. Micromelo undatus (miniature melo; Mollusca, photo G. sp Giribet), Flabellum 17. luzmurubeae intestinalis credits, otherwise. whereImage indicated Greg except Rouse, Brief Communications Relationships Genetic Background Alters Dominance between Letter to Editor Volvox, Rolling out from under the Shadow of Chlamydomonas with Support from the AGA David Roy Smith Announcements Strong Population Structure and Shallow Mitochondrial Structure and Shallow Mitochondrial Strong Population Guitarfish, Phylogeny in the Banded thermophila Sujal S. Phadke, Tiago Paixão, Tuan Pham, Stephanie Pham, and Rebecca A. Zufall and Characterization of Microsatellite Loci Repeat Density in the Gooseneck Barnacle, Pollicipes elegans, Using Next Generation Sequencing Louis V. Plough and Peter B. Marko (Jordan y Gilbert, 1880), from the Northern exasperata (Jordan y Gilbert, Mexican Pacific Sosa-Nishizaki, Jonathan Ana Castillo-Páez, Oscar Galván-Magaña, María-del-Pilar Sandoval-Castillo, Felipe Rocha-Olivares Blanco-Parra, and Axayácatl Patterns of an Alpine Plant, Phylogeographical Rhodiola dumulosa (Crassulaceae), Inferred from Chloroplast DNA Sequences Yan Hou and Anru Lou to the How Much Does Inbreeding Contribute Reduced Fitness of Hatchery-Born Steelhead (Oncorhynchus mykiss) in the Wild? Marine, Mark R. Christie, Rod A. French, Melanie L. and Michael S. Blouin At Risk of Population Decline? An Ecological Palm and Genetic Approach to the Threatened Species Brazil Alison G. Nazareno and Maurício S. dos Reis 1 19 28 39 48 60 70 82

Erebia alberganus Neonympha mitchellii mitchellii Lontra canadensis) in Central North America genomic and evolutionary diversity genomic and An international journal of journal of An international Perspective (GIGA): Developing Community Resources (GIGA): Developing Original Articles for Conservation Genetics and the Implication Satyr Recovery of the Endangered Mitchell’s The Global Invertebrate Genomics Alliance The Global Invertebrate to Study Diverse Invertebrate Genomes to Study Diverse Invertebrate Scientists GIGA Community of Butterfly, Christopher A. Hamm, Victoria Rademacher, Christopher A. Hamm, Victoria Rademacher, Douglas A. Landis, and Barry L. Williams Out of the Alps: The Biogeography of a Out of the Alps: The Biogeography of Disjunctly Distributed Mountain Butterfly, the Almond-Eyed Ringlet Impact of Population Expansion on Genetic Diversity and Structure of River Otters (Lepidoptera, Satyrinae) and Dirk Louy, Jan Christian Habel, Werner Ulrich, Thomas Schmitt ( Ammer, Jessica R. Brandt, Adam L. Brandt, Frank K. Alfred L. Roca, and Thomas L. Serfass Phylogeography of the Dark Fruit-Eating Bat Phylogeography of the Dark Fruit-Eating Artibeus obscurus in the Brazilian Amazon Nascimento Wallax Augusto Silva Ferreira, Bárbara do Atanaena Borges, Symara Rodrigues-Antunes, Fernanda Souza Gonçalves de Andrade, Gilberto Ferreira de Aparecida Aguiar, José de Sousa E Silva-Junior, Suely Marques-Aguiar, and Maria Lúcia Harada (Passer Range Expansion of House Sparrows domesticus) in Kenya: Evidence of Genetic Admixture and Human-Mediated Dispersal Aaron W. Schrey, Andrea L. Liebl, Christina L. Richards, and Lynn B. Martin Genetic Variation and Population Structure in the Endangered Hermann’s Tortoise: The Roles of Geography and Human-Mediated Processes Melanie 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 The Elephants of Gash-Barka, Eritrea: Nuclear and Mitochondrial Genetic Patterns Adam L. Brandt, Yohannes Hagos, Yohannes Yacob, Victor A. David, Nicholas J. Georgiadis, Jeheskel Shoshani, and Alfred L. Roca journal of journal HEREDITY Journal of Heredity The American Genetic Association Editor-in-Chief www.theaga.org C. Scott Baker, Oregon State University

President Managing Editor Robin Waples Anjanette Baker, American Genetic Association

Past President Editorial Board Kerry Shaw Ernest Bailey, University of Kentucky Adalgisa Caccone, Yale University President Elect Andrew J. Crawford, Universidad de los Andes, Colombia Katie Peichel Jerry Dodgson, Michigan State University Robert C. Fleischer, Smithsonian Institution Executive Vice President Susan Gabay-Laughnan, University of Illinois Brad Shaffer Stewart Grant, Alaska Dept of Fish & Game Secretary J. Perry Gustafson, University of Missouri Oliver Ryder Fangpu Han, Chinese Academy, China Scott Hodges, University of California, Santa Barbara Treasurer Jennifer Jackson, British Antarctic Survey, United Kingdom Kim Hughes Prem P. Jauhar, USDA-ARS, North Dakota State University Warren E. Johnson, Smithsonian Conservation Biology Institute Council F. Andrew Jones, Oregon State University David Baum Stephen A. Karl, Hawaii Institute of Marine Biology Anne Bronikowski Susan J. Lamont, Iowa State University Lila Fishman John Logsdon, University of Iowa Steve Karl Jose V. Lopez, Nova Southeastern University Oceanographic Center Carlos Machado Therese Markow, University of California, San Diego Maria Orive William S. Modi, American Genetic Association Elaine Ostrander William Murphy, Texas A&M University Robert Wayne Mohamed Noor, Duke University Kelly Zamudio Taras Oleksyk, University of Puerto Rico Kenneth Olsen, Washington University, St. Louis Elaine Ostrander, American Genetic Association Reid G. Palmer, USDA, Iowa State University Jill Pecon-Slattery, Smithsonian Conservation Biology Institute Roger H. Reeves, Johns Hopkins University Howard Ross, University of Auckland, New Zealand Oliver Ryder, San Diego Zoo Horacio Schneider, Universidade Federal do Para, Brazil Hector Seuánez, Instituto Nacional de Cancer, Brazil Brad Shaffer, University of California, Los Angeles William Sherwin, University of New South Wales, Australia Christopher I. Smith, Willamette University Tomoko Y. Steen, The Library of Congress John R. Stommel, USDA-ARS, Beltsville Agricultural Research Center William Tracy, University of Wisconsin David B. Wagner, University of Kentucky Robin Waples, Northwest Fisheries Science Center, NMFS Robert Wayne, University of California, Los Angeles

JHERED10501M Journal of Heredity 2014:105(1):1–18 © The American Genetic Association 2013. All rights reserved. doi:10.1093/jhered/est084 For permissions, please e-mail: [email protected] The Global Invertebrate Genomics Alliance (GIGA): Developing Community Resources to Study Diverse Invertebrate Genomes

GIGA 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]. Downloaded from

*Authors are listed in the Appendix

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3 Journal of Heredity

-<67%57=,)?#0#.26:%-)$57),)?26+,:%+0A%#6),)?26+,%A2"#$42.=% +4% 5)4429,#% +0A% .7#% 5$+6.26+,% ,2-2.+.2)04% )* % .7#% 202.2+.2"#B% D2"#0%.7#%,+$?#%5)5<,+.2)0%42/#4%)* %-+0=%20"#$.#9$+.#%45#K 62#4:% 6),,#6.2)0% )* % +% 4<**262#0.% 0<-9#$% )* % 20A2"2A<+,4% -+=% #J+-20#A%6<$$#0.%57=,)?#0#.26%7=5).7#4#4%+0A%4#,#6.#A%G#=% 20"#$.#9$+.#4%<4<+,,=%20"),"#4%*#>#$%A2**26<,.2#4%>2.7%$#?+$A%.)% 20"#$.#9$+.#% 45#62#4% .7+.% 45+0% .7#% 57=,)?#0#.26% A2"#$42.=% 5#$-2.4%.7+0%6),,#6.2)0%)* %"#$.#9$+.#%.244<#4B%T)>#"#$:%4)-#% +0A%-)$57),)?26+,%A245+$2.=%)0%.7#%+02-+,%.$##%)* %,2*#%14##% 20"#$.#9$+.#%.+J+%5)4#%"+$2)<4%,)?24.26%+0A%.#670),)?26+,%67+,K ,#0?#4% *)$% >7),#K?#0)-#% 4#@<#0620?O% -+0=% 45#62#4% ,2"#% 20% $#"#+,%0)"#,%4#@<#06#4%>2.7%4<**262#0.%57=,)?#0#.26%42?0+,%.)% $#,+.2"#,=%20+66#4429,#%7+92.+.4%1#B?B:%+4%5+$+42.#4:%20%.7#%A##5% $#4),"#%,)0?4.+0A20?%@<#4.2)04B 4#+:%)$%?#)?$+5726+,,=%$#-).#3%)$%+$#%.))%4-+,,%.)%=2#,A%4<*K K Developmental Biology ,#0?#4% >2,,% 9#% 6)042A#$#A% >2.7% ).7#$% 6$2.#$2+% +4% 4#@<#0620?% !0#%)* %.7#%-)4.%#J62.20?%+55$)+67#4%.)%<0A#$4.+0A20?%.7#% 5$)H#6.4%+$#%A#"#,)5#A%+0A%5$2)$2.2/#AB )$2?204% +0A% #"),<.2)0% )* % +02-+,% A2"#$42.=% 24% A#.+2,#A% 6)-K F02-+,%92)A2"#$42.=%24%0).%6)04.$+20#A%9=%5),2.26+,%9)<0AK 5+$+.2"#%$#4#+$67%)0%#-9$=),)?26+,%A#"#,)5-#0.%12B#B:%#"),%DEDF%C#9%42.#%>2,,%9#%<4#A%.)%*)4.#$%6))5#$+K A2,#--+B%C7),#K?#0)-#%4#@<#0620?%5$)"2A#4%.7#4#%A+.+%20%

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4 (*("$PNNVOJUZPG4DJFOUJTUTr(MPCBM*OWFSUFCSBUF(FOPNJDT"MMJBODF (*("

Table 1

Species Major group Subgroup Size URL Citation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

Downloaded from 0'#-)*$)4'.-<' N<026+.+ 7..5OII>>>B9$)+A204.2.<.#B)$?I +00).+.2)0I62)0+I D'E#%(,>")*2'#'9) N<026+.+ 7..5OII>>>B?#0)46)5#B604B*$I !2G)5,#<$+

89"#%#")*2'.'/'F,") F0.7)/)+ 7..5OII-+$20#?#0)-264B)24.BH5I http://jhered.oxfordjournals.org/ G,+)/#$/,(()* F0.7)/)+ 7..5OII?#0)-#BH?2K54*B)$?I 4,9/,-$'$ 6<2")* T=A$)/)+ 7..5OII>>>B0692B0,-B027B?)"I +).-'%)%'(()/) !)%&-')*%>(,7 H-,+'#%$'$*(,'2<' W)9+.+ 7..5OII$#4#+$67B07?$2B027B?)"I -0#-2)5424I \#4#+$67%E04.2.<.#% 1<05<9,247#A%A+.+3

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5 Journal of Heredity

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7 Journal of Heredity

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.+$?#.%)$?+024-%1 http://jhered.oxfordjournals.org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02.=:%4<67%+4%.7)4#%$#6)--#0A#A%9=%D#0)-264%Q.+0A+$A4% 0).%*<,,=%$#4),"#A%1*)$%$#6#0.%$#"2#>4%4##% Downloaded from 7..5OII?#046B)$?I3B 3B%Q#"#$+,%$#6#0.%4.+.#$% 4)04:%9#4.%5$+6.26#4%+0A%4.+0A+$A4%>2,,%9#%#4.+9,247#A%*)$% http://jhered.oxfordjournals.org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

6+0% )0,=% 9#% A#.#$-20#A% )06#% 0)092,+.#$2+0% $#,+.2)047254% by guest on December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

9 Journal of Heredity

K K ,2-#.#$4%+6$)443%9#0.726%-+$20#%+02-+,4%.7+.%$#4#-9,#%+%*,+.% 62,2+.#A%A246%13B%!0,=%)0#%45#62#4%7+4%9##0%*)$-+,,=% A#46$29#A:% 9<.% .7#% 57=,<-% ,2G#,=% 24% -)$#% 45#62)4#% 1c)2?.% 67+$+6.#$2/#A%9=%.7#2$%+92,2.=%.)%-),.%.7#%6<.26,#%A<$20?%.7#2$% ,2*#%6=6,#:%+0A%*)$%7+"20?%$#A<6#A%#52.7#,2+,%62,2+.2)0:%>7267% 5)42.2)0%+-)0?%0)092,+.#$2+04%24%6<$$#0.,=%<0$#4),"#A%1#B?B:% $#@<2$#4%,)6)-).2)0%"2+%-<46<,+$%+6.2)0B%N7#=%206,

Heteroscleromorpha (5341) Downloaded from Demospongiae Haploscleromorpha (1084) Myxospongiae (138)

bath sponges Keratosa (558)

Hexasterophora (513) glass sponges

Hexactinellida http://jhered.oxfordjournals.org/ Porifera sponges Amphidiscophora (177)

Oscarellidae (19) Homoscleromorpha flesh sponges Plakinidae (68)

Calcinea (167) Calcarea calcareous sponges Calcaronea (514)

Placozoa (1) by guest on December 15, 2013

comb jellies Ctenophora (192)

corals, sea anemones Anthozoa Hexacorallia (3578) soft corals, sea pens, gorgonians Octocorallia (3549)

Myxozoa Malacosporea (4) Cnidaria Myxosporea (2180)

Polypodiozoa (1)

stalked jellies Staurozoa (48) Medusozoa marine & freshwater jellies Hydrozoa Trachylina (157) Hydra, siphonophores etc. Hydroidolina (3395)

box jellies Acraspeda Cubozoa (40) true jellies Scyphozoa (186)

Bilateria (other animals)

Figure 2. :% :%+0A% 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+.% W)-% B%C7#0%+"+2,+9,#:%6)--)0%0+-#4%+$#%>$2..#0%+9)"#%.7#%9$+067#4B

10 (*("$PNNVOJUZPG4DJFOUJTUTr(MPCBM*OWFSUFCSBUF(FOPNJDT"MMJBODF (*("

sea spiders Pycnogonida (1323) Chelicerata horsehoe crabs Xiphosura (4)

spiders, scorpions, harvestmen, ticks, mites. Arachnida (110615)

centipedes Chilopoda (3100) Myriapoda garden centipedes Symphyla (197)

Pauropoda (835)

millipedes Diplopoda (7753)

Mandibulata Mystacocarida (13)

Oligostraca seed shrimps Ostracoda (7882)

fish lice, tongue worms Ichthyostraca (339)

fairy shrimps, water fleas, clam shrimp etc. Branchiopoda (1184)

Pancrustacea Downloaded from barnacles Thecostraca (1424)

crabs, lobsters, amphipods, isopods, krill Malacostraca (40065) Vericrustacea copepods Copepoda (15976) http://jhered.oxfordjournals.org/ Remipedia (18) Xenocarida horsehoe shrimp Cephalocarida (13)

Insecta (1020007)

Hexapoda Entognatha (9734)

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11 Journal of Heredity

Cirratuliformia (539)

spaghetti worms etc. Terebelliformia (995)

Canalipalpata earthworms, leeches etc. Clitellata (9485)

lugworms, spoonworms etc. Capitellida (1757)

mud-blister worms etc. Spionida (792)

featherworms, pogonophores etc. Sabellida (1705)

paddleworms, bloodworms etc. Annelida Phyllodocida (4615) palolo worms, bobbit worms etc. Aciculata Eunicida (1444)

fireworms Amphinomida (230) Myzostomida (?155) peanut worms Sipuncula (148)

solenogasters Neomeniomorpha (280) Aplacophora Downloaded from Aculifera Chaetodermomorpha (130)

chitons Polyplacophora (989)

Mollusca octopus, squids etc. Cephalopoda (835) http://jhered.oxfordjournals.org/ Monoplacophora (30)

tusk shells Conchifera Scaphopoda (578)

snails, slugs etc. Gastropoda (62000)

clams, oysters, etc. Bivalvia (9532)

ribbon worms Nemertea (1285) by guest on December 15, 2013 lamp shells Brachiopoda (388)

horseshoe worms Phoronida (18)

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Conclusions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

12 (*("$PNNVOJUZPG4DJFOUJTUTr(MPCBM*OWFSUFCSBUF(FOPNJDT"MMJBODF (*("

Pterobranchia (27)

acorn worms Harrimanidae (41) Hemichordata acorn worms Ptychoderidae (37)

Torquaratoridae (6) Ambulacraria acorn worms Spengelidae (20)

featherstars & sea lilies Crinoidea (635)

sea stars and sea daisies Asteroidea (1858) Echinodermata brittle stars etc. Ophiuroidea (2073) Eleutherozoa sea cucumbers Holothuroidea (1688)

urchins etc. Echinoidea (1076)

lancelets Cephalochordata (33)

vertebrates Craniata

Chordata larvaceans Appendicularia (67) Downloaded from sea squirts Aplousobranchia (1526)

doliolids Doliolida (25)

Tunicata salps Thaliacea Salpida (45)

pyrosomes Pyrosomatida (8) http://jhered.oxfordjournals.org/ sea squirts Phlebobranchia (330)

sea squirts Stolidobranchia (1024)

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by guest on December 15, 2013

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13 Journal of Heredity

Funding .7+.% 5$)"2A#A% .7#% 5$2-+$=% *<0A20?% *)$% .7#% -+2A#0% DEDF% Acknowledgments K

DEDF%>)$G47)5%E%,)?24.264B%S)$%7#,5*<,%*##A9+6G%+0A%6)--#0.4%)0%#+$,=%A$+*.4:% Downloaded from $#6)--#0A+.2)04%#J5$#44#A%20%.724%-+.#$2+,%+$#%.7)4#%)* %.7#%+<.7)$4%+0A%A)%

!6#+0)?$+5726% E04.2.<.#% +.% S,)$2A+% F.,+0.26% e02"#$42.=:% S.% http://jhered.oxfordjournals.org/ Appendix

by guest on December 15, 2013 References )* %!"#$#%&'()*+,()-#.)$/," CB %\)<4#%1Q6$2554%E04.2.<.2)0%)* %!6#+0)?$+57=:%e02"#$42.=%

14 (*("$PNNVOJUZPG4DJFOUJTUTr(MPCBM*OWFSUFCSBUF(FOPNJDT"MMJBODF (*("

+55$+24+,% )* % .7#% .).+,% 92)-+44% +0A% +00<+,% 5$)A<6.2)0% )* % F0.+$6.26% G$2,,B% K K &PLX&LMB K B9&'$/#$#+)*+)-$#-' A$+*.%?#0)-#%)* %0'#-)*'-/,$/'-)('$O%2042?7.4%20.)%67)$A+.#%+0A%"#$.#9$+.#%)$2K Downloaded from K H#-/)$/"),)* F)4,#()/) http://jhered.oxfordjournals.org/ K K #"+,<+.20?%A#%0)")%-#.7)A4%)* %?#0)-#%+44#-9,=%20%.7$##%"#$.#9$+.#%45#62#4B%

by guest on December 15, 2013 K 03*,(,.)-$ K I#(<%#2'>+* &<2"'F#"+,:% I#(<%#2'>+* K &<2"'F#"+, 42"#%?#0)-#%)* %!)%&-')*%>(,7 )* %.7#%>)$,AU4%20"#$.#9$+.#4B%W)0A)0O%f)),)?26+,%Q)62#.=%)* %W)0A)0B 5#$45#6.2"#4%)0%.7#%,)0?K.#$-%+94#06#%)* %4#J<+,%$#5$)A<6.2)0%20%+02-+,4B%

15 Journal of Heredity

K K K 6)"#$=%)* %0#>%-+$20#%0+.<$+,%5$)A<6.4%*$)-%20"#$.#9$+.#4%)"#$%.7#%,+4.%.>)% A<$20?%.7#%6#04<4%)* %-+$20#%,2*#%A#6+A#%+0A%9#=)0AO%+%5$)5)4#A%A##5K)6#+0% N9'2,$*9#"2)/>$ Downloaded from K A,/")-<9&>$* >"/'9),:% +% 6+0A2A+.#% *)$% K

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16 (*("$PNNVOJUZPG4DJFOUJTUTr(MPCBM*OWFSUFCSBUF(FOPNJDT"MMJBODF (*("

K 0",(()* ,(,.)-$ K Downloaded from K K http://jhered.oxfordjournals.org/ .7#%-+$20#%67)$A+.#%D'E#%(,>")*2'#'9) 0"<%/)$/,"'-) A2?2.2*#$+%?#0)-#%.)%<0A#$4.+0A%6)$+,%$#45)04#4%.)%#0"2$)0-#0.+,%67+0?#B% ?#0)-#%$#"#+,4%+06#4.$+,%#<-#.+/)+0%?#0#%$#5#$.)2$#%+0A%?#0)-26%)$?+02K by guest on December 15, 2013 K K K !5.2-2/+.2)0% )* % 5$#4#$"+.2)0% +0A% 4.)$+?#% .2-#% )* % 45)0?#% .244<#4% .)% B/"#-.<(#9,-/"#/>$* %>"%>")/>$B% Q62#06#B% A"'9&#%()7 B9&+'2/,)*+,2'5 8+%&'+,2#-* /,"")-,) K>,,-$()-2'9) B/"#+1>$*.'.)$:%$#4#+$67%20%.7#% K

17 Journal of Heredity

F"+2,+9,#%*$)-O%% .7#%5#+$,%)=4.#$%I'-9/)2)*F>9)/)O%+%5,+.*)$-%*)$%<0A#$4.+0A20?%92"+,"#%92),)?=B% K ,20#% A#-)45)0?#% 8$/"#$9(,")* ;'((,<)-) F0% +0+.)-26+,% A#46$25.2)0% )* % +% -202+.<$2/#A% +6)$0% >)$-% 17#-267)$K Downloaded from A$2"#0% 5T% A#6$#+4#% 20% 6)$+,4O% 0#>% 2042?7.4% *$)-% .$+046$25.)-#% +0+,=424B% http://jhered.oxfordjournals.org/ K K -)$5726%?#0)-#4O%+,?)$2.7-4%+0A%+55,26+.2)0%.)%0'#-)*$)4'.-<'B%D#0)-#%\#4B% Received July 15, 2013; First decision September 3, 2013; by guest on December 15, 2013 Accepted October 18, 2013 0(#-#"9&'$*$'-,-$'$ Corresponding Editor: Stephen Karl