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A total evidence cladistic analysis of the Haliotidae (Gastropoda: Vetigastropoda)

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A total evidence cladistic analysis of the Haliotidae (Gastropoda: Vetigastropoda)

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Content A TOTAL EVIDENCE CLADISTIC ANALYSIS OF THE HALIOTIDAE (GASTROPODA: VETTGASTROFODA) by Daniel Ludwig Geiger A Dissertation Presented to the FACULTY OF THE GRADUATE SCHOOL UNIVERSITY OF SOUTHERN CALIFORNIA in Partial Fulfillment of the Requirement for the Degree DOCTOR OF PHILOSOPHY (BIOLOGY) December 1999 Copyright 1999 Daniel Ludwig Geiger Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UNIVERSITY OF SOUTHERN CALIFORNIA. THE GRADUATE SCHOOL UNTVERSItrPARK LOS ANGELES. CALIFORNIA 90007 This dissertaHortr ivritten by Daniel Ludvig GEIGER under the direction of his. Dissertation Committee^ and approved by aU its member^ has been presented to and accepted by The Graduate School in partial fulfillment of re quirements for the degree of DOCTOR OF PHILOSOPHY Dam of Graduate Studies Date N'ovember 2 2 , 1999 DISSERTATION COMMITTEE % Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. A Total Evidence Cladistic Analysis of the Haliotidae (Gastropoda: Vetigastropoda). Daniel L. Geiger (Co-Advisors: R. L. Zimmer and J. H. McLean) This total evidence cladistic analysis is partitioned into five chapters: nomenclature, fossils, coding strategies, biogeography, and combined data analysis. All genus- and species-level taxa are reviewed. Fifty-sbc described species plus ten subspecies are considered valid. For all these taxa the geographic distribution and sources o f published illustrations are given. All species are figured. The taxonomy o f the rare fossil abalone is problematic and their use in phylogenetics is questionable. Abalone first appear in the Maastrichian, are unknown in the Paleocene, reappear in the Eocene/Oligocene, and are regularly found firom Nfiocene to Recent. A list o f fossil records is given. Fossil and Recent abalone both lived in the shallow, rocky sublittoral in tropical and temperate climates. No on-shore/oflP-shore pattern was detected. In cladistic analyses all observations are equivalent. A gap should be coded as a fifth character state. Global alignment o f large sequences violates character independence, mandating the here-introduced minimal fiagment alignment. (Questionable alignments must be occluded or contracted; other methods conflict with the test o f conjunction and/or contradict the original observatiorL Highly dissimilar sequences may be represented using the here-mtroduced block or stretch coding. The distribution o f all species is documented. The three models for the family's origin are evaluated. The area cladogram is m general agreement with an unrooted taxon cladogram. The basal node o f the area cladogram is a fairly large polytomy uniting rather distant provinces such as the north Pacific, Australia, and Afiica and is in closest Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. agreement with, the Indo-Pacffîc model for the origin- A major hiatus o f approximately 2 0 0 million years can account for the discrepancy between the earliest fossils and the biogeographical reconstruction^ as well as for the large basal polytomy in the biogeographical analysis. The phylogenetic analysis reveals three consistent groupings: north Pacific, Australian endemics, and Indo-Pacific. Morphological data do not add information to the molecular hypotheses. Mssing data is identified as a source for the numerous most parsimonious resolutions. Recommendations for the use of genus-level taxa are given. [348 words, 350 maximum] Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS E ntire dissertation My dissertation commitee, composed o f co-chairs Jim McLean and Russ Zimmer, and members Dave Bother, Bob Lavenberg, and Loren Smith, provided guidance when needed, but otherwise permitted me to pursue my work with minimal interference. I would like to thank all the staff at museums, who helped to make stays at their institutions successful, helped with library research, and made specimens available for study: Henk Mienis (HUJ), Kathie Way, Julia Freeman, Joan Pickering, David Reid, and John Todd (BMNH), Alison Trew (NMW), Yves Finet and Claude Vaucher (MHNG), Roger Pickery (KBIN), Raye Germon, Jerry Harasewych, and Alan Rabat (USNM), Jim McLean and Lindsey Groves (LACM), Paul Scott and Henry Chaney (SBMNH), Terry Gosliner (CASIZ), David Lindberg (UCMP), Paula Mikkelsen (DMNH & AMNH), Gary Rosenberg and Ned Gilmore (ANSP), Peter Jung (NHB), Philippe Bouchet, Pierre Lozouet, and Philippe Maestrati (MNHN), Margaret Gosteli (NMBE), K. Boss (MCZ), Winston Ponder and Ian Loch (AMS), Shirley Slack-Smith (WAM), and Hike Neubert (SMF). Additional specimens were made available by the following persons: Ted Baer (Lutry, Switzerland), Dominik and Verena Brantschen-Geiger (Bern, Switzerland), Franziska Brantschen (Randa, Switzerland), Anuschka Faucci (Basel, Switzerland), Marc Girona (France), Mark Jones (Auckland, New Zealand), Peter Schuchert (MHNG), Solly Singer (Rehovot, Israel), Brian îfayes (South Afiica), Joan Koven (Astrolabe Inc., Washington, D. C.), Roger Pickery (WiMjk, Belgium), Katharine Stewart (Carmel, Cal- ifi>mia). Buzz Owen (Gualala, California), and Wolfgang Wranik (University o f Ros tock, Germany). u Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The systematics discussion group at the Los Angeles County Museum o f Natural History was instrumental in shaping my thoughts on cladistics: Bob B e ^ , Brian Brown, Kirk Fitzhugh, Guillermo Herrera, John Heynig, David Kizirian, Bob Lavenberg, Don Reynolds, and Christine Thacker. The staff at Hancock Library at USC and the Research Library o f the Natural His tory Museum o f Los Angeles County helped to locate many obscure references. The work was supported by the following grants and fellowships. The visits of DMNH and ANSP were made possible through a DuPont Merck Scholarschip in Mala cology, and a Jessup Fellowship, respectively. Research grants &om the Hawaiian Mala- cologfcal Society, the Western Society for Malacology, the Lemer Gray Fund for Marine Research (AMNH), Sigma Xi, and a Trojan League Grant are kindly acknowledged. USC granted a one year Dissertation Feilowiship. This dissertation was written in Word 5.1 and QuarkXPress 4 with Times New Roman as justified font and Courier as monospaced font Illustrations were made in Photoshop 3, and 4, MacDraw H, Powerpoint 4, QuarkXPress 3.1, and MacCIade. C hapter 1: Recent taxa This work would not have been possible without the critical remarks by several colleagues, who also made then knowledge and data generously available to me: Phil Colman (Sydn^, Australia), Yoim-Ho Lee (Pasadena, California), Mark Jones (Auck land, New Zealand), Roger Pickery (W ilrijk, Belgium), Benjamin Singer (Rehovot Israel), Scoresby A. Shepherd (Adelaide, Australia), Katharine Stewart (Carmel, Cali fornia), Victor Vacquier (Sar^ Diego, Califomia), Rick Fallu (Canberra, Australia), Buzz Owen (Gulala, Cali&mia), and Luiz Shnone (Sao Paulo, Brazil). Jim McLean (Los Angeles, Califomia), Veronica Nfiller (Los Angeles, Califomia), Scoresby A. Shepherd. Ill Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. José Leal (Sanîbel, Florida), and two anonymous reviewers read the manuscript, includ ing earlier drafts, and made valuable comments. Gert Lindner (Hamburg, Germany) pointed out a typographical error in Geiger (1998a). This chapter has been slightly updated from Geiger (1998a), however, no new taxonomic decision have been intro duced, because a dissertation is unsuitable for such procedures. C hapter 2t Fossil taxa This chapter is a collaborative effort by DLG and Lindsey T. Groves (LACM Mala- cology/lnvert. Paleo). LTG has contributed numerous Neogene locality citations, pro vided obscure references and some figured specimens, and reviewed various ms drafts particularly for paléontologie accuracy. DLG has written and rewritten in excess 90% o f the text and produced the figures and plates. This chapter has been published in Jottr - nal o f Paleontology (Geiger & Groves, 1999). Dave Bother (USC), Jim McLean (LACM), and Richard Squires (California State University Northridge), David Lindberg (UCMP), David Dockery, Laurie Anderson, and Tim Hazen critically reviewed various drafts o f the manuscript. Atrfbfm Matsukuma (Kyushu University) kindly helped with some o f the obscure references. Darius Stram- ski (USC) and Anders Wàrén (Swedish Museum of Natural History) translated a Polish and a Swedish paper, respectively, for us. Trta Rodica (Muzeul de Istorie Naturala “(jrigore Antipa”, Bucharest) drew out attention to a Romanian publication and trans lated the relevant parts. Additional information was provided by Claus Hedegaard (UCMP), Hans-Dietrich Laatsch and Manfred Jakubeit (Wolfsburg, Germany), and Padermsak Jarayanbhand (Thailand). IV Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. C hapter 3: DNA sequence alignment This chapter is a collaborative effort between DLG and Kirk Fitzhugh (LACM). The chapter was build on the abductive nature o f cladistics worked out by KF. The application o f abductive principles to molecular data with the comparison to practices in mophoiogy was carried out by DLG including the development o f the new coding strategies. Between 80 and 90% o f the text was written by DLG. This chapter is in review with Cladistics since May 1999. A number o f people are thanked for engaging discussions on the above topic. The mention o f their names shall not signal their agreement with the content o f the chapter. They have been instrumental, however, in sharpening the arguments: Verena and Dominik Brantschen-Geiger, Mark Dawson, Paul Flook, Gonzalo Giribet, Guillermo Herrera, Bob Lavenberg, Jim McLean, Gavin Naylor, Jan Pawlowski, Mason Posner, M ark Siddall, Petra Sierwald, Ellen Strong, and Ward Wheeler. Brian Brown, Doug Eemisse, Jaqueline Reich, and Don Reynolds read earlier versions o f the manuscript and made valuable suggestions. C hapter 4: Biogeography My appreciation goes to the Palo Alto Research Center o f Xerox for offering the ffee map site on the wold wide web. Jim McLean helped to improve the manuscript. The manuscript is in press with the Bollettino Malacologico since August 1999. L ast hn t not least The beginnings o f this dissertation go back to my highschool years. A number o f persons have been instrumental in futhering my interst in marine invertebrates, and par ticularly in moUusks. Bo Holmstead QCarolinska Institute, Stockholm), Yehuda and Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Fanny Mazur (Weizman Institute, Rehovot, Israel), and Prof. emeritus Nüesch (Univer sity of Basel) convinced my parents that my interest in snails was worthy o f encourage m ent And so my family—my mother Anneliese Geiger-CIoos, my father Urs-Peter Geiger, and my sister Verena Brantschen-Geiger—did support in the most admirable fashion the son/brother. Early mentors at the University of Basel include Lukas Hot- tinger, Peter Jung (NHB), David Senn, and Werner Stingeli. Continuing contact with fellow shell collectors not mentioned elsewhere is a joy as always: Yvonne Grimmer, Roland Hadom, Jean-Caude Caillez, Michael Trippner, Axel Alf. Support and friendship with many fellow graduate students were mission critical for this undertaking: Nick Appelman, Jay Vavra, Judy Doino-Lemus, Todd Zimmerman (UCLA), Ellen Strong (GWU), and Rob Guralnick (UCB). Other people responsible for keeping me going include Michel Burla, Janka Jarchow, Jasmin Joshi, Alexander Kocyan, Veronica Miller, Willi Rühl, Willy Rühl, Marlies Rflhl, Brigitte Wyss-Heiz. VI Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table of Contents INTRODUCTION CHAPTER 1: RECENT GENERA AND SPECIES O F THE FAMILY HALIOTIDAE RAFINESQUE, 1815 (GASTROPODA: VETIGASTROPODA) Introduction ....................................... j M aterials and methods ...............................................................................................4 Systematic afBnities and characteristics of the fa m % ............................................ 6 Problems pertaining to the taxonomy of abalone ................. 8 Tremata as a taxonomic character and teratological type specim ens...................... 8 H ybrids...................................................................................................................12 Genus-level taxa .............................................. 16 H aliotis, sensu stricto ............................................................................................16 Other genus-level taxa» and their type species ...................................................... 18 Use o f genus-level ta x a ......................................................................................... 21 Species-level taxa .................... .22 Use o f subspecies ................................................................................................. 22 Index .................................................................................................................... 25 N otes.......................................................................................................................39 C aribbean.............................................................................................................. 69 European and Senegalese....................................................................................... 70 South African ........................................................................................................70 Eastern African, Red Sea and Persian G u lf ..........................................................71 vu Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Indian Ocean and Tropical West P acific................................................................ 72 Temperate A ustralian............................................................................................. 73 New Z ealand.......................................................................................................... 7 4 Northwestern P acific..............................................................................................75 Northeastern Pacific ..............................................................................................75 Tropical Eastern Pacific......................................................................................... 76 Zoogeography....................................................... 76 CHAPTER 2: REVIEW OF FOSSIL ABALONE (GASTROPODA: VETIGASTROPODA: HALIOTIDAE) W ITH COMPARISON TO RECENT SPECIES. Introduction ............................................. .78 Diagnostic C haracters of the Famify ............ 78 Taxonomy ......................................................................................... .80 The shell as the basis of taxonomy ...................................................................... 80 Fossil abalone taxa ............................................................................................... 83 Fossil abalone in the phylogenetic context............................................................ 88 P reserv atio n ............................................................................................................... 89 Paleoenvironments .................. 90 Reef paleoenvnronments ....................................................................................... 90 Shailow-water rocl^ paleoenvironments..............................................................91 Time Range ............ ...9 2 Geographic D istribution .............................................. *94 vui Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CHAPTERS: DNA SEQUENCE ALIGNMENT IN THE U G H T O FTH E ABDUCTIVE NATURE OF CLADISTIC HYPOTHESIS GENERATION. Introduction .......................................................... I l l M olecular and Morphological C haracter Equivalence ..................................... .112 The observational phase........................................................................................113 The explanatory p h ase.......................................................................................... 116 Cladistics as science .................. 119 Observation: From Perception to Classification .................. 121 i: Special sim ilarity.............................................................................................. 123 ii: Positional correspondence -..................................................................123 Sequence alignm ent................................................................................* .............124 Observation and Explanation ............................................... 125 Primary and secondary h o m o io g r.......................................................................125 Separation o f pow er....................................................................................... 126 C haracter In d ependence........................................................................................ 128 Objectivity and Subjectivity .............. I l l Character selection and observation.....................................................................131 Alignment param eters..........................................................................................132 Manual editing .............................................................................................. 133 A lternative Coding Strategies ................................................................................ 133 E lision...................................................................................................................134 Case sen sitiv e...................................................................................................... 136 Nfissing d a ta ........................................................................................................ 137 IX Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Polymorphic ........................................................................................................ 137 Exclusion ............................................................................................................ 138 C ontraction.......................................................................................................... 138 Homology is Special SimOarity sensu Remane (1952) ........................... 139 Belief formation ..................................................................................................140 Rational for data exclusion ................................................................................. 142 HIghfy Dissim ilar Taxa ........................................................ 146 Stretch c o d in g ......................................................................................................148 Block co d in g ........................................................................................................ 149 Presence/absence co d in g ..................................................................................... 150 Comparison to practice in morphology .............................................................. ISO B etter alignm ent? ....................................... .152 Conclusions and Recommendations .................................................... 154 CHAPTER 4: DISTRIBUTION AND BIOGEOGRAPHY OF THE RECENT HALIOTIDAE (GASTROPODA: VETIGASTROPODA) WORLD-WIDE. Introduction .......................................................................................................156 M ateriab and Methods ........................................................................... .158 Sources o f Raw D ata............................................................................................158 Abbreviations o f collections ............................................................................... 159 Biogeographi(al regions ..................................................................................... 160 Biogeographical analysis.................................................... 162 Cladistic analysis o f ta x a ..................................................................................... 165 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Dîstribatfon .............................................................................................................. Distrîbutîoiial p attern s......................................................................................... 175 Taxonomic issues..................................................................................................177 Area cladogram s..................................................................................................178 Cladistic analysis o f ta x a ..................................................................................... 182 Illustrations o f species........................................................................................... 185 Biogeography and the fossil re c o rd .................................................................... 188 DISTRIBUTIONAL DATA AND MAPS .....................................................189 Mediterranean and West Aftican Species (Figures 4-4-14,4-146, 4-147, 4-149,4- 150,4-170,4-172,4-174) 189 H aliotis marmorata Linnaeus, 1758 (Figures 4-14,4-146,4-149) ................... 189 H aliotis stom atiaeform isRs& fc, 1846 (Figures 4-14,4-147,4-150).................. 192 H aliotis t. tuberculata Linnaeus, 1758 (Figures 4-4-9,4-12-14,4-172,4-174) . 192 South Afiican Species (Figures 4-14,4-142-145,4-148,4-151-156,4-159-162) 200 H aliotis midae Linnaeus, 1758 (Figures 4-14,4-142-145,4-148)...................... 200 H aliotis parva Linnaeus, 1758 (Figures 4-14,4-154-156) .................................201 H aliotis queketti Smith, 1910 (Figures 4-14,4-161,4-162) ...............................202 H aliotis spadicea Donovan, 1808 (Figures 4-14,4-151-153)............................ 202 H aliotis speciosa Reeve, 1846 (Figures 4-14,4-159,4-160) ............................ 203 East Afiican Species (Figures 4-14-23,4-130,4-133,4-157,4-158,4-169,4-171) 204 H aliotis mariae Gray, 1826 (Figures 4-14-16)................................................... 204 H aliotis pustulata Reeve, 1846 (Figures 4-14,4-17-20)..................................... 204 H aliotis rugosa Lamarck, 1822 (Figures 4-14,4-21,4-22).................................208 H aliotis squamosa Gray, 1826 (Figures 4-14,4-130,4-133) ............................ 208 X I Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis m ila tera lis Lamarck, 1822 (Figures 4-14,4-23,4-157, 4-158,4-169,4- 1 7 1 )................................................................................................................ 208 Indo-Pacific and Central Pacific Species (Figures 4-24-44,4-63,4-66,4-116,4-117, 4-119,4-120,4-122-129,4-136-141,4-163-166,4-175,4-176).................. 209 H aliotis asinina Linnaeus, 1758 (Figures 4-35,4-36,4-43,4-165,4-166)____210 H aliotis clathrata Reeve, 1846 {non Lichtenstein, 1794) (Figures 4-24,4-138.4- 141,4-175,4-176) 219 H aliotis crebrisculpta Sowerby, 1914 (Figures 4-24,4-25,4-28)...................... 222 H aliotis diversicolor Reeve, 1846 (Figures 4-24,4-40-42).................................222 H aliotis dissona (Iredale, 1929) fig u res 4-24,4-139,4-140) .......................... 224 H aliotis dohmiana Dunker, 1863 (Figures 4-24,4-120,4-123)........................ 225 H aliotisfatui Geiger, 1999 (Figures 4-43,4-116,4-117) ................................... 225 H aliotis glabra Gmelin, 1791 (Figures 4-26,4-27,4-29,4-30,4-43)................ 227 H aliotis jacnensis Reeve, 1846 (Figures 4-24,4-124-129).................................228 H aliotis ovinaGméihïy 1791 (Figures 4-31-33,4-43)......................................... 230 H aliotis planata Sowerby, 1882 (Figures 4-43,4-119,4-122) .......................... 236 H aliotispulcherrim a Gmelin, 1791 (Figures 4-38,4-39,4-44) ........................ 237 H aliotis rubiginosa Reeve, 1846 (Figures 4-24,4-136,4-137).......................... 239 H aliotis squamata Reeve, 1846 (Figures 4-24,4-63,4-66) ...............................240 H aliotis varia Linnaeus, 1758 (Figures 4-34,4-37,4-43,4-163,4-164)............ 241 Australian Endemic Species (Figures 4-45-67,4-131,4-132,4-134,4-135,4-183- 1 9 2 )................................................................................................................ 250 H aliotis brazieri Angjss, 1869 (Figures 4-45,4-48,4-67)...................................250 H aliotis coccoradiata Reeve, 1846 (Figures 4-47,4-50,4-67,4-18Q .............. 253 H aliotis cyclobates Pérou & Lesueur, 1816 (Figures 4-51,4-52,4-67,4-185) . 254 xn Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis elegans Philippe 1844 (Figures 4-53,4-54,4-67,4-192) .................... 255 H aliotis hargravesi Cox, 1869 (Figures 4-46,4-49,4-67,4-189)...................... 256 H aliotis laevigata Donovan, 1808 (Figures 4-57,4-60,4-67,4-184)................256 H aliotis roei Gray, 1826 (Figures 4-61,4-64,4-67,4-188) .............................. 257 H aliotis rubra Leach, 1814 (Figures 4-62,4-65,4-67,4-183) .......................... 260 H aliotis scalaris Leach. 1814 (Figures 4-55,4-56,4-58,4-59, 4-67, 4 -1 9 0 )... 263 H aliotis sem iplicata Menke, 1843 (Figures 4-67, .4-131, 4-132, 4-134, 4-135,4- 1 8 7 )............................................................................................................... 264 New Zealand Species (Figures 4-67-79,4-177-182)...........................................266 H aliotis australis Gmelin, 1791(Figures 4-67,4-68,4-71,4-177,4-178)..........266 H aliotis iris Gmelin, 1791 (Figures 4-67,4-74-79,4-179,4-180) .................... 268 H aliotis virginea Gmelin, 1791 (Figures 4-67,4-69,4-70,4-72,4-73,4-181,4-182) 270 Northwest Pacific Species (Figures 4-80-86,4-118,4-121) .............................. 272 H aliotis discus Reeve, 1846 (Figures 4-80,4-83,4-86) .....................................272 H aliotis exigua Dunker, 1877 (Figures 4-87,4-118,4-121) .............................. 276 H aliotis gigantea Gmelin, 1791 (Figures 4-81,4-84, 86) ...................................276 H aliotis madaka Habe, 1977 (Figures 4-82,4-85,4-86).....................................277 East Pacific Species (Figures 4-87-102,4-104-107,4-109,4-110,4-112,4-113,4- 167,4-168,4-173) 278 H aliotis corrugata Wood, 1828 (Figures 4-88,4-91,4-100).............................. 278 H aliotis cracherodii Leach, 1814 (Figures 4-89,4-92,4-100) .......................... 284 H aliotis dalli Henderson, 1915 (Figures 4-107,4-110,4-113) .......................... 289 H aliotisfitlgerts Philippi, 1845 (Figures 4-90,93,100,173) ............................ 290 H aliotis k. kamtschatkana Jonas, 1845 (Figures 4-87,94,97) .......................... 293 xin Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis kamtschatkana assim ilis Dall, IS78 (Figures 4-95,4-99,4-100)..........296 H aliotis roberti McLean, 1970 (Figures 4-107,4-109,4-112,4-167,4-168) . . 298 H aliotis n^escens Swainson, 1822 (Figures 4-96,4-99,4-100)........................ 298 H aliotis sorenseni Bartsch, 1940 (Figures 4-100,4-101,4-104).........................303 H aliotis walallensis Steams, 1898 (Figures 4-100,4-102,4-105) .................. 305 Western Atlantic Species (Figures 4-107,4-108,4-111,4-114.4-115).............. 307 H aliotis aurantiian Simone, 1998 (Figures 4-107,4-108,4-111)...................... 307 H aliotispourtalesii Dali, 1881 (Figures 4-107,4-114,4-115) ...........................307 CHAPTER 5: A TOTAL EVIDENCE CLADISTIC ANALYSIS OF THE WORLD-WIDE HALIOTIDAE. Introduction ..................... 324 M aterials and Methods .................................................. 325 Source o f specimens ........................................................................................... 325 SEM preparation..................................................................................................325 Cladistic m ethodology......................................................................................... 325 Outgroup comparison ......................................................................................... 329 AUo^me frequencies....................... 330 Outgroup comparison ......................................................................................... 332 Reanalysis o f allozyme data with comparison to the results o f Brown (1993) . 332 DNA coding strategies fo r I 6kD cDNA ......................................... 336 Outgroup comparison ......................................................................................... 337 Reanalysis o f the lysin d a ta .................................................................................337 XIV Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. P artial 16S mtONA (C harier Wray, onpabL data) ............................................J4 5 M orphological characters ............................................................................... 348 The radula in abalone ......................................................................................... 348 Outgroup comparison ........................................................................................ 351 Characters and character states ...........................................................................352 TheEpipodium ...................................................................... 356 Outgroup comparison ......................................................................................... 359 Characters and character states ...........................................................................359 Potential characters not u s e d ...............................................................................363 Shell morphology ................................................................................................363 Shell m ineralogy................................................................................................. 363 Anafyses with morphological characters .......................................................... 364 Classification ................ 369 H aliotis, sensus stric to ......................................................................................... 370 N ordotis................................................................................................................ 370 K otohaliotis..........................................................................................................371 Sanhaliotis............................................................................................................372 Use o f Genus-level ta x a .......................................................................................372 CONCLUSIONS C hapter I : Recent T a x a ............... - ................. *373 C hapter 2: Fossil T a x a ....................................................... *373 C hapter 3: DNA Sequence Alignment .......................................................... • • *373 C hapter 4: Biogeography ............................... 374 C hapter 5: Total Evidence Cladistic A nafysis.......................................- ............ 375 XV Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF FIGURES Figures 1-2 - 1-5. Shells of imperforate specimen and potential hybrid o f H aliotis bra - zieriT/L hargravesi. ...............................................................................................11 Figures 1-6 - 1-9. Shells of designated lectotypes for H aliotis m ultiperfbrata and H revela ta ................................................................................................................ 29 Figures I-IO - 1-15. Shells o f uncommonly illustrated abalone 1 ................... 53 Figures 1-16 - 1-21. Shells of uncommonly illustrated abalone H ............... 54 Figures 1-22 - 1-27. Shells of uncommonly illustrated abalone HI................ 55 Figures 1-28 - 1-33. Shells o f uncommonly illustrated abalone IV. ............ 56 Figure 2-1: Specimens o f fossil haliotids................................................................. 79 Figure 2-2: Histograms o f number o f open tremata for several species of H aliotis. 82 Figure 2-3. Localities o f fossil abalone listed in Appendix. .................................... 95 Figure 3-1. Treatment o f morphological and molecular characters in cladistics with respect to homology. .........................................................................................114 Figure 3-2. The effect of character coding on homology statements in questionably aligned DNA sequences, using hypothetical sequences o f four tax a................ 135 Figure 3-3. Coding strategies for a few highly dissimilar tax a............................... 147 Figure 4-1. Illustration o f the three biogeographical models proposed for the origin o f H aliotidae.......................................................................................................... 157 Figure 4-2. Area cladogram (Brooks parsimony) for the genus H aliotis. .............179 Figure 4-3. W dpoint rooted taxon cladogram o f the genus H aliotis. ................... 183 Figure 4-4-13. Shells o f the European-Mediterranean species o f H aliotis spp. . . .190 Figure 4-14. Dûtribution o f European-Mediterranean, west Afoican, and south A6 ican species o f H aliotis spp........................................................................................ 191 Figures 4-15-23. Shells o f East Afiican abalone..................................................... .205 XVI Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 4-24. Distributioii of Indo-Pacifîc species o f H aliotis sp. (I)...................... .211 Figures 4-25-42. Shells o f ludo-Pacific abalone..................................................... 2 \2 Figure 4-43. Distribution of Indo-Pacific species o f H aliotis spp. (II)................... 226 Figure 4-44. Distribution o f the central Pacific (Tuamotus) H aliotis pulcherrima. 238 Figures 4-45-60. Shells o f abalone endemic to Australia 1 ..................................... 251 Figures 4-61-66. Shells of abalone endemic to Australia II.................................... 258 Figure 4-8. Distribution o f endemic Australian and New Zealand species of H aliotis spp.......................................................................................................................265 Figures 4-68-79. Shells o f New Zealand abalone. ................................................ 267 Figures 4-80-85. Shells o f the northwest Pacific abalone....................................... 273 Figure 4-86. Distribution of northwest Pacific species o f Haliotis spp................... 274 Figure 4-87. Distribution o f east Pacific H aliotis k. kamtschatkana....................... 279 Figures 4-88-99. Shells o f the northeast Pacific abalone I . ....................................280 Figure 4-100. Distribution o f rast Pacific species o f H aliotis spp. (I)....................299 Figures 4-101-106. Shells o f Northeastern Pacific abalone I* and unidentified species. 304 Figure 4-107: Distribution o f east Pacific (H), tropical Pacific, and western Atlantic species o f Haliotis spp....................................................................................... 208 Figure 5-1. Trees generated firom allozyme fiequency data (Brown, 1993)........... 333 Figure 5-2. Trees generated firom data o f Lee & Vacquier (1995).......................... 338 Figure 5-3. Strict consensus topology o f 16S mtDNA sequences...........................346 Figure 5-4. Strict consensus tree firom all lysin, allozyme, and 16S mtDNA data com- bm ed..................................................................................................................347 Figure 5-5. Diagram illustrating the terminology used for the radular teeth found m the family Haliotidae. ...........................................................................................350 xvn Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-6. ülustratioa of radular character states for central field. Number on images refer to character number. ................................................................................353 Figure 5-7. Illustration of radular character states for lateral teeth 3-5...................354 Figure 5-8. Illustration of radular character states for marginal teeth..................... 355 Figure 5-9. Sample epipodia o f H aliotis spp...........................................................358 Figure 5-10. Example o f a hypobranchial gland..................................................... 362 Figure 5-11. Strict consensus tree o f680 MPRs firom combined analysis of allo^mie, lysiiu 16S mtDNA, and morphology................................................................ 365 Figtne 5-12: Strict consensus tree o f 32 ingroup taxa firom combined analysis of allozyme, lysin, 16S mtDNA, and morphology, without outgroups.................. 367 Figure 5-13. Strict consensus tree for combined analysis o f lysin, 16S mtDNA, and morphology, without allo^mie d ata................................................................. 368 xvui Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LISTOF'M LES Table l-L Genus-level taxa in the family BMiotidae................................................. 17 Table 2. Differences between the valid taxa of this study with their status in Wagner & Abbott (1978) and illustrations in Kaicher (1981)............................................... 26 Table 2-1: Comparison of upper Tertiary species with Pleistocene-Recent species from C alifornia............................................................................................................85 Table 2-2: Chromosome number in Recent Haliotis spp. as indicated in the respective source....................................................................................................................97 Table 2-3: List o f fessil abalone taxa. ................................................................... 100 Table 4-1. Data matrix of biogeographical analysis................................................. 163 Table 4-2. Recoded allo^mie frequency data from Brown (1993)...........................166 Table 4-3. Recoded sequence o f the lysin data o f Lee & Vacquier (1995).............. 168 Table 5-1 : Source o f specimens and data. .............................................................326 Table 5-2. Character states for the radular characters. .......................................... 349 Table 5-3: Character states for epipodial characters...............................................357 Table 5-4. Character states for hypobranchial gland characters.............................. 361 XIX Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Introduction This dissertatioa evaluates the evolutionary history o f the marine gastropod family Haliotidae, part o f the clade Vetigastropoda. Abalone are important for commercial fish eries» hence are relatively well-known. However, it is surprising that the assessment of its species-level diversity has remained untouched since the beginning of this century. After one hundred years o f neglect it seems appropriate to re-evaluate the alpha and beta taxonomy o f the family and to provide a hypothesis o f relationships. At the beginning o f any phylogenetic analysis, the ingroup taxa need to be evalu ated, because any phylogenetic tree is only as good as the names on the terminal nodes. In chapter I, I assess all the over 200 species-level and the 17 genus-level taxa. The evaluation is based on all the original descriptions as well as much secondary literature, both scientific as well as grey literature. Inspection o f US and European museum col lections adds to the in-depth study o f the taxonomic issues. The family has an evolutionary history reaching at least to the Upper Cretaceous, with some fossil representatives known firom many regions of the world. Can the fossils contribute to the understanding o f the evolution o f the group leading to to d y ’s diver sity? Chapter 2 investigates this question, which will also determine whether the fossil representatives can help to elucidate the phytogeny o f Haliotidae. The goal o f this study is a “total evidence cladistic analysis", which by definition utilizes multiple sources o f information. The treatment o f diftorent kinds o f data often gives rise to mtemai methodological conflicts. Some o f the most severe conflicts arise in the treatment o f DNA sequence data. In order to include DNA sequences in the analy sis, the treatment o f all observations is scrutinized, and a m ethodolo^ that can accom modate all types o f data is developed in Chapter 3. The new and rather unorthodox Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. coding strategies are based on a sound understanding of principle in philosophy o f sci ence. These strategies will be applied in the analyses presented in Chapters 4 and 5. One o f the basic biological parameters is the distribution o f species. This informa tion is mostly unavailable for members of the family Haliotidae. Chapter 4 presents dis tributional data for each species within the family, for the inference on distributional patterns wül be based on actual localities, as opposed to vague intuitive indications. The distributional data are subjected to a rigorous analysis using Brooks parsimony, where taxa are used as characters, and the distributional areas are used as taxa. This will help to evaluate the possible evolutionary scenarios and their implication for the distri bution o f abalone world-wide. Chapter 5 reviews published data and phylogenetic hypotheses, re-analyzes the data, and assesses the differences in the phylogenetic hypotheses due to altered character coding. The main re-coding strategies will convert alloqmie ftequencies to character state data and change the handling o f questionably aligned sequences. Morphological characters 6 om the radula, the epipodium, and the hypobranchial gland are added, and all data are analyzed using outgroups from all vetigastropod families. The effect o f recoding strategies and missing data on the ensuing hypotheses are evaluated. Recom mendations concerning the use o f genus-level taxa within the family Haliotidae, based on recurring groupings from the phylogenetic analyses are given at the end o f chapter 5, bringing to closure the cfrcle started in chapter I. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter 1: Recent Genera and Species of the Family Haliotidae Rafinesque, 1815 (Gastropoda: Vetigastropoda) INTRODUCTION Members o f the family Haliotidae occur in most tropical and temperate oceans, par ticularly in the shallow subtidal zone. The first mention o f abalone in the literature was made in the 4* century B.C. by Aristotle. In the 1 “ century AT), the name o f otia (little ear) was used by Pliny. In Japan abalone were mentioned as early as the 4“ * century A D . In the medieval literature o f Europe they were noted for the first time by Gessner in 1553. The first good illustrations were provided by Buonanni, Lister, Gualtieri, and Rumphius between 1681 and 1741, which were later cited by Linnaeus (1758). The pre- Linnean descriptions o f abalone have been dealt with more extensively in Crofts (1929), Cox (1962) and Muller (1984a). Taxonomic publications on the family began with Linnaeus (1758), who described the first seven species o f abalone using his system of binommal nomenclature. His work was continued and enlarged by Gmelin (1791), who added a further twelve taxa. Reeve (1846) described 43 new taxa in his monograph, which is one of the most important sources for the taxonomy o f the femify Haliotidae. In the late 19* century, three larger monographs were published by Sowerby (1882), Weinkauff (1883), and Pilsbry (1890), but only a few new taxa were introduced by these authors. Wagner & Abbott (1978) provided a list of taxa includmg tentative ^ o n y m ies. Kaicher (1981) illustrated all the species and subspecies she considered valid, providing the most comprehensive means available to identify the Recent Haliotidae. Pickery (1991) listed m ost abalone taxa chronologically, mcluding their references. Ubaldi (1993,1995) has started to publish a Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. series intended to cover, in alphabetical order, ail extant species o f abalone; to date, four species have been treated. The latest valid species described is H. roberti McLean, 1970. H aliotis coccinea canariensis Nordsieck, 1975, is the most recent taxon that has been described. All taxa have been based on shells; only the neotype of H im ilateralis Lamarck, 1822, and the types o f H. aurantium Simon, 1998, firom Brazil are complete specimens with the ani mal (Geiger, 1996; Simone, 1998). The alpha taxonomy of the majority of species has been uncertain, except for most of the important commercial species. Between 30 (Dauphin etaL , 1989) and 130 (Cox, 1962) o f the over 2 0 0 species-level taxa described have been considered valid species. Most authors have estimated the number of distinct species to be approximately 75 (Thiele, 1931; Pickery, 1980; Kaicher, 1981; Lindberg, 1992). The objective here is to re-evaluate this family, to critically review all the published information, and to include additional unpublished observations. This groundwork is necessary in the light o f a forthcoming phylogenetic analysis o f the entire family, because the working unit—the species—should be clearly understood so that the data matrix will not be obscured by unresolved taxonomic problems. The conclusions that form the substance of this paper are presented in three lists: Index, Notes, and Valid Species by Faunal Regions. The alphabetical index and the valid species by faunal regions are both cross-referenced to the notes. Species illus trated are those that are infiequently shown in other publications. MATERIALS AND METHODS This work is based on a number o f visits to major museums in Europe and the United States, where the available type specimens were «cammed. Museum material was sup Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. plemented with specimens from private collections, including those o f Katharine Stew art (Carmel, California), Don Pisor (La Jolla, California), Roger Pickery (Wilrijk, Bel gium), Mark Jones (Auckland, New Zealand), and Benjamin Singer (Rehovot, Israel). Every original description was carefully checked in the original language. An eSbrt was made to track all relevant secondary literature through later citations and the Zoological Record. Some results from ongoing studies o f the animals are mentioned where they help in the understanding o f taxonomic questions; a detailed coverage o f the characters o f the animals is beyond the scope o f the present chapter and will be provided else where. The radular terminology o f Geiger (1996) is used. The statistical analysis o f the shell morphometrical data was performed with STA TISTICAL Mac 4.1 (StatSoft, 1994). For the breakpoint regression, piecewise linear regression with Quasi-Newton estimation method and least-square loss function was employed. Linear regression on the data-sets on either side o f the breakpoint yielded the slope statistics. Abbreviations o f collections. ANSP: Academy o f Natural Sciences, Philadelphia; BMNH: The Natural History Museum, London; DMNH: Delaware Museum of Natural History, Wilmington, Delaware; HUJ: Hebrew University, Jerusalem; KBIN: Koniglich Belgische Institut for Natuurwetenschappen, Brussels; LACM: Los Angeles County Museum o f Natural History; LSL: Linnean Society London; MCZ: Museum o f Com parative Zoology, Harvard University, Cambridge, Massachusetts; MNHN: Museum Nationale d’Histoire Naturelle, Paris; MHNG: Muséum d’Histoire Naturelle, Genève; NMW: National Museum Wales, Cardiff; SAM: South Australian Museum, Adelaide; SBMNH: Santa Barbara Museum o f Natural îBstory, California; USNM: United States National Museum o f Natural History, Smithsonian Institution, Washmgton. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. SYSTEMATIC AFFINITIES AND CHARACTERISTICS OF THE FAMILY Haliotidae are part o f the prosobranch Vetigastropoda, having the nacreous shell, certain characters in the cleavage pattern (van den Biggelaar, 1996), a streptoneurous nervous ^stem , two bipectinate ctenidia (gills) served by two heart atria (Diotocardia), and ctenidial bursicles (Hickman, 1988; Haszprunar, 1987; 1993 for review). The row o f tremata in abalone and the slit o f pleurotomariids (Pleurotomariidae) are homolo gous structures, but are not diagnostic characters (Delhaes, 1909; Haszprunar, 1993). Historically, Haliotidae were considered to be closely related to Pleurotomariidae and Scissurellidae and were united with these families in the superfamily Pleurotomari- oidea. The rhipidoglossan radula was thought to be a common character for Pleuro tomariidae and Haliotidae. However, the radulae in the two families differs in many ways. The rows o f the radular teeth are almost symmetrical in abalone, but in the pleu rotomariids they are distinctly asymmetrical. The rachidian tooth is well-fbrmed in Hali otidae, but reduced in Pleurotomariidae. The fine outer marginal teeth in Haliotidae show denticulate cusps (Wu & Huang; 1989; Herbert, 1990; Geiger, 1996; Stewart & Geiger, 1999); in Pleurotomariidae, however, a fan o f articulated bristles is found (Hick man, 1984a; Harasewych & Askew, 1993; Anseeuw & Goto, 1996). A comparison of Pleurotomariidae and Haliotidae to Scissurellidae is not appropriate, because the latter are subjected to different evolutionary constraints due to their small size and their detri- tal diet (Fretter & Graham, 1976; Herbert, 1986), which is reflected in their radular structure. The radulae o f juvenile abalone were treated by Tong (1985) and Garland et al. (1985) and are similar to those o f small Trochidae (Ifickman & McLean, 1990) and Scissurellidae (Marshall, 1993). The independence o f radular m orpholo^ and feeding ecology has to be questioned due to the «ctensive morphological plasticity o f the radula in response to the feeding e c o lo ^ o f the respective animals. The radula o f Pleuro- Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tomariîdae and Haliotidae will not help to resolve their phylogenetic relationship, because the former is that o f a specialized spongivore, the latter that o f a strict macroal- gal herbivore. The coding o f paedo- and peramorphic structures adds further problems, as in the case o f the radular characters o f Scissurellidae and Haliotidae. Either stage- specific structures are considered for coding the character states, which overstate the degree of differentiation. The rachidian tooth in mature animals is coded as serrate in Scissurellidae, but bears a smooth cutting edge in Haliotidae. Alternatively, when hete rochronic processes are taken into account, characters with inapplicable character states are created. In this case the serrate rachidian tooth o f animals <5 mm unites Scissurelli dae and Haliotidae, but the rachidian characters for animals > 10 mm are inapplicable to Scissurellidae. The use o f the radula to resolve family-level relationships within Veti gastropoda is, therefore, questionable (see also Haszprunar, 1993). The nearly symmetrical body plan o f H aliotis has been cited as being plesiomor- phic, which puts the abalone very close to the root of the fiexoglossate prosobranchs (Fleure, 1904; Salvini-Plawen & Haszprunar, 1987). This view is supported by the prim itive sperm ultrastructure (Lewis et aly 1980; Healy, 1988; 1990; Healy & Harasewych, 1992). However, several left parts of the paired body structures, e.g., the gonad and the kidney, are reduced o r modified in abalone (Crofts, 1929; Haszprunar, 1988a). The nervous system is close to the primitive condition o f Pletavtomariay but in abalone the additional structures o f the epipodium and the osphradium are hmervated by pleural and visceral elements, respectively. Haliotidae and Trochidae have synapomorphic osphradial characters (Haszprunar, 1985; 1993), but the Trochidae are clearly separated fiom Scissurellidae, Pleurotomariidae, and Haliotidae by the lack o f the right ctenidium and associated organs (Salvini-Plawen & l^szprunar, 1987). Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The three families within the old Pleurotomarioidea are now placed in their nominal superfamilies. Le., Haliotidae are now in Haliotoidea, on the same level as Pleurotomar ioidea, Scissurelloidea and the remaining Vetigastropoda (Haszprunar, 1988b; Healy & Harasewych, 1992). PROBLEMS PERTAINING TO THE TAXONOMY OF ABALONE Tremata as a taxonomic character and teratological type specimens Abalone can be easily recognized by the depressed shell and the tremata, the row o f holes used for respiration, release o f gametes, and defecation (Ino, 1952; Tissot, 1992; Voltzow & CoUin, 1995). One may potentially confuse them with some members o f the Stomatellinae, a trochid subfamily (Hickman & McLean, 1990; Pickery, 1995). Some taxa with imperforate shells described in the genus H aliotis are actually Stomatellinae {e.g., H impertusa Burrow, 1815, and are not dealt with here. Stomatellids that resem ble haliotids (Gena, Stomatella and M icrotis) are rather small (< 40 mm), have no right ctenidium, a flat shell, no tremata, and no spiral sculpture. They might suggest imperfo rate specimens o f juvenile H asinina Lirmaeus, 1758, but the latter have several dis tinct spiral ridges (see Kaicher, 1981), which are no longer formed as the shell grows larger than 3.5 to 4 cm. Specimens o f the trochid genus Granata have been erroneously identified as imperforate K cyclobates Peron, 1816 (Geiger, pers. obs.). In most descriptions o f abalone the number o f open tremata is indicated, erroneously suggestmg that it is o f value for the identification o f a given species. However, the num ber o f open tremata changes durmg the growth o f the shell (Hemphill, 1907; Sinclair, 1963). The larval shell has no tremata at all; the & st one is formed at a size o f approxi mately 1-3 mm (Crofts, 1929; Murayama, 1935; Bonnot 1940; Ino, 1952; Shibui, 1971; Mu e t aL, 1976; Beveiander, 1987). Figure 1-1 shows the size dependent change o f the Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. m m CD C D o in in in C O CO to C M in in CO t c number of open tremata Figure l-L Plot o f shell length versus number o f open tremata in K tuberculata from numerous Mediterranean populations (n = 433). Note the positive correlation between the two parameters, fbr which a loganthmic curve has been fitted. Note the intersection with the x-axis around 2 mm shell length, the size at which the & st perforation is formed- The first specimens with five open tremata are found at 7 mm, those with six open tremata at 15 mm, and those with seven open tremata at 24 mm. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. number o f open holes for the European K tuberculata Linnaeus, 1758. Linear break point regression (r^= 0.69) estimated the breakpoint at 5.5 open holes, corresponding to a size o f approximately 33 mm, where the slope changes from 3.8 (SE = 0.53) to 0.0017 (SE = 0.56). The number o f open tremata varies greatly within a species and between populations (HempfaiU, 1907; Schepman, 1909; Sinclair, 1963; Talmadge, I960; Geiger & Groves, 1999). Therefore, the notion that the number o f open tremata or total number o f tremata is fixed, is misleading. Taxa based solely on the number o f open tremata have to be rejected as in the case o f H. m ultiperforata Reeve, 1846 (note 32), or K cracherodii Leach, 1814, with the subspecies bonita Orcutt, 1900, and califom iensis Swamson, 1822. Specimens from some populations on Guadalupe Island off Baja Cali fornia have the H. cracherodii califom iensis morphologr. These specimens have more and sm aller tremata, which are also more closely spaced than in specimens from the mainland. Specimens with the califom iensis morphology have been cultured on the mainland o f central California and the size and spacing o f the tremata changed to those o f a typical K cracherodii cracherodii (B. Owen, pers. comm.), indicating that these characters are under environmental control. Imperforate abalone shells have been found (Smith, 1893; Marquand, 1906; Hemphill, 1907; Dall, 1919; Leighton, I960; Geiger, pers. obs.; Figure 1-2), but are rather rare. Even rarer are specimens with a double row o f tremata (Smith, 1888; ffamadfl, 1982). A somewhat larger number of specunens with a continuous slit instead o f the row o f tremata are known IfL laevigatai Gray, 1856; H. asirtma, ff. cracherodii, HI parva Lmnaeus, 1758, K planata Sowerby, 1882, H. rvfescens Swamson, 1822, K tuberculata: Geiger, pers. obs.). New tremata are formed at the anterior margin o f the shelL and posteriorly the mantle eventually seals them when they are no longer used. Occasionally one to several tremata are closed out o f the sequence just described 10 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 1-2 - 1-5. Shells of imperforate specimen and potential hybrid o f H aliotis bra - zieri x hargravesi. 1-2. H. cf. diversicolor Reeve, 1846. 28 mm. MNHN [not num bered]. No locality data. An imperforate specimen o f H aliotis. 1-3. H brazieri - hargravesi. 31 mm. K. A. Stewart collection. Solitary Island, Cofis Harbour; New South Wales, Australia. This specimen begins growth with the smooth morphology o f H. bra - zieriy but midway develops to the spual ridges typical o f H. hargravesi. 1-4 - 1-5. H. brazieri AngaSy 1869.29 mm. MHNG [not numheredj. Australia. II Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (Geiger, 1991: fig. 8; K parva, H. pulcherrimaGva&^xi, 1791: Geiger, pers. obs.). Only in H. elegans Philippi, 1844, is this phenomenon found regularly and becomes the rule in large specimens (see Wilson, 1993: pi. 3, figs. 2A, 2B). H aliotis im perforata Dali, 1919, H. lusus Finlay, 1927, and H. holzneri Hemphill, 1907, are based on imperforate type specimens. It is clear that these are teratological shells not warrantmg taxonomic recognition (note 6 6). Three type specimens show further deformities. H aliotis sieboldii Reeve, 1846 (note 64) has been described firom a distorted shell, in which the row o f tremata is located abnormally close to the periphery of the shell. The same applies also to K whitehousei (Colman, 1959) (note 55), although the distortion is not as pronotmced as in /£ sieboldii. This type o f a deformation is also known firom some specimens o f H. cracherodii (LACM 23452; SBMNH 13522; USNM 199890). H aliotis diegoensis Orcutt, 1900 (note 65) has an extremely thick and stout shell, a growth form that had been induced by boring organisms, most likely sabellid polychaetes. H ybrids The occurrence o f hybrids may be challenging to the biological species concept, but can be better understood firom the standpoint o f the evolutionary species concept. As long as the two species that hybridize keep their identity, they remain intact; otherwise a case o f reticulate spéciation is found (c f Vfiley; 1981). When discussing hybrids, it is assumed at the outset that the two parent specimens belong to two discrete species, which may not be the case. We may also have a case o f unrealized mtraspecific vari ability o f a single species. Hybridization among the Californian species is well established and has been stud ied on the basis o f the shell, the epipodium, and by using mimimological techniques. 1 2 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The hybrid specimens with intermediate shell and epipodial characters occurred with a ftequency o f approximately 1:500 (Owen et al.^ 1971). These authors also found evi dence for back crosses o f hybrids with some o f the parental species, demonstrating that the FI hybrids are fertile. These results were later corroborated by laboratory rearings (Leighton, 1988). The identity o f the recognized species was confirmed with DNA sequence data (Lee & Vacquier, 1992; 1995). Although hybrids involving H. cracherodii were successfully reared in the laboratory (Leighton, 1988), they are markedly absent in the field, but this can be accounted for by two factors. First, H. cracherodii has an entirely smooth and uniformly black shell, whereas all the other California species show some sculpture and may have a color pattern in the shell. As it is much more difficult to generate a new character (sculpture, color pattern), than to modify an existing one, the simple condition in H. cracherodii prevails in hybrid specimens. This condition is termed ‘overshaddowing eS ect’ Second, H. cracherodii is the only Californian abalone found in the intertidal region. The species is ecologically separated firom the remaining species. Within the remaining six Californian species, 13 o f the 15 possible hybrid com binations have been documented (Talmadge, 1977b; Leighton, 1988). The two missing hybrids both involve H fidgens Philippi, 1845, which has been shown in the phyloge netic analysis o f DNA sequence data to be slightly more distantly related to the other California species (Lee & Vacquier, 1995). Pre-fertilizing barriers (analogous to pre mating barriers in broadcast spawners), such as spawning season and vertical distribu tion o f the species, have to be taken into accotmL Unfortunately, no information on the fertility o f F2 and backcross specunens is available, although many profound changes affecting the fertility may take place after the FI generation (see Kmg, 1993 for review). In a second case, two Australian species fT. laevigata and H. rubra Leach, 1914, form hybrids that occur in nature (Anonymous, 1973; R. Fallu, pers. comm.). These 1 3 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. two species are well-known, and have been shown to be distinct species (Brown, 1993; Lee & Vacquier, 1995). In this case, hybrids have been identified on the basis o f en^nne electrophoresis data. The hybrids occur in varying frequency between 0% and 2% in several populations studied. Good evidence has been presented for the presence of back- crosses of hybrids with either parental species. The hybrids are not believed to repre sent a starting point o f reticulate spéciation because o f the different habitat requirements o f the two species, which is strongly reflected particularly in the thickness o f the shell (Shepherd, 1973; Brown, 1995). Hybrids are reported from two sympatric populations of the two subspecies of K discus Reeve, 1846, K discus discus and H. discus hannai Ino, 1952 (Fujino et air 1980; Sasaki et a i, 1980). Although a case o f sympatric subspecies may seem a contra diction in terms, it can be seen as a necessary step o f gradual, sympatric spéciation (see Futuyma, 1986). As the two subspecies are very difficult to distinguish, en^nne elec trophoresis was used to identify them with polymorphic loci o f several emymes. Later the same data and method were used to identify hybrids between these two taxa. It is not clear whether there was rather extensive variability in one biological species, or whether true hybrids were fbtmd; no data on the identification of the specimens studied using alternative methods had been mentioned. Yet another case is the least clear and has not been discussed in the literature to date. It involves the following taxa from south eastern Australia: H. brazieri PiUges, 1869, K ethologus (fredaie, 1927), K hargravesi Cox, 1869, and H. m elculus (fredale, 1927), all uncommon to rare. I regard K melcuhts as a synonym o f K brazieri^ and H. ethologus a synonym o f Æ hargravesi (notes 62,66, and 72). Spiral ribs showing exten sive mtraspecific variation in respect to number and elevation are found on the dorsal side o f the shell o f K hargravesi H aliotis brazieri (Figures 1-4 - 1-5) has no spiral 14 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. sculpture at ail, only an uneven but smooth surface. Intergrading specunens, ie ., poten tial hybrids, show slight spiral grooves, which, however, are much less distinct than any form within the range o f K hargravesi. Some specimens are known in which the early whorls lack spiral cords, but on the body whorl such ridges appear abruptly (Figure l- 3). These patterns suggest a highly variable species with a few conchological forms, which would not warrant taxonomic separation. M. Jones (pers. comm.) noted that H. brazieri and H. hargravesi are separated by their vertical distribution. H aliotis brazieri occurs from the lower reef surface to 40 m and H. hargravesi firom 40 m downwards o f the slope o f the reef, where they can be found under coral heads and boulders. Coleman (1981:86) confirmed a deep water habi tat (“ 15 to 40 m”) o f H. hargravesi as compared to other abalone species, though the depth range indications o f these two sources do not overlap. Hybrids are most often found around 40 m, the depth where the vertical ranges o f the two species overlap. On one occasion a H. brazieri and a H. hargravesi were found under the same boulder. The epipodium o f the hybrid specimens showed intermediate characteristics (M. Jones, pers. comm.), though my own investigations show very little difference in the two taxa. It is well known that abalone show gregarious spawning behavior (Murayama, 1935; Shepherd, 1986) and it is, therefore, not too difficult to imagine hybrids being formed under the circumstances described above. Although Vacquier and co-workers (Vacquier et o/., 1990; Lee & Vacquier, 1992; 1995; Vacquier & Lee, 1993; Lee et al., 1995) have demonstrated that the protein lysin in the head of the acrosomal vesicle in the head o f the sperm strongly promotes species-specific fertilization, it does not fully prevent inter specific fertilization. Although no good data on the fisquency o f the Australian hybrids are available, it is evidently rather low. It is open to discussion whether we observe here a case o f an erratic fertilization pattern o f two distmct species, or whether two recently 15 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. diverged species still occasionally form hybrids, or whether we stand at the cradle o f a reticulate spéciation event. GENUS-LEVEL TAXA A list of all supraspecific taxa of the family Haliotidae is given in Table l-l. Some comments are warranted on a fow taxa and are given here. Haliotis^ sensu stricto The subsequently designated type species (Montfort, 1810) o f the genus Haliotis, H asinina^ has been accepted by the majority o f authors (e.g., Iredale, 1910; Kennard e t a t., 1931; Wenz, 1938; Talmadge, 1963a; Ubaldi, 1985; 1993). Other species have been cited as type species, but these are erroneous (e.g., Cossmann, 1918: H. tubercu - lata. Thiele, 1931; Cotton, 1943; Cox, 1962: H. midae Linnaeus, 1758. Children fid e Kennard et al., 1931: H. iris Gmelin, 1791). The type designation by Montfort (1810) was unfortunate, as the most atypical species within the family was chosen as the type species. The two presumed type specimens o f H. asinina are held in the LS, and corre spond with what is currently known as this species. The type locality has been desig nated as Amboina (= Ambon), Indonesia (Talmadge, 1963a). Cotton (1943:176) discussed the status o f H. asinina as type species and following Adams & Adams (1853- 1858) stated “that asinina is the genotype o f the H aliotis o f Montfort 1810 and not o f the true H aliotis Linnaeus 1758,” a statement in contradiction with the original text. Montfort (1810:115) only introduced PadoUus as a new genus: “... nous avons cru pou voir en former un genre particulier” (... we have thought to be able to form foom it a distmct genus). A similar statement cannot to be fotmd with H. asinina, which is only designated as type o f the genus H aliotis on page 120: “Sous la denomination dforeille 16 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 ( O ' 3 . 3 " CD CD ■ D O Q . C a o 3 " O o CD Q . ■ D CD C/) C/) Thxon Valid type species OriRinal typo species (if different) EuhaliQtisyfmz, 1938 Eurotis Habe & kosuge, 1964 Exohaliotis Cotton & Godfrey, 1933 Haliotis Linnaeus, 1758 Marinauris Iredale, 1927 Miohaliotis Itoigawa & Tomida, 1982 Nfohaliotis Cotton & Godfrey, 1933 Nordotis Habe & Kosuge, 1964 Notohaliotis Cotton & Godfrey, 1933 Ovinolis Cotton, 1943 Padollus Montfort, 1810 Pam Fleming, 1952 Sanhaliotis Iredale, 1929 Schismotis Gi«y, 1856 Adams, & Adams 1854 Thinotis Adams & Adams, 1854 Timtis Fischer, 1885 Usahaliotis Hste & Kosuge, 1964 midae Linnaeus, 1758 lodl tuberculata Linnaeus, 1758 (m) cyclobates Pérou & Lesueur, 1816 |m) asinina Linnaeus, 1758 [sd: Montfort, 1810) brazieri Angas, 1869 amabiUs Itoigawa & Tomida, 1982 jmj scalaris Leach, 1814 (od| gtganrea Gmelin, 1791 |odJ rubra Leach, 1814 ov/nn Gmelin, 1791 (odj parva Linnaeus, 1758 /r/s Gmelin, 1791 (ml varia Linnaeus, 1758 (od) laevigata Donovan, 1808 tuberculata Linnaeus, 1758 asinina Linnaeus, 1758 |mj asinina Linnaeus, 1758 (obj. syn. of Haliotis) cracherodii Leach, 1814 |m) melculus Iredale, 1927 jsd: Wcnz, 1938) naevosa Martyn, 1784 (od) rubicundus Montfort, 1810 jmj excisa Gray, 1856 (not available) incisa Reeve, 1846 (sd: Cossmann, 1918) de mer, tous les anciens conchyliologues avoient déjà reconnu ce genre.... Nous avons choisi pour type, au lieu de l’haliotide commun (haliotis vulgaris, haliotis tuberculata [sic]), celui auquel on a donné le nom. d’oreille d’âne” (Under the name o f ear o f the sea, all the old conchologists have already known this genus .... We have chosen as type, instead o f the common abalone (^Haliotis vulgaris, H aliotis tuberculata\ the one, for which one has given the name of donkey’s ear [H. asininaj). Montfort clearly desig nated the type species o f Linnaeus’ Haliotis, and not a monospecific genus for H asin - irta because two other taxa are included under H aliotis. The fact that a common name is used in the text o f the designation (see ICZN Article 12c) does not invalidate it. Mon- fort (1810:119) used the following titles for the description o f the species: “Espèce ser - vont de type au genre. Haliotide orielle d'ane. H aliotis asinirtus." (Species serving as type o f the genus. Haliotid donkey’s ear. H aliotis asininus). First, the type species is clearly designated, and second, the association between common name and scientifîc name is unequivocal. The name H aliotis stems from the two Greek words halios (the sea) and ous, otis (the ear). The gender o f the genus is feminine because Linnaeus used the feminine end ing for the species-level taxa derived from an adjective, particularly for H. asinina, the type species o f the genus by subsequent designation o f Montfort (1810). Other genus^Ievel taxa, and their type species One objective synonym o f Haliotis has been introduced. Le., a taxon that is based on the same type species, H. asininai Teinotis Adams & Adams, 1854. Note that The G enera o f Recent M ollusca by Adams & Adams (1853-1858) has as an inside cover publication cfcite 1858, but in Vol. 2:661 the actual publication dates o f the various parts 18 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. were indicated. For part 14 on Haliotidae, June 1854; therefore, this latter is the correct date for Adams & Adams* abalone taxon. The genus Padollus Montfort, 1810, with the type species by monotypy, P. rubicun - dus Montfort, 1810 (non H. rubicunda Roding, 1798: see note 20), is problematic Halt - Otis scalaris (Leach, 1814) has been listed as the type species o f Padollus {e.g., Knight et al., I960; Wagner & Abbott, 1978; Lindberg, 1992:16), but P . rubicm dus Montfort is a synonym o f H. parva and not of H. scalaris (see note 20); therefore, the valid type species for Padollus is H. parva. The fact that the type species in the original descrip tion o f Padollus is no longer invalid does not invalidate the description (ICZN Recom mendation 67B). H aliotis rubicunda (Montfort) can be unequivocally identified and the senior synonym is the correct, valid type species. bfeohaliotis Cotton & Godfi%y, 1933, has been synonymized with Padollus on the basis o f having the same type species: H. scalaris (see above: Pickery, 1991). However, as I have shown that H. rubicm dus Montfort is not a synonym o f H. scalaris but o f H. parva, the synonymy o f Neohaliotis with Padollus has to be rejected. The type species o f Sulculus hdasns & Adams, 1854, is H. incisa Reeve, 1846, which is a synonym o f H. tuberculata (see also note 16). As H. tuberculata has also been chosen as the type species o f Eurotis Habe & Kosuge, 1964, the latter is now a subjective, junior synonym o f Sulculus. Most genus-level taxa had their type species designated by the original author. Des ignations made in the 19* century may not meet modem conventions for designation of a type species in respect to the specificity o f the language used. However, I (unlike Fleming, 1952) interpret “explicitly designated** (ICZN Article 67b) somewhat more generously. 1 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Adams & Adams (1853-1858) gave the genus name, followed by its diagnosis and the mention o f one particular species as * ‘£x.” (example = type species). This example cannot be confused with the species considered to belong in that genus, which were listed, further below in smaller print and in two columns. This form had been utilized for all the genera {Haliotis Linnaeus, Teinotis H. & A. Adams, Padolliis Montfort), but not for the subgenus SulculuSj for which no example was given. I consider this con struction as an explicit designation (ICZN Article 67b), which cannot be confused with the exeption mentioned imder ICZN Article 67c 1 (“mention o f a species as an example o f a genus or subgenus”) in conjimction with the example thatfbilows in the Code. The narrow reading o f the word “example” should not obscure the clear intentions o f the authors. Fleming (1952) did not accept the designation fbr Teinotis and attributed the subsequent designation to Cossmann (1918). M arinauris was described by Iredale (1927) without any perceptible intention to designate either M. melculus or M. ethologus as the type species; the genus was intro duced in an extremely casual fonn. Fleming (1952:229) made claims for the designa tion (“Type (here designated)”), but had overlooked the clear designation by Wenz (1938:172). Schism otis Gray, 1856, has been listed by some authors (Pickery, 1991; Ubaldi, 1993). Gray did not intend to introduce a new name for the specimens he discussed as monstrosities o f H. albicans Quoy & Gaimard, 1834 (= K laevigata)^ but only indi cated a suitable, hypothetical name. The name is a nomen nudum and is not available (see also note 70). H aleotis Binkhorst, 1861, has been cited as an objective synonym o f HaliotiSy Le., is treated as an available emendation o f H aliotis {e.g., Knight e t al., I960). An emenda tion according to ICZN 33b(0 and 33b(iii) must be “demonstrably intentional,” other 2 0 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. wise it is aa incorrect subsequent spelling (ICZN 33c), which, is not available. No intent by the author to change the original spelling can be found; the name is unavailable. Tinotxs Fischer, 1885, however, is available as a clear intent to change the name is given (Fischer, 1885:845): ""Tinotis H. et A. Adams, em. (T einotis)” It is an objective syn onym of H aliotis. Use of genus-level taxa Some authors (e.g., Talmadge, 1963a; Habe & Kosuge, 1964; Hara & Fujio, 1992) have used several genera in the family Haliotidae. Pickery (1991) provided a list and all references for the Recent taxa, which have been ranked as genera or subgenera (Table l-l). As pointed out recently (Geiger, 1996), I consider the usage o f these genera to be unjustified at this time for the following reasons: 1) In the descriptions o f the one fossil (Itoigawa & Tomida, 1982) and the 17 Recent supraspecific taxa, only the Qfpe species had been assigned, occasionally with selected species. O f the 200 species-level taxa only approximately 83 have ever been assigned to any supraspecific taxon, and 2 2 o f those to more than one group, demonstrating the problematic supraspecific taxonomy o f haliotids. The descriptions o f these genus-level taxa are entfiely Qrpological and no author has attempted to provide serious di% rential diagnoses. 2) Only two studies have utilized systematic methods other than shell morphology to determine the relationships o f abalone species. Brown (1993) studied 17 species using enzyme electrophoresis and Lee & Vacquier (1995) used cDNA sequences o f the sperm acrosomal protein lysm (see Vacquier & Lee, 1993 for review) o f 22 haliotids. The nominal supraspecific taxa and the limited number o f associated species are not in accordance with the groups hypothesized by these more recent studies. However, the 21 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. type species o f H aliotis, H asinina (see above for discussion), was not included in either study, making any sound taxonomic decisions impossible. 3) The only inferable consensus in the literature (McLean, 1966:151-153; Lindberg, 1992) as well among abalone researchers (Workshop “Evolutionary Biolo^f and Genet ics o f Abalone’ ^ during the Second International Symposium on Abalone Biologr, Fish eries and Culture, February 1994, Hobart, Tasmania) was not to use any genus-level taxa other than H aliotis. Yet there was an equally, strong consensus among the work shop participants that the diversity within the family may well justify the recognition of several genus-level taxa. However, monophyletic groups will have to be determined feom a phylogenetic study treating the majorify o f all abalone species. SPECIES-LEVEL TAXA Use of subspecies Subspecies are defined as allopatric populations with a fixed character (Futuyma, 1986). They do not yet represent discrete, evolutionary lineages. Interbreeding at the periphery o f these populations is not necessarily a sign o f the erroneous application of the subspecific classification, but may show that the populations are not yet independ ent species or discrete lineages. Only in very few shallow-water, broadcast-spawning gastropods have subspecies been described, but in H aliotis these have been invoked a number o f times (Lindberg, 1992), Le., in the following groups: H. disctts, H. mariae Wood, 1828, H. pustulata Reeve, 1846, H rubra, H. scalaris, H. tuberculata, H. varia, H. virgjnea Gmelin, 1791, and the populations o f Californian abalone occurring on Guadalupe Island off Baja California, Mexico, hi most cases the subspecific division is found along a temperature gradient; Le., subspecies are described fiom different lati tudes and not firom different longitudes. Most continental coast lines run m a north- 2 2 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. south, direction, which may explain the above observation. The exceptions to the north- south pattern are two southern Australian species, H. scalaris and H. rubra. On the other hand, the Lado-Malayaa Archipelago offers rich opportunities for the developent o f isolated populations, but no subspecies have been proposed for species with a wide east-west range such as H. asinina (Thailand - Fiji), H. clathrata Reeve (East Africa - Samoa), H. ovina (Maldives - Tonga), or H. planata (Thailand - Fiji). In H. discus, the karyotypes differ between the two subspecies (Nakamura, 1985), although their populations interbreed at their common boundary (reviewed in Fujino, 1992). In this case the two taxa are more likely to represent subspecies than ecomorphs. The two formerly recognized subspecies o f H. tuberculata—H. tuberculata tuber - culata and H. tuberculata lam ellosa Lamarck, 1822—have identical caryotypes (Colombera & Tagliaferri, 1983) and have been recently shown to have identical sequences o f the lysin protein (Lee & Vacquier, 1995; see also note 5). In New Zealand, four geographically separated subspecies o f H. virginea Gmelin are reported (note 60): H. virginea virginea, H. virginea crispata Gould, 1847, H. vir - ginea m orioria Powell, 1938, and H. virginea huttoni Filhol, 1880 (Kaicher, 1981; Ubaldi, 1986). These subspecies show gradual changes in several morphological char acters used to distinguish them due to environmental parameters associated with geo graphical location based on Geld observations as well as collection records (Talmadge, 1957a). M. Jones (pers. comm.) has found some other forms on the very remote islands off New Zealand that seem to be more stable and distmct than the subspecies mentioned above. In these two cases the described subspecies seem to be ecomorphs not justifying taxonomic separation. However, as no hard data is available, and because they are widely used, the subspecies o f H. virginea are conservatively retained. 23 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. None o f the Californian subspecies is retained because those cases for which data is available (Æ cracherodii caltfom iensis, H corrugata diegoertsisz see above and note 65) have been shown to be mere ecomorphs or teratological specimens. One former species has been allocated at the subspecies rank {K kam tschathm a assim ilis Dali, 1878: see McLean, 1966; Owen et aL, 1971; note 67). Two pairs of subspecies are found in southern Australia. These are separated along an east-west axis: the eastern H. r. rubra with the western H. rubra conicopora Péron, 1816 (see note 53), and the eastern H. scalaris emmae Reeve, 1846, with the western K s. scalaris (see note 56). Information other than shell morphology (Shepherd, 1973; Brown, 1993) suggests one polymorphic species in both instances. However, the shells can be distinguished reasonably well and the geographic distribution o f the morphs is more or less disjunct The available data suggests subspecies status o f the respective populations. Due to the limited information currently available, the last three cases are provi sionally studied as follows: K martae dentata Jonas, 1846, is easily recognized by the deep furrows resulting in the denticulated anterior margin o f the shell, which are miss ing in the nominate subspecies. The biogeographical data on the species is scant. No assessment o f geographic variation is possible. I retain the two subspecies of H. mariae. H aliotis pitstulata cruenta Reeve, 1846, has a reddish coloration, a very flat shell, and is found particularly in the Red Sea. Haliotis pitstulata pustulata on the other hand is more sculptured, usually dark green to mud-colored, and is found along the east AMcan mainland. There are no strikmg differences between the animals, and molecular data on the two morphs is not yet available. 1 retain the two subspecies tentatively. How ever, as their relation to H rugosa Lamarck, 1822, is currently unresolved (see note 24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 12), they may eventually be treated as subspecies o f H. rugosa or may be united in a single species without further division into subspecies. For H. varia, seven subspecies are regularly found in the literature, but none are recognized here. As the name suggests, this species is extremely variable. I have been unable to find any geographic pattern and intermediate specimens abound. These taxa are clear results o f typological thinking of the 19* century. Index In this section I give my opinion o f the taxonomic status o f every taxon o f Recent abalone. Spelling mistakes, erroneous dates, and incorrect taxon-author combinations have not been included, unless widespread confusion exists. Controversial opmions and new assessments o f the taxonomic status o f various taxa are indicated in the “^Notes” that follow. The notes are arranged according to their appearance in the section '"Valid Species by Faunal Region” below. Le., they are grouped by valid species. The two most recent, but brief, taxonomic assessments o f the family Haliotidae were provided by Wagner & Abbott (1978) and Kaicher (1981). All differences between the present study and the opinion o f these authors are listed in Table 1-2. An analysis o f all the species-level taxa reveals that only 27% of described taxa are still considered valid species. The status o f the subspecies (5%) is very much debated, because the unit of a subspecies is somewhat vaguely defined as a geographically lim ited population with a certain character, raismg the percentage o f all valid taxa to slightly less than one third (see also discussion below). Taxa that had originally been described as eidier forms or varieties are «ccluded fiom modem taxonomy and represent what we generally call ecomorphs. I do not use Latin fi>im names in an informal feshion (contra Reid, 1996), as all available evidence for abalone suggests that such taxa do not consti- 25 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 2. Differences between the valid taxa o f this study with their status in Wagner & Abbott (1978) and illustrations m Kaicher (I98I). Subspecies rank is indicated by inden tation under the respective species. The following epithets were regarded as valid by Wagner & Abbott (1978), but are here variously synonymized (see Index for details): canariensis, (ùingii, gibba, hanleyU howensis, japonica, kraussi, tuberculata lamellosOy m elculus, m ultiperforata, sulcosa, vixlirata, whitehouseL Forms listed by Kaicher (1981) have been ignored unless specifically mdicated. The following epithets shown by Kaicher (1981) are either variously synonymized here, or were illustrated by speci mens that cannot be identified (see Index for details); bistriata, corrugata oweni, dis - sona, elevata, gemma, multiperforata, sepiculata, whitehousei 26 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. This study Wagner & Abbott (1978) Kaicher f 1981) b r e a i e r i 1869 c r a c h e r o d i f L e a c h , 1814 c l a d a r a a d i v e r s i c o l o r Reeve, 1846 d o h m i a n a Danker, 1863 p l a n a t a Sowerby, 1882 h a r ^ a v e s i Cox, 1869 j a c n e n s i s Reeve, 1846 k a m ts c h c u k a n a Jonas, 1845 a s s i m i l i s Dali, 1878 m a r i a e d e n t a t a Jonas, 1846 m a d a k a (J^aiSoa, 1977) m a r m o r a ta Linnaeus, 1758 ovma Gmeiin, 1791 p u s t u l a t a Reeve, 1846 c r u e n ta Reeve, 1846 r o e i Gray, 1826 ruAügmara Reeve, 1846 ruéra Leach, 1814 c o n i c o p o r a Pérou, 1816 r u g o s a Lamarck, 1822 s c a l a r i s e m m a e Péron, 1816 s p e c i o s a Reeve, 1846 s q u a m o s a Gray, 1826 s to m a ti a e f b r m i s tu b e r c u l a t a Linnaeus, 1758 c o c c i n e a Reeve, 1846 u r d l a t e r a l is Lamarck, 1822 varût Lmnaeus, 1758 vo^gmeaGmelm, 1791 c r i s p a t a Gould, 1847 h u t t o n i FOhoi, 1880 m onona Powell, 1938 b r a z i e r i c r a c h e r o d i i as r u b r a c l a t h r a t a not mentioned “no information” p l a n a t a h a r g r a v e s i j a c n e n s i s k a m ts c h a tk a n a a s s i m i li s not mentioned not mentioned undetermmed species a v i n a p u s t u l a t a not mentioned r o e i synonym o f v a r i a r u b r a c o n i c o p o r a synonym o f t u b e r a d a t a e m m a e synonym o ff u lg e n s s q u a m o s a v a r i a t u b e r a d a t a c o c c i n e a u n i l a t e r a l i s v a r i a v v r g in e a as form o f v ir g i n e a as form o f v ir g i n e a as form o f v ir g j n e a also as m e lc u lu s includmg all ssp. also as c r e b r is c u lp t a auct and v e n u s ta also as a q u a t i li s and all ssp d o h m ia iu z also as g f a y a n a also as e th o lo g u s also as h a n le y i also as a u la e a a s s im U is m a r ia e d e n ta ta not mentioned d & g u in e e n s is shows c y c l o b a t e s also as r e v e l a t a c r u e n ta also as s u lc o s a as v a r ia r u b ig in o s a , h o w e n s i s also as a t K i l e c o n ic o p o r a as p u s t u l a t a form ( d t e m a t a e m m a e s p e c io s a c f . d i v e r s i c o lo r shown not mentioned also as l a m e l lo s a also as z e a l a n d i c a vom shown also as w d l a t e r a l i s , c ù ’i n g i i and ssp. p a p u l a t a p u s tu lif é r a , v i r i d i s a ls o d s g i b b a c r i s p a ta h u tto n i m o r io r ia 27 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tute discrete gene pools. The invalid taxa—synonyms (46%), homonyms (5%), nomina dubia (6 %), nomina nuda (2%), ecomorphs (6%), unavailable names (3%)—account for over two thnrds o f all published species-level taxa. The status o f some taxa is still unclear; some may never be folly resolved due the loss o f type material, or there is currently too little material available to make a defini tive assessment. Unresolved cases are mentioned here. The type of K canaliculata Fis cher, 1907, was lost prior to 1872 (Ivanov & Kantor, 1991) and its synonymy with H. parva is uncertain (note 19). The following taxa can only be tentatively synonymized, because the type material remains to be located and the original description and/or illus- tration do not allow a clear assessment: crenata Swainson, 1822, glabra Swainson, 1822, scutulum Reeve, 1846 (notes 47,48), sepiculata Reeve, 1846, and sinuata Perry, 1811 (note 22). Seven taxa cannot be identified and are treated as nomina dubia: acbiat - ica Nardo, 1847, bistriata Gmelin, 1791 (notes 5, 14), imperfbrata Gmelin, 1791 {not Dali, 1919) (note 6 6), interrupta Valencieimes, 1831 (note 73), parm a Valenciennes, 1831 (note 7 3 \p lic a ta Gmelin, 1791, and rotundata Perry, 18II (note 22). The taxa maculata Küster, 1840 (note 18), maculosa KQster, 1840 (note 18), modesta auct. (note 75), secem enda Monterosato, and schroeteri Menke (note 76) are treated as nomina nuda, because th ^ could not be traced to the original source. The deposited type speci men o f H. victoriae Brazier is, in the absence of an original description, a nomen nudum (note 77). H aliotis stom atiaeform is Reeve, 1846 = neglecta Philippi, 1848 (Geiger & Owen, m prep.), is tentatively resurrected but the material available is very limited and restricted to a Ihnited number o f shells with some some preserved anhnals (notes 5 , 8, 9). H aliotis exigua Dunker, 1863, is tentatively retained as a valid species (note 63). Lectotypes are here selected for two taxa: H multiperforata Reeve, 1846 (note 32), and/X: reve/ntoOeshayes, 1863 (note 12). The figured type specimen o f unknown prove- 28 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 1-6 - 1-9. Shells o f designated lectotypes ïo t Haliotis m ultiperforata and H revelata, 1-6 - 1-7, H. m ultiperforata Reeve, 1846.63 mm. BMNH. Mus. Cuming. Lec- totype here selected. 1-8 - 1-9. H, revelata Deshayes, 1863. 58 mm. MNHN. Bourbon. Lectotype here selected. 29 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. nance of K multiperforata, which, is here designated as lectotype (Figures 1-6 - 1-7), is clearly a specimen o f H. rugosa Lamarck (note 12), however, with rather weak spiral sculpture. The two other specimens in the lot are H. tuberculata. The number of open tremata o f these specimens is somewhat higher than usual, but not beyond the expected and documented range o f the latter species (data not shown). Consequently a taxonomic separation is not Justified. H aliotis revelata Deshayes, 1863, was described fiom lie de Bourbon (= Reunion Island). The MNHN holds three lots o f type material o f this species with a total o f six specimens. One lot with a snigle specimen is labeled as “ex auteur,” another o f three specimens is labeled as “synlypes,” and a third lot with two specimens is called “type.” From the labeling o f the specimens, it is not clear which may be the holotype; there fore, all six specimens are presumed to be syntypes. Deshayes’ original illustrations (plate 36, figs. 1&2), were meticulously drawn, but cannot be matched to any o f the shells in the MNHN. In feet, the illustrations do not resemble any of the species found on Reunion Island, which may be attributable to excessive artistic liberty. The rather long description lacks much necessary detail. Only two quantitative indications are of some help. The length o f the largest specimen (61 mm) and the cited six open perfora tions are applicable to one specimen in the thfid lot, but this specimen has irregular growth on the columella that is clearly not illustrated or mentioned. The illustration shows only a weak growth line at the level o f the penultimate hole, which is also found in the second specimens o f the third lot. This specimen is here designated lectotype (Figures 1-8 - 1-9), and the remainder become paralectotypes. H aliotis revelata is iden tified as H. rugosa Lamarck, a species common at the type locality. The status o f H rugosa Lamarck itself is not resolved (see also note 12). 30 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis fa tu i has only recently been validly described (Geiger» 1999a), based on an unpublished name of the late Dr. H. Rebder (USNM) given to specimens that somewhat resemble H. varia Lirmaeus, 1758. Ubaldi (1993) listed H. fa tu i with the authority “Rheder [sûr], 1981 ?”. This listing is a nomen nudum (see also note 71). In the index that follows, the taxa with their status and/or the currently recognized valid species name are given. The taxa in bold are species currently recognized as valid species and subspecies with the original genus given if different from H aliotis s.s. The numbers in brackets refer to the notes that follow. Abbreviations: juv.: juvenile shell; s.L i sensu lato; ssp.: subspecies; syn.: synonym; var.: variety; form; ?: uncertain state ment (status, identifrcation). adriatica Nardo, 1847: nomen dubhan albicans Quoy & Gaimard, 1834: laevigata Donovan, 1808 alfiedensis Bartsch, 1915: speciosa Reeve, 1846 aliéna (Iredale, 1929) [in Sanhaliotis\i varia var. papulata Reeve, 1846 aleata Rôdmg, 1798: australis Gmelin, 1791 altem ata Sowerby, 1882: rugosa Lamarck, 1822 [31] ancile Reeve, 1846: juv. rubra Leach, 1814 [52] aquatilis Reeve, 1846: diversicolorResvCy 1846 asinina Linnaeus, 1758: valid [4] asinum Donovan, 1808: asinina Linnaeus, 1758 ossi mi Bs Dali, 1878: ssp. of kamt schat kana Jonas, 1845 [67] astricta Reeve, 1846: var. o f varia T.innaeus, 1758 aulaea Bartsch, 1940: kamtschatkana assim ilis Dall, 1878 [67] aurantium Sm ione, 1998 [1] 31 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. australà Gmelm, 1791: valid barboiai Foster, 1946: varia Linnaeus, 1758 [1] bistriata Gmelin, 1791: nomen dubitan, tuberculata Linnaeus, 1758, s.L [5, 14] bistriata Costa, 1829: homonym, van, syn. o f tuberculata var. lam ellosa Lamarck, 1822 bisundata Monterosato: tuberculata var. lamellosa Lamarck, 1822 [15] bonita Orcutt, 1900: cracherodii var. califom iensis Swainson, 1822 brazieri Angas, 1869: valid [34,35] cae/ora Roding, 1798: oviha Gmelin, 1758 californiana Valenciennes, 1831: rufescens Swainson, 1822 califi>miensis Swainson, 1822: var., syn. o f cracherodii Leach, 1814 canaliculata Fischer, 1807: parva Linnaeus, 1758 ? [19] canaliculata Lamarck, 1822: homonym, parva Linnaeus, 1758 canariensis Nordsieck, 1975: tuberculata coccinea Reeve, 1846 capensis Dunker, 1844: midae Linnaeus, 1758 carinata Swainson, 1822: parva Linnaeus, 1758 cingulata Rdding, 1798: parva Linnaeus, 1758 clathrata Lichtenstein, 1794: elegans Philippi, 1844 [25] clathrata Reeve, 1846: valid (homonym) [24,25] coccinea Reeve, 1846: ssp, of tuberculata Linnaeus, 1758 [17,18] coccoratUata Reeve, 1846: valid [49] concinna Reeve, 1846: varia Linnaeus, 1758 conicopora Péron, 1816: ssp. of r« 6ra.Leach, 1814 [53] coreanica Weinkauf^ 1883: nomen rtudum [6 8] corrugata Wood, 1828: valid [26] 32 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. costata Swainson, 1822: australis Gmelin, 1791 cracherodii Leach, 1814: valid crebrisculpta Sowerby, 1914: valid [25,36] crenata Swainson, 1822: scalaris Leach, 1814 ? crispata Gould, 1847: ssp. of virginea Gmelin, 1791 [60[ cruenta Reeve, 1846: ssp. o f pustulata Reeve, 1846 [10] curminghami Gray, 1826: rubra conicopora Péron, 1816 [27,53] (yclobates Péron, 1816: valid dalli Henderson, 1915: valid [2] decussata Philippi, 1850: marmorata Linnaeus, 1758 [7] dentata Jonas, 1846: ssp., var. of mariae Wood, 1828 diegoensis Orcutt, 1900: van o f corrugata Wood, 1828 [65] discus Reeve, 1846: valid [62] dissona (Iredale, 1929) [in Sanhaliotis\% valid [36,39] diversicolor Reeve, 1846: valid [37] dohm iana Danker, 1863: valid [40] ebingii Reeve, 1846: varia Linnaeus, 1758 [45] dubia Lamarck, 1822: nomen dubhan [69] echinata Sowerby, 1882: jacnensis Reeve, 1846 [31,41] elatior Pilsbry, 1890: var. o f midae Lmnaeus, 1758 elegans Philippi, 1844: valid [23,50] elevata Sowerby, 1882: squamata Reeve, 1846 [31,57] emmae Reeve, 1846: ssp. o f scalaris Leach, 1814 [56] ethologus (Iredale, 1927): hargravesi Cox, 1869 [34] excavata Lamarck, 1 822: cyclobates Péron, 1816 33 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. excisa Gray, 1856: unavailable, laevigata Donovan, 1808 [70] exigua Danker, 1877: valid [63{ expansaTeixaaàgCy 1954; cracherodii Leach, 1814 fa tu i Geiger, 1999 valid [71] ficfo rm islA evke, 1844: jpad/cea Donovan, 1808 fu lg en s Philippi, 1845: valid fu m b ris Reeve, 1846: squamata Reeve, 1846 [58] gemma Reeve, 1846: varia Linnaeus, 1758 [46] gf66a Philippi, 1846: v ô ^ e a Gmelin, 1791 [61] gigantea Chemnitz, 1788: unavailable, gigantea Gmelin, 1791 [59, 64] gigantea Gmelin, 1791: valid [64] gigas Rôding, 1798: gigantea Gmelin, 1791 glabra Chemnitz, 1788: unavailable, glabra Gmelin, 1791 [59] glabra Gmelin, 1791: valid glabra Swainson, 1822: homonym, laevigata Donovan, 1808 ? granti Pritchard & Gatliff, 1902: rubra conicopora Péron, 1816 [53] ffranulata Rdding, 1798: varia Linnaeus, 1758 grayana Sowerby, 1882: planata Sowerby, 1882 [31] gruneri Philippi, 1848: var., syn. o f diversicolor Reeve, 1846 guadalupensis Talmadge, 1964: var. o f fidgens Philippi, 1845 guineensis Gmelin, 1791: marmorata Linnaeus, 1758 hanleyana Sowerby, 1882: nomen dubhan [31 ,7 2 % hanleyi AncQr, 1881: jacnensis Reeve, 1846 [41] banaat Inc, 1953: ssp. o f t& cus Gmelm, 1791 hargravesi Cox, 1869: valid [34,35] 34 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. hattorii Bartsch, 1940: var. oîntfescens Swainson, 1822 holzneri Hemphill, 1907: var. o f cracherodii Leach, 1814 [66] howensis (Iredale, 1929) [in SanhaliotisY rubiginosa Reeve, 1846 [43] huttoni FflhoL 1880: ssp. of virginea Gmelin, 1791 [60] imperforata Gmelin, 1791: nomen dubium [66] imperforata Dali, 1919: homonym, cracherodii Leach, 1814 [66] improbula Iredale, 1924: syn., var. of rubra Leach, 1814 mcrra Reeve, 1846: tuberculata Lwaaeus, 1758 [16] rnferrwpfa Valenciennes, 1831: nomen dubium \73] rrft/tf Karsten, 1789: unavailable, Gmelin, 1791 [74] iris Gmelin, 1791: valid [54] jacnensis Reeve, 1846: valid [41] janus Reeve, 1846: tuberculata coccinea Reeve, 1846 [17] japonica Reeve, 1846: tuberculata Linnaeus, 1758 [16] jousseaum i Mabüle, 1888: pustulata Reeve, 1846 [11] kam tschatkana Jonas, 1845: valid [62,67] ^owsrzTurton, 1932: parva Linnaeus, 1758 laevigata Donovan, 1808: valid lamellosa Lamarck, 1822: var. o f tuberculata Linnaeus, 1758 [5,13] W /abrir Philippi, 1848: ovma Gmelm, 1791 [8] lauta Reeve, 1846: semiplicata Menke, 1843 [49] lucida Requien, 1848: tuberculata var. lamellosa Lamarck, 1822 lusus Finlay, 1927: var. o f cracherodii Leach, 1814 [66] maculata Küster, 1840: nomen dubium [18] maculosa Küster, 1840: nomen dubium [18] 35 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tm daka (Habe, 1977) ]^N o rd o tis\i valid [64] mariae Wood, 1828: valid [26] marmorata Linnaena, 1758; valid [4,5,6] marmorata Reeve, 1846: homonym, virginea Gmelin, 1791 melculus (Iredale, 1927) [in Marinaurisy. brazieri Angas, 1869 [35] midae Linnaeus, 1758: valid [04] modesta auct: nomen nudum ?, midae Linnaeus, 1758 [75] morioria Powell, 1938: ssp. of virginea Gmelin, 1791 [60] m ultiperforata Reeve, 1846: rugosa Lamarck, 1822 [32] naevosa Philippi, 1844: rubra Leach, 1814 [54] nebulata Reave, 1846: n/gosa Lamarck, 1822 [33] neglecta Philippi, 1848: stomatiaeformis Reeve, 1846 [5, 8,9] nodosa Philippi, 1845: corrugata Wood, 1828 ovina Gmelm, 1791: valid oweni Talmadge, 1966: var. o f corrugata Wood, 1828 papulata Reeve, 1846: var. o f varia Linnaeus, 1758 parma Valenciennes, 1831: nomen dubium [73] parva Lmnaeus, 1758: valid [4] parva Risso, 1826: homonym, tuberculata var. lamellosa Lamarck, 1822 pellucida von Salis, 1793: tuberculata var. lamellosa Lamarck, 1822 pertusa Reeve, 1846: rugosa Lamarck, 1822 [33] picta Rddmg, 1798: glabra Gmelin, 1791 planata Sowerby, 1882: valid [42] planilirata Reeve, 1846: fidgens Philippi, 1845 [47] plicata Karsten, 1789: unavailable, australis Gmelin, 1791 [74] 36 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. plicata Gmelin, 1791: nomen dubium ponderosa Aàsaos^ 1848: rufescens Swainson, 1822 pourtalesU Dali, 1881: valid [1 ,2 ,3{ pulcherrtm a Gmelin, 1791: valid pustulata Reeve, 1846: valid [12] pustulifera Pillsbry, 1890: varia Linnaeus, 1758 queketti Smith, 1910: valid reticulata Reeve, 1846: tuberculata Linnaeus, 1758 [16] revea Bartsch, 1940: fulgens, nomen nudum revelata Deshayes, 1863: rugosa Reeve, 1846 [12] roberti McLean, 1970: valid [2] roedingi Menke, 1844: squamosa Gray, 1826 [29] roei Gray, 1826: valid [27] rosacea Reeve, 1846: marmorata Linnaeus, 1758 rosea Orcutt, 1900: cracherodii Leach, 1814 rotundata Perry, 1811: nomen dubium [22] rubicunda Rôding, 1798: parva Linnaeus, 1758 [20] rubicunda (Montfort, 1810) [in Padollus]: homonym, parva Linnaeus, 1758 [20] rubiginosa Reevev 1846: valid [43] rubra Leach, 1814: valid [25,51] rufescens Swainson, 1822: valid rugosa Lam arck, 1822: valid [12] rugosa Reeve^ 1846: homonym, tuberadata Linnaeus, 1758 rugosoplicata Chemnitz, 1788: unavailable, australis Gmelin, 1791 [59] rugosoplicata Reeve, 1846: australis Gmelin, 1791 [59] 37 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. sanguinea H an l^, 1840: spadicea Donovan, 1808 [2IJ scalaris (Leach, 1814) ^ P a d o iïu s \i valid [56] scabricostataM & vkt, 1843: roef Gray, 1826 schroeteri Menke: nomen dubiian [76] scutulum Reeve, 1846: varia Linnaeus, 1758 ? [47,48] secemenda Monterosato: tuberculata van lamellosa Lamarck, 1822 sem iplicata Menke, 1843: valid sem istriata Reeve, 1846: varia Linnaeus, 1758 sepiculata Reeve, 1846: diversicolor Reeve, 1846 ?, tuberculata Linnaeus, 1758 ? sieboldiiReevCy 1846: gigantea GtnsUii, 1791 [64] sinuata Perry, 1811: spadicea Donovan, 1808 ? [22] sm ithsoni Bartsch, 1940: kamtschatkana assim ilis Dall, 1878 sorenseni B artsch, 1940: valid spadicea Donovan, 1808: valid [21] speciosa Reeve, 1846: valid splendens Reeve, 1846: julgens Philippi, 1845 splendidula Williamson, 1893: cracherodii Leach, 1814 squam ata Reeve, 1846: valid squamosa G ray, 1826: valid [27, 28] stom ariaeform is Reeve^ 1846: valid [39] striata Linnaeus, 1758: tuberadata van lamellosa Lamarck, 1822 [4,13] strigata Wemkauff, 1883: marmorata Linnaeus, 1758 Wemkauf^ 1883: vErgwea Gmelm, 1791 sidcosa Philippi, 1845: roei Gray, 1826 supertexta Lischke, 1870: van, syn. o f diversicolor Reeve, 1846 [38] 38 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tayloriana Reeve, 1846: van, syn. of diversicolor Reeve, 1846 tricostalis Lamarck, 1822 : scalaris Leach, 1814 tricostata Wood, 1828: scalaris Leach, 1814 tuberculata Linnaeus, 1758: valid [4,5,13] tubifera Lamarck, 1822: gigantea Gmelin, 1791 turveri Bartsch, 1940: var. o f fulgens Philippi, 1845 unilateralis Lam arck, 1822: valid [30) varia Lmnaeus, 1758: valid [4] varia Risso: homonym, tuberculata van lamellosa Lamarck, 1822 [4,44] venusta Adams & Reeve, 1848: clathrata Reeve, 1846 [24] victoriae Brazien nomen itudum ?, rubra Leach, 1814 [77] virginea Gmelin, 1791: valid [60] vù^gmea Reeve, 1846: homonym, mnmorara Linnaeus, 1758 viridis Reeve, 1846: varia Linnaeus, 1758 vixlirata (Cotton, 1943): rubra conicopora Péron, 1816 [53] vulgaris da Costa, 1778: tuberculata Linnaeus, 1758 walaUensis Steam s, 1898: valid whitehousei (Colman, 1959) \mSanhaliotis]: rubra Leach, 1814 [55] zealandica Reeve, 1846: coccinea Reeve, 1846 [17] 2K30C Reeve, 1846: g/ohra Gmelin, 1791 Notes 01. H aliotis barbotari Foster, 1945, has been a very controversial species. It was described form a single beach shell from the coast o f BraziL It has been hypothe sized that it is either a distmct species or a mislocated specimen o f either H. pour - 39 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. talesii Dali, 1881, from the Caribbean ^ o s , 1975) or the only Polynesian species H. pulcherrim a Gmelin, 1791 (Cox, 1962; Rios, 1985, 1994). H aliotis barboari has not been treated in the discussion o f Brazilian abalone by Simone (1998). Klappen- bach (1968) mentioned a living specimen taken off Brazil, refuting the claim the species represents specimens left by tourists (jcf. Cox, 1962). Specimens o f H povr - talesii have been indicated from the G ulf o f Mexico to as far south as Venezuela and Brazil (Henderson, 1915; Foster, 1946; Aguayo & Jaume, 1947; Harry, 1966; Guice, 1968; Klappenbach, 1968; Sarasua, 1968; Nijssen-Meyer, 1969; Titgen & Bright, 1985; Ode, 1986; Martinez & Ruiz, 1994). The more southern reports o f H povrtalesii including Klappenbach’s (1968) live specimens can be attributed to con fusion with H. aurantium (Simone, 1998). Inspection of the type of H. barbouri (MCZ 152469) revealed its true identity as a somewhat aberrant form o f H. varia Linnaeus, 1758. At a size of 22 mm it is a rather small specimen for the species. In general, small specimens o f abalone tend to be rounder than larger ones (cf. Stewart & Geiger, 1999: fig. 4). Hence, based on the overall shape of the shell one may be led to place the type o f H barbouri in the vicinity o f H pulcherrim a. However, the sculpture differs greatly. Most signifi cantly, in H. pulcherrim a a narrow spiral band adjacent to the row o f tremata and towards the suture is found, which is devoid o f oblique radial folds or other ele vated shell structures. A similar structural element is present in H. jacnensis Reeve, 1846 (of. Figure 1-16, note 41). In H. varia and the type o f K barbouri this bare space is not seen. The type o f H barbouri reminds one somewhat o f the type speci mens o fH . gemma Reeve, 1846 (= K varia: see note 46) in terms o f size, rotundity and sculptural elements. 40 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 02. H aliotis dalli Hoiderson, 1915, and H roberti McLean, 1970, are two smaU, insu lar species found of6hore 6om Central America at the Galapagos Islands {H dalli) and Cocos Island {H. roberti). The two species are fairly similar, but can be distin guished as indicated by McLean (1970). Radulae o f both species share a very nar row lateral tooth I and concentric rings on the cephalic tentacles. These characters are shared only with H. pourtalesU Dall, 1881, and H. aurantium from Brazil (Simone, 1998), but are not seen in any o f the other 30 abalone species studied so far (Geiger, pers. obs.). 03. The neotype o f H. pourtalesU Dall, I88I, is now broken in many pieces. The origi nal illustration o f the neotype (Henderson, I9I5: pis. 45-46) showed an intact sheU. 04. At the time when Linnaeus (1758) introduced the first seven haliotids {H. asinina, H. marmorata, H. midae, H. parva, H. striata, H. tuberculata, H. varia), the con cept o f type specimens was not yet established. The LSL holds specimens in the collection, and I agree with Talmadge (1977a) that most correspond well with the current concept o f the respective species; Talmadge (1977a) noted a single speci men o f H tuberculata, whereas I found six specimens with a note by S. P. Dance from 1963 also referring to six specimens. No specimens are currently designated as types. As indicated by Talmadge (1977a) there is no specimen o f H. parva in the LSL. As it is a improblematic species, a designation o f a neotype is not necessary aCZN Article 75b). 05. The H. tuberculata-ffoup contains the following main taxa: H. tuberculata Lin naeus, 1758, lam ellosa Lamarck, 1822, coccinea Reeve, 1846, bistriata Gmelin, 1791; this group may additionally contain H. marmorata Linnaeus, 1758 (cf. Tal madge, 1963a) (Figures I-I8 - I-I9) aadH. stom aticform is Reeve, 1846 (Figures I- 20 - I-2I). H aliotis tuberculata was the earliest to be named o f the well-known taxa 41 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. in the group, hence the group name is attributed to this species. All the species are found in Einope and northwest Africa (Mediterranean, Brittany to Sao Tomé and Gabon, Canary Islands, Azores). cDNA analysis o f the protein lysin (Lee & Vac- quier, 1995) has shown, that tuberculata and lamellosa from the Mediterranean are the same species. The tuberculata and the lamellosa forms are usually found within one population (Geiger, pers. obs.). The taxon lamellosa, therefore, refers to a vari ety o r ecomorph o f K tuberculata. The taxon coccinea was shown to be closely related to H. tuberculata (Lee & Vacquier, 1995). It differs form H. tuberculata in only four positions o f the 132 amino acid sequence o f lysin whereas most species differ in approximately 15-20 amino acid positions. Despite the small difference in amino acid sequence, the shells are readily distinguishable and coccinea is foimd only at the Canary Islands. The two conditions for a subspecifrc status for H tuber - culata coccinea are found: distmct character and specific geographic location. H ali - Otis bistriata seems to be a variation o f ff. tuberculata, sensu lato, based upon shell morphological observations (see note 14). H aliotis marmorata Linnaeus, however, seems to be distinct in terms o f shell morphology (Figures 1-18 - 1-19), but neither anatomical characters nor biochemical data are available. Its status in relation to H. tuberculata remams unresolved. 06. The specimens labeled H. marmorata Linnaeus, 1758, are conspecific with what is best known as H. rosacea Reeve, 1846, which has been synonymized with H. guineensis GtnsUa, 1791 (Talmadge, 1963b; Ubaldi, 1987) (Figures 1-18 - 1-19). Despite the possibüîty of exchange o f material (K. Way, pers. comm.), three lines o f evidence suggest that the specimens are likely to represent the species as described by Linnaeus: 1) Sowerby (1882) alrearfy mdicated the synonymy between H. m ar - morata Lmnaeus, H. rosacea, and H. gumeensis (explanation to plate 11, figs. 88, 42 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 89); 2) some old specimens o f this species labeled H. marmorata Linnaeus have been found in collections {e.g., DMNH 011036); 3) S. P. Dance stated on a label from 1963 that “from several undocumented shells I have selected one that matches specimens in B rit Mus. (N at Hist) labeled rosacea Reeve.’ This is almost cer tainly the example mentioned by Hanley as present.” Therefore, the correct name for the continental west African species is H. marmorata Linnaeus. Talmadge (1977a), in his discussion o f Linnean haliotids, erroneously synonymized H. mar - morata Reeve {non Linnaeus) with H. virginea Reeve {non Gmelin). H aliotis mar - m orata Reeve is clearly H. virginea Gmelin from New Zealand and not H. marmorata Linnaeus (= H. virginea Reeve) from west Africa. 07. H aliotis decussata Philippi, 1850, was described in Philippi’s (1847-1851) third volume, with the date o f the volume given as 1851. However, the pages with the description o f Haliotis are dated April 1850, which is the correct date. 08. Philippi referred to figure 4 for his H. latilabris Philippi, 1848, and figure 5 for H. neglecta Philippi, 1848; this is obviously an error and one should consult figures 5 and 4 for these species, respectively. Pickery (1991) indicated 1851 as publication date, but the species were already described by Philippi (1848:16). 09. Philippi’s (1847-1851) H. neglecta Philippi, 1848, is a junior synonym o f H. stoma - tiqform is Reeve, 1846, as recognized by Sowerby (1882:27): “... H. neglecta of Philippi... is undoubtedly identical with our present species [H. stom atiaeform isY (see also note 39). Ubaldi (1987) indicated this species [as H. neglecta] as being distinct, occurring on islands close to Sicily, M alta and Lampedusa, but without illustrating any specimens. Some specimens, which correspond very well to the fig ured specimen have been located: one in MNHN (Sicily and Palermo), four m HUT (6313a, b: Lampedusa and Giardini), one in DMNH (097371: near M alta Island), 43 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. and one in my personal collection (Malta: Figures 1-20 - 1-21); additional speci mens were obtained by Buzz Owen (pers. comm.) and were made available for examination. H aliotis tuberculata Linnaeus, 1758, with, the form lamellosa Lamarck, 1822, is the only native, well known, and highly variable species in the Mediter ranean. H aliotis stom atiaefbrm is might Just represent aberrant specimens o f H. tuberculata {cf. Weber, 1928). However, the specimens I have seen, including two preserved anim als, are very distinct and unlikely to be only a variation o f H. tuber - culata. I consider H. stomatiaeformis to be a valid species. Geiger & Owen (in prep.) will discuss this taxon in more detail. Geiger (1998a) erroneously synonymized H stom atiaeform is with H. squamata Reeve, 1846. Interestingly, a sunilar case is found in the Confdae. Conus ventricosus Gmelin, 1791, is the species predom inantly living throughout the Mediterranean. In the Sicily Channel area C. vayssieri Pallary, 1906, is found (Villa, 1985), but this species has also been regarded as a mere form o f C. ventricosus (Poppe & Goto, 1991). 10. H aliotis pustulata Reeve, 1846, migrated from the Red Sea into the Mediterranean Sea through the Suez Canal. It has been found along the Israeli coast (Talmadge, 1971; Fainzilber, 1984) and in Lybia (Giannuzzi-Savelli et aL, 1994). One alcohol- preserved specimen without the shell from Greece has an epipodium more or less identical to that o f K pustulata from the Red Sea. The epipodia of H. pustulata and H. tuberculata differ markedly. It is, therefore, possible that H. pustulata has advanced frnther into the eastern Mediterranean. The few reports in the literature o f H. pustulata in contrast to other species—e.g., Strombus decorus (Rôding, 1798) (see e.g., Fischer, 1993; Lindner, 1993)—may be due to the feet that the shells o f H. pustulata often are not strikmgly different from the native H. tttberculata. 44 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. IL The geographic provenance o f H, Jousseaumi Mabille, 1888, was not given in the original description (Mabille, 1888), but the label o f the specimen held in the MNHN cites the Red Sea (“M et Rouge”). The name has only been used once thereafter by Talmadge (1956), who correctly synonymized it with H. pustulata Reeve, 1846. 12- A lectotype for K revelata Deshayes, 1863, is here selected (see main body o f text. Figures 1-8, 1-9). The specimens are conspecific with what is better known as H. rugosa Lamarck, 1822. Herbert (1990) speculated about the synonymy between H. pustulata Reeve, 1846, and H. rugosa^ Miich had already been indicated indirectly by Wagner & Abbott (1978) by the synonymization o f K alternata Sowerby, 1882, with K pustulata I have not seen any material that has intermediate characters and question the synonymy between the two taxa. The taxa cruenta Reeve, 1846, and pustulata may eventually be treated as subspecies o f K rugosa, pending further clarification based on animals o f the these taxa, but I tentatively consider them as distinct species (cf. Herbert, 1990; Geiger, 1996). 13. H aliotis striata Linnaeus, 1758, belongs in the Æ tuberculata Linnaeus, 1758 group (see note 5). I agree with Talmadge (1977a) that it corresponds with what is well known as K tuberculata var. lamellosa Lamarck, 1822 (Figures 1-28 - 1-29). If Æ tuberculata is a synonym o f H. lamellosa, then H. striata is also a synonym o f H. tuberculata H aliotis tuberculata is preferred over H striata as the valid name for the species because it is the established name. However, if a taxonomic distinction between tuberculata and lam ellosa is desired, then striata has priority over lam el - losa. Potentially, a suppression o f striata would be advisable as already suggested by Pilsbry (1890:87). 14. Weber (1928) mentioned the extensive variability o f H. bistriata Gmelin, 1791, fiom Tenerife, Canary Islands; some specimens correspond with its origmal descrÿ- 45 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. don o f K bistriata, some are typical K tuberculata Lmnaeus, 1758, and concluded that the two taxa are most likely conspecific. He did not mention H. coccinea Reeve, 1846, and possibly was synonymizing too much under one taxon (see also note 9). The taxon bistriata is mostly encountered in the old literature, and a few specimens firom old collections labeled K bistriata were be foimd in the BMNH and ANSP. Most o f the specimens that show some resemblance to old illustrations {e.g.. Reeve, 1846: pL II, fig. 33) originated fiom the Canary Islands and the Azores. The speci mens are characterized by strong radial groves and a flared aperture. A separation of K bistriata firom other members of the H. tuberculata group (see note 5) cannot be accepted. The localities, illustrations, and specimens labeled as H. bistriata do not show any coherent pattern. H aliotis bistriata should be regarded as a growth form o f either H tuberculata coccinea for specimens fiom the Canary Islands and the Azores, or o f H. tuberculata or H. marmorata in the case o f specimens fiom the East Atlantic mainland. As bistriata most likely refers to three different species- level taxa, 1 prefer to treat it as a nomen dubium, which further provides stability to nomenclature, because the populations fiom the Canary Islands are well known under the epithet coccinea. 15. H aliotis bisundata Monterosato is listed as a variety o f H. tuberculata Linnaeus, 1758, in Priolo (1948) and Ghisotti (1964), but cannot be traced. 16. H aliotis incisa Reeve, 1846, H. japonica Reeve, 1846, and H. reticulata Reeve, 1846, are all reported fiom Japan. This type locality is doubtful and the type speci mens look exactly like the Mediterranean H. tuberculata Linnaeus, 1758, as also indicated by Reeve (1846) and discussed in Dunker (1882), Weinkauff (1883), McLean (1966), and Kaicher (1981: card no. 2882). Weinkauff (1883:59) neverthe less reported fiir H. japonica similar specimens “without any doubf* fiom Ji^an, 46 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. but not giving the locality more credibility, hi modem times no similar specimens have been mentioned from the well studied area of Japan. H aliotis japonica and K tuberculata var. lamellosa Lamarck, 1822, have also been synonymized erroneously (Pilsbry, 1890:87) with H aquatilis Reeve, 1846, this latter a synonym of H. diver - sicolor Reeve, 1846. 17. The epithet coccinea Reeve, 1846, is assigned as a subspecies o f Æ tuberculata Linnaeus, 1758. This is here established on the basis of cDNA sequencing data (Lee & Vacquier, 1995), distinct shell characters, and its isolated geographic occurrence on the Canary Islands (see also note 5). Haliotis Janus Reeve, 1846, is a color form o f H tuberculata coccinea (Talmadge, 1958a). 18. H aliotis macidata KOster, 1840, and K maculosa Küster, 1840 (spelling variations ?) are mentioned by Sowerby (1882:36, pi. 9) and Weinkauff (1883:83) as senior synonyms o f H coccinea Reeve, 1846. The name is based on figure 137 m Martini & Chemnitz (1769) (non-binominal), which shows very clearly an H tuberculata coccinea (see note 17). The original source could not be traced, and Küster’s taxa were not listed in Sherbom (1922; 1932). According to Weinkauff, KOster’ s name has priority over Reeve’s, an opinion not followed here, because the citation o f KQster’s work cannot be located. 19. The type specimen o f H canaliculata Fischer, 1807, was lost prior to 1872 (Ivanov & Kantor, 1991). Fischer (1807) indicated the presence of a deep, spfral canal and a brownish shell with many spiral cords in this small species. These characters point toward H parva Linnaeus, 1758, a species to vdiich he did not refer. The synonymy is tentative. 20. H aliotis rubicwtda (Montfort, 1810) has also been attributed erroneously to Gray (1826) (see also note 27). Gray clearly referred to Montfort. M ontfort used this 47 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. species as type species for his genus PadoUus and did not refer to any other author, which makes it likely to be a new taxon introduced by him and not K rubicunda Rôding, 1798 (synonym o f K parva Linnaeus, 1758). H aliotis rubicunda (Mont fort) was inflected to the masculine by Montfort for his new genus Padollus (see Knight era/., I960; Pickery, 1991; also Schremp, 1981:1125). The type locality is the “Anican coasts.” The key feature of the species is the spiral ridge on the shell; a character seen only in H. parva among the Aftican species (Iredale, 1927:334). Padollus rubicundus Mont&rt has also been put into synonymy with H scalaris (Leach, 1814) (see also note 56), a species that, however, occurs only in Australian waters. Because H. rubicunda Rôding and P . rubicundus Montfort refer to the same species, they must belong to the same genus, for which the species have to take the correct generic ending o f the adjectival species name, causing P . rubicundus Mont fort either way to become a secondary homonym. Note, that although Padollus is based on an invalid name, it does not make Padollus itself invalid. 21. H aliotis sanguinea Hanley, 1840, was reported to have been described in 1808 (cf. Muller, 1986), in the same year as the senior synonym H. spadicea Donovan, 1808. Muller (1986) discussed the date o f publication o f H sanguinea at length and con cluded it was actually described in 1840. 22. The descriptions o f H rotundata Perry, 1811, and H sinuata Perry, 1811, given by Perry (1811: pi. 52) are very brief and general, and his figures are rather stylistic. Pilsbry (1890) listed both as unidentified species. The undulation of the apertural m argin as well as the general shape of the shell o f H sinuata is reminiscent o f K spadicea Donovan, 1808. 23. H aliotis clathrata Lichtenstein, 1794, which has so far been considered a nomen dubium (V%gner & Abbott, 1978), would cause K clathrata Reeve^ 1846 to be an 48 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. invalid, junior, primary homonym. H aliotis clathrata Lichtenstein is discussed by Geiger (1998b) and identified as H. elegans Philippi, 1844. Geiger & Stewart (1998) have petitioned the International Commission on Zoological Nomenclature to sup press H clathrata Lichtenstein so as to preserve H. elegans and H. clathrata Reeve. This position, pending decision by the Commission, is adopted here (ICZN Article 80). 24. H. venusta Adams & Reeve, 1848, has so far been considered distinct from H. clathrata Reeve, 1846, due to the absence of radial lamellae on the type specimens. Several characters can be observed on both series o f type specimens o f K clathrata Reeve and H. venusta (1 holotype, 2 paratypes each in BMNH): I) a pattern of approximately 4 o f 5 faint growth lines followed by one stronger one; 2) tremata only slightly oval, but rather large in proportion to the shell; 3) all shells o f orange and white color, despite some fading; 4) in both series, some specimens with spire fiilly visible in ventral position, some only partially; 5) numerous spiral cords; 6) usually three or four o f these cords stronger than others; stronger cords regularly spaced between suture and row o f tremata; 7) the Indo-Pacific type localities for both taxa: Baclayon, Island o f Bohol, Philippines for K clathrata Reeve, and East ern Seas for H. venusta. The holotypes o f K clathrata Reeve is distinguished fiom K venusta by the discrete, numerous radial lamellae. The lamellae o f H. clathrata Reeve are formed along the stronger growth lines, £.e., every four or five faint growth Imes. The lamellae of K clathrata Reeve represent elevated, strong growth lines in H. venusta, hence the two structures are homologous. Radial lamellae are known to appear randomly in populations o f a smgle species o f H aliotis. The best documented case is that o f H tuberculata Lmnaeus, 1758, firom the Mediterranean, where the lamellate form is well-known as H lamellosa Lamarck, 1822, H tuberculata lamel - 49 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. losQ y or H, tuberculata form, lamellosa (Ubaldi, 1987; Poppe & Goto, 1991). In a large seriœ o f over 400 specimens, all intermediate fonns could be found within any given population (Geiger, unpubl. data). The two taxa have been shown to have an identical karyotype (Colombera & Tagliaferri, 1983) and an identical cDNA sequence o f the acrosomal sperm protein lysin (Lee & Vacqtiier, 1995). Although not as much material is available tor H, clathrata Reeve and IL venusta as for the Mediterranean Æ tuberculata, I am convinced that a similar range of variation can be expected for the Indo-Pacific H. clathrata and H, venusta. I, therefore, syn- onymize H. venusta under H. clathrata Reeve. 25. H aliotis crebrisculpta Sowerby, 1914, has been synonymized with H. clathrata Reeve, 1846 (Kuroda & Habe, 1952). It has also been listed as a subspecies o f K rubra Leach, 1814 (Talmadge, 1957b). Both opinions are rejected; the validity o f the species is discussed in Stewart & Geiger (1999) and the reader is referred to this work for an in-depth treatment (see also note 36). 26. H aliotis corrugata Wood, 1828, has also been attributed to Gray. However, Wood (1828) published a figure with the name o f K corrugata in the supplement to his Index Testaceologicus. Gray apparently supplied some o f the materfel, on which the figures were based (p. iiv o f Supplement), but Wood actually published the figure; hence, he is the author o f the taxon. 27. Serious confusion about the date o f publication o f Gray’s (1826) work is found in the literature. The publication dates mdicated are 1826 and 1827. The w ork was edited in two volumes, o f which volume I is dated 1827, but one copy o f volume 2 that I have seen is dated 1826 and another copy is dated 1827. Generally, 1826 is accepted as the correct date, because it is the earliest credibly supported date o f publication. 50 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 28. Haliotis squamosa Gray, 1826, was erroneously described from Australia, but actu ally occurs in a restricted area in southern hfedagascar. The species was rediscov ered in Madagascar by Dautzenberg (1932) and again by Stewart (1984). 29. Haliotis roedingi Menke, 1844, was described by Menke (1844:97) in “2 neue Hali - Otis Arten, beschrieben vom Herausgeber” (2 new species of H aliotis, described by the editor). According to Menke, the species was already known and named by Chemnitz; Menke gave the name “ 'H aliotis Roedingi, Chemn.” It is clear that the species was known to Chemnitz, but was described by Menke. I agree with Stewart (1984) that Menke, 1844, is the author, and that H roedingi is a synonym of H squamosa Gray, 1826, based on the description as well as the locality (jrf. Pilsbry, 1890:112, note 28). 30. Haliotis unilateralis Lamarck, 1822, has been much disputed (Figures 1-32 - 1-33). Geiger (1996) designated a neotype and discussed the taxon in detail. Menke (1830:88) listed this species. The credibility o f the mention in Casto de Elera (1896) in his catalog o f shells firom the Philippines is very low. He also mentioned species known at the tune not to be found on the Philippine Islands. The best example is H mariae Wood, 1828, an endemic species to Oman. 31. Sowerby’s abalone taxa from the Thesaurus Conchyliorum have been dated 1883 (Abbott & Dance, 1983) or 1887 (Pickery, 1991). The individual volumes were issued in a number o f parts ( c f British Museum (Natural History), 1915). Volume 5 was published between 1882 and 1887. Haliotids appeared in the first part of Vol ume 5 in 1882, which is the correct date for these taxa. It is not a printing date as opposed to a publication date, as most taxa are listed in the Zoological Record of 1882 (Martens, 1882). 51 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 32. A lectotype for H m ultiperforata Reeve, 1846, is here selected (see main body o f text. Figures 1-6 - 1-7). The synonymy with K spadicea Donovan, 1808, (Wagner & Abbott, 1978), is certainly wrong. 33. H aliotis pertusa Bartsch, 1915, and H nebulata Turton, 1932, are indicated in Barnard (1963) as synonyms o f H. spadicea Donovan, 1 8 0 8 .1 assume that Barnard intended to mdicate K pertusa Reeve, 1846, sensu Bartsch (1915) and H. nebulata Reeve, 1846, sensu Turton (1932). The types o f K pertusa and H nebulata are clearly referable to H rugosa Lamarck, 1822 (see also note 12). 34. H aliotis hargravesi Cox, 1869, and H. ethologus (Iredale,-1927) are uncommon to rare in southern Queensland and northern New South Wales. Whitehead (1981:5) specifically noted for H. hargravesi that “known localities are as for Ü ethologus” The only characters distinguishing these two taxa are the numbers of spiral ribs and their elevations. However, these characters seem to be rather variable, pointing to extensive intraspecific variability and indicating presence o f a smgle polymorphic species. The two taxa represent slight variations within the morphological range when a large enough sample is examined. 35. The type o f H m elculus (Iredale, 1927) is severely chipped, has very elevated trem ata and hardly any spiral ribs: is reminiscent o f H. brazieri Angas, 1869. Wil son (1993) figured a specimen with many thin spiral threads as H melculus. 1 regard it as a specim en in the H. hargravesi Cox, 1869—brazieri continuum (see also note 34 and Hybrids section above). 36. For H crebrisculpta Sowerby, 1914, three tyntypes are known in the BMNH, the NMWi and the USNM. The specimen m the BMNH is very similar to that figured in Sowerby (1914) and has been has been designated as the lectotype (Stew art & Geiger, 1999). The specimens in the NMW and the USNM represent H. clathrata 52 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ; Y » :.■-•’ - n ^ T î ' ; ; I ■ ■ Figures 1-10 - 1-15. Shells o f uncommonly illustrated abalone I. H , dissona Qhedale, 1929). 33 mm. R. Pickery collection. New Caledonia. 1-12 - 1-13. H , diversicolor Reeve, 1846.30 mm. ANS? 319655. Bali Beach HoteL 1-14 -1-15. H. exigua Dunker, 1877.21 mm. NMW 1955.158.2133. Japan. 53 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures I-I6 - 1-2L Shells o f uncommonly illustrated abalone IL 1-16 - 1-17. K ja c - nensis Reeve, 1846.12 mm. R. Pickery collection. No location data. 1-18 - 1-19. K marmorata Linnaeus, 1758. 63 mm. LSL. Mare qfricanus^ 1-20 - 1-21. H. stomatiae - form is Reeve, 1846.29 mm. Collection Geiger AAB 51a. Malta Island. 54 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. J ^ 1 & ■ ■■ n . ,1^ ' ; - M .Y 'i m m Im m » * - . . X » . \ - : Figures 1-22 - 1-27. Shells o f uncommonly illustrated abalone UL 1-22 - 1-23. H. planata Sowerby, 1882.1-22. 39 mm. NMW 1955.158.2124. Philippines. 1-23.33 mm. NMW 1955.158.2125. Guadalcanal. 1-24 -1-25. K qiieketti Smith, 1910.33 mm. NMW 1955.1582129. Pondoland. 1-26 - 1-27. K rubigm osa Reeve, 1846.25 mm. USNM 791422. Lord Howe Island. 55 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 1-28 - 1-33. Shells o f uncommonly illustrated abalone IV. 1-28 - 1-29. H. strf - ata Linnaeus. 1758. 56 mm. One o f the six specimens in LSL. Mare europaeus. 1-30 - 1-31. H, speciosa Reeve, 1846.31 mm. NMW 1955.158.2126. Algoa Bay. 1-32 - 1-33. K unilateralis Lamarck, 1822. 1-32.23 mm. Collection S. Singer. Red Sea, Gulf o f Aquaba. 1-33.24 mm. Collection Geiger AAB 48a. Elat, Red Sea. 56 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reeve, 1846 (see notes 24,25). H aliotis dissona (Iredale, 1929) is here considered a valid species {cf. note 39; Figures 1-10 - 1-11) although Whitehead (1981) listed it as a synonym o f H. crebriscidpta. H aliotis dissona has also been synonymized by Wagner & Abbott (1978) with H diversicolor Reeve, 1846, and its form s (Figures 1-12 - 1-13), an opinion I strongly oppose. The characters common to H dissona and H. diversicolor (more or less smooth shell with spiral ridges) are due to the type specimen of H. dissona being badly worn and rather small. The distinct elongated shape o f the shell of H. dissona with rather deep spiral ridges and grooves in fresh specimens sets it apart from H. diversicolor. The illustration o f Iredale (1929) exag gerated the sculpture o f the shell to a great extent; it may be considered a recon structive drawing o f a fresh shell from a worn specimen. 37. H aliotis diversicolor Reeve, 1846, is well known from the temperate northwestern Pacifrc, from central to southern Japan (Lindberg, 1992) and somewhat further south. The species has now been found in a small number o f independent lots from Bali and New Caledonia, but only from localized upwelling areas (Figures 1-12 - 1-13). Upwelling areas are thought to exist on many of the surrounding islands that may provide habitat for this temperate species (S. A. Shepherd, pers. comm.). The shells are more elongated in general shape than typical ones and are as highly arched as shells o f K squam ata. Additionally, they are mostly o f uniform, dark sepia col oration, which may be interpreted as a case o f melanism. Melanistic shells are well known in some gastropods such as in the Cypraeidae, and New Caledonia is known for a higher frequency o f such dark-colored specimens. Whether these tropical pop ulations o f K diversicolor represent réfugia or have only been colonized in geolog ically recent tune is not known. A taxonomic separation seems inappropriate. 57 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 38. Two spelling variations are found in the literature: H supertexta Lischke, 1870, and H. supratexta. The original spelling was Hi supertexta. 39. Talmadge (1962) treated H. stom atiaeform is Reeve, 1846, as a subspecies o f H. varia Linnaeus, 1758. 1 disagree with his conclusion after inspection o f the type material: H. stomatiaeformis is the valid name for one Mediterranean species (see note 9). Specimens normally identified as H. stomatiaeformis are usually referable to H. dissona (Iredale, 1929) {cf. note 36). 40. H aliotis dohmiana Dunker, 1863, is a little known species with distinct affinities to H varia Linnaeus, 1758, as already indicated by Dunker (1870:7). The epipodium has distinct characteristics (Geiger, pers. obs.). 4L H aliotis hanleyi Ancey, 1881, and H. jacnensis Reeve, 1846, have been considered to be distinct species (e.g., Talmadge, 1963a; Kaicher, 1981). The distinguishing characters were thought to be the general shape of the shell and the extent to which the shell is corded. These characters are highly variable and the two taxa represent slight variations within the morphological spectrum o f one biological species. The geographical distribution o f the two taxa is congruenL The smooth, dorsal part of the shell proximal to the row o f tremata is a synapomorphy with H. pulcherrim a Gmelin, 1791 (see note I) and a diagnostic character of the species (Figures 1-16, 1-17). 42. Sowerby indicated Carpenter as authority for H. planata Sowerby, 1882. Weinkauff (1883:76) could not find the source; he mdicated ^ubVT (Latin for ‘where?’). Today Sowerby is usually credited with the authorship o f the taxon. H aliotis planata is occasionally confused with H. varia Linnaeus, 1758.1 agree on the basis o f the epipodia with Talmadge (1963a) that the two species can be separated. 58 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 43. H aliotis rubiginosa Reeve, 1846, has most often been used as a synonym, form, or subspecies o f H varia Linnaeus, 1758 {e.g., Kaicber, 1981). H aliotis howensis (Iredale, 1929) was described ftom and is endemic to Lord Howe Island. Old speci mens labeled as H rubiginosa from Lord Howe Island were located in the BMNH (K. Stewart, pers. comm.) and in the HUJ (12557). The types of the two species are bigbly similar, and I agree with K. Stewart that the two names refer to the same species. H aliotis rubiginosa bas priority over H howensis, despite the latter being better known (Figures 1-26 - 1-27). 44. The type locality o f H varia Linnaeus, 1758, is Philippine Islands as designated by Iredale (1910). 45. It is unclear whether H dringii Reeve, 1846, represents a distinct species or a form o f H varia Linnaeus, 1758. As H varia is such a variable species, I am inclined to synonymize H dringii under H. varia. Many specimens in collections identified as H. cùingii are actually K jacnensis Reeve, 1846 (Figures 1-16 - 1-17). 46. Haliotis gemma Reeve, 1846, is quite certainly a color form of H varia Linnaeus, 1758, as seen from the series of four type specimens in the BMNH, o f which one specimen is marked “type ” The synonymization is not entirely certain as the shells are rather small; juvenile abalone are notoriously difBcult to identify, and H. varia is an extremely variable species. The selection o f the specimen labeled “type” is somewhat doubtful because the measurements o f the shells and the indications in the description o f the species do not match. The illustration of H. gemma is “magni fied double” and the illustration is 19.5 mm long; hence, the corresponding speci men should be a little less than 10 mm long. The specimen labeled “type” measures 20.5 mm, which is twice the size o f the original specimen. It is believed that the ref- 59 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. erence to “magnified double” in the original description was incorrect. Kaicher (1981) illustrated another, unidentifiable specimen as H. gemma. 4 7 .1 found no type specimens for H. planilirata Reeve, 1846, or H. scutulum Reeve, 1846, in the BMNH. Talmadge (1964) indicated a type o f H. planilirata was in the BMNH. 48. From the description “waved, here and there larger.” the coloration “olive brown ... dotted and spotted with green,” and the irregular spiral ridges with occasional thick enings to be seen in the figure 64 o f Reeve (1846), I tentatively identify H. scutu - lum Reeve, 1846, as H. varia Linnaeus, 1758 (cf. note 47); 49. H aliotis lauta Reeve, 1846, was tentatively placed in the synonymy o f H coccora - diata Reeve, 1846, by Talmadge (1960). After inspection o f the type specimens o f both species in the BMNH, it is clear that the two taxa are not synonymous. 1 agree with other authors that H. lauta represents H. sem iplicata Menke, 1843. 50. The authorship o f H. elegans has also been indicated as “Koch in Philippi” (e.g.. Wells & Bryce, 1985:34). The date printed on the description page o f H elegans is June 1844. This is the correct date. Philippi (1842-1845) indicated Koch as a refer ence; however, it is apparent that Philippi actually wrote the description and that only a short note was taken directly from Koch. Accordingly, 1 consider Philippi, 1844, the author. 51. The taxon as described by Leach (1814: pL 23) is H. rubsLr The correct inflected spelling for this adjectival species epithet, however, is H. rubm - Both spellmgs are found in the literature. 52. H aliotis andle Reeve, 1846, has been shown to be a juvenile H. rubra Leach, 1814 (Geiger, 1996; Stewart & Geiger, 1999), and not a green form o f K pustulata Reeve, 60 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1846 {cf. Talmadge, 1956), or H Japonica Reeve, 1846 {cf, Wagner & Abbott, 1978) (note 16). 53. H aliotis conicopora Péron, 1816, and H. rubra Leach, 1814, bave been shown to be closely related on the basis o f allozyme frequency data (Brown, 1993: 430): "''Halt - otis conicopora clustered with the three populations o f H rubra^ with a maximum D o f 0.018. Similar genetic distances were found between all conspecific popula tions studied (0.003 in H. roei and 0.014 in H. laevigata)” Further on pages 430- 431: "'Haliotis conicopora appears to represent an allopatric population (Western Australian) o f H rubra, as originally suggested by Shepherd (1975) and noted by Brown & Murray (1992a).” However, the two taxa can readily be distinguished by the shell thickness (dependent upon wave energy o f the habitat), coloration (a poten tial function o f food availability), and the presence of strong spfral cords (somewhat variable in both taxa) on a bumpy dorsal shell surface in H rubra. Additionally, H. rubra is foimd in southeastern Australia, whereas H. conicorpora occurs in south western Australia. The genetic data suggest conspecifity, but the shell and the dis junct geographical distribution mdicate distinct taxa. Thus, there is some justificatioa for subspecies recognition o f conicopora under H, rubra, although this remains to be resolved. 54. H aliotis iris is credited either to Martyn (1784: cf. Sinclair, 1963) or to Gmelin (1791: cf. Pickery, 1991; Lindberg, 1992). H aliotis pulcherrim a has mostly been attributed to Gmelin accept by Cotton (1943), but H. naevosa exclusively to Mar- ^ n . All three taxa are figuied and named in Linnean binommal nomenclature by MartyiL However, the work o f Martyn has been invalidated by ICZN opinion 456 (ICZN, 1957). Martyn’ s taxa are, therefore, not available and are now credited to the author who thereafter mentioned the species for the frrst time, which is Gmelin 61 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (1791) for ü iris, and Philippi (1842-1845:147) for Hi naevosa. Philipprs Volume I is dated 1845, but the page with the description o f H. naevosa is dated “Juli 1844;” therefore, the latter is the correct date for H naevosa. 55. Only a single specimen, i.e., the holotype, o f H. whitehousei (Colman, 1959) is known. It is considered by P. Colman (pers. comm.), who described the taxon, as a terato logical specimen o f H. rubra Leach, 1814. Despite regular collecting at the type locality, no second specimen has ever been found (P. Colman, pers. comm.). 56. Intermediate specimens o f H. scalaris (Leach, 1814) and H emmae Reeve, 1846, are well known from western South Australia, indicating a close relationship between the two taxa (Shepherd, 1973; Wilson, 1993). The variation in the speci mens seems to be caused by the interplay o f environmental factors such as wave energy, depth, and temperature (S. A. Shepherd, pers. comm). As the morphological gradient between the two taxa occurs only in a small geographical area, with the majority o f the distribution showing only one o f the morphologies, subspecific sta tus o f the taxon emmae under H. scalaris is indicated. 57. The type specimen o f H. elevata Sowerby, 1882, has not been found. The specimen was not figured in the standard dorsal and ventral positions, but was tilted. How ever, it is evident firom the dorsal sculpture and the excentric spfre that a small spec imen o f H squamata Reeve, 1846, was illustrated. According to Sowerby ( 1882:27), H. elevata differs firom H. stom atiaeform is Reeve, 1846 (see note 9), by having strong, scaly ridges, a character rather typical for juvenile specimens of H. squa - mata (Talmadge, 1955). 58. H aliotis fim ebris Reeve, 1846, was synonymized by Hedley (1914) with H diversi - color Reeve, 1846, and K tcQfloriana Reeve, 1846. My inspection o f the type mate- 62 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. rial in the BMNH shows that K fim ebris is clearly a synonym o f K squamata Reeve, 1846 (see Pilsbry, 1890:92). 59. H aliotis gigantea Chemnitz, 1788, H. glabra Chemnitz, 1788, and H, rugosoplicata Chemnitz, 1788, are names that stand only from the two first words o f the descrip tions and, therefore, cannot be interpreted as binominal names: the work and the taxa therein are not available (ICZN, 1954). The names were validated by Gmelin (1791: Æ gigantea, H glabra) and Reeve (1846: H rugosoplicata). 60. H aliotis virginea Gmelin, 1791, has been divided into four, geographically sepa rated subspecies: H. virginea virginea, H virginea crispata Gould, 1847, H. virgina huttoni Filhol, 1880, and H. virginea morioria Powell, 1938 (Kaicher, 1981; Ubaldi, 1986). Conflicting opinions are expressed as to whether these are valid subspecies (Powell, 1979; Ubaldi, 1986), or represent variations caused by differences in water temperature (Talmadge, 1957a). No good data are available to support either of the hypotheses; I retain usage o f subspecific taxa. 61. H. gibba Philippi, 1846, is given as a synonym o f H virginea Gmelin, 1791, by Suter (1913), supported by the figures in Reeve (1846) and Weinkauff (1883). 62. Old specimens o f H discus hannai Ino, 1952, from Japan are usually identified on the original label as the North American H. kamtschatkana Jonas, 1845, to which it is strikingly similar. H aliotis discus hannai and H k. kamtschatkana differ at the species level (Owen et al., 1971; Brown, 1993; Lee & Vacquier, 1995). 63. H aliotis exigua Dunker, 1863, is usually encoutered in material from the Ryukyu Archipelago (Figures 1-14 - 1-15). It has morphological af&iities to both H. varia Linnaeus, 1758, and H diversicolor Reeve, 1846. Pilsbry (1895) regarded H. exigua as a juvenile o f H. diversicolor. The fiat shells are rather nondescript, have an irreg ular sculpture, and are usually o f a dark, mudtfy coloration. Whether these speci 63 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. mens deserve recognition at the species level is unclear. Inspection o f preserved material would, certainly help to resolve the status of this species. 64. The type specimen o f H. sieboldii Reeve, 1846, is an aberrant specimen, as indi cated g.g., by Dunker (1882:148): “Haec species mihi est valde d u b ia,... credam eam nil representare nisi confonnationem Haliotis giganteae monstruosam.” (This species is very doubtful to m e,... 1 believe it does represent nothing but a grotesque form of H aliotis gigantea). The types o f H sieboldii and H. gigantea Gmelin, 1791, refer to the same species (Habe, 1983). The taxon sieboldii had long been used for a further species eventually named H. madaka (Habe, 1977). The two species can be distinguished as follows. In H. gigantea a line drawn through the apex o f the shell and the last perforation results in very unequal areas o f the shell, whereas such a line drawn in H. madaka divides the surface area o f the shell approximately in half. Haliotis gigantea Menke was mentioned in Wemkauff (1883:25) as published in Menke (1843), but Sherbom (1922) considered 1830 to be the year o f publication of this taxon. Menke (1830:87; 1843:31) referred to H. gigantea Chemnitz, 1788, an unavailable name first mentioned thereafter by Gmelin (1791:3691), who listed this species from Australia (^Habitat rarissima a d novam Holandiam” [lives very rarely in Australia]). However, the type specimen o f H gigantea Gmelin, 1791, figured in Habe (1983) clearly shows the well-known northwest Pacific species. Due to the erroneous type locality, H. gigantea has been incorrectly synonymized with the common, commercially exploited H. rubra Leach, 1814, from southern Australia (e.g., Menke, 1843). 65. The type specimen o f H. diegoensis Orcutt, 1900, is a monstrosity induced by bor ing organism s, most likely sabellid polychaetes (see also Oakes & Fields, 1993). 64 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 6 6 . Adult specimens with missing tremata have been reported occasionally in the litera ture. These represent deformations (see Leighton, I960), rather than valid taxa. H aliotis im perforata Gmelin, 1791, may belong in this category or may be a misidentified, non-haliotid species. In the case of K im perforata Dali, 1919 (non Gmelin, 1791), H lusits Finlay, 1927, was proposed as a nomen novum. Finlay (1927), however, had overlooked H. cracherodii holzneri Hemphill, 1907, which has priority and precludes the establishment o f an nomen novum. 67. H aliotis assim ilis Dali, 1878, is a subspecies o f H. kamtschatkana Jonas, 1845, as discussed by McLean (1966). 6 8 . H aliotis coreanica Weinkauff, 1883, is mentioned in Habe & BCosuge (1964) and Habe (1983) as a synonym o f H. gigantea Gmelin, 1791. This synonymy is based, however, on a misunderstanding o f the German text o f Weinkauff by these two authors. Weinkauff (1883:27-28) wrote (translated from German): “From Nfc Paetel 1 received on a loan basis a species, which was labeled H. coreanica A. Adams, which, however, could not be separated from H. discus, which would be considered a further variety. However, 1 cannot find where this species is described.” Although the second and the third statements contradict one another to a certain extent, my interpretation o f the text is that Weinkauff considered the specimen labeled H. core - anica A. Adams to be the same as his H gigantea Var. 1 = if. discus Reeve, 1846. This particular specimen was not figured by Weinkauff ruling out the possibility that an illustration could serve the purpose o f a valid description. The name is a nomen nudum. 69. The type specimen o f H. dubia Lamarck, 1822, is not in MHNG or the MNHN. Most o f Lamarck's types are otherwise deposited in the MHNG (cf. Mermod & Binder, 1963). 65 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 70. H aliotis excisa Gray, 1856, is not listed in Sowerby (1882) or Weinkauff (1883), but is indicated in Abbott & Dance (1983) and in Pickery (1991) without indication o f page number under Gray (1826) and not Gray (1856). Schism otis excisa was men tioned in Gray (1856:148), but as a hypothetical name for the teratological speci mens with slit tremata he had at hand: “When 1 first saw the shell, I was inclined to regard it as a monstrosity; but when I considered the uniformity... in the specimens ..., I thought that it might be the type of a new form, for which Schismotis excisa would be a good nam e.... I am inclined to believe that the slit in the specimens i s ... caused by the eroded and evidently diseased state o f the specimens.” Clearly, no new taxa had been described and the genus and species are unavailable. 71. H aliotis fa tu i Rheder [s/c], 1981, is mentioned m Ubaldi (1993d l 3-1). The date is most likely based on Kaicher’s (1981: card no. 2902) statement: “This subspecies [of H varia Linnaeus, 1758] is currently under study by Dr. Harald Rehder (USNM) and will probably be described in detail before the end o f the year (1981).” The name was mentioned a second time in the same year (Anon., 1981). H aliotis fa tu i has been validly described by Geiger (1999a). 72- Haliotis hanleyana Ancey, 1881, has some affinities with H clathrata Reeve, 1846 (see also notes 24,25). However, this identification is highly tentative. 73. Haliotis califam iana Valenciennes, 1831, Æ interrupta Valenciennes, 1831, and H parma Vaiencieimes, 1831, were described fiom “America.” The author stated that it was unknown Wiether the specimens were obtained fiom the east or west coast. The specimens were 36 mm and larger, making it unlikely that they could have been the small H pourfa/esn Dali, 1881, Æ dh/fi Henderson, 1915, o tH rohertf McLean, 1970. H aliotis parm a has markings fiom the shell muscle m the shell (\Menciennes, 1831), which additionally suggests that it represents one o f the large California 66 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. species. If H. ccdifomiana, H interrupta^ or K parm a should be synonyms of either K fitlg e n s Philippi, 1845, H. kamtschakana assim ilis Dail 1878, H. w alallensis Steams, 1898, or H. sorenseni Barsch, 1940, then Vaiencieimes^ name would have priority; the other Californian species (fll cracherodiU Leach, 1814, H. corrugata^ Wood, 1828, K rufescens Swainson, 1822) were described before 1831. H aliotis interrupta has been synonymized with H. cracherodii, and H. califom iana with H. ntfescens (Carpenter, 1864:521 fid e McLean, 1966:156, 159); the synonymy oiH . parm a is unresolved and the taxon is treated here as a nomen dubium. If any of Vaiencieimes^ taxa should eventually be shown to be senior synonyms, then an appli cation to the ICZN for the suppression of Valenciennes’ taxa would be advisable. 74. Karsten (1789) published two names for H aliotis, K iridis Karsten, 1789, and H plicata Karsten, 1789. These two taxa are synonyms of H iris Gmelin, 1791, and H. australis Gmelin, 1791, respectively. Karsten’s taxa would be the senior syn onyms, but Rosenberg (1996) has formulated a case to suppress Karsten’s work that is currently pending with the ICZN. His argument, based on ICZN Article 80, is accepted here. 75. H aliotis modesta auct was mentioned by Menke (1845:194) as a juvenile specimen o f H. capensis Dunker, 1844 (= H midae Linnaeus, 1758). The origmal source of H. modesta is unknown to me. Menke (1845), as the only author, also put K semi - plicata Menke, 1843, into tynonymy with H. capensis. I do not accept his opinion for the following reasons. H. sem iplicata occurs exclusively in Australian waters, whereas H. midae is endemic to South AMca. Comparisons o f the shells o f adult H. sem iplicata and juvenile K midae, which are o f the same size, reveal the following characters. H aliotis sem iplicata has distinct spûal cords, Wiereas in H. midae they are only weakly developed. Spûal undulations form nodes in the middle of the shell 67 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. o f K semiplicata^ which, give rise to a slight, but fairly distinct, spiral ridge, whereas K midae tends to have a spiral depression approximately one third from the suture towards the row of holes. The shape o f H. semiplicata is much more elongated than the rotund shell o f K midae from a size o f approximately 2-3 cm onwards. 76. H aliotis schroeteri Menke was mentioned by Weinkauff (1883:83) as a taxon of doubtful status. Weinkauff indicated a potential synononymy with H. scutulum Reeve, 1846 (= H. varia Linnaeus, 1758 ?, cf. note 48). Weinkauff did not know of the original publication, and it is not listed in Sherbom (1922; 1932). I regard it as a nomen dubium. 77. A specimen labeled as type o f H. victoriae Brazier is known from the SAM (R. Pickery, pers. comm.); however, no formal description has been located so far, which renders the name unavailable (ICZN Article 12c). The specimen can be identified as a H. rubra Leach, 1814. Any description o f H. victoriae is very likely to have been published after 1814, as the only abalone species had been described by Brazier in 1878. In case a published description o f H. victoriae should be found, the taxon would then only become a further junior synonym of H rubra. Valid species by faimal regions For each taxon considered valid here the broad zoogeographical distribution is indi cated below. In the case o f a minor overlap the species is listed only in the major provmce. The format o f the entries is as follows: taxon. + syonynms. (Figures herein). Geographic distribution (illustrations) [Notesj. Nomina dubia, nomma nuda and unavailable names are not included tmder the synonymies. Tentative synonyms are mdi cated by a question m ark after the taxon, and the author o f a ^ o n y m is only indicated in case o f homonymy (see Index to Species-Level Taxa for details). 68 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The distributions o f the species are mainly according to Macnae & Kalk (1958), Kira (1962), Habe (1964), McLean (1978), Muller (1984b), Ubaldi (1986), Dharma (1988), Herbert (1990), Vrilson (1993), Geiger (1996), Stewart & Geiger (1999), and various collection records in museums and private collections. Species occurring in more than one province are marked with an asterix and are cross-referenced; their dis tribution is indicated for the area within the respective province. The list o f illustrations is not exhaustive, but focuses on the more recent publica tions. The following numerical code has been used: 1, Abbott (1954); 2, B C ira (1962); 3, Habe (1964); 4, Keen (1971); 5, Hinton (1972); 6 , Dance (1974); 7, Hinton (1978); 8 , Powell (1979); 9, Eisenberg (1981); 10, Kaicher (1981); 11, Bosch & Bosch (1982); 12, Kilbum & Rippey (1982); 13, Abbott & Dance (1983); 14, Sharabati (1984); 15, Stew art (1984); 16, Wells & Bryce (1985); 17, Springsteen & Leobrera (1986); 18, Richards (1987); 19, Dharma (1988); 20, Drivas & Jay (1988); 21, Salvat et al. (1988); 22, Poppe & Goto (1991); 23, Barash & Danin (1992); 24, Wilson (1993); 25, Giannuzzi-Savelli et al. (1994); 26, Geiger (1996); 27, Stewart & Geiger (1999); 28, Simone (1998); 29, Geiger (1999a). Species that have seldom been Ggured (Æ dissona (Iredale, 1927), H. exigua^ H. jacnensis Reeve, 1846, H. marmorata Linnaeus, 1758, H. planata^ H. queketti Smith, 1914, Æ rubiginosa Reeve, 1846, K speciosa Reeve, 1846, H. stomatiaeformiSy H. uni - lateralis) are here illustrated in Figures 1-10 -1-33 along with some noteworthy speci mens (K diversicolor Reeve, 1846, from Bali, H. striata Linnaeus, 1758). All species are illustrated m Chapter 4 (= Geiger, m press). Caribbean auranthan Simone, 1998. Venezuela to central Brazil (28) [note 1 % . 69 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. povrtalesii Dali, 1881. S Florida to Surinam. (10,13) [notes 1 ,2, 3]. European and Senegalese marmorata Linnaeus, 1758. + decussata, guineensis, rosacea^ strigata, virginea Reeve {non Gmelin). (Figures 1-18 - 1-19). Central W Africa (10 [as H. guineensis]^ 13 [as K rosacea\) [notes 4 ,5 ,6,7 ]. * pustulata cruenta Reeve, 1846. Israel and Lybia; see also east African province (10 [as K cruenta], 14,23,25) [notes 10, II, 12]. stomatiaeformis Philippi, 1848 (Figures 1-20 - 1-21). Isl. S o f Italy [notes 5, 8,9 ]. tuberculata tuberculata Linnaeus, 1758. + bistriata Gmelin, bistriata Costa, bisundata, incisa, Janus, japonica, lam ellosa, lucida, parva Risso {non Lvameus), pellucida, reticulata, rugosa Reeve {non Lamarck), secemenda, striata, varia Risso {non Lin naeus), vulgaris. Mediterranean, Brittany to Morocco (10 [as H. lamellosa], 13,22) [notes 4 ,5 ,1 3 ,1 4 15,16]. tuberculata coccinea Reeve, 1846. + canariensis, zealandica. Canary Isl., Azores (10, 22 [both as H. coccineaj) [notes 5,13,17,18]. South African midae Linnaeus, 1758. + ctqtensis, elation S t Helena Bay to WTranskei (6,10 , 12,13, 18) [note 4]. parva Linnaeus, 1758. + canaliculata Fischer ?, canaliculata Lamarck, carinata, cingu - lata, kraussi, rubicunda ROdin^ rubicunda (Montfbrt). False Bay to East London (10,12,13,18) [notes 4,19,20]. queketti Smith, 1910 (Figures 1-24 - 1-25). Transkei to S Mozambique (10). 70 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. spadicea Donovan, 1808. -^fic^ormiSy sanguinea, sinuata ? Partridge Point, Cape Penin sula to N Natal (9, 10,12,13,18) [notes 21,22]. speciosa R esve, 1846. + alfredensis. (Figures 1-30 - 1-31). Port Alfred to WTranskei (10 [also as H. speciosa form alfredensis], 13,18). Eastern African, Red Sea and Persian Gulf * clathrata Reeve, 1846. + venusta. Kenya, Madagascar, Mascarene Isi., Rodrigues IsL, Aldabra, Seychelles, Chagos Arch., Maldives; see also tropical Pacific province (5 & 19 [both as H. crebrisculpta], 10 & 27 [also as H. venusta\) [note 24,25]. mariae mariae Wood, 1828. Oman (10, 11) [note 26]. mariae dentata Jonas, 1846. Oman (10 [as H. mariae form dentata\) [note 26]. pustulata pustulata Reeve, 1846. + jousseaum i. N South Afiica to Persian Gulf (10) [notes 10, 11, 12]. * pustulata cruenta Reeve, 1846. Particularly Red Sea; see also European and Sene galese province (10 [as H. cruenta], 14,23,25) [notes 10,12]. squamosa Gray, 1826. + roedingL S Madagascar (15,27) [notes 27,28,29]. unilateralis Lamarck, 1822 (Figures 1-32 - 1-33). Central E Africa to Red Sea, Aldabra, Madagascar, Mascarene Isl. (14 [as Sanhaliotis pustulata], 20 [as H. varia], 26) [note 30]. rugosa Lamarck, 1822. + altem ata, m ultiperforata, nebulata, pertusa, revelata. Mas carene Isl., Madagascar, central E Afiica (6 & 20 [both as H. pttstidata], 10 [as H. pustulata form altem ata^ [notes 11,31,32,33]. 71 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. bidian Ocean and Tropical West Pacific asirima Lmnaeus, 1758. + - asiman, S-most Japan to Sydney, Andaman Isl. to New Cale donia (2 ,5 ,6 , 7,10 (juvenile and adult], 13 16,17,19,24) [note 4]. brazieri Angas, melculus. (Figures 1-4 - 1-5). S Queensland to Jervis Bay, New South Wales (7, ID, 13,24) [notes 34,35]. * clathrata Reeve, 1846. + venusta. Andamans to American Samoa, S Japan to Sydney; see also east Afiican province (5 & 19 [both as K crebrisculpta], 10 & 27 [also as H. venusta]) [note 24,25]. crebrisculpta Sowerby, 1914. New Caledonia (10,27) [notes 25, 36]. * diversicolor Reeve, 1846. + aquatilis, gruneri, supertexta, tcQ/loriana. (Figures 1-12 - 1-13). China, Taiwan, Bali; see also northwestern Pacific province (2 [as H. super - texta], 3,10 [also as forms gruneri, tayloriana, supratexta, and H. aquatilis^) [notes 37,38]. dissona (Iredale, 1929). (Figures 1-10 - 1-11). Queensland, New Caledonia, Tonga (10) [note 36,39]. dohm iana Dunker, 1863. New Caledonia, Tonga (10,21,27) [note 40]. fa tu i Geiger, 1999. Tonga, Marianas (29) [note 71]. glabra Gmelin, 1791. + p icta ziczac. Pbi%pines, Maluku, Lesser Sunda Isl. (9,10,13, 17,19 [as K planatd^i. hargravesi Cox, 1869. + ethologus. S Queensland to N New South Wales (7, 13, 10 & 24 [both also as K ethologus\) [notes 34,35]. jacnensis Reeve, 1846. + - echinata, hanleyL (Figures 1-16 - 1-17). S-most Japan (S o f Amami Isl.), Philippines, New Caledonia, Marianas, Nficronesia (3,10 [also as H. hanleyi^ [notes 31,41]. 72 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ovina Gmelin, 1791. + caelata, latilabris. Maldives to Tuamotus, SW Japan, Philip pines, Vietnam, Queensland to Western Australia (2,5, 7 ,9 ,1 3 , 16, 17, 19,21, 24) [note 8J. * planata Sowerby, 1882. -^grayana. (Figures 1-22 - 1-23). Philippines to Fiji, North ern Territories; see also northwestern Pacific province (10 [also as H. grqyana]) [notes 31,42]. pulcherrim a Gmelin, 1791: Tuamotus, Henderson Isl. (9, 10,13). rubiginosa Reeve, 1846. + howensis. (Figures 1-26 - 1-27). Lord Howe Isl. (10 [as H. varia rubignosa and H. howensis], 7 & 13 [both as H. howensis]) [note 43]. varia Linnaeus, 1758. + aliéna, astricta, barbouri, concinna, dringii, gemma, granu - lata, papulata, pustulifera, scutulum , sem istriata, viridis. Sri Lanka to Tonga, S Japan, Philippines, central W Western Australia to Sydney (2, 5, 7, 9, 10 [also as ssp. astricta, papulata, pustulifera, viridis, and H. unilateralis], 13, 16,19) [notes 1,4,44,4 5 ,46 ,4 7 ,4 8 ]. Temperate Australian coccoradiata Reeve, 1846. New South Wales to E Victoria (7,9,10,24) [note 49]. cyclobates Péron, 1816. + excavata. South Australia to central S Western Australia (9, 10 [also as H. ovina], 13,16,24). elegans Philippi, 1844. + clathrata Lichtenstein (non Reeve). Western Australia (7, 9, 10,13,16,24) [notes 23,50]. laevigata Donovan, 1808. + albicans, glabra Swainson? (non Gmelin). Victoria to SW Western Australia (6,7,10,13 16,24). roei Chay, 1826. + scabricostata, sulcosa. ^ cto ria to central W Wèstem Australia (6 ,7 , 10 [also a s K sulcosa], 13,16,24) [note 2 7 % . 73 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. rubra rubra Leach, 1814. + ancile, mprobulay naevosa, whitehouseL New South Wales to South Australia, Tasmania (7 & 13 [both as H. ruber], 10 [also as ruber, n improb - ulum, ancile]) [notes 25,27,51, 52, 53, 54, 55]. rubra conicopora Peron, 1816. + cunninghami, granti, vixlirata. Victoria to Freemantle SW Western Australia. 7,16, 24 [all as H. conicopora], 10 (also as H. c. vixlirata) [note 53]. scalaris scalaris (Leach, 1814). + crenata ?, tricostalis, tricostata. W South Australia to central W Western Australia (7,9, 10,13,16,24) [notes 24, 56]. scalaris emmae Reeve, 1846. Victoria to W South Australia (7, 10, 13, 24 [all as emmae]) [note 56]. sem iplicata Menke, 1843. + lauta. SW Western Australia (7, 13,24) [note 49]. squam ata Reeve, 1846. + elevata,funebris. Central W Western Australia to Northern Territories, BaH (7 ,9 ,1 0 ,1 6 ,19 ,2 4 ) [notes 31,39, 57,58]. New Zealand australis Gmelin, 1791. +a/eafo, costata, rugosoplicata. New Zealand (8,10,13) [note 59]. iris Gmelin, 1791. New Zealand (6 , 8,10,13) [note 54]. virginea virginea Gmelin, 1791. + gibba, marmorata Reeve (non Linnaeus), subvir - ginea. S South Isl. & Stewart IsL, N South Isl. to S North Isl. (6 , 8, 10) [notes 60, 61]. virginea crispata Gould, 1847. NE North Isl. (8, 10) [note 60]. virginea huttoni Filhol, 1880. Auckland Isl., (Zhapman Isl. (8,10) [note 60]. virginea m orioria Powell, 1938. Chatham Isl. (8,10) [note 60]. 74 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Northwestern Pacific * dîversîcolor Reeve, 1846. + ■ aquatilis, gnm eri, supertexta, tayloriana. (Figures 1-12 - 1-13). S Hokkaido; see also tropical Pacific province (2 [as H. supertexta\, 3, 10 [also as forms gnm eri, tc^loriana, supratexta, and H, aquatilis^ [notes 37,38]. discus discus Reeve, 1846. Honshu, Kyushu, Shikoku (10) [note 62}. discus hannai Ino, 1953. Korea, Hokkaido, NE Honshu (2,10,13) [note 62]. exigua Dtmker, 1877 (Figures 1-14 - 1-15). S Japan (Okinawa) (10) [note 63]. gigemtea Gmelin, 1791. sieboldii, tubifera. Honshu (2,10) [note 64]. madaka (Habe, 1977). S and central Honshu, Kyushu (2, 10 [both as sieboldii\) [note 64]. * planata Sowerby, 1882. -^grayana. (Figures 1-22 - 1-23). S o f Yakushima; see also tropical Pacific province (10 [also as K grayana\) [notes 31,42]. Northeastern Pacific corrugata Wood, 1828. + diegoensis, nodosa, owenL Central to central Baja California (1 ,6,9 ,1 0 [also as ssp. aweni], 13) [notes 26,65,66]. cracherodii Leach, 1814. + bonita, californiensis, expansa, holzneri, imperforata Dali {non Gmelin), lusus, rosea, splendidula. N California to central Baja California (1, 6,9 , 1 0 [as K cracherodii californiensis and H. c. cracherodit\, 13) [note 66]. folgens Philippi, 1845. + guadalupensis, planilirata, splendens, turverL Central Califor nia to central Baja California (1 ,6,1 0 [also as ssp. guadalupensis\, 13) [note 47]. kamtschatkana kamtschatkana Jonas, 1845. Alaska to Point Conception, S California (1,10,13) [notes 62,67]. kamtschatkana assim ilis Dali, 1878. + aulaea, smithsoni. Central to S California (1,9, 10,13 [all as K assim ilis^ [notes 62,67]. 75 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. rufescens Swainson, 1822. califom îana^ hattorii, ponderosa. N California - Central Baja California (1 ,6,13). sorenseni Bartsch, 1940. Point Conception to central Baja California (10). walallensis Steams, 1898. S Washington to S California (10, 13). Tropical Eastern Pacific dalli Henderson, 1915. Galapagos Isl.. Gorgona Isl. (4, 10, 13) [note 2]. roberti McLean, 1970. Cocos Isl. (4, 10) [note 2j. Zoogeography The species from South Africa, New Zealand and the northeastern Pacific are all endemic, and with the exception o f one questionable report (Macpherson, 1953: the South African H. spadicea Donovan, 1808, in Australia) no wide spread species have been found in these areas. The East Afncan and the Indo-Pacific provinces are also more or less isolated from each other; Geiger (1996) discussed the apparent faunal bar rier around India. Only K clathrata Reeve, 1846, occurs in both areas (Stewart & Geiger, 1999). Indications particularly o f H. varia in the East African province, and H. piistulata in the central Indo-Pacific are erroneous (Geiger, 1996; pers. obs.). In the Australian province the separation o f tropical and temperate species is not very sharp; considerable overlap has to be noted for a few species. Only a limited number o f species are widespread over several faunal provinces, /.e., H. asinma^ K clathrata Reeve, H. o v im and H. varia. A much more detailed, specimen based account o f the distribution o f all abalone species will be provided elsewhere (see Chapter 4). Three biogeographical models have been introduced and are reviewed in Geiger & Groves (1999 = Chapter 2). Talmadge (1963a) proposed the Pacific Rim hypothesis 76 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. with multiple centers o f radiations along the Pacific islands fiom Japan to Australia and New Zealand. Lindbei^ (1992) pointed to the highest diversity o f abalones being found in the central Indo-Pacific, which may (or may not) indicate the origin o f the group there. Based on published chromosomal data Geiger & Groves (1999) suggested a potential origin o f abalone in the Tethys Sea, which is also discussed by Lee & Vac- quier(l995). 77 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter 2: Review of Fossil Abalone (Gastropoda: Vetigastropoda: Haliotidae) with Comparison to Recent Species. INTRODUCTION Recent members of the family Haliotidae, with “abalone” as their common name, are well-known. Due to their economic value, living species have received much scien tific attention (e.g.. Shepherd et al. [1992,1995]; Fleming & Hone [1996]), for only rel atively few and isolated accounts of fossil abalone are found in the literature, with Lindberg (1992) supplying a limited overview. A more extensive review on is presented what little is known about fossil abalone to stimulate further work. DIAGNOSTIC CHARACTERS OF THE FAMILY Shell morphological characters clearly separate abalone from any other family of fossil as well as extant gastropods (Figure 2-1). Abalone shells are easily recognized by their flat, limpetlike shape and row of tremata toward the left periphery. This row of tremata represents the subdivided selenizone found in Pleurotomaroidea, Scissurel- loidea, and Fissurelloidea (Knight et al., I960; Bandel, 1998; Geiger, 1998a; McLean & Geiger, 1998). The extremely hypertrophied epipodium is diagnostic character for the anatomy of Haliotidae, but such characters do not apply to fossil representatives, and are not further discussed here (see Geiger, 1998a). Some Paleozoic Bellerophon- toidea possessed shells somewhat resembling those of abalone. The former, however, are involute, have the row of tremata along the median of the shell, have cross lamellar aragonite, and have a muscle scar is more similar to that of Fissurelloidea than to Hali otidae (McLean, 1984a). 78 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 2-1: Specimens o f fossil haliodds. I, H aliotis rufescens. Pliocene, Saugus For mation. Simi Valley, Ventura County, California, LACMIP loc. 12660, internal mold showing muscle scar. 209 nun. 2, H aliotis sp., Miocene, Topanga Canyon Formation, Moorpark, Ventura County, California, LACMIP loc. 16896, internal mold, LACMIP 12659, 116 mm. 3, H aliotis tuberculata volhynica, “Post Pliocene? West Indies? Europe?”, internai mold with dendrites, AMNH 45571,62 mm. 4, H aliotis walallensis. Pliocene, San Diego Formation, hills south ofTguana River, San Diego County CA, LACMIP loc. 16817 (ex LACMIP loc. 305c), specimen with shell preserved, 113 mm. 79 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Several genera in the trochid subfamily Stomatellinae (Hickman & McLean, 1990; Pickery, 1995) have shells that loosely resemble abalone. Stomatellids are found in the late Triassic?, and from the Pliocene through Recent (Knight et a i, I960). They have rather small (^ 40 mm), flat, oblong shells that lack tremata or spiral sculpture, and may be mistaken for imperforate specimens of juvenile H aliotis asinina Linnaeus, 1758. The latter, however, have several distinct spiral ridges that become obsolete as the shell grows larger (> 35 ram). Specimens of the living genus Granata (Trochidae: Eucycli- nae) have been erroneously identified as imperforate H cyclobates Peron, 1816 (Geiger, 1991, 1998a). Imperforate specimens of abalone have been found in the Recent but are very rare (see Geiger, 1998 a, for review). For Recent as well as for fossil specimens, it is unlikely that an imperforate shell with a depressed, flaring shell is an imperforate abalone. Although Trochotomidae (Pleurotomaroidea) are superficially similar to abalone, most trochotomid species have a distinctly trochiform shell and only one trema on the shoulder of the last third of the body whorl. In addition, their early Triassic to Late Jurassic geologic range does not overlap with the known range of Haliotidae (Knight et oL, I960). TAXONOMY The shell as the basis of taxonomy As with most fossils, discrmiination o f taxa in abalone is based on then hard parts. The shell o f abalone however, is extremely plastic in Recent species and, therefore, can be mferred to be plastic m fossil congeners by application o f uniformitarian principles. Some examples o f morphological plasticity in Recent taxa are discussed below to illus trate the problems using a limited number o f spechnens to deSne taxa. 80 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The most striking example of variability is the number of open tremata» which has been considered a constant and diagnostic character by previous workers. The above is particularly true for Recent species (e.g.» Kaicher, 1981; Abbott & Dance» 1983), but only to a lesser extent for fossil ones (but see e.g., Sohl, 1992), because the incomplete state o f most specimens is recognized. In Figure 2-2» the number o f open tremata is plotted for several Recent species for which sufBcient data is available. The number of open tremata varies within species, and the range for each species overlaps to a great extent with ranges o f the other species shown. Therefore, this character is not diagnos tic (cf. Geiger 1998a). Sculpture has been used to separate Recent “species”. For example, the European “/T. lam ellosa^ Lamarck, 1822, and tuberculatcT Linnaeus, 1758, which are now considered forms (Geiger, 1998a) of the single, variable species H tuberculata^ are dis tinguished by the presence or absence o f obliquely radial lamellae. As with the number o f open tremata, there is a great variability for this character. If a large number of spec imens Grom any population is examined, entirely smooth to highly lamellar shells can be fotmd (Geiger, tmpublished data). Shell outline, particularly roundness, is to some extent under environmental control as documented by transplant experiments o f a Japanese species (Ino, 1952). Stewart & Geiger (1999) showed for the tropical K clathrata Reeve, 1846, that shell roundness also changes through ontogeny. Therefore, this character is o f limited use for species discrimination. Other variable shell characters include (a) the degree o f coverage o f the shell with a particular sculptural element (e. g., scales in H. ja cn en sis Reeve, 1846: Geiger, personal observation), (b) the stength o f spiral cords for H. rubra Leach, 1814 (see Geiger, 1998a), and (c) shell Gamess m K o vim Gmelin, 1791, for which extremely 81 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H . cracherodif 'californiensis ~ n = 305 H. tubercuiata L > 15 mm n = 403 L -i H . cracherodii juveniles n = 87 H,ins n = 442 I H . cracherodii californiensis n = 305 H . cracherodii adults n = 30 ..Il i-J I 4 6 7 9 TOIT 12 13 T4 15 16 Number o f open perforations 5 6 7 8 9 10 Number o f open perforations Figure 2-2: Histograms of number o f open tremata for several species o f H aliotis. n = sample size. Sources o f data as follows. H aliotis tuberculata: only Mediterranean popu lations: Geiger (unpublished). H aliotis coccoradiata: Talmadge (1960); the number o f half open tremata was split equally between the neighboring integers. H aliotis iris: Sin clair (1963). H aliotis cracAeroc/iff juvenile: Hemphill (1907). Haliotis cracherodii cali - fo m ien sis: Hemphill (1907). H aliotis cracherodii adult: Hemphill (1907). Note the bell-shaped distributions, with large differences in the modal class. Further note the shift o f the modal class between juvenile and adult H cracherodii. The large number o f specimens with 16 open tremata in the plot of H cracherodii californiensis stems from a priori selection from the original stock o f shells. 82 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. flat, Vietnamese specimens contrast with towering forms from the Philippines (Geiger, personal observation). Hence, sound taxonomic decisions are impossible on the basis o f a limited number o f shells. Usually for each fossil taxon few specimens are available, which in many instances show only slight differences between the nominal taxa. Nevertheless it would also be unwise to synonymize all fossil taxa, and the taxonomy o f fossil abalone will probably remain highly typological. It is the intention o f this contribution to demon strate that character variability within living taxa must be considered when describing new fossil taxa. Multivariate statistical techniques might eventually help, but to date have not been used for fossil haliotids and only in a single study o f a Recent species (McShane et a i, 1994). Fossil abalone taxa At least 35 fossil abalone species have been described, excluding reports o f Recent taxa with a fossil record. It is unclear whether all these taxa are truly distinct species. A critical and comprehensive taxonomic revision of fossil abalone has not been attempted and is not feasible because o f the limited material. Most fossil abalone have been described from single specimens (cf. Vbkes, 1978), the exceptions being the two speci mens o f K kurosakiensis Kotaka & Ogasasawara, 1974, from the Miocene o f Japan, four specimens o f H. saldanhae Kensley, 1972, from the Pliocene o f South Africa, and ten specimens o f H. antillesensis from the Maastrichian o f Puerto Rico and Jamaica (Kensley, 1972; Kotaka & Ogasawara, 1974; Sohl, 1992). Several fossil species have shells similar to thoses o f modem represoitatives withm Hialtotis. It is unknown whether these fossil forms are ancestors, conspecifrcs, or share similarities due to convergent evolution. A list o f fl)ssil taxa, expanded from Lindberg (1992), is contained in Table 2- 83 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3, where all records are listed using the original taxa. Nevertheless, it was tried to shed some light on potential synonymies and similarities between certain taxa. H aliotis lomaensis Anderson, 1902, from the Late Cretaceous (Maastrichian) o f San Diego County, California, has been compared to the extant H. iris Gmelin, 1791, endemic to New Zealand. H aliotis antillesensis Sohl, 1992, from Upper Cretaceous (Maastrichian) rocks o f Puerto Rico and Jamaica is similar to the extant Australian species H tyclobates Peron, I8 I6 (Sohl, 1992). Due to the magnitude o f the temporal hiatus and the large geographical distances involved in these two species pairs, a close relation is doubtful in either case. Additionally, the fossil H lomaensis is only 13 mm in size, and is a Juvenile specimen by the standard o f all northern Pacific species as well as o f i ï iris. Identification o f Juveniles is extremely difficult. Juvenile H. iris, in partic ular, have a very distinct morphology and are often confused with, adult H. virginea Gmelin, 1791, from New Zealand, a species readify distinguishable from H. lomaensis by its pronounced sculpture. The cited similarity between H. lom aensis and H iris, therefore, must be viewed with much caution. Shell morphological similarities between certain pairs o f species from the upper Tertiary (Miocene and Pliocene) and the Quartenary (Pleistocene-Recent) from Califor nia and Japan have been noted (Hertlein, 1937; Talmadge, 1964; Hatai et a i, 1970; Mulliner, 1984) and are listed in Table 2-1. A close evolutionary affinity can be consid ered in each case because species are morphologically similar, are reported from the same area, and are separated by relatively small time spans. H aliotis poweUi Fleming, 1952, from the Miocene and Pliocene o f New Zealand, was thought by Talmadge (1963) to belong to the group o f H. clathrata Reeve, 1846, H. nibiginosa Reeve, 1846 [as H. hawensis (Iredale, 1929)], and H. coccoradiata Reeve, 1846. H aliotis clathrata Reeve, 1846 (non Lichtenstem, 1794), has recently received 84 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. § I I Û I 4 > I ! u g e s C Û I I es O m T T 00 s :§ « I e s u es S â. CL O e n C - I •- W î il 5; ïi « r v e s G \ I I 1 I es u s § §• oo Os 00 c / i î g s j - î ai I ai O e - i I a; 1 1 a; a; a; a; Table 2-1: Comparison o f upper Tertiary species with Pleistocene-Recent species from California according to Hertlein (1937), Talmadge (1964), Hatai e t aL (1970), and Mulliner (1984). 85 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. farther attentîoa elsewhere (Geiger, 1998a, 1998b; Geiger & Stewart, 1998; Stewart & Geiger, 1999). H aliotis clathrata Reeve, 1846, H. rubiginosa, and H. coccoradiata do not occur on New Zealand, and none o f the New Zealand species—Recent or fossil— has been recorded outside these islands. In addittion, Stewart & Geiger (1999) disagreed with Talmadge^s (1963) opinion on the basis of discrete morphological differences and diqunct geographical distribution of these taxa. H aliotis clathrata Reeve, 1846, is mentioned from Fiji (as H. tuvuthaensis Ladd in Ladd & Hoffineister, 1945) and Guam (Ladd & Hoflhneister, 1945; Ladd, 1966). Stew art & Geiger (1999) have synonymized the Fiji record listed as H tuvuthaensis under H clathrata Reeve, 1846, because the type of H tuvuthaensis can not be distinguished from H clathrata Reeve, 1846. Talmadge (1963) listed H clathrata Reeve, 1846, as H crebrisculpta Sowerby, 1914, a highly controversial but distinct species for which a lec- totype has since been designated (Stewart & Geiger, 1999). The taxonomic states o f H barbadensis Trechmann, 1937, from the Pleistocene of Barbados, and the Recent H. pourtalesii Dali, 1881; H aliotis aurantium Simone, 1998; H. dalli Henderson, 1915; and H. roberti McLean, 1970, bears mentioning. These are small species for the genus with a maximum size o f approximately 2 cm. They live in the Caribbean (H barbadensis, H. pourtalesii), on the Atlantic coast of Venezuela and Brazil (Æ aurantium), and in the eastern Pacific outliers of Isla del Coco and Islas Gala pagos (H. dalli, H. roberti). The living species are found at a depth o f60 - 400 m (Hen derson, 1915; Bartsch, 1940; Foster, 1946; Aguayo & Jaume, 1947; Harry, 1966; Jung, 1968; Klappenbach, 1968; Sarsua, 1968; Nijssen-Meyer, 1969; McLean, 1970; Kaicher, 1981; Titgen & B ri^ t, 1985; Odd, 1986; Finet, 1993; Martinez & Ruiz, 1994; Simone, 1998). Using the species concept o f mterbreeding populations, the Recent H. roberti and H d a lli versus K pourtalesii are clearly distinct species because they occur on 86 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. opposite sides o f Centrai America. However, the geological closure of the Isthmus of Panama in the middle Pliocene (Coates & Obando, 1996) complicates the situation for the fossil species considered here. The open waterway could have provided a means for gene flow within a single amphipanamic species. The question arises, when the modem species became distinct, and where to draw the line between the fossil species. Con flicting opinions are expressed by authors cited above as to whether H, barbadensis and K pourtalesii are endpoints of a morphological range within a single species or are two valid species. Due to the scarcity o f material for both species and the fairly extended time period separating these taxa, are considered them distinct Strausz (1966) is followed here and all fossil European taxa {H. anom iaeform is Sacco, 1896; K benoisti Cossmaim, 1895; H. lam ellosa Lamarck, 1822; H. lam el - losoides Sacco, 1896; K monilifera Bonelli, 1827; K n ew illei Bial de Bell, 1909; H. ovata Bonelli, 1827; K tauroplanata Sacco, 1897; H. torrei Ruggieri, 1989; K tuber - culata Linnaeus, 1758; K volhynica Eichwald, 1853) are referred to H. tuberctdata vol - hynica because the Atlantic and Mediterranean populations o f the Recent species {H. t. tuberculata) are known to be extremely plastic in its shell morphology. Most illustra tions and material o f European fossil specimens (e.g.. Figure 2-13) fall within the range o f variation within the Recent species. The time lapse may justify a separation on the subspecies level. All fossil taxa are o f Miocene or younger age; with the exception o f the Oligocene records (Lozouet, 1986) o f the nominal taxon H. benoisti. The following differing opinions on the taxonomic states o f these taxa have been offered. Hômes (1856) synonymized H. volhynica with H. ovata. Delhaes (1909) referred all six o f the European taxa he mentioned to H. tuberculata. Krach (1981) discussed the European taxa and retained two subspecies o f H. tuberculata. Specimens rotmder than any Recent H. tuberculata, sensu lato, were considered ff. tuberculata volhynica (Krach, 1981: par- 87 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ticularly figs. 2 and 3). Krach. (1981 : figs. 4-7) also figured Æ tuberculata tauroplanata^ which showed a typical representative o f the Recent K tuberculata. Lozouet (1986) separated H. benoisti firom the French Oligocene firom H. tuberculata. However, the material in the Muséum Nationale d’Histoire Naturelle in Paris does not justify a sepa- ratation o f those specimens fix)m H. tuberculata, sensu lato (Geiger, personal observa tion). The remaining European taxa have not received any attention beyond a simple mention in the sources cited in Table 2-3. Fossil abalone m the phylogenetic context A phylogenetic study of fossil abalone either alone or integrated in the fimnework of Recent taxa (cf. Smith, 1994 for review o f conceptual approaches), is unfortunately fraught with problems. As with most fossil material, fossil abalone with soft-part preser vation are unknown. The morphological plasticity in shell characters, outlined above, also makes phylogenetic analysis problematic. In addition, most fossils are preserved as internal and/or external molds, which limits the suite of potential characters to the shell sculpture characters. The predominance o f moldic preservation is unfortunate. The pris matic layer o f abalone shells has been reported to exist in three fimdamentally different mineralogical types (calcitic, aragonitic, admixed calcMc and aragonitic: Mutvei et a i, 1985; Dauphin e t cd., 1989; Dauphin & Denis, 1995; Shepherd e t a i, 1995), and may be taxonomically infbimative. These authors used Feigel’s stain to identify aragonite, although this stain also shows high-magneshim calcite (C. Hedegaard, personal com mun.). Therefore, mineralogical composition o f abalone shells needs reexamination. However, the phylogenetic character states “Feigl-stainin^ and “non-Feigl-staining” may also be usefiil without an explicit asstunption o f stain mineralogical specificity. The spatial sampling scale in X-ray diffraction studies is limited by the beam size to I- 88 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2 mm (e.g., Hedegaard & Wenk, 1998), which is too coarse to reveal mineralogical pat terns, particularly admixed aragonitic elements o f 2-5 pm width and 5-10 pm length m the calcitic external shell layer (Dauphin e t a i, 1989). Additionally, it would only be possible to investigate rarely preserved shell material. PRESERVATION Abalone are rarely encountered as fossils, although roclqr shores, which abalone inhabit, particularly in temperate regions, are thought to have occurred widely along the west coast o f North and South America. However, rock-inhabiting organisms are rarely preserved (Carter, 1972), because the habitat is destructive to the shells (Woodring, 1931; Kotaka & Ogasawara, 1974; Parsons & Brett, 1991). Many tropical abalone species occur in reef environments and tend to hide in cracks or underneath boulders. Upon death, the shells are likely to remain in the cryptic habitat and will eventually be incorporated into the biogenic limestone o f the reef. Tectonic uplift or eustatic sea level change may expose these ree6 above sea level, making them accessible (e.g., the Pleistocene record o f K pusttdata cruenta Reeve, 1846, from the Red Sea [Hall & Standen, 1907]). A major constituent o f abalone shells is aragonitic platelets in the nacreous layer. This type o f microstructure is very prone to disintegration, dissolution, and diagenetic change (Dodd, 1966). Disintegration accounts for the general scarcity o f complete abalone specimens in the fossil record. The most frequent mode of abalone preservation is as internal and/or external molds (Figures 2-1.1 - 2-13), occasionally with, some attached shelly remains. Preserved shell fragments are all calcitic, indicating some dia genetic effects (cf. Dodd, 1966), because the inner nacreous layer o f the shell is always composed, o f aragonite (cf. Dodd, 1966). fri addition, Dudiam (1979b) observed recrys- 89 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tallîzatîoa in K lomaensis,Tb& effects o f diagenesis make the wide application o f shell m ineralo^ in the taxonomy o f fossil abalone doubtful, although results on the mineral - ogy o f shells from Recent abalone are promising (see also previous section on phy- logeny). PALEOENVmONMENTS The habitat o f abalone, with their large aperture, is the rocky shores, as is the case for morphologically similar groups such as limpets. Not surprisingly, fossil Haliotidae inhabited comparable habitats. Below supporting evidence is presented. Reef paleoenvironments Both fossil and Recent abalone are found in similar microhabitats within tropical reefs. H aliotis antillesensis has been found in a radist-framework bioherm in the Upper Cretaceous o f Puerto Rico (Sohl, 1992), part o f the shallow tropical Tethys Sea (Kauff man & Sohl, 1974; Sohl, 1987). Associated species included abundant neritid gas tropods, indicators for an intertidal to shallow subtidal hard substrate in the tropics. H aliotis pustulata cruenta has been found in a Pleistocene raised coral reef o f the Red Sea area [Newton, 1900; Hall & Standen, 1907 (as H unilateralis Lamarck, 1822); Brattwer et a i cited in Yaron, 1983] in association with the typical, moiluscan reef fauna found m the Red Sea. The reconstructed paleoenviromnent is fully congruent with the modem habitat o f this species (Tforon, 1983; Geiger, 1996). For European sites, Davi- daschvili (1937) and Krach (1981) proposed a nearshore shallow-water reef environ ment for then H volhynica based on a community analysis. 90 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Shallow-water rodqr paleoenvironments Fossil abalone also are reported from deposits associated with rocky shorelines, a common habitat for Recent members of the family. In the Cretaceous of California, K lomaensis was found with an oyster, a serpulid worm, and encrusting calcareous algae, which together indicate a shallow-water, rocky environment (Durham, 1979b). In the Miocene o f California, K palaea Woodring, 1931, was discovered with other moilus can genera indicative o f rocky substrate, i.e., Tegulctr Area, Crassadoma [as H înnites\, Lima, and M ytilus (Woodring, 1931). Only the limids may be freely mobile as adults and associated with sandy substrates. H aliotis elsm erensis Yokes, 1935, from the Pliocene of California, however, was found admixed with infaunal bivalves (Chione) as well as with rocky shore bivalves (Area), This mixed fauna indicates some transport (Yokes, 1935). Similarly, K eraeherodii Leach, 1814, from the Pleistocene o f north western Baja California (Addicott & Emerson, 1959). H aliotis rufeseens Swainson, 1822, has been found in a Pliocene shallow-water channel deposit with rock-dwelling species, as well as infaunal species (Groves, 1991). Additional, indirect evidence o f California abalone paleohabitat stems from the type specimen o f the Late Pleistocene vermetid gastropod Petaloeonehus anelltim (Mdrch, 1861), which is attached to a shell o f an unspecified abalone (Grant & Gale, 1931). Yermetids are common in shallow, rocky habitats. In Japan, H kurosafdensis Kotaka & Ogasawara, 1974 (upper Miocene); H, fujiokai Hatai, Kotaka, & Noda, 1937 (middle Miocene); and H. koehibei Hatai & Nisiyama, 1952 pliocene), have been found in strata containmg Qfpical rock-mhabitmg moiluscan genera such as Ostrea, Area, and Aemaea admixed with sand-dwellers, such as Maeoma and Lueinoma (Hatai e t aL, 1970; Kotaka & Ogasawara, 1974; Noda et a i, 1995). As 91 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. with K elsmerensis, some transport of the original fauna must be assumed, but a roclqr shore habitat for abalone is indicated. In siunmary, available evidence indicates that the ecology o f the fossil abalone is comparable to that o f Recent species (see e.g., Crofts, 1929, Cox, 1962, Shepherd, 1973). Particularly, no difference in the depth distribution (= on-shore/off-shore pattern) can be detected. This pattern has been documented during the geological history o f a number o f invertebrate taxa (Hickman, 1984; Bottjer & Jablonski, 1988). It may seem farfetched to consider an on-shore/off-shore pattern in a largely herbivorous organism, however, one abalone species (K cracherodii) is known to feed on bacterial mats as adults (Stein, 1984). Juvenile abalone feed on diatoms and the associated biofilms (Kawamura et aL, 1998, for review). Accordingly, the potential to exploit deep water habitats directly or by neoteny is present in abalone. TIME RANGE Fossil abalone have been found from the Late Cretaceous through the late Pleis tocene, with a conspicuous hiatus in the early Paleogene (see Appendix for details). A controversy has long existed regarding the true identity and systematic affinities o f some o f the earliest records (Late Cretaceous: Maastrichian) in this family: H. antiqua Binkhorst, 1861, from the Netherlands; K cretacea Lundgren, 1894, from Sweden, and H. lom aensis from San Diego County, California (Binkhorst, 1861; Lundgren, 1894; Anderson, 1902). Davies & Eames (1971) considered none o f these records to represent abalone. Woodring (1931) doubted that £ T . lom aensis is an abalone, because o f poor preservation; the only diagnostic characteristic preserved was the flat shell. In addition, Knight et aL (1960:221) cited the Cretaceous record o f H aliotis with a query. Yokes (1935) considered H lom aensis to be an abalone, but most likely a young specimen. 92 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Equally, Sohl (1992) discussed these three Cretaceous taxa and concluded that only H. lom aensis is an abalone, as did Durham (1979b) and Lindberg (1992). More recently, Sohl (1992) described a second Late Cretaceous abalone from the Maastrichian of Puerto Rico and Jamaica, H. antillesensis. Only H. lomaensis and H. antillesensis are considered here to be Cretaceous abalone, both 6om Maastrichian rocks. Within the Tertiary, Delhaes (1909) and Beu & Maxwell (1990) reported abalone from the Eocene. However, the Standard European Stages in Delhaes (1909) are Miocene (Lindberg, 1992), and consequently only one record, firom New Zealand, is of Eocene age. Three factors may explain the hiatus in the fossil record between the Maas trichian and the Miocene. First, Paleogene rocky-shore habitats are hardly preserved (Lindberg & Squires, 1990). Second, in general there is a taphonomic bias against rock- clinging moUusks (Sohl, 1992). Third, by comparison to ecologically similar taxa such certain limpets (Acmaeidae), which are known from the Triassic through the Recent (Knight e t aL, 1960), Cretaceous abalone can be inferred as rare with just two localities. K-T extinction may have reduced the abundance o f abalone to such low densities that they do not show up in the fossil record o f the Paleocene. However, complete extinction at the K-T boundary with convergent evolution of the same shell m otpholo^ by another gastropod group in the Eocene-Oligocene is unlikely. Cretaceous and Tertiary/Recent specimens share many similarities, such as the row o f tremata, the flattening o f the shell, and the high translation value o f the spire. This combination o f structural ele ments is not known fiom any other gastropod, living or extinct. These similarities are best explained by common descent, rather than two separate radiations. 93 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. GEOGRAPHIC DISTRIBUTION In general, one might expect the Neogene distribution of taxa to be fairly congruent with its present-day distribution. However, the very few recorded fossil abalone speci mens from the central Indo-Pacific (Figure 2-3, Appendix) is surprising, because this region houses the highest diversity o f Recent Haliotidae (Delhaes, 1909; Lindberg, 1992). This discrepancy may be associated with the greater paieotologicai effort being carried out in the United States and Europe as compared to the Indo-Pacific. In the few, more extensive studies in tropical settings [Hall & Standen, 1907 (Red Sea); Ladd & Hofrneister, 1945; Ladd, 1966 (Indo-Pacific)] abalone specimens have been recovered. Therefore, more extensive searches for fossil abalone in the Indo-Pacific, as well as in the Australian- region, might close both this noticeable geographical gap for fossil abalone, as well as narrow the Cretaceous to late Eocene hiatus in the abalone record. Geiger is currently preparing a phylogenetic analysis o f the Recent species o f the family, and the biogeographical investigation will be useful in evaluating the proposed models of radiation, summarized below; 1) Living abalone are most diverse in the Indo-Pacific, which may indicate that this is their center o f radiation. However, this is certainly not a safe conclusion, as discussed by Lindberg (1992). It is unclear whether several small radiations or a single large-scale radiation took place within this family. 2) Talmadge (1963) proposed a “Pacific Rim” model, in which he postulated that abalone were originally found on an island arc from Japan to northern Australia and radiated fi»m this area to California, southern Australia, and the Indo-Pacific. This pro posal is similar to one o f the two models, in Lee & Vacquier (1995). 3) A third scenario is based on chromosomal data, which have been documented for 10 species (Table 2-2). Because the basal gastropods have a very low diploid numbw o f 94 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 2-3. Localities o f fossil abalone listed in Appendix. The two inserts provide details for California, and Honshu, Japan; the location of each area on the larger map is shaded. Overlapping symbols indicate different geological times are represented at the same locality.. 95 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3 B- o îi: p o Z .2? O O u t 0 _ - I I S U sa 96 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Taxon H D 0 Source H aliotis tuberculata 14 28 EM Colombera and Tagliaferri, 1983 28 EM Arai and Wilkins, 1986 H lamellosa^ 14 EM Colombera and Tagliaferri, 1983 H aquatilis^ 17 34 IP Nakamura, 1985 H diversicolor aquatilis 16 32 IP Nakamura, 1985, 1986 H diversicolor 32 IP Arai et a l. 1988, Yang et al., 1998 H exigua* 32 IP Arai era/., 1988 H. planatc^ 32 IP Araieffl/., 1988 H asinina 32 IP Jarayanbhand et a l, 1998 H ovina 32 IP Jarayanbhand ef a/., 1998 H asinina 32 IP Jarayanbhand et oL, 1998 H varia 16 32 IP Nakamura, 1986 H cracherodii 36 NP Minkler, 1977 H discus discus 36 NP Arai et a l. 1982 H discus hannai 36 NP Arai et a l, 1982 H madakcf 36 NP Nakamura, 1986 Table 2-2: Chromosome number m Recent H aliotis spp. as indicated in the respective source. H, Haploid number; D, Diploid number; O, Geographic occurrence; EM» Euro pean-Mediterranean; IP, bido-Pacifîc; NP, North PaciGc. ^Haliotis lam ellosa has been shown to be a synonym/ecomoroph o f H . mberculma (Lee and Vacquier, 1995). ^As K diversicolor aquatilis in Nakamura (1985), but as K aquatilis in Nakamura (1986). ^As H. Japonica. *As H planata. *As H. varia. *As H gigantea. 97 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. chromosomes (18-20: Patterson, 1967; Haszprunar, 1988), a model o f progressive increase in chromosome number can be postulated, 6 om a low of diploid number o f 28 in K tuberculata from the Mediterranean, to 32 in Indo-Pacific species, to 36 in North Pacific species. This would suggest that H. tuberculata is a relict species from the ancient Tethys Sea, and that abalone dispersed eastward, which is in agreement with the eastward dispersal pattern in the Pacific (Briggs, 1995). This is in marked contrast to the westward dispersal o f other molluscs (Squires, 1987). In westward dispersing fami lies moiluscan families such as Cypraeidae, Volutidae, and Ranellidae, however, larvae are teleplanktic. Abalone, on the other hand, have a limited dispersal capability, the pre cise extent o f which is currently debated (Tegner & Butler, 1985; Prince et aL, 1987; McShane et a i, 1988; Wells & Keesing, 1990; Shepherd e t a l., 1992). Note that, under this model, the Upper Cretaceous abalone from California and the Caribbean would not be early representatives o f the family and that the root o f the family would have to be placed at a much earlier time. A similar model has been proposed by Lee & Vacquier (1995) and is preferred by Eagle (1996). None o f these models has more than anecdotal evidence. The phylogenetic trees published to date (Brown & Murray, 1992; Brown, 1993; Lee & Vacquier, 1995) are all unrooted, and species suggested to be in a basal position may not be truly basal. It is possible that m the Pleistocene, species o f H aliotis migrated between Asia and North America (Grant & Gale, 1931, p. 96). Biochemical data support this hypothesis. In a tree based on the cDNA sequences o f the fertilization protein lysin (see Vacquier & Lee, 1993, for review) o f 22 abalone species, all California species cluster closely together with the west Pacific species H discus Reeve, 1846, and H. gigantea Gmelm, 1791, whereas another species o f the latter region (H. V ersico lo r Reeve, 1846) is far removed from the western Pacffic species group (Lee & Vacquier, 1992, 1995). This 98 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. pattern has been corroborated with allozyme frequency data from 17 abalone taxa (Brown, 1993). The California species have the most derived character states, which suggests an east-to-west progression. All three o f the above models imply favorable conditions for dispersal of abalone between the east and west coasts o f the north Pacific, with subsequent reproductive isolation o f the amphipacific populations. Glaciation events could facilitate dispersal across the Pacific as sea level dropped, because the dis tance between the two shores at any latitude would be somewhat shortened (see also Lee & Vacquier, 1995). On the other hand, the distance across the Pacific is extensive, hence, dispersal across the Pacific must be termed a rare chance event. Accordingly, one may debate the possible contribution o f glaciation to dispersal. Migration at north ern latitiudes along the Bering Bridge seems unlikely due to the absence o f any Recent species generally beyond 5 0 ® north or south (60® N for H. k. kamtschtkana). 99 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 2-3: List o f fossil abalone taxa, as indicated in the original source, by first occur rence in the fossil record and locality compiled firom the literature and museum speci mens. General locations such as ‘California’ have been omitted if more precise information is available for the same age. Taxa preceded by an asterisk (*) are living species, u: upper, m: middle. I: lower. BC: Baja California, Mexico. CA: State of Cali fornia, USA. NZ: New Zealand. Pref.: Prefecture. AMNH: American Museum of Nat ural History, New York. ANSP: Academy o f Natural Sciences Philadelphia. BMNH: The Natural History Museum, London. LACMIP: Natural History Museum o f Los Angeles County, Invertebrate Paleontology. MNHN: Muséum National d’Histoire Naturelle, Paris. Circumpacific geologic time intervals have been correlated with Euro pean ones according to Kennett, and Siinivasan (1983). ^Haliotis sp. is similar to H, waitemataensis (Ben & MaxweU, 1990) and the illustration strongly resembles H emmae, HTie Shigarami Formation is interpreted as two different ages by the four publications cited. 3 : Correlation of stages accon^g to Beu & MaxweU (1990:8-9). ^OriginaUy reported as H aliotis sp. but corresponds exactly to the type o f H. pow elli Ulustrated in Beu & Maxwell (1990). H)elhaes (1909) interpreted these finds as Eocene. ^Haliotis tuvuthaensis has been synonymized with H. clatfirata Reeve, 1846 (Stewart & Geiger, 1999). TIate 2, figure 5 Ulustrated a fknly round specimen superfi- ciaUy more similar to H aliotis ovina than H clatfirata Reeve, 1846. However, this spec imen is only 15.9 mm long. SmaU specimens o f H clatfirata Reeve, 1846, are much more round than adult ones, which makes the identification of Ladd (1966) valid. T he Miocene o f Durham (1979a) has been revised to late Pliocene by Hickman & Lipps (1985). ’Durham re-dated the coral rock in which the abalone was found firom Jung’s PUocene to Pleistocene. "The identification of the specimen by Suter (1913) as H aliotis iris was rejected by Eagle (1996). too Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. C D ■ D O Q . C g Q . ■ D CD C/) C/) 8 ■ D 3. 3 " CD CD ■ D O Q . C a O 3 " O o CD Q . ■ D CD C /) C /) Table 2-3 Taxon miilfesensis lomaemis sp. Geologic Time Interval Maastrichian Maastrichian Eocene, 1 sp, aff. waHetmtaenis Oligocene/Miocene benoisti Oligocene, u: Chattien Adour Oligocene, u; Chattien Adour Miocene, 1 Miocene, 1 Miocene, I Miocene, I Miocene, I Miocene Miocene, m Miocene, u sp sp amabilis kurosakiensis Miocene, m Miocene, u Locality CRETACEOUS Pueito Rico and Jamaica San Diego County, CA TERTIARY; Eocene NZ Oligocene Cookson Volcanics, NZ St.-Paul-Lès-Dax, France Belus “Marcon", Landes, France St.-Martin-de-Hinx, France Cabanes, St.-Paul-Les-Dax, France Meilhan, Landes, France Lariey, Saucats, Gironde, France Merignac, France Gironde, France Mirabeau, Indra and Loire, France Beugnon Miocene Gifu Pref., Japan Aomori Pref ,Japan Literature Source and Museum Specimens Sohl, 1992 Anderson, 1902; Woodring, 1931, 1932; Bartsch, 1940; Cox, 1962; Durham, 1979b Beu and Maxwell, 1990; Eagle, 1996 Beu and Maxwell, 1990' Cossmann, 1895; Delhaes, 1909; Lozouet, 1986 MNHN no # MNHN no # MNHN no ff MNHN no # MNHN no# Cossmann, 1918 Delhaes, 1909 MNHN no # MNHN no # Itoigawa and Tomida, 1982 Kotaka and Ogasawara, 1974; Itoigawa and Tomida, 1982; Lindbeig, 1992 C D " O O Q . C g Q . ■ D CD C/) C/) 8 ■ D CD 3. 3 " CD CD ■ D O Q . C a O 3 " O o CD O . ■ D CD C /) C /) Table 2-3, continued kochlbei Miocene, I Miocene, m Miocene, m - u Pliocene Pliocene Miocene, 1 koikei glabrosa koyamai notoemis Pliocene Pliocene, 1 Miocene, 1 - m Pliocene, I Pliocene, I - m Miocene; Pliocene, Miocene, m Miocene, m Kanagawa Pref., Japan Ibaragi Pref, Japan Ibarabi Pref Japan Ishikawa Pref, Japan Ibaraki Pref, Japan Kanagawa Pref, Japan Kanagawa Pref, Japan S Fossa Magna, Japan Shizuoka Pref, Japan Shizuoka, Pref, Japan South Fossa Magna, Japan Nagano Pref, Japan Ishikawa Pref, Japan Fukui Pref,Japan Hatai and Nisiyama, 1952; Hatai e( a i, 1970 Kotaka and Ogasawara, 1974; Itoigawa and Tomida, 1982 Hatai and Nisiyama, 1952 Lindbeig, 1992 Shibata, 1957; Nodaern/., 1995 Hanzawa et a i, 1961; Hatai et a i, 1970; Itoigawa and Tomida, 1982 Kotaka and Ogasawara, 1974 Tomida, 1996 Nomura and Niino, 1932; Hanzawa et a i, 1961; Hatai and Nisiyama, 1952; Hatai et a i, 1970; Kotaka and Ogasawara, 1974 Itoigawa and Tomida, 1982 Tomida, 1996 Makiyama, 1927; Hatai and Nisiyama, 1952; Hanzawa et a i, 1961; Hatai et a i, 1970; Kotaka and Ogasawara, 1974; Itoigawa and Tomida, 1982 Masuda, 1966; Kotaka and Ogasawara, 1974; Itoigawa and Tomida, 1982 Ozawa e ra/, 1986 8 CD " O O Q . C s Q . ■ D CD C O o ' 3 O 8 " O ( O ' 3 . 3 " CD CD ■ D O Q . C a O 3 ■ D O CD Q . ■ D CD ( / ) ( / ) g Table 2-3, continued moniwaensis Miocene Miyagi Pref., Japan Kotaka and Ogasawara, 1974; Itoigawa and JutJokai Miocene, m Miyagi Pref., Japan Tomida, 1982 Hatai et al., 1970; Kotaka and Ogasawara, *japonica Miocene, 1 Miyagi Pref, Japan 1974; Itoigawa and Tomida, 1982 Hatai and Nisiyama, 1952; Lindberg, 1992 kolioki Miocene, 1 Santa Barbara County, CA Hertlein, 1937; Bartsch, 1940; Keen and lasia Miocene, u San Luis Obispo County, CA Bentson, 1944; Woodring, 1932; Bartsch, 1940; Keen and palaea Miocene Los Angeles County, CA Bentson, 1944 Woodring, 1931; Bartsch, 1940; Keen and flemingt Miocene, 1 Kawau Isf, NZ Bentson, 1944 Powell, 1938; Eagle, 1996 Pleistocene Whakalane, NZ Beu and Maxwell, 1990; Lindberg, 1992 *iris Miocene Cape Rodney, NZ Harris, 1897 Miocene NZ Suter, 1.913»' Pliocene, u - Recent NZ Beu and Maxwell, 1990 Pleistocene Lyttleton, NZ BMNH 98048 tmthesommis Miocene, 1 Leigh, Rodney District, NZ Eagle, 1996 powelli Miocene, 1 Mt Luxmore, Fiordland, NZ Fleming, 1952; Lee e/o/. 1983^'^ Pliocene Komako, NZ Carter, I972\ Beu and Maxwell, 1990; Pleistocene Bay of Plenty, NZ Eagle, 1996 Fleming, 1952 waiiematensis Miocene, 1 Kawau Isf, NZ Powell, 1938'; Eagle, 1996 moorabolensls Miocene Moorabool River, Australia McCoy, 1876* Miocene Victoria, Australia Harris, 1897 : : o CD ■ D O Q . C g Q . ■ D CD C/) o ' 3 8 " O CD 3. 3 " CD CD ■ D O Q . C a o 3 ■ D O CD Q . ■ D CD C/) o ' 3 Table 2-3, continued Miocene,u mevosoides Miocene Miocene, u Pliocene Pliocene Pliocene Pliocene Miocene, u Miocene, 1 Miocene, 1 ; f Miocene, 1 ; f ♦ clathmta Reeve, 1846 Plio-, Pleistocene Miocene, lowest; Aquitan Miocene, 1 ; Elveziano Miocene Miocene, I: Elveziano Miocene, 1 ; Elveziano Miocene tubercuhfa lamellosoides oyinoides sp, sp. tuvuthaensis neuvlllei moniU/era anomiaefomis tuberoulata var. ovata Pliocene Miocene Miocene Miocene Flemington, Australia Flemington, Victoria, Australia Flemington, Australia Adelaide, Australia Adlaide and Melbourne, Australia Royal Park, Melbourne, Australia Flemington, Australia Victoria, Australia New Caledonia Fiji Fiji Miocene, u; g Guam Gironde, France T\irin, Italy T\irin, Italy T\irin, Italy Turin, Italy Miodoborow, W. Ukraine Miocene Piémont, Italy Italy Tétényer Plateau, Hungaiy Vienna Bassin, Austria BMNH G 1934 Lindbeig, 1992* BMNHG1935 Cotton, 1952; Ludbrook, 1954 Ludbrook, 1956 BMNH GG2373 McCoy, 1876; Harris, 1897 McCoy, 1876 Eagle, 1996 Ladd and HoBmeister, 1945; Ladd, 1966 Ladd and Hoflmeister, 1945 * Ladd, 1966 Tinian Ladd, 1966’ Ladd, 1966’ Delhaes, 1909 Delhaes, 1909 Gilbert, 1962 Delhaes, 1909 Delhaes, 1909 Davidaschvili, 1937; Krach, 1981 Cserhàt Mountains, Hungaiy f i d e Strausz, 1966 Delhaes, 1909 Delhaes, 1909 ; Ruggieri, 1990 Strausz, 1966 H D, Laatsch, personal commun. CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 ■ D CD 3. 3 " CD CD " O O Q . O 3 ■ D O CD Q . ■ D CD C/) C/) Table 2-3, continued tubercuhia volhynicQ Miocene Miocene Miocene S torrei 8p. sp. sp. sp. * gigmtea giganioides ♦ discus * dmrsicolor Torton/Saratian Miocene, 1 Miocene Tertiaiy Miocene Miocene, u Tertiaiy ? Miocene, u Miocene Miocene Miocene, u (?) Miocene Pliocene Pleistocene, 1 Pliocene Pliocene, u Pliocene, u Roztocze Lubelskie, Poland Poland and Italy Austria, Romania, Bulgaria, West Ukraine Buituri, Romania Austria and Hungary Cserhàt Mountains, Hungary Gaudemdorf, Austria Malta Phoukasa, Cyprus Lebanon Asti and Turin, Italy Sicily, Italy Mallorca, Balearic Islands, Spain Cyprus Cyprus T\irkey (Asia minor) Pliocene Ishikawa Pref, Japan Ishikawa Pref., Japan Okinawa Pref. Japan Ishikawa Pref, Japan Okinawa Pref, Japan Krach, 1981 Ghisotti, 1964; Ruggieri, 1990 Krach, 1981 Moisescu, 1955 Delhaes, 1909 Strausz, 1966 Hômes, 1856 BMNH 27501 BMNH GH3252-5 BMNH no U Homes, 1856 Ruggieri, 1990 H. D. Laatsch, personal commun. Hertlein, 1937 H. D. Laatsch, personal commun. Hertlein, 1937 Hatai et o/., 1970 Itoigawa and Tomida, 1982 Yabe and Hatai, 1941; Hanzawa et a i, 1961; Hatai et a i, 1970; Kotaka and Oga sawara, 1974 Itoigawa and Tomida, 1982 Itoigawa and Tomida, 1982 CD " O O Q. C g Q. ■ D CD ( / ) Table 2-3, conllnued 8 ■ D C Q ' 3- 3 CD 3. 3 " CD CD ■ D O Q. O 3 ■ D O CD Q. O C % % O 3 sieboMU * pourtalesi elsmerensis *Ju(gens craçhemdii Plio-, Pleistocene Pliocene, u Pliocene, early Pliocene, early Pliocene, 1 Pliocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pliocene Pliocene/Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Okinawa Pref., Japan Japan Riverside County, CA Los Angeles County, CA Los Angeles County, CA NWBC Los Angeles County, CA San Diego County, CA San Clemente Island, CA Orange County, CA BC BC Sur CA Punta China, BC San Diego County, CA Santa Barbara Islands, CA Santa Rosa Island, CA San Nicolas Island, CA MacNeil, 1960 Itoigawa and Tomida, 1982 Schremp, 1981 Yokes, 1935; Bartsch, 1940; Keen and Bentson, 1944 Woodring, 1931; Grant and Gale, 1931; Yokes, 1935 Rowland, 1968, 1972 Arnold, 1903; Grant and Gale, 1931; Woodring, 1931; Woodring era/., 1946 Arnold, 1903, Grant and Gale, 1931; Woodring, 1931; AMNH 12409; AMNH 12411 Lipps, 1967 Kanakolf and Emerson, 1959 Hertlein, 1937 Jordan and Hertlein, 1936; Emerson, 1980; Emerson enil., 1981 Lindbeig, 1992 ANSP 31487 Webb, 1937; Gilbert, 1962; this study: LACMIP loo. 12015; AMNH 12409 this study: LACMIP Iocs. 326, 329, 5068 Orr, 1960 Yeddcrand Norris, 1963 À CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 ■ D Table 2-3, continued 3. 3 " CD CD ■ D O Q . C a O 3 " O O CD Q . ■ D CD C/) C/) ru/escens Pleistocene, u Pleistocene Pleistocene Pleistocene Pleistocene Pliocene Pliocene Pliocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene San Clemente Island, CA Los Angeles County, CA Orange County, CA San Luis Obispo County, CA NW BC, Mexico Humboldt County, CA Ventura County, CA Los Angeles County, CA San Quintin, BC, Mexico Monterey County, CA Santa Barbara Island, CA Santa Rosa Island, CA Santa Clemente Island, CA San Nicolas Island, CA Anacapa Island, CA San Luis Obispo County, CA Los Angeles County, CA Susuki and Stadum, 1978 Chace and Chace, 1919; Willett, 1937; Woodring e/a/., 1946; Marincovich, 1976; this study: LACMIP loc. 10439 Kanakolf and Emerson, 1959 Valentine, 1962 Addicott and Emerson, 1959; Herlein, 1934; Valentine, 1957; this study: LACMIP Iocs. 23161,22715-22718. 8220- 8224, 10131.23162. Woodring, 1931 Woodring, 1931; Grant and Gale, 1931; Groves, 1991 Vokes. 1978 AMNH 12410 Gilbert, 1962 this study: LACMIP Iocs. 5066-5067 Orr, I960 Lipps, 1967 Vedder and Norris, 1963 this study: LACMIP Iocs. 24381,24383 Valentine, 1958 Chace and Chace, 1919; Grant and Gale, 1931; Woodring el a i, 1946; Valentine, 1962; Marincovich, 1976 à CD ■ D O Q . C g Q . ■ D CD C/) W o " 3 O 8 ■ D CD 3. 3 " CD CD ■ D O Q . C a O 3 ■ D O CD Q . ■ D CD C/) C/) Table 2-3, continued o 0 0 Pleistocene Orange County, CA Kanakolf and Emerson, 1959 Pleistocene San Diego County, CA Webb, 1937; this study: LACMIP loc. 11701 Pleistocene NWBC Valentine, 1957 Pleistocene BC, Mexico this study: LACMIP loc. 22723,22717 ♦ cf ru/escens Pliocene Isla Cedros, BC Jordan and Hertlein, 1926; Woodring, 1931 Pleistocene Isla Cedros, BC Jordan and Hertlein, 1926; Grant and Gale, 1931 Pleistocene Los Angeles County, CA Woodring, 1931 * corrugata Pliocene Ventura County, CA Woodring, 1931 Pliocene, m Ventura County, CA Grant and Gale, 1931 Pleistocene Orange County, CA Kanakolf and Emerson, 1959 Pleistocene San Diego County, CA Webb, 1937 * assimilis Pliocene CA Lindbeig, 1992 * walallemis Pliocene San Diego County, CA Hertlein and Grant, MS sp. Pliocene Los Angeles County, CA Woodring, 1931 sp Pliocene Ventura County, CA Woodring, 1931; Kern, 1973 sp. Pliocene San Diego County, CA Woodring, 1931 santaorwsensis Pliocene, u Santa Cruz, Islas Galàpagos Durham, 1979* *ovim Pliocene, Pleistocene Guam Ladd, 1966 Pliocene — Talmadge, 1974 Pleistocene Tonga: Tongatapu Osteigaard, 1935 * cf. varia Pliocene Kankasanturai, Sri Lanka BMNH G51083 ♦ (uberculata Pliocene Piémont, Italy Delhaes, 1909; BMNH G32151 Pliocene Palermo, Italy BMNH 31287 CD ■ D O Q . C g Q . ■ D CD C/) C/) Table 2-3, continued 8 " O CD 3. 3 " CD CD ■ D O Q . C a O 3 " O O CD Q . ■ D CD C/) C/) g Pleistocene Cyprus Fischer, 1993 Pleistocene Sicily, Italy Gilbert, 1962 Pleistocene Ravagnee, Calabria, Italy BMNH G31514 Pleistocene Panchia di Livorno, Italy BMNH G11529 Pleistocene Malta BMNH no # ? Favignana Isl. and Taranto, Italy Philippi, 1844 ? Sicily, Italy Philippi. 1844; Weinkauff, 1868 * cf. tubercuhia Miocene, m Loire Bassin, France Gilbert, 1962 sp 1 Pliocene / u Miocene Milazzo, Sicily, Italy H D. Laatsch, personal commun safdanhae Pliocene Langebaanweg, South Africa Pleistocene, Kensley, 1972 * australis Pleistocene, u - Recent NZ Beu and Maxwell, 1990 ? Gisborne, North Island, NZ Smith, 1977 * vlrghea Pleistocene, u - Recent NZ Beu and Maxwell, 1990 * cyolobates Pleistocene Fleurieu Peninsula, Australia Lindbeig, 1992 * emmae Pleistocene Victoria, Australia Lindbeig, 1992 * laevigata Pleistocene Victoria, Australia Lindbeig, 1992 ♦ rubra Pleistocene Victoria, Australia Lindbeig, 1992 ♦ hamtschatkana Pleistocene San Nicolas Island, CA Vedder and Norris, 1963 * c f sorenseni Pleistocene San Nicolas Island, CA Vedder and Norris, 1963 ♦ c f cracherodll Pleistocene San Luis Obispo County, CA Valentine, 1962 sp. Pleistocene Ventura County, CA Woodring, 1931 Pleistocene Santa Barbara County, CA this study Pleistocene NW BC, Mexico Valentine, 1957 *Julgens guadalupemlsPkisioceno Isla Guadalupe, Mexico Lindbeig g/ c;/., 1980 â 7 3 C D T3 S Q . C g Q . ■ D C D 8 T3 C 5 - C D C D " O S Q . C O T3 S C D Q . O C " O C D Table 2-3, continued barbadfnsis Pleistocene lamellosa Pleistocene Pleistocene Pleistocene sp. Pleistocene /subfossil ^(uberculata Pleistocene * (uberculata cooclnea Pleistocene * midae * unilateralis ' pustulata cruenta Pleistocene Pleistocene and Recent Pleistocene Pleistocene Pleistocene Pleistocene Pleistocene Whitehaven, Rugged Point and Spring Hall, Barbados Camargue, France Monte Pellegrino, Italy Sicily, Italy Lanaica, Cyprus Lanzarote, Canary Isl. Tenerife, Canary Isl. Algoa Bay, South Africa Port Sudan, Sudan Zanzibar, Tanzania Gulf of Suez, Egypt Gharib Lighthouse, Red Sea Mugarsim Isl., Sudan Zanzibar, Tanzania Trechman, 1937; Jung, 1968; Durham, 1979a; BMNH GG4018-26; BMNH GG1913; BMNH GG4001-6; BMNH GG3977; BMNH GG9012-16 » Paulus, 1949 Ghisotti, 1964 Malatesta, 1960; Gilbert, 1962 H. D. Laatsch, personal commun. Lecointre e/^A, 1967 TalaveraernA, 1978 Barnard, 1963 Hall and Standen, 1907 BMNHG41Î36 Newton, 1900 BMNH G24386 BMNH G35.128 BMNH G41537 Â Chapters: DNA Sequence Alignment in the Light of the Abductive Nature of Cladistic Hypothesis Generation. ... so wiederholt sich fOr das engere Problem der Homologiefeststel- lung dasselbe, was fiir die gesamte vergleichende Morphologie und die Systematik gilt, es setzte eine Période intensivster wertvoUer Arbeit ohne vorherige Klârung der Arfaeitsprmzipien ein. (... so happens the same for the more narrow problem o f the establishment o f homology that applies for the entire comparative morphology and systematics, a period o f most intensive, valuable work started without prior clarification o f the working principles.) Remane (1952:31 Chapter 2: The term homology and the criteria o f homology). INTRODUCTION The section quoted above firom Remane (1952) has a modem ring when applying his words to DNA sequence data. A wealth o f molecular characters has been acquired, but the debate as to how to use, or interpret, this information in the context o f organis- mal relationships is unsettled. Shall we use maximum likelihood, neighbor joining, or parsimony (e.g., HïlTîs e t aLy 1994)? Is transition/transversion weighting appropriate (Kluge, 1997a)? W ithin the cladistic paradigm, sequence alignment is a crucial step because it establishes some concept o f‘hom olo^’ m one of several competing incarna tions currently en vogue. Sequence alignment is performed by a number o f different computer programs (e.g., CLUSTAL ofHiggms & Sharp, 1988; MALIGN o f Wheeler 111 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. & Gladstein^ 1994) and, alternatively, also manually. Wheeler (1996) claimed to have developed an alignment-ftee phylogenetic analysis program. However, it is only align ment free in so far as alignment and phylogenetic reconstruction are implemented in a single computer program. The advantages and disadvantages o f certain alignment pro cedures have been considered in the framework o f minimising gap cost (Waterman et aL, 1991), maximizing computational efiSciency (Waterman et o f, 1991), and evolu tionary relevance (Gatesy et a l, 1993). Alignment has not, however, been discussed explicitly and rigorously {contra Mindell, 1991) in the context o f establishing primary homologies. The latter have also been termed ‘putative’ or ‘weak homologies’, ‘posi tional correspondences’, or ‘topographical identities’ (de Pinna, 1991; Brower & Schawaroch, 1996). These terms often indicate marginally to substantially different view points applied to the same observational qualities. hi the following the process o f DNA sequence alignment is illuminated in a bottom up approach. This necessitates the introduction o f some philosophical concepts on the nature o f observation, as well as a limited discussion o f the mode o f infrrence employed in cladistics as a whole, because the observational phase needs to be framed relative to the entire process o f phylogenetic inference. The homology concept serves as a crucial reference point, because it is one o f the central, mdeed probably the most important, tenets o f phylogenetics. It will become cl«ir upon closer inspection that most current practices in DNA sequence alignment can not be upheld, appropriate alternatives are suggested throughout the chapter and are summarized at the end. MOLECULAR AND MORPHOLOGICAL CHARACTER EQUIVALENCE An hnportant question to start with is whether morphological and molecular data, are equivalent and whether these data «m, must, or must not be treated in a comparable 111 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. fashion. Figure 3-1 illustrates the pathway by which data—fflorphological as well as molecular—are treated. Here the commonalities o f the different data types in terms o f observation and explanation are emphasized. The observational phase From an epistemological standpoint, the acquisition of basic observational knowl edge can be characterized as a three-step psychological process: sense perception, per ceptual belief formation, and classification (Audi, 1998). Using the example in Figure 3-1, the general aspects o f observation are outlined. Individual specimens comprise the source o f all observations (Figure 3-1: abalone moUusk at top). Regardless o f specimen preparation, original observations manifest themselves in the form o f sensory perceptions. At its most basic level, sensory percep tions are registers o f our surroundings. As such, one does not perceive individuals per se, but particular properties expressed by those individuals. This modus operandi of perception accounts for our practice o f describing organisms in terms of'character states.’ In contrast, living and inanimate objects only possess characters, not states. It would be quite impossible to have, say, a character 'gills’ separate fiom the mdividual subsidiary components that make up a 'branched gill’ as opposed to a 'filamentous gill.’ The latter is nothmg more than a specifier or a predicate o f the former. For the sake of historic consistency, character and state here are used here. In our example (Figure 3-1), a scanning electron micrograph (SEM) o f a radula fi:om an abalone is used as a generic place holder for morphological data, and a ficti tious read-out fiom an automated sequencer or the manual reading o f a gel stands for sequence data. The direct or indirect sense perceptions lead immediately to perceptual beliefo; the properties observed are independent o f the observer as opposed to being 113 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Specimen Observation Perception % BeUef formation special similarity positional similarity I 001101001001101 assifîcation 0 0 0 1 0 1 0 0 1 0 0 1 0 0 1 0 0 0 1 1 1 0 0 1 1 0 0 0 0 1 Explanation Elementary hypotheses specimen preparation AACGTACGTACCG special similarity I positional similarity AACGTACGTACCG AA-GTACGAACCG AACGTTCGTACCT parsimony Composite hypothesis Figure 3-L Treatment o f morphological and molecular characters in cladistics with respect to homologr. For details see main body o f texL 114 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. hallucinatory events produced by the observer. It is on the basis o f our perceptual beliefs that we claim particular properties, i.e., states, to exist In other words, we engage in the process o f naming those states by way of a predicate language. The act o f naming prop erties observed o f organisms is a base level act o f classification. Here the term ‘classifi cation’ is used not in the conventional systematic sense o f ranking taxa, but in the sense o f grouping observations. The step fiom perception to classification within the observa tional phase is the process of belief formation using auxiliary information (see below). Are morphological and molecular observations equivalent as observations? One might make a distinction between the two on the following grounds. 1) Molecular data do not consist o f transformation series. Bases can not show intermediate characteristics, for which reason they can not be treated like morphological data. The following, non- molecular examples, which show no intermediate character states, invalidate this argu ment: chromosome duplication, doubling o f perianth, addition/loss o f segments. 2) Ordering character states in sequence data is impossible. It has been argued that more explicit hypotheses o f primary homologies are investigated, as the information con tained in a fixed sequence o f character-state transitions is higher than that in an unordered set o f character states (Wise & Strong, 1997). The use o f ordered characters may be viewed as a powerful tool, but it also harbors the potential for introducing notions on the evolution o f character states that lack an empirical basis (Emberton, 1994; Sundberg & Hylbom, 1994; Wise & Strong, 1997). The above distinctions are viewed as unremarkable and not warranting a discrete morphological-molecular dichotomy. Consequently, all data as perceived shared similarities stemming fiom observations can and must be treated in an equivalent form, assuming our goal is the causal explanation o f the phenomenon o f shared similari^. As a corollary, commonali ties o f all observations need to be highlighted. 115 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Note that the perception and the classification o f a property do not convey a homol ogy statement The gray-scale values o f each pixel in a SEM TIFF file is as little a pri mary homology statement as the bases in the original sequence read-out o f any one taxon. The comparison o f a set o f SEM micrographs or a number o f sequences in order to determine similarities still does not invoke primary homology, as no causal explana tion is sought y et To equate basic, named shared similarity (te., ciassificatory events) with homology is to remove any notion of causality. And, to remove causality is to remove any need for a term such as ‘homology’. The explanatory phase From the observed distribution of named states o f any single character, an elemen - tary explanatory hypothesis o f homology is inferred. That homology hypotheses are explanatory resides in the Darwinian accounting o f shared similarity by way o f com mon ancestry, which incidentally, is nothing more than a replacement o f Owen’ s earlier causal accounting by way o f archetypes. The very process o f reasoning from some set o f observed effects to an hypothesized cause, has typically been referred to as abductive inference, or “inference to the best explanation” (Harman, 1965; Fann, 1970). Unfortu nately, the specifics o f inference with respect to homology, as well as cladistics in gen eral, have been at best poorly characterized. The importance her^ however, is simply to point out that the inference o f explanatory hypotheses is neither a deductive nor an inductive form o f reasoning (Kluge, 1997b:92). This because the very structure o f such hypotheses invokes unobserved (not necessarily unobservable) entities, in this case common ancestors, to causally account fer what is observed, a particular character state distribution. 116 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The claim that mmimum-Iength cladograms are hypotheses with maximiim explana tory power has been a fundamental tenet held by many advocates o f parsimony. The abductive nature o f homology hypotheses is not only consistent with this view, but in fact, the claim o f explanatory power is completely dependent upon the abductive infer ence both from observed shared similarity to homology (= elementary hypothesis), and from homology to cladogram(s). In other words, cladograms are composite explanatory hypotheses based on the conjunction o f all elementary hypotheses. Note that the (Dar winian) inference o f homology is the association o f common ancestry, i.e., cause, with shared similarity, or effects. The inference of cladograms from homology is nothing more than an additional abductive procedure wherein all elementary hypotheses are treated together applying the principle of common cause (cf. O’Hara, 1998). Abductive inference, or a logic of discovery {sensu Reichenbach, 1951), is used to infer first ele mentary and subsequently composite hypotheses, hence, nested abductive reasoning is employed (Josephson & Josephson, 1995; Richter, 1995; Fitzhugh, 1997; Fischer, 1997:375; Moser et a l,, 1998). The question asked during the explanatory process is, ‘why are the character states distributed as they are?’ A primary homology hypothesis may be shown to be a ‘confirmed’, ‘strong’, or ‘secondary homology’ explained by common ancestry when considered in light of all shared similarity in need o f explana tion by way o f common ancestry. Otherwise a case o f homoplasy has been invoked (Farris, 1983). As an additional outcome o f the generation o f a composite explanatory hypothesis, one may even say a by-product, a pattern o f relationships among taxa is revealed. One must be keenly aware that relationships among organisms are only a short hand form fi)r the causal accounting o f the character-state distributions among taxa. Unfertunately, it is the explanatory basis o f cladistics, as well as hom olo^, which seems to have been overlooked by proponents o f methods other than parsimony. 117 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Interestingly, the basis for the parsimony criterion in cladistics is not to minimize homoplasy, but to apply as consistently as possible the very causal notion o f common ancestry used to infer homology. Any inference from effects to hypothetical cause(s) requires the association o f those effects with some causal theory. To insist that homol ogy, as well as cladistics, function in some theory-free, or model-neutral realm removes the ability to explain and, therefore, takes homology and cladistics out o f the realm of science. The inference o f a composite explanatory hypothesis requires an inference rule that is consistent with the major premise, or theory, used to infer elementary hypothe ses. General evolutionary theory relative to inheritance and spéciation justifies the use o f parsimony as the criterion for hypothesis selection. This is nothing more than the application o f the principle o f the common cause (e.g.. Sober, 1988). To do otherwise would be to call into question the very homology hypotheses one has at hand, which would obviate cladistics altogether. If multiple sources o f mformation are at hand, all of which require explanation by common ancestry, then they are combined in a ‘ total evi dence’ approach to obtain the (composite) hypothesis o f relationship with the ‘maxi mum explanatory power’ for the character-state distributions in the form o f a cladogram (Kluge, 1989; Kluge & Wol^ 1993). Unfortunately, the issue o f‘total evidence’, as well as ‘explanatory power’, have lacked any lucid philosophical underpinnings in the cladis tics literature, except in the case o f Kluge (1989) for the former. These matters will be treated elsewhere (Fitzhugh, in prep.). To propose similarity perceptually^ and to explain similarity with unobservables (Le., ancestors), thus hypothesizmg prnnary homology, are entirely separate inferential events, though, iu practice, they are treated as ^simultaneous to form a cohesive unit. An accepted similarity statement autom atical^ opens the path to causally account for those similarities by way o f common ancestry, as primary homologies. All observations 118 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. entering an analysis must be treated in an equivalent manner, as all observations are subject to the same goal in cladistics: the causal explanation o f shared similarity. Obser vations that do not satisfy the criteria o f similarity (special similarity, positional corre spondence: see below) are not included in the ensuing parsimony analysis. If the treatment o f observation conflicts with the concept o f homology, then these observa tions can not be included in a cladistic analysis unless the empirical basis can be estab lished that one should not trust one's perceptual beliefs. On the other hand, to make such an assertion impinges not on cladistics, but on the very basis o f inferring homol ogy. Cladistics as science Cladistics does not follow the traditional Popperian concept o f hypothetico-deduc- tivism (contra Freudenstein, 1998:97). As no hypothesis is initially available, it can nei ther be confirmed, nor rejected, nor statistically tested. “Accordingly, there is no point in engaging in such fashionable academic probabilify games as musing about the prob ability o f phylogenetic trees...” (Mahner & Bunge, 1997:48). Kluge (1997b), arguing that cladistics embodies Popperian tests, used the term ‘test’ in two different senses: A) provide reasons to select the most parshnonious tree against the hypothesis o f a bush, which is not a Popperian test; B) evaluate an established phytogeny with new data, which again can not constitute a test since no test can rest solely on the very effects the hypothesis it is intended to explain. In contrast to Kluge’s position, the actual evidence Jbr so-called Popperian corroboration resides not in character-state distributions among term inal taxa, but must be those independent, subsidiary effects that must exist as a result o f the specific initial causal conditions. Le., the character states in the postulate ancestors, that lead to shared similarity. For the most part, thetestmg o f historical expla- 119 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. nations was a matter o f little concern to Popper. The time spent with continued attempts to force Popper’s views o f science into history would be better served by investigating the philosophical studies of explanatory hypothesis testing. Either use o f‘test’ in cladis tics does not constitute a deductive test as demanded by Popper. Interestingly, Kluge (I997b:92) also pointed out that “[h]ypotheses can never be proven true, as inductivists seek to do, nor proven false, as deductivists claim to be able to do.” If cladistics employs neither induction nor deduction, indeed neither provides the inferential ability to gener ate explanatory hypotheses, then only abductive inference can be employed to address observations in need o f causal explanation by which an hypothesis o f relationship pro viding the maximum explanatory content is generated. As the ultimate goal o f science is to causally account for observed phenomena (Popper, 1979:191; Moser et a l, 1998; Salmon, 1998), abductive, i.e., non-deductive, reasoning employed in cladistics is fully compatible with phylogenetic inference being a branch of science. Figure 3-1 smnmarizes several o f the points made above on the nature of cladistics, comparing explicitly the treatment o f morphological and molecular characters. It is nec essary to properly understand the entire sequence o f actions during cladistic analysis, even if only a small part is treated here in detail. The following points are of particular concern for the discussion o f sequence alignment presented here: • Objects are perceived as sets o f properties; • All character states are comparable as observations; • Homology is abductively inferred; • Sequence alignment is the process establishing shared similarities (see below); • Observation leads from perception to classiScation by way o f perceptual belief for mation (see below). 120 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. OBSERVATION: FROM PERCEPTION TO CLASSIFICATION As noted earlier, observation can be separated into two distinct phases: sensory per ception and classification connected by belief formation. Perception is applied in the sense o f uninterpreted, sensory input, which has also been termed ‘sensation’ by Mah ner & Bunge (1997). The matter is treated in a simplistic fashion here and do not address the question o f whether perception/sensation itself is selective and, therefore, an inter pretation o f facts; see e.g., Campenhausen (1993) and Josephson & Josephson (1995) for discussion. The two phases of perception and classification are linked by the mental process o f belief formation (Audi, 1998). One readily recognizes a picture o f a flower by comparison to one’s mental library, but a computer would have difflculties to iden tify a flower flx)m a graphics file. We arrive at the conclusion that this assembly o f pix els represents a flower by forming a belief using auxiliary information (see below). Indeed, it has been recognized for some time that belief formation is itself an applica tion o f abduction (Devitt, 1997). By labeling entities with a name, which is a classifica tion process, one proposes shared similarity. Similarly, with DNA sequences, the original read-out—the sensory input—must be converted using auxiliary information before the individual bases can be labeled shared similarities. Sequence alignment is part o f the process o f observation leading fiom perception to classification, hi that process gaps, which represent the absence of bases, are introduced in many instances. Nelson & Platnick (1981) have pointed out that the absence o f a character state can not be observed. One does not note the absence o f legs in snakes, but the continuity o f the bo<fy wall. Corresponding arguments can be applied to gaps. What we can observe m DNA sequences is the adjacent position o f two bases, or their homologues being set apart by inserted bases. Acknowledging Nelson & Platnick’s (1981) point, the conventional use o f ‘absence’ and ‘gap’ is continued here, despite 1 2 1 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. being somewhat imprecise. As gaps are introduced during the observational phase, gap^ can not be viewed as missing data, but are observations o f absence of a base as the char acter state: gaps should be treated as a fifth, character state. The same procedure has long been employed for morphological characters. Consider the legless condition o f snakes as compared to other reptiles. If snake taxa were coded for legs, then legs would be postulated as one of the possibilities during character state optimization, which is in conflict with the original observation. The Tegless’ condition is a clear observa tion, if for no other reason than being a statement as to body form, which must be coded as such using a separate state. The same reasoning applies to gaps. Determining character states and identifying the character to which the states belong are common to both morphological as well as molecular data. These two steps do not exist seperately, but are predicated upon one another. The two are part of an elementary classification system where various observations (= states) are united under the collec tive term o f character. More formally, a character is nothing but a collective set o f states (Goodwin, 1994). Note the necessary unity o f the classification process: one can not identify a character without an underlying observation; and, what we observe are states, not characters. This process establishes shared similarify, which is open to explanation in terms o f primary homology via common ancestry. Examples: A) character state known, position in question: this structure is yellow. In which position is this structure found? In the position o f the petal (relative to other structures we identify a p rio ri as flower-like properties). The petal is yellow. This DNA base is a G. In which position do I find it? It is in position 213. Base 213 is a G. Or, alternatively B) position known, character state in question: this is a petal. Which color does it have? This petal is yel low. This is position 213. Which base is found in this position? Position 213 is a G. The process o f identifying stalK o f particular characters is fundamentally one and the same 122 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. process for any observation. Petals do not exist separate from the colors (and other properties) that allow for recognizing petals. Bases are only meaningfiil when classified in a particular position. ‘Organism X has a G’ is pointless, but ‘organism X has a G in position 213’ carries information. Observed similarities leading to primary homologies are established using various additional information. Although the supplemental information used may depend on the type o f data, the common theme is that auxiliary information is used for the identifica tion o f these similarities (see Remane, 1952). Two pathways can be identified. i: Special sim ilarity Perceptual similarities are based on special, associated properties being shared. The associated properties are used to establish the similarity o f the observations, which results in grouping states in characters. This special similarity argument is the most fre quently used means for identifying similarities of morphological characters. Such an assessment is trivial with DNA sequences because the four bases are identical to their respective molecular structures. n: Positional correspondence Remane (1952) used the agreement-in-position o f a (morphological) character within the fiiamework of the organism as the argument for similarity (see also Hawkins ef fl/., 1997). This is the only applicable argument that can be extended to DNA-sequence alignment (Efillis, 1994; Swof&rd e t aL^ 1996). Although the concept o f positional cor respondence was origmally develop»! for morphological characters, its application to other types o f characters, including DNA sequences, is straightforward (Brower & Schawaroch, 1996). 123 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Sequence alignment Three processes relevant to the context o f sequence alignment can lead to differ ences between two sequences: mutation and insert/deletion (= indels). The goal o f sequence alignment is to distinguish between these two sets o f events in order to deter mine instances o f shared similarity (c f B C Iu g e, I997a:352). Four major methodological approaches can be considered. I) Mutation or indels only. All evolutionary change is attributed to either o f the mechanisms to the exclusion o f the other. Such models are considered extremely unlikely and are not used. It is mentioned here for the sake o f completeness. 2) Gap-cost function. A cost ratio for mutation to indel is applied and the least costly alignment is considered the most ‘likely’ (Li & Graur, 1991; Waterman et aL, 1991). 3) Parsimony-based. Parsimony is applied to &id the alignment requning the fewest number o f character state transitions by employing a gap-cost ratio as well (Wheeler & Gladstein, 1994). 4) Manual alignment. The investigator matches the bases as well as possible by eye. The exact arguments for manual alignment can not be speci fied, but may well be a mncture o f gap-cost function and parsimony. Much debate has centered on which alignment parameters are the most appropriate, because it is widely appreciated that different parameters yield different alignments, i.e., propose different shared similarity (e.g:, Gatesy et a/., 1993; Hillis, 1994; Bridge et a f, 1995; WSgele & Stanjek, 1995; Wheeler et o f, 1995; Wheeler, 1995). One might prefer m anual a lignm ent as it forces one to consciously and critically evaluate every observation. Manual alignment also forces one to recognize areas o f problematic alignment An initial computer-facilitated alignment o f any sort may reduce the manual labor, but as the critical work is done subsequent to the rough estimate, it becomes irrelevant which method o f initial computer-facilitated alignment is chosen. 124 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. The pertment issue is, however, the philosophical and epistemological bases for align ment as a similarity function. OBSERVATION AND EXPLANATION Homology is inherently comparative by the very fact that such statements are explanatory hypotheses, i.e., homology can not be applied to data from a single speci men, except in terms o f individual ontogeny. Homonomy (= serial “homology”) is not considered here and the discussion is restricted to interorganismal homology. The com parison o f sequences, hence, sequence alignment, leads to elementary explanatory hypotheses in the form o f primary homology statements. As Hawkins et a i (1997) have pointed out, the data matrix can be viewed as a set o f assembled, primary homology statements. Primary and secondary homology Primary homologies are shared similarities causally accounted for through the pos tulate o f a common ancestor at the level o f each elementary hypothesis. At this level, homoplasy is an irrelevant issue. Secondary homologies are also shared similarities explained though a common ancestor, but in the context o f a composite hypothesis. The latter, in the form o f a cladogram, is not a simple summary statement o f all elementary hypotheses, but a new hypothesis generated from all relevant evidence (cf. Wenzel, 1997). Relevance relates directly to the issue o f total evidence, as only the total relevant evidence must be considered in non-deductive inference. Note that total evidence and relevance do not apply to deductive inference. Secondary homologies refer then to a restricted set o f explanations, i.e., those that are explamed by a common ancestor in the composite hypothesis. Homoplasies, however; despite being explanatory (Farris, 1983; 125 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Siddall & Kluge, 1997:317), do not invoke a common ancestor at the point where they are shown to be homoplasious. Hence, the crucial point o f homology is the explanation o f shared character states by means of conunon ancestors at that particular leveL The terms primary and secondary homologr fiilfiil these conditions at their respective lev els, hence, their use in phylogenetics is appropriate. Separation of power Observation, and explanation of observations, need to be independent o f each other (Hawkins et o f, 1997). The observational phase must not dictate a preconceived expla nation not evident in the observational process itself upon the observations. Otherwise the observational phase would interfere with the explanatory phase of the very observa tion. For morphological characters this problem has long been recognized in coding character states for a structure, such as 0: small, 1: large, 2: secondarily reduced. If the original observation ^smalT is divided into two subcategories 'sm all' and 'secondarily reduced’ despite no differences between the two conditions being discernible, then an explanation o f the condition 'secondarily reduced’ by way o f reversal is included. Clearly, such coding practices are unacceptable and are based on preconceptions about the distribution o f the data. The preconception o f introducing an explanatory element into the observation may either originate outside the data (e.g., following the “estab lished” classification scheme), or may be derived fiom a cursory glance at the data matrix where some pattern is visually detected. Leglessness of some lizards and snakes must be coded the same, even though we in&r fiom a wealth o f other observations that it is a separately derived condition in the lizards. Otherwise we take the data beyond the observation o f the absence o f legs, and already include an explanation for the distribu- 126 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. don o f character states. (One may argue for the exclusion o f this particular character, because it has already been explained, but this is a separate issue; see below.). Confusion about the separation o f observation and explanation in phylogenetics, includmg the necessary priority o f observation over explanation, dates back to Huxley, and this confusion continues to this day (Brady, 1994). For instance, it has been argued that sequence data afford the unique possibility to combine the two steps—establishing shared similarities with the explanatory equivalents of primary homologies (= elemen tary hypotheses) and to explain them in a composite hypothesis—into one single proce dure. In one case, sequence alignment is said to be abandoned altogether by a direct form o f character state optimization (Wheeler, 1996; Wheeler & Hayashi, 1998). In a second approach, parsimony is used as the optimality criterion to align sequences (MALIGN: Wheeler & Gladstein, 1994) or to determine the sequence input order (Knight & Mindell, 1995). What is overlooked is that the establishment o f primary homologies as elementary hypotheses is necessarily prior to the composite hypothesis, and the establishment o f shared similarities must be independent for each character (see below), whereas the explanation o f observed shared similarity at the level o f the com posite hypothesis is carried out m the framework o f all data under consideration. Mindell (1991) argued that, because the ‘test’ o f congruence through parsimony minimizes homoplasy and respectively maximizes secondary homologies, the same maximizing methodology can also be applied to the establishment o f primary homolo gies in the form o f sequence alignment Although the establishment o f shared similari ties and their subsequent explanation at the level o f a composite hypcth^is are contained in one logical sequence, the two steps must remam separate. The same criterion (parsi mony) can not be used to find shared snnilarities, using as arguments for the determina tion o f particular similarities all positions across all taxa o f a particular sequence, and to 127 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. causally explain the very same observations, using again all positions across all taxa of that particular sequence, because then circular reasoning ensues (Brady, 1994; Wheeler, 1995). This problem applies only if all data entering the observational phase are the sole bases & r the explanatory process. This is the case with studies using a single, par simony-aligned sequence to the exclusion o f any other information, which is then ana lyzed using parsimony (e.g., Wagele & Stanjek, 1995; Wheeler, 1995). Using parsimony to align sequences is comparable to the above mentioned cursory glance at the data matrix, only that the entire data matrix is relied upon for the establishment of shared sim ilarities. The data matrix is the source o f the bias. However, as soon as a single synapomorphy is added to the parsimony-aligned sequences, circular reasoning is no longer an issue. The parsimony argument at the explanatory level provides the best explanation for a more inclusive data set, not only for the parsimony-aligned sequence. However, the problems with character independence are still valid. CHARACTER INDEPENDENCE One o f the central tenets m character coding is character independence. The term ‘independence’ is used here in the sense o f potential of changes in character states not being imder the influence o f any other state; independence is distinguished &om the existence o f one state being predicated upon the existence o f a second, the latter being better described as the problem o f inapplicables. Separate characters as groupings o f sets o f states should only be considered if the set o f states in each character has the potential for independent evolution (e.g., Kluge, 1989; Brower & Schawaroch, 1996; Hawkins e t o f, 1997; Luckow & Bnmeau, 1997). The co-varmtion, or congruence, o f multiple characters expected from common ancestry is not an indication for non-inde pendence in the above sense; but is a matter o f causal association by at least common 128 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ancestry. This distinction can also be characterized as the difference between an inter - active fo rk (non-independence) and a conjunctive fo rk (co-variation due to common ancestry: Sahnon 1998:296). As coded characters are independent, the inference of pri mary homologies for each individual character must be based on relevant auxiliary information (special similarity and positional correspondence: see above). Consider a problematic bone in the skull o f some vertebrate. To establish its shared similarity we only consider features of the skull, but we do not take into consideration the hand or the rib cage. The latter are found beyond certain landmarks such as the non-arbitrarily iden tifiable neck, shoulders, and elbow. On the other hand, we can use position, structure, histology, etc., o f a skull bone and its surrounding components to determine similarity, thus homology. In more abstract terms, the auxiliary information permissible to estab lish primary homologies is restricted in width, but potentially open in depth. In molecular data the only available auxiliary information is positional, i.e., adja cent regions o f the particular nucleotide provide the basis to postulate shared similarity. It is inappropriate to use auxiliary information beyond a secure anchoring point such as conserved regions o f DNA. Therefore, characters beyond a conserved DNA region are ineligible to influence the establishment o f the stretch within. It is well-known that different genes are optimally aligned with different alignment parameters (gap weights: see below for further detail), so also various regions in one and the same gene may align “best” under different alignment conditions. To ferce one single set o f alignment parameters on the entire sequence may deny opthnal alignment to variable regions between conserved ones. From a process perspective, the same evo lutionary conditions (e.g., mutation rate-, transition/transversion bias) are assumed to hold for the enthe sequence. The parameters that produce an overall optimal alignment may be suboptimal for some regions, regardless o f whether the minimization function 129 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. is gap-cost or parsimony. In other words, process independence for each region is ill considered, resulting in a violation o f character independence. The alignment process should pertain to the smallest fragments between adjacent conserved regions, which may be termed minimal fragment alignment (MFA). Alternatively, one might ask what is comparable to a global minimization with morphological characters. There is no com parable procedure. Wheeler & Hayashi (1998) argued for using multiple gene sequences and even mor phological data to help with sequence alignment. They pointed out that congruence is a well established decision making argument in phylogenetics, hence, the congruence between different data sets helps to establish similarities. The transition / transversion / gap ratio resulting in the best Mickewich-Farris metric between the data-sets was used. Congruence, indeed, has a sound grounding in cladistics, but at the level o f the clado gram construction, i.e., the explanatory phase. The establishment o f similarity, however, is carried out in the observational phase. Second, the issue of relevance arises. At the level o f cladogram construction all character-state distributions with their similarities already identified are relevant. However, when the similarities are established only the appropriate auxiliary information is relevant. Clearly, morphological characters are entfrely irrelevant when considering sim ilari^ issues at the level o f DNA sequences. How are conserved regions to be found? To identify them, indeed, a larger part o f the sequence must be scrutmized, which stül is not an argument to use global alignment for individual bases. We encotmter the hierarchical nature m the relation o f objects and their parts (part-whole relationship). We first need to identify the similarities o f more inclusive structures (e.g:, gene, skull), then intermediate ones {e.g., conserved region, neck), before detailed questions can be addressed (e.g:, base 213, bone A). The auxil iary information appropriate for each hierarchical level is chosen, keeping in mind the 130 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. scope o f the problem addressed. The auxiliary infbrmatioa deemed appropriate can not be determined with mathematical accuracy. However, some information is clearly inap propriate such as variable sites beyond a conserved region when aligning individual positons. Mindell (1991) asserted that primary homologies established probabilistically through sequence alignment are o f binary nature, i.e., qualitatively related so that struc tures are either considered to be homologous or not, but that none is labeled ‘87% homologous.’ Mindell’s rationale for the sudden transition from a probabilistic to a binary statement can not be followed, but it is taken at face value for the sake of the argument The question arises o f what would have to be done with morphological data to ensure equivalent treatment o f all data? The morphological matrix would have to be aligned using parsimony (with the characters arranged anterio-posterior or dorso-ven- tral?). The interpretation o f potentially resulting gaps would be challenging. As the approach is nonsensical for morphological data, the comparable treatment of all data is lost. Mindell’s (1991) point is rejected. OBJECTIVITY AND SUBJECTIVITY Character selection and observation Sets o f molecular data are acquired by the use o f certain pairs o f primers. This choice o f prhners is a willful act, therefore; inherently subjective, as is the case with observa tion m general. The subjective choice is not limited to the stretch o f DNA as an entity, but includes every single base. Character sets in need o f explanation are chosen for morphology as well as in the case of genes or gene regions (cf. Salmon, 1998:306 on the relativity o f relevant information). The choice o f characters whose shnilarity is to be determined is related to the part-whole relationship o f objects (Skull: dermatocra- 131 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. niinn: bone A. Gene: gene region: base 213). Features for which, shared similarity is not clear (see below: Belief formation), hence, that can not reasonably be suspected to be sim ilar between taxa, do not need to be explained. A character state distribution is already accounted for by virtue o f recognized non-similarity, and consequently non- homology, and is no longer in need of explanation at the level o f cladogram construc tion. The particular point is further explored when addressing the problem o f character inclusion and exclusion below. Al^nment parameters With molecular data, character states are given as a sequential reading o f bases. The goal o f alignment is the assigmnent o f a limited number o f shared character states to linearly arranged characters. The set o f alignment parameters (gap weight with or with out different extension costs: see Waterman et al., 1991) is an assumption entering the analysis. These alignment parameters can also be viewed as the the quantifiers o f an evolutionary model. As alignment parameters or the underlying evolutionary model can not be observed directly from DNA, they are extraneous to the data. Like any other weights (character weight, transition/transversion weight) they constitute assumptions, are inherently subjective, and generally are non-empirically Justifred. The advantage of computer-facilitated over manual alignment is the explicitly stated assumption in the form o f specific alignment parameters (Gatesy et aL, 1993). The comparison to mor phological observations will be taken up in the section ^Belief formation’ below. Objectivity seems to be some general goal in molecular phylogenetics (Messenger & McGufre, 1998:93) and is briefly discussed in Moore & Willmer (1996). It is claimed to be obtained in sequence alignment through a so-called sensitivi^ analysis (Wheeler, 1995). Hereby, a number o f parameters are applied in order to investigate whether or 132 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. when the alignment changes. I f a wide range o f parameters results in the same align ment, it is said to be more ‘robust’, giving results the guise o f objectivity. If a change of parameters results in a different alignment then the operator is faced with the question o f which parameters / alignment to choose. An important question in sensitivity analy sis is at which interval and over which range the alignment parameters are used. Inter vals may be subjectively chosen as integer numbers on a linear scale, or on a log scale with bases such as 2, e, or 10 (Wheeler, 1995). The range o f values for gap weights has a logical lower limit o f 0.5 because of the triangle inequality (Wheeler, 1993), but has an open upper bound. The only logical upper limit is infinity; i.e., change by mutation only. Most workers would argue that the infinity boimdary is unrealistic. Restricting the range o f alignment parameters used to less than the objective 0.5 to infinity automati cally introduces subjectivity (Gatesy et aL, 1993). Whether the range is explicitly restricted in computer facilitated alignment or while performing manual alignment is irrelevant to the question of the subjective choice of parameters. Manual editing In many studies, a compute- generated alignment has been subsequently edited man ually (e.g., Collins et aL, 1994). Such a practice can not be justified in the context o f objectivity. A single alteration o f a computer-generated alignment instantaneously for feits the advantage o f the explicitly specified assumptions. ALTERNATIVE CODING STRATEGIES Unambiguous alignment is always unproblematic. The clearest case is with con served regions harboring uninformative states. As soon as regions o f unequal length are compared, we may face questionable or ambiguous alignment. Statements o f shared 133 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. similarity, thus primary hom olo^ hypotheses, may depend upon the alignment algo rithm and parameters chosen. Even with a particular setting, multiple, equally parsimo nious or costly alignments may be found, although not many programs report more than one. Ambiguous alignment can be addressed using various, flexible coding strategies. Each strategy is considered and the consequences for homology are illustrated with simple example. This example consists of four hypothetical sequences (Figure 3-2: Original data). The sequences are characterized by initial and terminal conserved regions o f five bases each, adjacent to variable regions spanning two positions, i.e., six and seven. In the variable region, taxon 1 shows a CC, and for taxon 2 a double gap (— ) is found. Taxa 3 and 4 show one base each, C and A, respectively, and one gap. The absolute and rela tive positions o f base and gap are unresolved for the last two taxa (Figure 3-2: Four possible alignments). Two problems relating to homology wül be encoimtered throughout Figure 3 -2 .1) If any character state in one taxon has more than one homologous state in another taxon, i.e., the latter is coded in more than one column, then the test of conjunction is failed (de Pinna, 1991). This situation is indicated with underlined positions. 2) Contradic tions in possible character state optimizations with original observations are flagged by the position in italics. Elision Finding a consensus o f multiple alignments resulting firom the choice o f different alignment parameters has been addressed by Wheeler et al. (1995), who proposed a method called 'elision.' The alternative alignments for each taxon (Figure 3-2: Four possible alignments) are appended to one another (Figure 3-2: Elision) and analyzed 134 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Origmal data A T C T G C C A C Q T A C A IC T G A C G T A C A IC T G C A C S T A C A IC T G A A C G T A C Four possible alignments A T C T G C C A C G T A C A T C T G C C A C G T A C A T C T G C C A C C T A C A T C E G —A C C T A C A TCtG —A C G T A C A T C T G —A C C T A C A T C T G -C A C G T A C A T C T G C -A C G T A C A T C T G -C A C G T A C A T C T G -A A C G T A C A IC T G -A A C G T A C A T C T G A -A C G T A C A T C T G C C A C G T A C A T C T G —A C G T A C A T C T G C -A C G T A C A T C T G A -A C C T A C Elmon ATCTGCCACCTACATCTGCCACGTACATCTGCCACGTACATCTGCCACCTAC ATCTG— ACGTACATCTG— ACGTACATCTG— ACGTACATCTG— ACGTAC ATCTG-CACGTACATCTGC-ACGTACATCTGC-ACGTACATCTG-CACGTAC ATCTG-AACGTACATCTG-AACGTACATCTGA-ACGTACATCTGA-ACGTAC Case sensitive: a,c=A/T/G/0- Symptesioinorpfues removed CCCCCCCC — CC— C— c -A-AA-A- Recoded: A T C T G C C A C G T A C A T C T G —A C G T A C A T C T G O -A C G T A C A T C T G -àA C G T A C Opnmizadons: ATCTGCCACGTAC ATCTGCCACGTAC ATCTGCCACGTAC ATCTGCCACGTAC ATCTG— ACGTAC ATCTG— ACGTAC ATCTG— ACGTAC ATCTG— ACCTAC ATCTG— ACGTAC ATCTGC-ACGTAC ATCTGC-ACGTAC ATCTG— ACGTAC ATCTG=iACGTAC ATCTG=ACGTAC ATCTG-CACGTAC ATCTG-CACGTAC Missing data: ?=A/T/G/0 Recoded: A T C T G C C A C C T A C A T C T G —A C G T A C A T C T G ??A C C T A C A T C T G ?? A C C T A C Optimizations: A T C T G C C A C G T A C A T C T G —A C G T A C A T C T G —A C G T A C A T C T G C C A C G T A C A T C T G C C A C C T A C A T C T G —A C G T A C A T C T G = A C G T A C A T C T G C C A C G T A C A T C T G C C A C G T A C A T C T G C C A C G T A C A T C T G —A C G T A C A T C T G —A C G T A C A T C T G C C A C C T A C A T C T G C C A C C T A C A T C T G —A C C T A C A T C T G —A C C T A C A T C T G Œ A C G T A C A T C T G £ £ A C G T A C ATCTG&tCtgPAC A T C T G C C A C G T A C Polymorphic codmg: i = o-; 2 =a/- Recoded: Optimizations: A T C T G C C A C C T A C A T C T G —A C C T A C A TC TG llA C C TA C A T C T 622A C C T A C Exclusion Recoded: A T C T G A C G T A C A T C T G A C 6 T A C A T C T G A C C T A C A T C T G A C G T A C A T C T G C C A C C T A C A T C T G C C A C C T A C A T C T G C C A C C T A C A T C T G —A C C T A C A T C T G —A C C T A C A T C T G —A C C T A C A T C T G C C A C G T A C A T C T G —A C G T A C A T C T G C S ^ C C T A C A T C T G —A C G T A C A T C T G =A C G T A C A T C T G = A C C T A C Contraction: i=o-;2=A /- Recoded: Optimizations: A T C T 6 C A C C T A C A T C T G C A C G T A C A T tZ T G C A C G T A C A T C T G -A C G T A C A T C T G -A C G T A C A T C T G -A C G T A C A T C T G IA C G T A C A T C T G C A C G T A C A T C T G -A C G T A C A T C T G 2 A C C T A C A T C T G -A C C T A C A T C T G -A C C T A C Figure 3-2. The effect o f character coding on homology statements in questionably aligned DNA sequences, usmg hypothetical sequences o f four taxa. Uhderiming: viola tion o f test o f conjunction. Italics^ character states not found m origmal data. Gray back ground: switching positions o f the respective bases results in further possible optimizations with corresponding consequences for homology. For details see main borfyoftexL 135 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. together usmg parsimony. Essentially, elision is a weighting procedure, giving charac ters with unequivocal position more weight than those for which the specific similarity statement is uncertain. Those characters with unambiguous alignment are found multi ple times, whereas any particular type o f a column in regions o f ambiguous alignment is included with a lesser frequency (four times versus once in Figure 3-2). Character weighting is highly controversial, because it introduces subjectivity into the analysis (Wheeler, 1986). Elision also has effects on homology (Wheeler e ta i, 1995: 5-6): “The elided data... have the disturbing property o f assigning multiple primary homologies to the same datum .... the implications for homologr are unsettling, since individual bases must have individual histories, but are not treated as such.” To rephrase their finding, the test o f conjunction is failed for all positions with variable alignment. All other sin gle positions, which occur more than once, are regarded as instances of character weight ing and not failure o f the test o f conjimction. Case sensitive PAUP (Swoflford, 1993) can be mstructed to treat characters in a case-sensitive man ner. Unequivocal primary homologies are shown in upper case, those with uncertain alignment in selected taxa are in lower case. Lower case states are then treated as miss ing data (coded ‘?’), but preserve the original information in a convenient form (Figure 3-2: Case sensitive. Recoded). The first problem with case-sensitive coding is the inabil ity to express a gap in lower case, for which reason an a priori alignment must be cho sen subjectively, unless gaps are specified as missing characters. The latter is clearly inappropriate, as pointed out above; in the present example it would result in a CC opti mization for all taxa. Le., render the character uninformative (not shown). Assuming gaps are treated as a ffîth character state, lower case codmg becomes pomtless, as a par- 136 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ücular posMoa for the lower case characters must be chosen to begin with. Addition ally, the lower case (= missing) characters can now take on any state (A, T, G, C, -). Character-state optimization will only consider existing states: C and -. The observed state A in taxon 4 will never be found in the optimization; only base C. If in taxon 3 or 4 a gap should be optimized, the test o f conjunction is failed (Figure 3-2: Case sensi tive, Optimizations). Case sensitive coding confuses the uncertain expression o f a state with the uncertain position of an observed state in the sequence. Missing data If a region o f ambiguous alignment is coded as missing data (= ?; e.g., Whiting et a i, 1997), the parsimony algorithm is offered more possibilities to optimize the states for these missing data entries. Such a strategy seemingly offers flexibility but comes at a high price, because the same problems with character-state optimization as discussed above apply. One additional type o f inconsistency can result in taxon 4, when CC is optimized; it fails the test o f conjimction and contradicts the original observation at the same time (Figure 3-2: Missing data. Optimizations: italicized and underlined). Polymorphic Regions with ambiguous alignment can be coded as polymorphic, restricting the possible states a particular position may exhibit (see Wiens, 1995, for overview). Posi tions sbc and seven for taxon 3 are coded as 1 = C or -, and for taxon 4 as 2 = A or - (Figure 3-2: Polymorphic, Recoded). The possibility o f assigning unobserved character states to a position is barred, but still only existing states (C or -) will be optimized. Taxon 4 will always show a double gap, hence, fails the test o f conjunction consistently. 137 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Taxon 3 will also fail the test o f conjunctioa in two o f the four possible optimizations (Figure 3-2: Polymorphic, Optimizations). Exclusion Questionably aligned regions may be excluded from analysis (Figure 3-2: Exclu sion, Recoded. £.g., Gatesy et a i, 1993; Cerchio & Tucker, 1998). In this example used, this method will result in the example used here in a sequence with only uninfor mative characters, leaving the relationship o f the four taxa unresolved. The exclusion o f characters leaves the hom olo^ concept intact. Specifically, some observations are dis regarded because they are already accounted for as being not due to common ancestry. Therefore, these observations can be o f no cladistic interest. The other strategies dis cussed above lead to the same unresolved topology using elaborate coding schemes, which as a consequence contradict the observation and/or fail the test o f conjimction during character-state optimization. The only way to include observations whose simi larity is highly doubtful is to classify them as entirely different entities. This translates to the introduction o f additional character states. In its most extreme form, or from the perspective o f a skeptic, such practice will lead to a data matrix composed only o f autapomorphies. Contraction This less stringent method provides hypotheses o f homology at a higher level o f generality. The questionably aligned positions six and seven are combined mto a single character. Taxa 1 and 2 are coded straightforwardly as C and -, respectively. For taxa 3 and 4, polymorphic codmg is employed again (Figure 3-2: Contraction, Recoded). As mentioned above, character state optimization is restricted to existing states. Taxon 4 138 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. will always show a gap, taxon 3 either a C or -, The test o f conjunction is failed in nei ther case, nor do the optimized character states contradict the original observations: homology is intact (Figure 3-2: Contraction, Optimizations). A conceptually similar approach has been proposed by Wagele (1994). The reduction o f two or more positions to a single one may raise questions regard ing weighting issue. The problem o f character weighting has only bearing if two other wise equivalent coding schemes are compared. The latter is not the case here. Data contraction is carried out because o f problems relating to representation o f observa tional similarities. Hence, the character weighting argument, despite in general being of legitimate concern, has no force in the current context. HOMOLOGY IS SPECIAL SIMILARITY SENSU REMANE (1952) Cladistics is based on characters that share special similarities as opposed to overall similarities. Does every observation qualify as special similarity? Hardly so, as it is well-known from the classic insect wing - bird wing example. Nevertheless, Gatesy et al. (1993: 156) have used the total evidence argument to retain all available data at any cost: "... Kluge's (1989) notion o f ‘total evidence’ should be extended to the use o f scrambled alignment regions in phylogenetic reconstruction.” The total evidence argu ment, however, refers to characters as used in cladistics, i.e., those harboring special shnilarity, hence, does not justity indiscriminant inclusion o f observations. The inclu sion o f highly ambiguously aligned sequences should be avoided. The difficult question o f where to draw the line between unequivocal and ambiguous alignment will be dis cussed m the following section. 139 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Belief formation Character ‘selection’ is a critical part in any cladistic study^ because we select those states that require explanation by way o f common ancestry. Observations o f structures considered homologous are by definition explained first in elementary hypotheses of primary homology and subsequently in the composite hypothesis. Shared similarity must be evaluated at each level o f generality and follows the division o f whole and part. Although a skull (whole) is readily recognized as a homologous structure in all verte brates, each bone of the skull (parts) has to be evaluated individually. Conversely, how ever, if all bones of the skull (parts) are considered to be homologous, then the skull (whole) must be homologous as well (see Mahner, 1998, on the intransitive nature o f the part-whole relationship). Similarly, each base in a homologous gene must be criti cally assessed. The assessment comprises both the identification o f the character state (red/green/blue, A/G/C/T/-), as well as their classification in a character (petal color, position 213), termed ‘character-state identity’ and ‘topographical identity’ by Brower & Schawaroch (1996), or ‘prepositional belief formation’ and ‘objectual belief forma tion’ in epistemology (Audi, 1998). This characterization is fiilly compatible with the concept o f the predicate language for observations discussed above. The character is the noun, the state its predicate. In morphology, the categorization of the individual condi tions into discrete states is more problematic than with molecular features, but the iden tification o f the character is a greater problem with molecular characters. We readily classify the observed appendage o f a tetrapod as a leg (topographical identity), but struggle with the description o f it as stout or slender (character state identity). With molecular data, the identity o f the character states (A, G, C, T, -) are unequivocal, but the position (topographical identity) on the sequence is to be determined. Character state identity and topographical identity must be satisfactorily assessed m order for an 140 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. observation to be considered worthy o f an explanation as shared similarity. The equiva lent treatment o f all characters is maintained, although different problems regarding the establishment as shared similarities are encountered (Brower & Schawaroch, 1996). In cladistics, the characters o f interest to us are restricted to those which we con sider worthy of the explanatory effort. The cut-off point, whether or not one may still suspect certain structures as similar and to be explained in terms of primary homology in the elementary hypothesis, depends on the trust in one^s own observations (Ax, 1989). Remane (1952:103) phrased it clearly: “Because of this methodology it follows th at... a transitional area exists, in which a carried out or doubted hoinologization depends on the optimistic or pessimistic temperament o f the researcher” [translation from German}. This position is not unique to phylogenetics, but a general, philosophical/psychological principle. It is in the nature of belief formation that “[p}eople differ markedly in the beliefs they form about the very same things they each clearly see” (Audi, 1998:17). The boimdary between recognized and doubted shared similarity is not sharp and no hard and fast rules exist as how much uncertainty is sufficient during the process o f belief formation to argue for the a priori non-inclusion o f a morphological character (Remane, 1952) or the a posteriori exclusion o f a molecular character (see also Gatesy et o f, 1993) relative to the time of data acquisitioiL The treatment o f the a priori exclu sion o f morphological characters and the a posteriori exclusion o f molecular characters is equivalent; the perceived difference is inherent to the mode of data acquisition and is unrelated to the establishment o f primary homologies. The entire process o f belief formation using auxiliary information is but another form o f inference to the best explanation, i.e., abductive (Devitt, 1997). Abduction can not f u rn is h an explanation m the form o f newly postulated past ancestors for the present observations that is certain to be true, even if all the premises are true; as a form o f 141 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. hypothetic reasoning, abduction is ampliative and not truth-preserving. This is in marked contrast to valid deductive reasoning, where any conclusion must be true if the prem ises are true (non-ampliative, truth-preserving). The very nature o f explanatory hypothe ses is the inclusion of non-observed causal entities, in this case common ancestors, to account for observed effects, shared similarity. Any conclusion in the form o f an hypoth esis from an abductive inference, elementary as well as composite, will always be ten tative. Rational for data exclusion To exclude characters after their acquisition was characterized by Wheeler (1986:108) as “to give up” with a certain, highly homoplastic data set if even reluc tantly applied weighting did not provide better resolution. In the same sense, if multiple weighting schemes in sequence alignment lead to difforent similariQr statements, then we may well give up these characters, i.e., exclude them. From an explanatory point of view, these observations are judged not worthy o f explanation. Although unsatisfactory, it is more honest to admit the failure to recognize shared sim ilari^, than to make unfounded assertions. A primary bom olo^ statement indicates that two or more prop erties are considered by an investigator sufficiently similar in terms o f structure and position as to be tentatively accounted for by common ancestry. As a result, responsibil ity and accountabili^ is bestowed back upon each practicmg systematist. It is not the computer that proposed a particular hypothesis o f relationship—and many ‘intriguing’ phylogenies have been published particularly with molecular data—, but informed investigators stand behmd them. Some o f the more eggregious examples from molecu lar studies include the following: non-monophyly o f Tracheata ^ allard et aL, 1992; see Wagele & Stanjek, 1995; Farris, 1998); sperm whale as sister taxon to baleen whales: 142 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (Milinkovich, 1995; see Heyning, 1997; Messenger & McGuire, 1998; Cerchio & Tucker, 1998); position of Pogonophora (McHugh, 1997; see Siddall et aL, 1999). It has been argued that the exclusion o f molecular characters is a cardinal sin in cladistics (Gatesy et aL, 1993). What is the comparable procedure with morphological characters? It would be the non-inclusion o f available characters. As an example, in a class-level analysis body color of the exemplar taxa is not found in the data matrix {e.g., Whiting et aL, 1997), although this character is readily observed. Why is this character not included? The inexplicit answer is that the primary homology o f the character ‘color’ is questionable at the class level, although it is useful at the genus/species level {e.g., Westemeat, 1993; Swenson & Bremer, 1997). From an observational point o f view, the observer considers the superficial similarity unconvincing, so that no case for special similarity can be made. From an explanatory point o f view at the level o f the elemen tary hypothesis subsequent to the observational phase, the character-state distribution is already explained as being due to some causal event(s) other than common ancestry. The decision o f inclusion or exclusion o f a character is based on the investigator’ s will ingness to accoimt for shared similarity by way of common ancestry, by way o f some event other than common ancestry, by way o f not trusting their own observations, being unsure o f what they observe, or by way o f their inabili^ to characterize what they observe. With morphological characters the arguments for non-inclusion are hardly ever spelled out, unless a previously used perceived similarity can no longer be accounted for by common ancestry at the level o f the elementary hypothesis, i.e., is no longer con sidered a primary hom olo^. Similar practices can be fotmd with sequence data. The use o f the ITS region for population studies and species-level investigations is common {e.g., Vanherwerden et aL, 1998; Mes et aL, 1997; Nakasone & Sytsma, 1993). Expanding the scope o f such a 1 4 3 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. study to phyla w ill result in unalignable sequences o f approximately 25% similarity, which is tantamount to the random similarity o f sets composed o f four elements (Li & Graur, 1991). It is more sensible to select a more appropriate gene/gene-region that will allow for better alignment, than to argue for inclusion of all available data. Hence, we should try to avoid the problem that Wenzel (1997:37) characterized pointedly as “Garbage in, garbage o u t” The example chosen is extreme, but illustrates the point to be made at any level o f taxonomic inclusiveness. Here then another perceived, major advantage o f sequence data is put into perspective: the large number of characters obtained from sequences. As only synapomorphies are of explanatory interest, variable regions furnish, such characters. However, it is also variable regions where the problem atic alignments, i.e., designation of shared similarity, are prevalent, therefore, where the determination o f primary homolo©f is much more uncertain. The exclusion of question ably aligned regions also reduces the number o f explanatorily relevant, what most refer to as ‘ ^informative*, characters significantly, which may also lead to a loss o f resolution in the cladogram. The potential to increase resolution has led to the use o f methodolo gies in clear conflict with the philosophical basis o f homology; Wheeler et aL (1995:3) noted that “... the elided result was much more resolved”, but also came to the conclu sion that, “[cjlearly, these are not the best o f data to resolve insect relationship.” The resistance to exclude characters can be followed, but does not provide a justification to include characters o f doubtful primary homology, because the goal o f cladistics is expla nation o f character state distributions, and not maximmn resolution o f its representa tion, tree topology. Note, that there is no clear-cut argument for the inclusion or exclusion o f any particular character, because this decision relates to the process of belief formation and is mherently subjective (see previous section). There will never be a panacea for this problem, but being aware o f this difficulty may help to avoid some 144 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. major pitfalls. It is important to realize that an uncertain explanation o f character-state occurrence shown as an unresolved part o f a tree is increasing our knowledge about the cause o f the observed distribution. By not knowing something causally, we automati cally do know something: we recognize our ignorance, hence, we have further direction for our research (cf. Wenzel, 1997). With molecular data, the imcertainty o f topographical identity can be quantified. Positions that are not afiected by any set of alignment parameters are identified as unequivocal similarities. Those positions that are stable over a wide range of alignment parameters are less equivocal than those, which change positions with any alignment parameters. Hence, a minimum range (e.g., gap cost ratios fiom two to ten) over which the alignment must be unambiguous can be defined. Those observations remaining unambiguous are accepted as shared similarities, the remainder are excluded. The con ditions (gap-cost ratio two to ten) are explicit, but still subjective. As long as the evalu ated alignment parameters do not cover the entire spectrum o f possible values (0 5 to infinity) the alignment is based on assumptions which are inherently subjective. Whether it is more appropriate to use an explicit, but rigid exclusion argument, than to «cclude characters with implicit, but flexible exclusion arguments is open to debate. After all, what we are interested are observations in need of causal explanation by means o f com mon ancestry. One may argue that as the number o f taxa in an analyst increases, the portion o f the sequence with questionable alignment increases. Hence, all characters need to be included as eventually any position will be questionably aligned. The above is an over simplification o f two distinct cases. When taxonomic sampling density is increased, more landmarks wüI be recognized, which will aid m positionmg the variable regions, because the additional taxa provide more auxiliaty m&rmation to be used in the process 145 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. o f belief formation. These additional data fomish new arguments for the alignment of a given set of sequences; both resolution o f previously questionable alignments as well as flagging o f ‘new’ uncertain primary homologies are possible (Eemisse, 1997). With increased taxonomic sampling width, however, the new auxiliary information will not provide the landmarks needed to position variable regions. The tentatively identified similarities can not be upheld at that particular level, which is related to the explanatory relevance o f information. HIGHLY DISSIMILAR TAXA What is the most appropriate action if only one or a few taxa are extremely dissimi lar as compared to the remainder (Figure 3-3)? Often ‘dissimilar’ is equated with ‘diver gent’, although the latter implies an explanation at the outset. Exclusion or contraction o f data may eliminate much important information on the relationships o f the majoriQr o f taxa and would reduce the data matrix to the lowest common denominator. One may consider missing-data coding for that particular stretch in highly dissimilar taxa (Figure 3-3; taxa 6 and 7), because essentially one does not know anything about the specific homologies in these taxa. However, as pointed out above, such a coding strategy is inappropriate because it confuses the uncertam expression of a state (= character state identity) with the uncertain position (— topographical identity) of an observed state in the sequence and will create problems during character-state optimization. The two cod ing strategies introduced below address the question of how to best represent our obser vations. As belief formation is a psychological problem beyond scientific mechanics, there is no conclusive answer. 146 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I I 1 I o o I —( t— c o o o o o rH rH o r 4 I o o O O o r— 1 o o o o o r 4 1-4 o o o o o r —i o o o o o rH i“4 o o u C J C J C J CJ 3 S g g g 3 C D C D C D C D C D C D C D C J C J Ü C J p C J CJ S < cw < 3 3 o C J o * cw CW C D C D C D C D C D ' c w B B B B cw CW P p P P P c w c w P Q < < < < < CW cw U o O C J C J O Ü s a 3 3 g g 3 C D C D C D C D C D C D C D O C J O C J C J C J C J < < < < < 4 4 CJ C J 1 1 < VO r- u C J I I 3 VO p » C D C D C D C D C D VO B B B B VO p - C D C D C J C J O VO E 4 B B B B VO p - < 5 < < < < VO p * « — 4 CM c n in 4 4 C Mm m B B o O C J P C J C J C J S S S g g 3 C D C D C D O C D C D C D C J C J C J C J C J P C J < < < < < 4 4 o o I 1 < VO VO C J C J 1 1 2 VO VO C D C D C D C D C D VO VO Bi B B B B VO VO C D C D C J C J CJ VO VO B B B B B VO VO < < < < < VO VO u o C 3 O C D C D C D CJ U Figure 3-3. Coding strategies for a few highly disshnilar taxa 1-7 shown in rows 1-7. Original data shows one o f the many possible alignments o f this particular data set. Stretch coding and block codmg illustrate two alternative coding strategies compared to presence/absence codmg discussed in the main body o f the text. 147 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Stretch coding [a Figure 3-3, the observed dissimilarity is a piece o f available information about taxa 6 and 7 to be represented as such. This information can be expressed using new character states, i.e., the entire unalignable region can be coded as a single state 6 (Fig ure 3-3 : stretch coding). In this fashion it can be shown that taxa 6 and 7 are both highly dissimilar and show synapomorphies for the first seven characters. Some may argue that the statement o f high dissimilarity should just be coded in a single character, essen tially contracting the seven positions. Others may say that major diSerences are seen in all seven characters for which reason stretch coding shows the condition appropriately. It may seem as if a weighting issues arises. It has been shown above (section Contrac tion) that the representation o f observational similarities in agreement with homology takes precedence over considerations o f weighting. Minor differences between taxa 6 and 7 could be coded using an additional charac ter state for taxon 7 (not shown). One caveat applies: as additional state(s) must be specified for a particular character, it implies a homology statement with the characters in taxa I to 5, which is not feasible. Explicit homology statements o f T and A in taxa 6 and 7 are made, disregarding alternative alignment possibilities, e.g., the T may either be found in position 2 as mdicated or in position 4; the relative position of T may not be the same for taxa 6 and 7. One may argue that shared character states need to be explained and that the classification o f the character states under a particular character is o f lesser importance, because the homology statements are restricted to within the blocks o f taxa I to 5 and 6 and 7 (Figure 3-3: boxes). Then the information is suffi ciently represented by stretch coding. 148 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Block coding To circumvent the problem o f unspecifiable homology statements across blocks within a stretch (Figure 3-3: Original data: characters 1-7, taxa 1-5 and characters 1-7, taxa 6-7), one may consider treating states in taxa 1-5 and 6-7 separately (Figure 3-3: block coding: right boxes), inserting autapomorphies for the corresponding taxa in the other block (Figure 3-3: block coding: left boxes). There are two blocks with autapo morphies only (characters 1-2, taxa 1-5: upper left box; characters 3-9, taxa 6-7: lower right box), and two blocks containing the sequence information (characters 3-9, taxa 1- 5: upper right box; characters 1-2, taxa 6-7: lower left box). The homology statements o f the sequence information in each block are unconnected to one another. No homol ogy statements are made between the T in taxa 6-7 with respect to any T (in position 2 or 4 o f original data) in taxa 1-5. However, problems with character weighting arise because the information from the ftrst 7 characters is now coded in 9. The above dis cussed issue o f homology statements within the lower block also apply here, i.e., are the T’s and the A’ s in taxa 6-7 homologous or not? One may argue that the autapomor phies in respective blocks should be coded the same within each block but different ftom the information-bearmg entries. Such coding would accentuate the between-block differences b ^ond the original observations, which is not supported here. Either stretch- or block-coding strategies have theoretical advantages and disadvan tages, resulting in a classical trade-off situation inherent to the psychological process o f belief formatioiL Block coding may seem somewhat preferable for the following rea sons, When considermg taxa 1-5 and 6-7 separately, then the homology statements are clear. Only by combining the data might problems arise. Therefore, the information from the two blocks should be coded separately. As the other block does not contribute 149 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. any information to the data found in the block under consideration, the ‘empty' blocks should be assigned uninformative autapomorphies. Presence/absence coding Block coding is somewhat similar to the established procedure to code gaps in a supplemental presence/absence (p/a) matrix (e. g., Baum et a i 1998) (Figure 3-3: pres ence/absence). The two differ twofold. I) The characters with gaps in the p/a strategy are left in the data matrix, whereas block coding recodes the characters within the sequence portion. Issues with character weighting are reduced to a m inim um with block coding. 2) The gaps in the sequence part o f the p/a matrix are treated as missing charac ters, whereas no missing character states are introduced with block coding. As any miss ing character state misrepresents an actual observation as the result o f belief formation^ any use o f ‘?’ as an indicator o f uncertam assignment of observed character state (= character-state identity) to a specific character (= topographical identity) is positively misleading. P/a coding is also at odds with the classification process in the observa tional phase. By establishing separate bins (characters) for observations actually belong ing in one and the same bin, arguments for the actual existence o f separate classifications are introduced: parallel «cplanatory universes are in effect advocated and thK results m a schizophrenic view o f the real world. Devitt (1997) and Mahner & Bunge (1997) dis cussed in much detail the need o f a scientist to be a realist. Comparison to practice in morphology Comparison o f the coding strategies (particularly stretch coding) to practices in morphology is favorable. In morphology, a particular state recognized to be classified in a particular character that is highly disshnilar in a particular taxon is coded as a sep 150 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. arate state and is not forced into an existing state. One practical difference between morphological and molecular characters must be discussed. A single observation can be coded as an additional character state in morphology but not with molecular data. With sequence data the problematic part in the process o f belief formation is the establishment o f topographical identity, therefore, a single posi tion can not harbor ambiguous alignment The minimum number of characters required for ambiguous alignment is two adjacent positions. If the auxiliary, positional informa tion places a base in a particular position (= topographical identic), the identification o f the character state (= character-state identity) is not an issue. For morphological charac ters, in contrast the problematic part in the process o f belief formation is character state identity. Hence, for any single set o f observations classified in a character, any particu lar observation may not be classifiable in one o f the other states. An observation that can not be classified in an existing state is given a new state. The common denominator is, that whenever one o f the two conditions needed to postulate shared similarity is not met, topographical identity or character-state identity, then additional character states are introduced. In neither case are the particular characters coded as missing data, because otherwise the origmal observation can be contradicted during character-state optimization. The distinction between topographical identity and character-state identity has fur ther bearmgs. hi morphology, a property o f taxon X that can not be identified as a shared similarity, is given a new state, but remains classified in its origmal character. The new state mtroduced and found only in a single taxon do% not indicate a relationship with the other taxa, except for X not sharing a most recent common ancestor with any other taxon* at the level o f the elementary hypothesis, ha essence, a questionable property is given a new state. 1 51 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. With molecular data, an identified state that can not be classified is assigned to a new grouping (position). The new character introduced shall only carry information for the taxon or taxa in which it occur(s). For the remainder of taxa, it should remain unin formative with respect to the expressed relationship at the level o f the elementary hypothesis. This is best achieved using autapomorphies for ail except the highly dissim ilar taxon/taxa. For molecular data, then, a questionable position is given a new posi tion. Hence, the introduction o f a new position in block coding is related to the problematic topographical identity o f readily recognizable states and is not a fundamen tal difference as compared to practices in morphology. BETTER ALIGNMENT? One could ask whether the above alignment strategies produce “improved represen tation o f homology”. The catch phrase in itself lets the old molecular jargon o f ‘87% homologous’ resurface, overlooking the bmary nature o f hom olo^; is homologous, is not homologous. Hillis (1994: 339-340) was correct in stating that “molecular biolo gists may have done more to confound the meaning o f the term homology than have any other group o f scientists Why this confusion of terms [homology versus simi larly] has arisen in molecular biology is not clear; perhaps the term hom olo^ is thought to make the work sound more like science Today a probabilistic notion has been added to the hom olo^ concept, particularly under maximum likelQiood, an issue to be discussed elsewhere. The question arises o f how to evaluate the homologies. A number o f avenues may be considered, vAich will be addressed below. 152 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Comparisons to ‘known’ phylogenies: I f a phytogeny would be known, then why bother trying to build a hypothesis using an ampHative, non-truth preserving mode o f inference? One could only conclude that the mode o f inference is properly character ized. It does not address the perceptional question in any form. Conferred bootstrap, jackknife or similar support: It has been widely realized that these measures are fraught with problems of which Wenzel (1997) provided an overview. To Justify one alignment over an other by means of a questionable metric is imtenable. Determining nodal support indices for an entire tree has not even been attempted. All these metrics are based on the entire data matrix or permutations thereof, whereas justification o f the alignment applies for every single position individually, respecting character independence and relevance of auxiliary information. Accordingly, any approach using support indices o f any sort is misguided. If one would consider to use a character support index such as the rescaled consistency index, then the question arises, for which position the comparison is made, because the topographical identify of a perception is at stake. Value o f alignment score: To evaluate one alignment using other alignment parame ters or other methods o f alignment also misses the point. One would only compare the underlying models as characterized by the alignment parameters and could conclude that, indeed, they difEer. The same problems with overall metrics and character-specific metrics arise as discussed in the previous paragr^h. As shown above. Justification for a given alignment procedure can not come from the inferred hypothesis or any metric associated with the alignment. The Justification needs to come finm the factors surrounding the goal to be achieved, namely findmg similarities worthy o f explanation. It is accomplished by elimination o f conflicts with the cladistic methodology at large: character independence and relevance o f auxiliary 153 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. information, as well as contradiction with observations and violation o f the test o f con junction during character state reconstruction. These are the avenues that have been pursued, and on which the arguments have been built. Any criticism needs to address these issues. CONCLUSIONS AND RECOMMENDATIONS The following points can be extracted to form a guideline to DNA sequence align ment and character coding. • Global alignment is inappropriate, because it conflicts with character independence. After conserved regions are identified, only characters between two adjacent, con served regions should used to establish primary homologies o f bases within: mini mal fiagment alignment (MFA) should be practiced. • Objective sequence alignment is inherently impossible; some level o f subjectiviQr is always introduced. Due to MFA, computer-facilitated and manual alignment each have their discrete advantages. The former allows for explicit specification o f the assumptions in the form o f alignment parameters used, and the latter forces one to critically justify every similarity statement in the data matrix. • Gaps should be coded as a fifth character state as they are mvoked during the process o f belief formation in the observational phase. • Flexible coding strategies (elision, case sensitive, missing data, polymorphic, pres ence/absence) all conflict with the test o f conjunction or have the potential to con tradict our original observations, hence, can not be justified in any explanatory context Only data exclusion and data contraction do not introduce such problems. 154 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. • Data exclusion o f highly ambiguously aligned regions is not in disagreement with the total evidence argument, as the latter applies only to special similarities sensu Remane (1952). • For highly dissimilar taxa, new characters need to be introduced to represent the available information most appropriately. Highly dissimilar regions should be recoded with block or stretch coding. Treating DNA sequences in such a fashion is fully compatible with coding strate gies used for morphological data and assures compliance with fundamental principles o f cladistic analysis, particularly the concepts o f causal explanation and homology. 155 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter 4: Distribation and Biogeography of the Recent Haliotidae (Gastropoda: Vetigastropoda) World-wide. INTRODUCTION Abalone are a family o f marine gastropods consisting o f 55 currently described species of world-wide distribution in tropical and temperate waters of both hemispheres. During revision work on this family I have recently evaluated all Recent taxa, 200 at the species level and 17 at the genus level (Geiger, 1998a). Some other taxonomic mat ters have been dealt with elsewhere (Geiger 1998b, 1999; Geiger & Stewart, 1998, Stewart & Geiger, 1999). Knowledge o f the distribution o f most species is sketchy, par ticularly due to the absence o f specimen-based accounts. Most indications regarding the distribution o f abalone in the literature may be termed intuitive (e.g., Ubaldi, 1993, 1995) with limited exceptions (Geiger, 1996, 1999; Simone, 1998; Stewart & Geiger, 1999). In this second, larger treatment o f the 6 m % Haliotidae, 1 concentrate on the dis tributional pattern o f all Recent species, including a biogeographical analysis. To date, studies on the biogeography o f the family are marked by their anecdotal nature. Three hypotheses (Figure 4-1) for the oiigm have been proposed and have been discussed by Geiger & Groves (1999) in a review o f fossil abalone. Model I (Figure 4-lA ): Pacific Rim. An arc spanning from Japan to northeastern Australia has been identified by Talmat^e (1963a) as a likely cradle of the family. From this nuclear distribution abalone then dispersed in a star-shaped pattern to the north western and then the northeastern Pacific; to Australia and the Indian Ocean. The basis o f this model was never made explicit. 156 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 96 Figure 4-1. Illustratioa o f the three biogeographical models proposed for the origia o f Haliotidae. A) Pacific Rim origni (Talmadge, 1963a). B) ludo-Malayan origin (Lind- berg, 1992). C) Tethys origin (Geiger & Groves, I999).The numbers indicate the diploid number o f chromosomes in the species for which data is available. 157 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Model 2 (Figure 4-IB): Indo-Pacific. The highest present-day diversity o f the fam ily is found in the Indo-Malayan area. Although it is well appreciated that the correla tion o f high present-day diversity with origin o f the group in question is highly problematic, it provides one possible center of radiation for the family, and was dis cussed by Lindberg (1992) and Briggs (1999). Model 3 (Figure 4 -lQ : Tethys. From published chromosomal-count data Geiger & Groves (1999) considered an origin in the Tethys Sea. Since other Vetigastropoda and basal gastropods have a relatively low chromosome number (2n = 18-20: Patterson, 1967; Haszprunar, 1988b) as compared to the values found in Haliotidae (2n = 28-36: see Geiger & Groves, 1999, for sources) a progression from low to high values was considered. Bieler (1992) also concluded that the low chromosome number o f patel- logastropods is the plesiomorphic condition within the archaeogastropod grade. It would suggest a radiation starting in the Tethys, today represented by the Mediterranean species H aliotis tuberculata Linnaeus, 1758, with 2n = 28, moving eastward to the Indo-Pacific (2n = 32), and finally reaching the North Pacific (2n = 36). Here the three proposed models are evaluated using specimen-based distributional data for all species. The results firom area cladograms are then compared to preliminary data on the phytogeny o f the femily. MATERIALS AND METHODS Sources of Raw Data Specimens in the natural history museums as listed below and some specialized pri vate collections were personally inspected. After the identity o f each specimen was ver ified, their localities were recorded. Additional records o f specific localities and range information were obtained &om the literature. Only those data were included that 158 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. seemed to be reliable. Le., from publications Ulustrating specimens, from species that are very hard to nusidentify, and from publications by authors, whom I know and trust with the identifications provided. Any arbitrary locality data have been excluded. High resolution maps in a rectangular projection were obtained from the internet site provided by the Xerox Palo Alto Research Center (http://www.parc.xerox.com/). The maps were further processed in Adobe Photoshop 3.05 (see Geiger, 1998c, for tech nical details). Localities stemmmg from collection specimens are indicated with a solid circle, those from the literature with a solid square. There may be fewer dots than listed localities because more than one localiQr may fall under the same symbol. Localities that seem very doubtful are identified with an adjacent question mark. Published range information is indicated with continuous lines for the core range, on which all the cited authors agree; broken lines indicate the peripheral range given by at least one author. Only specific indications such as “Point Conception” were taken into account as distri butional limits, not general terms such as “central California”. For widespread taxa, information on their distributions in any sub-regions were combined. Abbreviations of collections Institutional collections: AMNH, American Museum o f Natural History, New York; AMS, Australian Museum, Sydney; ANSP, Academy o f Natural Sciences, Philadelphia; BMNH, British Museum o f Natural History, London; CASIZ, Califomia Academy o f Science, Invertebrate Zoology, San Francisco; DMNH, Delaware Museum o f Natural History, Wilmington; LACM, Los Angeles County Museum ofNatural History; MHNG, Muséum d’Histoire Naturelle Genève, Switzerland; MNHN, Muséum National d’His- toire Naturelle, Paris, France; DMMF, Deutsches Museum für Meereskunde und Fis- cherei, Stralsund, Germany; NHB, Natural History Museum Basel, Switzerland; NM, 159 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Natal Museum, Pietennaritzbmg, South Africa; NMBE, Natural History Museum Bern, Switzerland; NMW, National Museum o f Wales, Cardiff Great Britain; SBMNH, Santa Barbara Museum ofNatural History; SMF: Senkenberg Museum, Frankfurt, a. M., Ger many; UCMP, University o f California: Museum of Paleontology, Berkeley; UR, Uni- versitiy o f Rostock, Germany; USNM, United States National Museum o f Natural History, Washington, D. C. Private collections: DLG, Daniel L. Geiger collection, Los Angeles, California; EN: Eike Neuberg collection, Frankfurt, a. M., Germany; JK, Joan Koven collection. Astro labe Inc., Silver Spring, Maryland; KAS, Katharine A. Stewart collection, Carmel Val ley, California; RP, Roger Pickery collection, W lrijk, Belgium; SBS, Solly (Benjamin) Singer collection, Rehovot, Israel. Biogeographical regions The biogeographical regions were identified according to Eckman (1953) and Briggs (1995). The following areas were used, where both the distinctness o f the area as well as the number o f species present in the area was considered. The west American coast, for instance, was subdivided into several areas as they are well defined and several abalone species occur therein, whereas, the eastern coast o f South Ameri«i was repre sented as a single area containmg two species. 1 - Northeast Atlantic: Eastern Atlantic from Normandy, France, to western Morocco (30® S). 2 - Mediterranean: Mediterranean Sea. 3 - West African: 30® S including Canary Islands to Cape Town, Republic o f South Africa. 160 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 4 - South Aôican: Cape Town, Republic o f South Afîica, to Kei River Mouth. 5 - Western Indian Ocean: Kei River Mouth, Republic o f South Aôica, to Karachi (67® E) without Red Sea, including Madagascar, Mascarenes, Seychelles, and Rodriguez Islands. 6 - Red Sea: North of Djibouti. 7 - Central Indian Ocean: Islands between Western Indian Ocean, Indo-Malayan province, and Australia: mainly Maldives, Sri Lanka, Chagos Archipelago. 8 - Indo-Malayan: Andamans, Malaysia, and Indonesia (to Man Jaya) to south o f Oki nawa, including Taiwan and Philippines. 9 - Central Pacific: East o f Indo-Malayan Province and Australia to Henderson Island. 10 - Sub-tropical Japan: Okinawa to 3 6 ® N. 11 - Eastern North Pacific: From 36 ® N of Japan to western Aleutians. 12 - Aleutian: From western Aleutians to Portland, Oregon. 13 - Oregonian: From Portland, Oregon, to Point Conception. 14 - Californian: From Point Conception to tip o f Baja California, to the exclusion of the Gulf o f Cali&mia. 15 - East Pacific: From the tip o f Baja California to Peru, including Gulf o f California, Cocos IsL and (jalapagos Isl. 16 - Caribbean: From the tip of Florida peninsula to Salvador, Brazil. 17 - Argentinian: From Salvador, Brazil, to Golfo de San Jorge, Argentina. 18 - Solanderian: Torres Straits to Port Jackson, Australia. 19 - Peronian: Port Jackson to Melbourne, including Tasmania. 20 - South Australian: Melbourne to Hopetoun. 21 - West Australian: Hopetoim to Shark Bay. 22 - Northwest Australian: Shark Bay to Torres Strait. 161 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 23 - New Zealand: North, and South Island; Kennadec, Chatham, Bounty, Antipodes, Campbell, Auckland, and Snares Islands. Biogeographical anafysis Unrooted area cladograms were constructed using PAUP 3.1 (Swofford, 1993) fix> m the binary area data-matrix (Table 4-1). All taxa-as-characters were equally weighted, and uniformative characters were excluded. Heuristic searches with random taxon addition, 1.000 replications, tree-bisection-reconnection (TBR), and searches on non-minimal trees with steepest descent and MULPARS were performed. The maximum length of the non-minimal length trees was limited by the maximum number o f storable trees (approximately 32,700). This limit is a PAUP implementation problem and not due to insufficient RAM. Skewness or g, values were obtained hom five replications of 1 0 0 .0 0 0 random trees with interval width o f one and random starting tree. Two taxonomic uncertamties were treated as follows. Specimens o f the southern range o f H, diversicolor Reeve, 1846 (Figures 4-40-42), and those of the northern range of H. squamata Reeve, 1846 (Figures 4-63,4-66), bear some striking resemblance. The question o f synonymy or separate species is currently unresolved. Separate analyses with the two taxa coded separately and with a single wide ranging species were per formed. The status of H, brazieri Angas, 1869 (Figures 4-45,4-48), and H. hargravesi Cox, 1869 (Figures 4-46,4-49), is questionable in the light o f their potential hybrids and spechnens in which the identic may change during ontogeny (Geiger, 1998a). The two taxa have identical distributions; hence, the question o f character weighing is of concern. D a one analysis the two taxa were both mcluded; in a second analysis one was excluded. 162 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 4-1. Data matrix of biogeographical analysis. A U 'in the matrix denotes the pres ence o f that taxon in the respective area. Areas-as-taxa as defined in text Taxa-as-char acters; subspecies are included in nommai species. I: asinina. 2: aurantium.3: australis, 4: brazieri, 5: clathrata, 6 : coccoradiata, 7: corrugata, 8 : cracherodii, 9: crebriscidpta, 10: cyclobates, II: dalli, 12: discus, 13: dissona, 14: diversicolor, 15: dohm iana, 16: elegans, 17: exigua, 18: fatui. 19: fitlgens, 20: gigantea, 21: glabra, 22: hargravesi, 23: a is, 24: Jacnensis, 25: kamtschatkana, 26: laevigata, 27: madaka, 28: mariae. 29: mar - morata, 30: midae, 31: ovina, 32: parva, 33: planata, 34: pourtalesii, 35: pulcherrima, 36: pustulata, 37: queketti, 38: roberti, 39: roei, 40: rubiginosa, 41 : rubra, 42: rufescens, 43: rugosa, 44: scalaris, 45: sem iplicata, 46: sorenseni, 47: spadicea, 48: speciosa, 49: squam ata, 50: squam osa, 51: stom atiaeform is, 52: tuberculata, 53: unilateralis, 54: varia, 55: virginea, 56: walallensis. 163 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 4 I O TO O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O TO O O O O O O O O O O O O O O O O O O T O O OO TO OOO TOOO OOO OOO O O O O O O IO TO O O O O O O O O O O O O O tO O O O O O O O O O tO O O T OOTQOOOtOOOttOOIOtOOOOOOOtOOOOtOOOOOOOOOtOOOOOOOOOOTOOOO OOOOOOOOOOOttOOtOtOOOOOOOOOOOOTOOOOOOOOOOOOOOOtOOOOOOOOO OOOQOOOTOOOOIOOTOIOOOOOOOOOOOOtOOOtOOOOOOOOOOOTOOOTTTOOI OOtOOOOTOOOOOOOTTOOOOOOOOtOOOOOOOOTOOOOOOTOtOOOOOOTTTOOI O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O TO O O O O O O O O O O O O O O O O O O O O O O tO O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O TO ooooooooooooooooootooooooooooooooooooooooooootoooooooooo lO O O O O O O O O TO OO TO OOO OOO OOO OOO OOO tO OOO OtOOO OO OOO OOTTOOO OOO tO O O O O O O O O TO O O TO O O O O O O O O O O O O O O O tO O O O O TO O O O O O O O O O TTO O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O TO O O O O O O O O O O O O O O O O O O O O O O O OOOOOOOOOOOOOOOOOOOOOOOOOOOOOtOOOOOOtOOtOOTOtOOOOOOOOOOO OOTQOOOOOOOOOOOOOOOOOOOTOtOOOIOOTOOttOOTOOtOTOOOOOOTOOOT OOtOOOOOOOOOOOOOTOOOOTOtOTOOOOOOtOOOOOtOOTttOOOtOOOIOOOt OOtOOOOTOOOOOOOOOOOOOOOIOIOOOOOOtOOtOOOtOltOOOOOOOOtOOOt O O tQ O O O O O O O O O O O O OO OOO OOIO IOOO OOO OOO OO OOO OOO OOO OOO OOtOO OO O O O tO O O O O O O O O O O O O O O O TO O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O OOOtOOTOTtOOOIOOOOOITOOOOttOtOOOOOOOOOOOOOOOOOOOOOOTOOOO OOOOOOOOIIOOOOOOOOOIOOOOtOtOOOOOOOOOOOOOOOOOOOOOOOOOOOOO O O O O T O O O O O O O O O O O O O O O O O O O O O O TO O O O O Û O O O O O O O O O O O O O O O O O O O O O O O O O O TT O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O IO O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O O M 8H ZZ XZ Mottos O Z ueTUoaea g t LX 9 t ofjToedt gt et Zt oTïTOBd: t t UBdBf %BOTdoa%-c(n8 oT ôTïTOBd 6 u eA B % B W opui 8 Utaaoo tiB fpui i B88 patt 9 UB8O0 UBtPUI UJt8t^88M 9 q:»nos v ueot^JV ï^ fl8 M e U88U8aa8%tp8W Z oTpttUBt^V 4888qqjoN t 5GK:ZT068A9G^eZT0g8A$S^eZT0g8A9aMZt068Z9S^eZT0g8/.9GyeZT GSSSSSSt't'VÏ'frt'M'M'eEEeeeEEeeZZZZZZZZZZtTTttTTTITOOOOOOOOO 88T08dS ( / ) ( / > CD Q . 7 D CD 2 Q . c o " G 3 " O 2 Q . 2 tr g > \ Û . 8 CD C O CO CD Q . 7 D 8 3 " O 2 Q . CD Q 1 Cladistic anafysis of taxa Taxon cladograms were constructed fit>m recoded data published by Brown (1993) and Lee & Vacquier (1995). Midpoint rooting was employed (cf. Williams, 1992), because outgroup rooting was impossible with the data at hand. Allozyme frequencies were recoded with frequencies > 0.05 coded 1 and those < 0.05 coded 0. Data from multiple populations o f any species were averaged (Table 4-2). The cDNA sequences o f the lysin gene o f Lee & Vacquier (1995) were recoded using data contraction and stretch coding as detailed in Geiger & Fitzhugh (in review: Table 4-3). These two coding schemes use a higher level o f generality and introduce additional character states for regions o f questionable alignment The published align ment was adjusted manually. Parsimony analysis was carried out in PAUP using heuristic search options with random taxon addition. Gaps were treated as fifth character state (GAPMODE=NEW- STATE), uninformative character were excluded, and branch swappmg (TBR) on non- minimal length trees was performed with 1,000 replications. All characters and all character state transitions were equally weighted. MULPARS, STEEPEST DESCENT and RESPECT CASE were in effect. The allozyme data were not run with the tradi tional 0/1 character states as shown in Table 2, but with A/G, because otherwise the PAUP implementation limit of 31 character states was exceeded. This recoding has no effect on the phylogenetic reconstructions. Nfissing data lines for selected taxa not used m either o f the studies are not shown in Tables 2 and 3. Abbreviations: MPRs: maximum parsimonious resolutions; Cl: consistency index; RI: retention index; RC: rescaled consistency index. 165 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 4-2. Recoded allozyme frequency data from Brown (1993). The enzymes are identified by their full name, their standard abbreviation in parentheses, and the electro- morphs in lowercase letters. I-11 : Aspartate aminotransferase (Aat-I) a-k. 12-24: Aspar tate aminotransferase (Aat-2) a-m. 25-36: GlyceroI-C-phosphate dehydrogenase (aGPD) a-l. 37-59: Glucosephosphate isomerase (Gpi) a-w. 60-66: Isocitrate dehydro genase (Idh-I) a-g. 67-71: Isocitrate dehydrogenase (Idh-2) a-e. 72-78: Lactate dehy drogenase (Ldh) a-g, 79-89: Malate dehydrogenase (M dh-I) a-k. 90-99: Malate dehydrogenase (Mdh-2) a-j. 100-118: Mannosephosphate isomerase (Mpi) a-s. 119- 138: Phosphoglucomutase (Pgm) a -t 139-151: Triosephosphate isomerase (Tpi) a-m. 166 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 6 (S I O O O O TO TO O tO O O O O O O TO O O O TO O O O O O O O O O O O O tO O O O O O tO O tO O O O O O O O O O O tO O OOOOOTO O TO O O O O O O TO O tO O O O O O O O O O O O O O O O O O O O O TO tO O O ttO O O O O O dtO O O Ô O O O O O O O TO O O O O O O O O O O O O O O TO O tO O TO O O O O O O O O tO O O O O O O O O O O O O tO O O O O O O flU O O O O O O O O T O O O O O O O O O O O O O O O T O O O O O O O O O O O O O O O O tO O tO O O O O O O O O T O O O O O O (ptf O O O O O O O O TO O O O O O O O O O TO O O O tO O O O O O O O O O O O O O O O O O T O O O tO O O O O O tO O O O O O A O Jr O O O O O O O O TO O O O O tO O O TO TO O O O tO O O O O O O O O O O O tO O O O tO O O O O O O O O O O O O O O O O pptt O O O O O IO O tO O O O O O O TO O IO O O O O TO O O O O O O O O O O O O O O O O TO O O O O TO O O O TO IO O O O O O O O O O O tO O O O O O O O O O O O O O O O O O tO O tO O O O O O O O O tO O O O O O O O O O O O O tO tO O O O O tWY O O O O O O O O TO O O O O O lO O O O O O tO IO O tO O O O O O O O O O O TO O O TO O TO O O O O O O O O O O O tO m O O O O O O O TO O O O O O O O O O O O O O O tO O tO O tO O O O O O O O O O O O O O O O O Û TO O O O O TO O O O O O rm j O O TO O O O tO O O O O O O O O O O O O O O O O O O O O tO O ttO O O O O O O O O O O O O O O O O O O O tO O O O O O »rP O TO O O O O O TO O O O O O O O O O X O O O O TO O X O O O O O O O O O O O O O O O O O O O tO O O ttO O O O O O O O O iC a T O O O O O O X O O O O O O O O O O O O O X O X O O X O O O O O O O O O O O O O X O O O O O O O O O O O O O O O O O O O X w o O O O O O O O X O O O O O O O O O O O O O O O O O O X O O O O O O O O O O O O O X O O O O X O O O O O O Ô O O O O O O 0X jWO O O O O O O O O O O X O X O O O O O O O O O O O O O O X O O X Ô O O O O O O O O O O O O O 0O O O O O O O O O O O X O O O ooo 0X00000000000000000000X0X00X000X000X000000X000X000000000000X0 BHtf ooxoooooooooxooooooooxoooxoooooooooooxoooooooooooooooooooooooxooooooooxoooooooooxooooooooo 0000X000X00000000X0X000000000X00X0000X000X00000000000000000X0X0X000000X00000000000000X0000 ooooxoooooooooooxoooooxooooooxooooooooooooooooooxoooxooooxxooxooooxoooooooooooooooooooxooo 00X00000000000000X0000X000X000X0000X00000X000X000000000000000X00000000X0000X00000X00000000 0000X000000000000X0X000000000X00X00000X0000000000000000000000X0000000000000X000000000X0000 xoxoooooooooooooooxooxoooxoooooooooooxooooooxoxooooooooooooooxoooooooooooooooxoooooooxoooo ooooxoooxooooooooxoxoooooooooxoooooooooxooooooooooooooooxooxoxooooooooxooooooooooxoooooooo ooooxooxooooooooxoooooxooooooxooooooooooooooooooxoooxxoooxooooooooxooooooooooooxoooxoooooo 000X00000000X00000000X0000000X0000000000X0000000X000000000000X0000X000000000000000X00X0X00 O O O O O O O O X O O dO O O O X O O O O X X O O O O O O X O O O O O O O O O O O O O O O X O O O O O O O O O O O O O O O X O O O O X O O O O O O O O O O O O O O O O O O O X O O O O O O O O O O O O O X O O O O O X O O O O O X O O O O O O X O O O O O O O O O O O O X O O O O O X O O O O O O O O O O O O X O O O O X O O O O O O O O O O O O O O O O O O O O O O X O O O O O O O O X O O O O O O X O X O X O O O O O O O O O X O O O Ô O O O O O O O O O X O O O O X O O X O O O O O O O O O X O O O O O O O O O O X O O O O O O O O O O O O X O O O O O O O O O X O O O O O O O O O O tO O O O O X O O O O O O X O O O O O O O O O O O O O O O O O O X O O O O O O O O O O O O O X O O O X O O O O O O O O O O O O T O O O O tO O O O X O O O O O X O O O X O tO X O O O O O O O O X O O O O O O X O O O O O O O O O O O O O O O O O O O O O O X O O O O X O O O X O O O O X O O O O O O O O O O O O O O O O O O O X O O O O O O O O O X O O O O O O O O O O X O X O O O O O O O O O X O O O O O O O O O O O O O O O O O X O X O O O O O O O O O O O X O O O O O O O O O X O O O O O O O O O X O O O O O O O O 00X00000000000000X000X00X00000000000000X000000000000000000000X00000000X000000000X000000000 p0ttf£jrrA B fO IftttÛ tl eaaoeaftvt ttJtqtit roojc aitppu e m BMBfitg ettottfp eotVttotoAû ffpOMtplfJO tf4»6nxjKio » O tf r t> W O 0 O O 9 ( / ) ( /) CD Q . " O CD 2 Q . c o 3 " O 2 Q . 2 ■ c Q . 8 ( /) ( /) CD Q . " O 8 3 " O 2 Q . CD q : Table 4-3. Recoded sequence o f the lysin data of Lee & Vacquier (1995). Special cod ing strategies were employed for the listed positions, \^ e re the position refers to the current coding. Position numbers in plain type face use ^data contraction', those in bold type face employ ‘stetch coding’, “a = IT ” indicates a portion void of indels. “a = con tains T ’ indicates portions with indels; gap position is not indicated as it would imply an understood homology relation to other positions. Open Reading Frame. 21: a = con tains G; b = contains K; c = contains B C K ; d = BCKKGK e = -----. 153: a = contains KFCGBC; b = contains B C FK G K ; c = contains KQGGK; d = contains KQKGK; e = con tains GQKGBC; f = GQKQGGK; g = contains K; h = contains KG; i = contains Q; j = contains GKG; k = contains DFFGK; 1 = contains OFF; m = contams GQ. 3’ Untrans lated Region. 5: a = TT; b = contains T ;c = — . 6 :a = —; b = contains G ;c = contains T; d = GGC. 20: a = contains TG; b = contains CG; c = contains GG; d = TATT; e= con tains TT; f = — . 45-46: aa = contains A. 75: a = contains T ;b = contams TCGTC; c = contains TAGTC; d = contains TCTTT; e = TCTTTCTT; f = contains CTTT; g = con tains CCTT; h = contains TCCT; i = contains TCACT. 101-104: aaaa = contains CT. 111-114: aaaa = contains AC. 132: a = TGT; b = T%T; c = contains TT; d = GTT; e = contains TC; f = TTG. 9 = (ECJ); 8 = (HCJ); 7 = (GR); 6 = (AE). 1 6 8 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 3: OPEN READING FRAME POSITION * l *21 47* a u s tr a tlis C K W W C W G C G G G V G C G -K bP K PV G K V Q V Q K G P D V G V K V Q W G G E D c o c c in e a CKW W CW G CCV G CG V G —e ------------ Q V G K Q F D V G V K V Q W G G P D c o rru g a ta CK W W CV FG C C G G W C G -K cPK FV G K K FV K K D PD V G V K V D W G G FD c ra c h e ro d ii C K W W C V FG C C G G W C G -K bPQ FV Q K Q FV K K G FD V G V K V D W G G FD c y c lo b a te s C K W W C W G C C V G C G V G -K bF K F G K G ÎC K V G K K F D G G V K V Q V W G FD d is c u s CKW W CVFGCCGGVGCG-KbPQPVKKQPVQK(^VGVKDDW GGFD d iv e r s ic o lo r C K V W V C W G C G V G C G G G V K a F G F V G K G K V Q K G P D V G V K V Q W G G P D fu lg e n s C K W W FV FG C C G G V G C G -K bFG FV K FK FV Q K G PD V G C K V Q W G G PD g ig a n te a C K W W C V E G C C G G V G C G -K bF K F V Q K Q P V D K G FD V G V K D D W G G PD i r i s C K W V FC W G C -V G V G G G P-aFG FG G G G PV G K G K V G G V K G Q W G G FD kam tschatkana C K V F W C V F G C C G G V G C G -K aF G F V Q G K P V Q K G P D V G V K V D W V K F D la e v ig a ta C K V C W C W G C C V G V G C C -K bP K P V G K G K V G K G F D V G V K V Q W G G P D m idae C K W W PV FG C W G C V D G PK dPPFV K K Q E V G K G FD V G V K V Q W G G FD o v in a C K W W C W G C C V G V G V G -K bF G FV G G G K V G K G K D V G V K V Q W G G P Q p u s tu la ta C K W W C W G G C V G C G C G -K bFG FV G K K K V G K Q FD V G V K V Q W G G FD r o e i C K W W C W G C C V G C G C G -K bFK FV G K K K V G K G FD V G V K V Q W G G FD ru b ra C K V W V C W G C C V G V G C G -K b F K F V G K K K V G K Q F D V G V K V Q V C G G F D ru fe s c e n s C K W W C V F G C C G G V G C G -K aF K FV D G K FV Q K G F D V G V K V Q W G G F D s c a la r is C K W W C W G C C V G V G C G -K bFV FG G K K K V G K Q FD V G V K V Q W G G FD s o re n se n i C K T V W C V F G C C G G V G C G -iaaF K F V G G K F V Q K G P D V G V K V Q W G G F D tu b e r c u la ta CKWWCWGCCVGCGy— e----------- svgkqfdvgvkvqwggfd v a r ia C K V W V C W G C G V G V G C G -K b F G F V G K K K V G C G K D V G V K V D W G G F Q w a le a ie n s is C K W W C V F G C C G G V G C G -K bF Q F V G G K D V Q K G F D V G V K O Q W G G F D POS *48 105* au s K K V G G F V G K K G K K V -Q G V Q K K G V F F V Q K K F C Q G K F Q G F C V F V Q K Q V G K V G K G G G D G D C O C K K V G G F V Q K K G K G V -G G V Q K K G V F F V Q K K F C Q G F F Q Q F C V F V D D K V K K V G K G G Q V Q D c o r K G W K FV K V K G G K V -G G 7 Q K K G V F F V Q K K F C Q G K F Q Q F C V F W K K G D G V G K G G W G D e r a K G W K FV K Q K G K G V -Q D Q Q K K W F F V Q K K F C Q G K F Q Q F C V F W K K G Q G V G K G G W G D eye K K V G G F V G K F G K Q V -G G V K K K 6 V F F F Q K K K C Q G K F Q 6 F C K F G D K 0 V K K V G K G G Q G Q D d is rgvqkfvggkgkgv- ggdqkkgvffvqkkfcqgkfqqfcvfwkkvdgvgkggwgd div K K V G G F V (n a C G G K V -G G V 9 K K G V F F V Q K K F V Q G K F G G F C V F V Q K K V G 6 V G K G G G G G D f u l K Q V G 6 F V K V K 6 R K V -G Q Q Q K R G V F F V Q K K F C Q G R F Q Q F C V F V K K K V K 6 V G K G G G V G D gig K G W K F V K V K G K G V -G G V Q K K G V F F V Q K K F C Q G K F Q Q F C V F W K K V D G V G K G G W G D iri K K yG G FV K K K G Q K \r-G SV Q K K G V FFV Q K X FC Q qK FK 6FC K FV K Q gC K W (^G D G G G D kam K K W G F V K V Q G G G V -G K V Q K K G V F F V Q K K F C Q G K F G Q F C V F V Q K K G D G V G K G G W G D la e K K W G F V Q K K G K K V -G G V Q K K G V F F V Q K K F C Q Q C F Q G F C V F V G K Q V G K V G K G G G V Q D mid K K V G G F V K K K G G K V K G G V K K K G V FF V Q K K F C O G K F Q gF C V F W K K V K gV G K G G G V G D OVi K K V G Q F V 6 K K G 6 K V -G 6 V Q K K G V F F V Q K K F C Q G K F G G F C V F V K K Q V G K V G K G G Q G Q D pus K K V G G F V G K K G G K \r-G G V Q K K G V F F V Q K K F C Q G V F Q Q F F K F V G K K V G K V G K G G Q V Q D ro e K Q V G G F V Q K K G R K 7 -G 6 V Q K K G V F F V Q IC K F C Q 6 K F Q G F C V F W K Q V 0 K V G K G G Q V G D ru b kkvtqfvgkkgkûv- ggvqkkgvffvqkkfcqgkfqçfcqfwkdvdkvgkggqvqd r u f K G W K F V K V K G K G V -G G V Q K K G V E F V Q K K F C Q G K F G Q F C T E V Q K K V D G V G K G G W G D s e a K K g V Q F V K K K G K F V -G g g O K K G V E F T Q K K F C Q G K F Q G g C V E W K Q V G K V G K G G G V G D s o r K G W K F V K G K G G G V -G K V Q K K G V F E V Q K K F C Q G K E G Q F C V F V Q K K G D G V G K G G W m tu b K K V G G F V Q K K G G G y -G G V Q K K G V F F V Q K K F C Q G g g Q Q g C V F V D D K V K K V G K G G Q V Q D v a r K K V D V EV G K K G G K T r-G G V Q K K G V gF V O K K g C Q G K F G Q F C V g V K K K V Q G g G K G G G G G D w al K G W K F V K G K G K G T ^atV Q K K G T F F V Q K K F C Q G K F G Q F C V F V Q K K V D G V C T G G T V G D 169 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 3: continued PCS *106 *153 aus E G G V G G D V G ÏC K V G T D V G -F G E V V K K Q W G K P K G F C G G V C G K K V Q D V G V ta C O C F G G V G G D V G K K V G V Q V G V E G V W K P Q W G P K V G P C G K W G V K V D D V G V I c o r F G K V G G D V G K K V D C G F F -F Q F V Q G K Q C V G K F V G F C D D V Q K C K G G D V G V e e r a F K G V G G D V G K K V D C G F F -F Q F V K Q K V C V G K F V G F C K K V Q Q C K G D D V G V f eye F G K V G G D V G iO tV G V D V C -F G F C V K K Q C V G K F K G F C G V W K K K K D D V G V i d is F K K V G G D V G K K V D C D F F -F Q F V K Q K Q C V G K F V 7 F C G 6 V Q K C G G K D V G V d d iv F Q Q V G G D V G K K V G C D V G -F G F W K K Q W G K F K G F C G D V C G K G G G D V D V j f u l F G K V G 6 D V G K K V D C V F F -F Q F V G G K K C V G G F G 6 F C G K 7 Q G V K G G D V G V d g ig F K K V G G D V G K K V D C D F F -F Q F V K Q K Q C V G K F C G F C D D V Q K C G G K D V G V t i r i F Q K V G G G V G K K V G V D V G -F G F W K K Q W G F K V G F C G K V C G K K G G D V G G j kam F K K V G G D V G K K V D C V F F -F Q F V G G K Q C V G K F C G F C D Q V Q K C K G G D V G V a la e F G K V G G D V G K K V G V D V G -F G F W K K Q W G K C K Q F C K D W G K G V Q D V G V g mid F G G V G G D V G K K V G V D V G -F G F W K K Q W G G F K G F C G K V C O T K G G D V G V g O V i F G K V G G D V G K K V G C D V Q -F G F W K K Q W G K F K Q F C G Q V C G K K V D D V G V i pus F Q G V G G D V G K K V G V D V G -F G F W K K Q W G G F K Q F C G Q W G K K V D D V G V g ro e F G F V G G D V G IC K V G V D V G -F G F W K K Q W G Q F K Q F C G Q V C G K K V D D V G V g rub F G K V G G D V G K K V G V D V G -F G F W K K Q W G K F K Q F C G Q W G K K T D Q V G V g r u f F G K V G G D V G K K V D C G F P -F D F V K D K Q C V G K F V G F C D D V Q K C K G G D V G V a s e a F G F V G G D V G K K V G V D V G -F G F W K K Q W G D C K Q F C K D V Q G K K T D D V G V b s o r F G K V G G D V G K K V D C D F F -F Q F V K G K Q C V G K F V G F C D D V Q K C K G G D V G V a tu b F G G V G G D V G K K V G V Q V G V F G V C V K F K W G K F K G F C G K W G V K V D D V G V k v a r F K Q V G G D V G K K V G V D V G -F G F W K K Q W G Q F K Q F C G D V C ratK V G D V G V j w al F G K V G G D V G K K V D C D F F -F Q F V K G K Q C V G K F V G F C D D V Q K C K G G D V G V b 170 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 3: continued: 3’ UNTRANSLATED REGION POSITION a u s tr a li s c o c c in e a c o rru g a ta c ra c h e ro d ii c y c lo b a te s d is c u s d iv e rs ic o lo r fu lg e n s g ig a n te a i r i s kam tscbatka la e v ig a ta m idae o v in a p u s tu la ta r o e i ru b ra ru fe sc e n s s c a la r is so re n se n i tu b e rc u la ta v e tria w 2G.eULlensis *1 5**6 *20 43* T A A A ab C A C & T G G A T A C A A aC G C C A ---------------------- T C C C TAAAcaCACATGOAACCAAaOGCAAr—— — — — — c a ------^ T G A A A C C A A b A A G a A G A A A G C T G C A T G C A C C C T c a ------T G A A A C C C & b A A G A A G A C A G C T G C A T G C A C C G T TAG-ca-AC-TGGAACCA&aCACAA----------------------C C G T c a ------T G A A A C C C A a A A G A A G A C A G C T G C A T G C A C C G T TAA-acCACGTTGA AGTaAaTGCAA----------------------C C G T c a ------ T G aA A C C C A c A A G A A G A C A G C G T C A T G C A C C G T c a ------ TGAAACCCaaaaGftAGaCAGCTGCaTGCacCGT T A A r-adC A C G T G G A G — f —— ——— —— — —C C G T c a ------T G A A A C C C A b A A G A A G A C A G C T G C A T G C A C C G T T A A -abC A C G T G G ftA C C A A aC A C A A ---------------------- C C G T T A A -abC A C G T G G A A C C A A aT A C A A ----------------------C C G T T A A -abC A C G A G G A A G T A A dC G C A C ---------------------- C C G T T A A -caC A T G T G C A A C C A A eC G C A A ---------------------- C C G T TA A r-abC A C G T G G ftA C C A A aC A C ftA ----------------------C C G T T A A -bbC A C G T G G A A C C A A aC A C A A ----------------------C C G T c a ------T G A A & C C C A a A A G A A G A C A G C T T C A T G O V C C G T T A A -abC A C G T G G A A C C A A aC A C & A ---------------------- C C G T c a ------ T G A A A C C C A a A A G A A G ftC A G C T G C A T G C & C C G T TA A -caC A C A T G G A A C C R A aG G C A A ----------------------C C G T T A G A b b C A C G T G G A A C T A A eC A C A A ---------------------- C A C G c a ------ T G A A A C C C A b A A G A A G A C A G C T G C A T G C & C C G T PCS **45-46 *75 98* au s T A T C T G & C G E A A A A A G & C G G & G & -T G G G G —d C A T C A A G C A C T G G A T G G & G C C -A . COC G A C A T G A T G T A A A A A G A C G G A .-------------------- hC A T C A A G C aC A G G A T G G & G C C -A c o r G -C A T G ----T C A A C A G A T A C A G A A A A C A ---- b C G T C A A A C A C G A G G ----- AGCT-G e r a GaaATG--- T C A A C A G A T A C A G A A A A C A ----b C A T C A A A C A C G G G G ----- AGCT-A eye G A C A T G A C G T A A A G A G A C G G A T G G G G G -d C A T C A A A C G C A G G A T G G G G C C -A d is G -C A TG--- T C A A C A G A T A C A G A A A A C A ---- a-------------------------------------- d iv G A C G T G G C G T A A A A A G A C A G A G A A C G G G A —a --------- G C A T T G G A T G G A G G C -A f u l G -C A T G ----T C A A C G G A T A C A G A A A G C G ---- cC A T C A A T T A C G G A G ----- AGCT-A g ig G -C A T G ----T C A A C A 6 A T A C A G A A A A C A ---- a C A T C A A G C A C G G G G ----- AGCT-A i r i G -C C C G ----T T A A C A G A A G G C A --------------iC A T C A A A C A C T T G G G A G C A A kam G -C A T G--- T C A A C A G A T A C A G A A A A C A ---- a C G T C A A A C A C G G G G ----- AGCT-A la e G A C A T G A C G T A A A G A G A C A G A G A -T G G G G G -d C A T C A A A T G C T G G A IG G G G C C -A m id G A C C C G A C C T A A A G A G A G A G A G A -T G G G G G G d C A T C A A G C G T T G G A T G G A G C C A A 6ACTT6AT6TA--------- CA6G6A-T6GAG—eC A T C A A G C A T T G G A T G G A G C C -A G A C A T G C C A T A A A A A —C G 6 A G A T T G G A fC A T C A A G C G T T G G A T G G A G T C -A G A C A T 6A C G T A A A G A S A C A G C C A -T G G G G T -d C A T C A A A T G C T G G A T G G G G C C -A S A C A T G A C eC A A A G A G A C A G A G A -T G G G G G -d C A T C A A A T G C T G aA T G G G G C C -A G -C A TG--- T C A A C A G A T A C A G A A A A C A a C G T C A A A C A C G G G G ---- AGCT-A s e a G A C A T G A C G T A A A G A G A C A 6 A G A -T G G G G G -d C A T C A A A T G C T G G A T G G G G C C -A s o r G -C A TG--- T C A A C A G A T A C A G A A A A C A ---- a C G T C A A A C A C G G G G ----- AGCT-A tu b G A C A T G A T G T A A A A A G A C G G A --------------- h C A T C A A G C A C A G G A T G G A G C C -A v a r T A C C T G A C G G A A A N N G A C A 6 G A A -T G G G G A -fC A T C A A G C A T T G G A T G A A G C C -A w al T G C A T G --- T C A A C A G A T A C A G A A A A C A ---- a C G T C A A A C A C G G G G ----- AGCT-A OVX pus ro e ru b r u f 171 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 3: coatmued PO S ****101-4****111-U 4 *132 149* aus T G A T G T -------C A T C C C A A C G C G G C G T -G C C A G A b---------T G C A C C C A A A A C O C T G & C ------------C A C G C A A T G C G G G A T G 6 A C A G A C -------- T A C A C C C A A T A c o r T G A C A G C A T C C A T C C C A T C A T G G G T T G G C T A T T a--------------------------- e ra T G A C A C ------ C A T C C C A A C A IG G -----------------a--------T G C A C G -------- eye T G A C G T -------C A T C C C A G C G C T G G G T G G C C A G A c--------TTCA CCA G A IA d i s ---------------- C A T C C T A T C A T G G G T T G G C T A T T a--------T G C A C G --------- d iv T G A C G T ---------- G C C C A G C A C G G G T T G G C C A G A c--------------G C C T A A T A f u l T G A C A C ------ C A T C G C A T C A T G G G T T G G C T A T T a---------T G C A C G C A A T G g ig T G A C A C ------ C A T C C T A T C A T G G G T T G G C T A T T a-------- T G T A C G -------- i r i T G A C A C ------ T A T C C C A A C A G G G G T T G G C C C T T f-------- A G C A C A A G A C A kam T G A C A C ------ C A T C C T A A C A T G G G TT G G C T A IT a-------- T G C A C G --------- la e G G A C G T ------ C A T C C C A G C G C G G G G T G G C C A G A b -------- T T C A C C A G A T A mid T G A C G T ------ C A T C C C A G C G C G G G G T G G C C A G A C -------- T T C A C C G G A T A Ovi T G A C G T --------G C C C A G C G C G G G G T G G C C A G A b ----------T G C C C C C A G T A pus T G A C G T -------C A T C A T A A T G C G G A A T G G C C A G A e---------G G C A C C C A A X A ro e T G A C G T -------C A T C C C A G C A C G G G T T -G C C A G A b---------T T C A C C A G A T A rub T G C C G T ------ C A T C C C A G C G C G G G G T G G C C A G A b ---------T T C A C C A G A T A ru f T G A C A C ——C A T C C T A A C A T G G G T T G G C T A T T a-------- T G C A C G --------- se a G G A C G T ----- C A T C C C A G C G C G G G G T G G C C A G A b -------- T T C A C C A G A T A s o r T G A C A C ------ C A T C C T A A C A T G G G T T G G C T A T T a-------- T G C A C G --------- tu b T G A C ---------- C A C G C A A T G C G G G A T G G A C A G A c ---------T A C A C C C A A T A v a r TCaaaa--------- a a aaA G A G C G G G A T G A C T A G A dC A G A G T T G C C C C C A G T A w al T G A C A C ------ C A T C C T A A C A IG G G T T G G C C A T T a-------- T G C A C G --------- PO S *150 194* aus T A T T C T A T T T A A C T C G T A A A C A T T 6 T A C A A T A G T G T A C A A T A G T C O C AITTGTAITTAACTGCTGAATAIT--------------T G T A C T C ------- c o r A T T T T T A A C A C A 6 A A A G A T T — ---------------------- e r a A T T T T T A A C A C G 6 A A A G A T T —----------------------------- eye T A T T C T A T T T A A C T C G T A A A T A T T — —— — ———— — — d is — — A IT T T T A A C A C G G A A A G A T T — — —— — — — — d iv T A A C C T A T T T G A C C C G T C C A T A T G --------- f u l C A C G A iC nT T A A C A C C G A A A G A T T - g ig ——A T T T T T A A C A C G G A A A G A T T - i r l C A C G A T T T T T A A C A C G G 6A A C A T T - k am A T T T T T A A C A C G G A A A G A T T - la e T A X T C T A T T T C T A A T G G A A A C A T T - mid A A X T C T A T T T A A C T C G T A A A C A T T — o v i T A T T C T A ITT A G C T C A T A C A C G T T --------- pus T A X T G T T T T T A A C T C G T A A C C A T T --------- ro e T A T T C T A T T T A A C T C G T A A A C A T T —— — rub T A X T C T A T T T A A C T C G T A A A C A T T - tu f ——A T T T T T A A A A C G G A A A G A T T — —— — se a T A T T C T A T T T C T A A T G G A A A C A T T — — — — s o r — — A T T T T T A A C A C G G A A A G A T T - tu b A T T T G T A T T T A A C T G G T G A A T A T T -------------- TG T A C T C - v a r T A T T C T G C A T A A C IC G C C A A C A T IT ------- w al A IT T T T A A C A C G G A A A G A T T — 172 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 3: continued POS *195 227* au s 6 T A C A A T A G T G A A X A A A T A C T T T T 6A C T A 6A T — COC G T A A A A T A G T G A A IA A A C A T T TT C 6A C A C A G T 6- c o r GTACAirCCAGAATAAATAGATrE-ACTA------- e r a G T A C A T T C C A G A A T A A A T A G A T T T -A C T A ------- cyc G T A C A A T A G T A A A T A A A X A C T T T T 6 A C T A 6 A T — d is G TA C A IT C C A G A A TA A A TA G A TT T -A C TA ------- d iv G T A C A A T A G T A A A T A A A IA A IT T T G A C T A G ------ f o l GTACAITCCAGAAIAAAIAGATTT-ACTA------- g ig G T A C A T T C C A G A A T A A A T A G A T T T -A C T A ------- i r i G T A C A IT C A G G A A T A A A T T G T T T T T A ------------ kam G T A C A !E T C C A G A A T A A A T A G A IT T -A C T A ------- la e G T T C 6A T A G T A A A T A A A T A C T T T T G A C T A ------- mid G T A C A A T A G T A A A T A A A T A T T T T T G A C T A G ------ o v i G T A C A A IA G T A A A T A A A T A A ÎT T T G A C T A G G T — pus G T C C A A X A 6TC A A IA A A X A A 6T T C TA C TC TA T G - ro e G T A C G A T A G T A A A IA A A IA C T T T T G A C T A ------- ru b G T A C A A T A G T A A A T A A A T A C T T T T G A C T A G A T — r u f G T A C A T T C C A G A A T A A A T A G A T T T -A C T A C A A A A sc a G T A C G A T A G T A A A T A A A T A C T C T T G A C T A 6 A C — s o r G T A C A T T C C A G ftA T A A A T A G A T T T -A C T A ------- tu b G T A A A A X A G T G A A T A A A C A T T T T C G A C A C A G T G - v a r G T N C A A IA G T G A A T A A A T A C T T T C G G ------------ w al G T A C A T T C C A G A A T A A A T A G A T T T -A C T A ------- 173 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. DISTRIBUTION The data of over 4,000 lots representing approximately 3,000 localities for the dis tribution o f all the 56 species in the family Haliotidae grouped by larger geographic region is given at the end o f the paper. On the order o f 10,000 lots were inspected for the study. A number o f reasons can be cited for exclusion o f a particular lot: no or unprecise locality data; impossible locality data (e.g., the Australian endemic K ele - gems Philippi, 1844, with label from the Mediterranean); redundant lot for any given combination o f species and locality. The number o f data points for each species range from 1 {K crebrisculpta Sowerby, 1914: only a single specimen known. See Stewart & Geiger, 1999) to 317 (H. varia Linnaeus, 1758). Seven species afford more than 100 individual localities (Æ asinina Linnaeus, 1758, K cracherodii Leach, 1814, H. kamtschatkana Jovtas, 1845, H. ovmaOmelin, 1791, H. rubra Leach, 1814, H. tubercu - lata, H. varia) and for 12 species less than 10 localities are available. Some species are known from a reasonable number o f specimens, but mostly from only a few localities. Striking examples are K elegans and K sem iplicata Menke, 1843, for which most specimens come from Freemantle and Rottnest Island. A mean of 55 and a median o f 28 localities per species are listed. The data are arranged in the following format: 1) Specimen data by country/larger area in a general east-west and/or north-south organization: COUNTRY/LARGER AREA: Specific locality (lot number, number of specimens). The second lot from the same locality is separated by a semicolon. No more than two lots are listed for any given locality. The term Complete’ indicates preserved animals. Parentheses mdicate speOmg variations and alternative names o f the localities, documented in the literature or by other, objective means. Brackets indicate spellmg variations and potential misspellings (erroneous copy m the past, problems with hand written labels) stemmmg fix>m my personal juc^ment. 174 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2) Specimen data from the literature by author: locality (Author, Year). Here only localities not documented under I) are listed Any o f these additional localities is listed once, despite potential, multiple mention in other publications. 3) Range data from the literature. Range indications with at least one specific delim- inators were included whenever possible. Ranges stemming from apparent misidentifi- cation and other erroneous indications have been omitted The following abbreviations are used: ex: separated fiom that lot o f non-conspe- cific specimens. GBR: Great Barrier Reef. Isl.: Island(s); N, E, S, W: North, East, South, West. WA: Western Australia. QLD.: Queensland. NT: Northern Territory. NSW: New South Wales. SA: South Australia. TAS: Tasmania. VIC: Victoria. PNG: Papua New Guinea. [?]: questionable identification. No #: no catalog number available. Ten figures show dot maps for each species. The maps are grouped by geographic area: Mediterranean, west Africa, south Afirica, and east Africa (Figure 4-14); Indo- Pacific (Figures 4-24,4-43,4-44); Australia and New Zealand (Figure 4-67); north western Pacific (Figure 4-86); northeastern Pacific (Figures 4-87,4-100,4-107); and western Atlantic (Figure 4-107). Distributional patterns Abalone are found along most roclgr shores in tropical and temperate waters. Most are encountered in the shallow subtidal between sea level and 30 m depth. The substrate requirement explains the absence o f representatives from northeastern America as well as from the western coast o f the Didian subcontinent because sandy and muddy condi tions prevail there. A striking absence on western South America has to be mentioned where not a single species o f abalone is found despite rocl^ shorelines and suitable food sources m the form o f kelp. This pattern is a small mystery particularly as abalone 175 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. are so prominent on. the west coast of North America down to the tip of Baja California, Mexico. Part o f the answer may be found in the warm-water barrier between the two temperate areas on either side o f the equator. However, the occurrence o f the two abalone species on Galapagos Islands {H. dalli Henderson, 1915) and Cocos Island (/£ roberti McLean, 1970) invalidate this argument to a certain extent, although ocean cur rents certainly facilitated dispersal to those particular islands. There are no abalone species o f global distribution. The largest areas are inhabited by some Indo-Pacific taxa. H aliotis clathrata Reeve, 1846 (East Afiica to Samoa: Stew art & Geiger, 1999), H. asinina and ff. planata Sowerby, 1882 (Thailand to Fiji), and H. ovina (Maldives to Tonga), are the most widespread taxa. AU other species have much more restricted ranges. Four discrete regions o f endemism can be identified. In South Afnca five species are encountered, in New Zealand three are found, the western North American coast harbors six species, and Australia has nine endemic species (excluding K rubiginosa Reeve, 1846, and treating H. hargravesi as a form o f H brazien). Within these areas the overlap with widespread species varies from none in western north America and New Zealand, to slight as shown by the mUequent Unds in the South Afiican province o f K pustulata Reeve, 1846, which has a core distribution in east Afiica, to substantial in AustraUa. In the last, several species o f tropical Indo-Pacific distribution can be found along the warmer coasts o f Australia (i.e., H. asinina^ K clashratay H. ovina, K rubigi - nosa, K squamatd). However, those species that seem to prefer more temperate waters {H. brazieri, K coccoradiata Reeve, 1846, H. cyclobates Péron & Lesueur, 1816, Æ elegans, K laevigata Donovan, 1808, K .ro ei Gray, 1826, H. rubra, K scalaris Leach, 1814, Æ sem iplicata) are endemic to the Australian continent. The Australian record (Macpherson, 1953) o f the weU-known Æ spadicea Donovan, 1808, a common and 176 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. recreationally fished species endemic to South Afirica, is rejected. Some areas contain single species such as the eastern hido-Pacific, i.e., Polynesia and surrounding islands {K ptdcherrim a Gmelin, 1791), the eastern Pacific with Galapagos (K dalli) and Cocos (Æ roberti)^ and west Afirica (Æ marmorata Linnaeus, 1758). BIOGEOGRAPHY Biogeographical areas Biogeographical areas are often somewhat vaguely defined owing to the continuity o f geographic space. The limits o f biogeographic areas are then somewhat artificial. Three particular problems were encountered in the present study. I) Oregonian province: The Oregonian province can be regarded as a transitional area between the Californian and the Aleutian province (cf. Eckman, 1953). The boundary area exists for vhtually every area, hence, the allocation o f the Oregonian area to the two others will only shift the problem of the boundary, but not solve it. Here the Oregonian province has been taken as a separate area. 2) Northwest Australia: this area is sometimes united with west Aus tralia, but is retained here as a separate un it 3) New Caledonia and Papua New Guinea: These two areas are tentatively included within the Central Pacific province. Taxonomic issues This work follows the general treatment o f abalone taxa as outlmed in Geiger (1998a) and references therein. Although the treatment is comprehensive, some problematic issues remain Two particular cases have ramifications in the present context: K diver - sicolor - K squamata (Figures 4-40-42,4-63,4-66): characters 14 and 50; K brazieri - ff, hargravesi ^ ig in es 4-45,4-46,4-48,4-49): characters 4 and 22 (Table 4-1). The former may be two distinct species in Japan and western Australia, both with sparse 177 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. records in the Indo-Malayan region, or it may be a single widespread species. A further option to entertain is an antitropical distribution o f two potential sister species with extinction in the tropical region (cf. Briggs, 1999). The latter case may also represent two distinct species with identical distribution or a single polymorphic species. To address these problems and their effects on the biogeographic analysis, the data matrix was manipulated as detailed in materials and methods. Geiger (1998a) tentatively resurrected a second Mediterranean species under the name H. neglecta Philippi, 1848, based on seven shells only (Figures 4-147,4-150). Recent efforts have produced additional material o f this species including some ani mals, which clearly show the species to be distinct from the better-known, Mediter ranean K tuberculata (Figures 4-4,4-5,4-7,4-8, 4-12,4-13,4-172,4-174). B. Owen (pers. comm.) pointed out that the correct name for this second, Mediterranean species should be H. stomatiaeformis Reeve, 1846, itself a highly confused taxon. I use the cor rect name for this species here, but we will provide a more extensive treatment o f all the taxa involved elsewhere. Area cladograms Area cladograms show the relationship between geographic areas using a cladistic approach. The areas are treated as traditional taxa (areas-as-taxa) and the distributions o f the species serve as the characters (taxa-as-characters: Table 4-1). Any species occur ring in more than one area helps to illuminate the relationship between the areas, hence, is an informatwe taxon-as-characten The nmnber o f informative taxa-as-characters was 41 out o f the 56. The strict con sensus trees o f the MPRs are shown in Figure 4-2. Two different consensus topologies were recovered (Figures 4-2: A, B). The data-matrix as shown in Table 4-1 produced 178 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. E Northwest Australian Sotanderian Peronian South Australian West Australian Indo Malayan Sub-tropical Japan Central Pacific Central Indian Ocean Eastern North Pacific Aleutian Oregonian Caltfonuan Tropical East Pacific Caribbean Argentinian New Zealand South African P Western Indian Ocean - Red Sea 1 West Afiican “ European Mediterranean North Atlantic 1 2 P U ■ z s § B Peronian South Australian West Australian Northwest Australian Solanderian Central Pacific Indo Malayan Sub-ttopirâl Japan Central Indian Ocean Eastern North Pacific Aleudan Oregonian Qlifbmian Tropical East Pacific Caribbean Argentinian New Zealand Red Sea « South African - Western Indian Ocean 3 WiKt African “ European Mediterranean North Atlantic P U g Figure 4-2. Area cladogram ^ro o ks parsimony) for the genus H aliotis. Strict consen sus all MPRs. EA: East Afiica. EM: European-Mediterranean. IP: Dido-Pacific. NZ: New Zealand. SA: South Afiica. W Atl. : western Atlantic. A) From data matrix as shown in Table I, and when H. brazieri (taxon-as-character 4) is excluded. B) From data matrix combinmg H. diversicolor and H. squamata mto one widespread taxon. 179 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. the first topolo©r (Figure 4-2: A: 6 MPRs, length = 67, Cl = 0.612, RI = 0.679, RC = 0.416). The same topology was found when H brazieri and K hargravesi where treated as being the same species, i.e., when taxon-as-character 4 was excluded (6 MPRs, length = 66, Cl — 0.615, RI = 0.688, RC = 0.423). The topology of Figure 4-2: B resulted when H. squam ata and H. diversicolor were treated as one widespread species (8 MPRs, length = 63, Cl = 0.635, RI = 0.716, RC = 0.455), and also when K brazieri and H. hargravesi were treated additionally as being the same species (8 MPRs, length = 61, Cl = 0.639, RI = 0.725, RC = 0.464). Branch swapping on non-minimal length trees did not recover any additional MPRs as compared to simple heuristic searches. All searches with branch swapping on trees two steps longer than the most parsimonious trees resulted in memory overflow (more than 32,700 trees). All consensus trees are charac terized by a fairly broad basal polytomy, with additional resolution in more specific areas. Skewness or gj was -0.77 (± SE = 0.0014, n = 5), which is highly significant accord ing to the values provided by Hillis & Huelsenbeck (1992:25 taxa, 50 characters for p = 0.05 g, = -0.10; for p = 0.01 g, = -0.11). These contradictory results—highly signifi cant gj versus broad basal polytomy and over 32,700 MPRs with only two additional steps—support the suspicions voiced by BCâliersjd et aL (1992) that g, does not measure information content o f the data matrbc. The common feature o f both consensus topologies are as follows. A European-west Afiican group, a south Afiica and western Indian Ocean group, a broad Indo-Pacific group, an eastern Pacific group, and a western Atlantic group are recognized in all ana^- ses. A number o f regions are united in a basal polytomy. The position o f the New Zealand and the tropical eastern Pacific provinces can be explained as due to ail the respective species being endemic, i.e, no wide-spread species occurring there (New 180 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Zealand: K australis Gmelin, 1791, K iris Gmelin, 1791, K virginea Gmelin, 1791. Tropical eastern Pacific: K dalli, H. roberti). These species constitute aut^m orphies for the regions, hence it is impossible for these regions to show relationships with other provinces. The same applies also to the western Atlantic provinces, i.e., the (Caribbean, Argentinian) group. The remaining areas are placed in the basal polytomy due to uncer tain associations with other areas. The differences between the two consensus topologies lies in details of the associa tion o f provinces within the Indo-Pacific group. Figure 4-2: A shows four o f the five Australian provinces united, and an (Indo-Malayan, subtropical Japan, central Pacific) group separated from these Australian provinces. The central Indian Ocean and north west Australia join basally. In Figure 4-2: B, however, only three of the five Australian provinces, i.e., the southern provinces, are united, whereas the warm-water Australian provinces are found more closely associated with the (Indo-Malayan, subtropical Japan, central Pacific) group. The eastern north Pacific is a basal off-shoot fiom the latter. Hence, the main difference is the affiniQr o f the warm-water Australian provinces with either the remaining Australian provinces (Figure 4-2: A) or with the Indo-Malayan province and associated areas (Figure 4-2: B). The basal placement of the eastern north Pacific is difficult to explain as it lies between the large polytomy representing a broad Indo-Pacific region (see below), and a subset thereof. I f one traces back all the faunal provmces to the large polytomy, the latter repre sents a broad Indo-Pacific region fiiom which all the more resolved groupings radiated. This pattern is evident in both consensus trees shown m Figures 4-2: A and B. The bio geographical analysis then best supports the Indo-Pacific hypothesis for the origin o f the family, as discussed by Lmdberg (1992), and by Briggs (1999) for other taxa. How ever, some contra-indications are found within the analysis. The Indo-Pacific origin IS l Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. would imply that the northeastern Pacific provinces should be derived fiom subtropical Japan, but these are instead found in a basal position. To see whether a single erratic position o f some provinces may be responsible for the large basal polytomy, a 50% majority rule consensus tree was calculated. It was identical to the strict consensus tree showing that there is widespread disagreement among the individual MPRs regarding the position o f the areas grouped in the basal polytomy. The basal position of the broad Indo-Pacific and the northeastern Pacific groups seems also compatible with Talmadge’s (1963a) Pacific Rim hypothesis. It would, however, imply an origin o f the Indian Ocean and eastern Atlantic provinces fiom within the broad Indo-Pacific group. However, the Indian Ocean as well as the eastern Atlantic groups stem fiom the basal polytomy as well. Accordingly, the best supported hypothesis for the origin o f Haliotidae is within a broad Indo-Pacific region. From the present data, the precise location o f the origin can not be pinpointed more precisely. Cladistic analysis of taxa In the total evidence cladistic analysis for 25 taxa and 531 characters, 285 were par simony informative. 98 MPRs (length = 904, Cl = 0.621, RI = 0.760, RC = 0.472, g ^ = -0.533 ± SE = 0.00277) were recovered in the initial search. Branch swappmg on non- minimal trees o f m a x im u m length=908 did not recover any more parshnonious topolo gies or any additional MPRs. A comparison o f the area cladogram (Figure 4-2) to the taxon cladogram (Figure 4- 3) show a general agreement between the two analytical approaches. Most o f the Aus tralian (Figure 4-3: Australia) and all the north Pacific species are united as larger monophyletic groups, with some additional groupings in the Indo-Pacific (Figure 4-3: IP) and the Mediterranean (Figure 4-3: EM) provinces. The north Pacific group, unitmg 182 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. r C roei scalaris laevigata rubra cyclobates coccoradiata pustulata coccinea tuberculata australis ovina varia diversicolor midae virginea iris rufescens sorenseni kamtschatkana walallensis ■gigantea § discus EA II NZ I SA NZ NZ I □ corrugata cracherodii — fulgens 'Z Figure 4-3. Midpoint-rooted taxon cladogram o f the genus H aliotis. Strict consensus tree o f 98 MPRs. EA: east Afiica. EM: European-Mediterranean. IP: Indo-Pacific. NZ: New Zealand. SA: south. Afiica. 183 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. both the eastern and western regions, is very strongly shown in Figure 4-3 (North Pacific) as well as in a separate analysis o f various sub-sets o f the underlying data (not shown, see also Brown, 1993; Lee & Vacquier, 1995). The most unsettling problem relates to the position o f the New Zealand species [H. australis, K iris, K virginea Fig ure 4-3: NZ), which are all endemic to New Zealand. They are distributed throughout the tree, an outcome that is seen even more pronoimced in analyses of sub-sets o f the data (not shown). These three species seem to result fiom individual colonizations of the island as contrasted with a small radiation in this rather isolated region. The basal split o f H. australis fiom the remaining Australian species makes intuitive sense. The position o f the other two taxa is more challenging to interpret. H aliotis iris is wedged in between the Californian H fiilg en s Philippi, 1845, and the root o f the cladogram. The root would then be traced to some general south Indo-Pacific region. The New Zealand H. virginea is found in a polytomy with Indo-Pacific species and the south Afiican H. midae Lirmaeus, 1758. A general southwest Indo-Pacific origin is implied fiom that polytomy. The root can be traced back to a general Indo-Pacific region. This rather suprising pattern o f multiple colonizations o f New Zealand agrees with finds fiom other groups o f organisms such as beeches (N otofagus), Afiican violets (Gesneriaceae), a caddis fiy genus {HydrobioseUa\ and pla^cercine parrots discussed by Humphries & Parent! (1986). The type species o f the genus HaliotiSy H. asinina (Figures 4-35,4-36,4-165,4- 166), was not inclu& d in either stucty (Brown, 1993, Lee & Vacquier, 1995). Accord ingly, any pertment taxonomic decision can not be made at this time. The discussion o f supraspecific taxa in Lee & Vacquier (1995) also suffers fiom this condhioiL Some o f the disagreement between the area cladogram and the taxon cladogram may be explained by the taxon sampling o f the taxon cladogram. Note for instance that 184 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. the European species K L tuberculata and K tuberculata coccinea Reeve, 1846, are removed from the South African H. midae. It may be due to the missing west African H. marmorata^ and only one o f the five South African species being included in the analysis. Hence, some minor disagreement between the biogeographical and the taxon analysis should not distract from the overall pattern pointing to a broad Indo-Pacific origin postulated by both analytical approaches. Dlnstrations of species. All 56 described species in the family Haliotidae are figured here, along with one species in the process o f being described (Owen e t aL, in review). Those species that have hardly been illustrated are shown in color, the better known species in black and white. Indications to published illustrations can be found m Geiger (1998a). Some species have distinct juvenile forms, which deserve particular attention. H aliotis iris (Figures 4-74-79): the adults o f this species are very well-known from New Zealand and difficult to misidentify. However, the subadults lack the characteristic striated muscle scar and the only slightly widened columella may easily be missed. Juveniles lack not only the muscle scar, but also the wide columella, and are often con fused with adult H. virginea (Figures 4-69,4-70,4-72,4-73). The latter, however, has a lighter shell, more subtle cording, and often an undulating dorsal surface. H aliotis midae (Figures 4-142-145,4-148): The adults o f this commercial species endemic to South Africa has two distinct color morphs. In the eastern part the dark red form is found, in the western part the shells o f adults are Write. The strong lamellae and the numerous, elevated, but small holes are common to both. The juveniles, on the other hand, have an entirly smooth shell, which is always dark earthy-red m color, the larger tremata are not elevated at all, and the overall shape is more elongated than the almost 18 5 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. '1 5 - round adults. Often the nacre in the sphe region has an orange tinge, which has led to misidentifîcations o f Juvenile H. midae as K spadicea (= K sangumea. Figures 4-151- 153). Juveniles o f H. spadicea^ however, have an arched and inflated shell contrasting to the flat shell o f Juvenile H, midae, H aliotis spadicea also shows a few distinct spiral cords, whereas Juvenile H midae show numerous, very fine spiral threads. H aliotis asinina (Figures 4-35,4-36,4-165,4-166): Adults are characterized by the very elongated, smooth, and relatively thin shell of green or brown color with white tri angular markings. Juveniles up to a size o f 3.5 - 4 cm show distinct spiral ridges, which display a crenulated color pattern o f alternating white, red, green, and sometimes even blue dots (Figure 4-166). Juveniles can not be confused with other abalone species, but often remain unidentified due to the discrete diiSerences to adult shells. Color variability is extensive in Haliotidae. I have selected a few particularly well- known cases, but those species that are only shown with a single specimen often show a similar range in their coloration (e.g., H varia). H aliotis jacnensis (Figures 4-124-129) exhibits variation in both sculpture as well as coloration. The major sculptural element are the hollow scales, which start at the suture and extend to a variable degree towards the row o f holes (Figure 4-126 vs. Figure 4-129), though a more or less distinct area devoid o f scales is located proximal to the row o f holes. For H. sem iplicata Menke, 1843, three shells are illustrated, vdiich differ both in overall tonality as well as in the color pattern. Differences in color pattern are abundant and are presented for H. glabra (Figures 4-26,4-27,4-30), K ovina (Figures 4-32,4-33), H. diversicolor (Figures 4-40, 4-42), and H. virginea (Figures 4-69,4-70,4-73). H aliotis parva fig u res 4-154-156) shows the usual variation m the mottled specimens with red and green tones, but a num ber o f specimens have a uniform bright orange shell. Such a coloration is now also known from two other South African species, H queketti and H. speciosa (B. Owen, 186 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. pers. comm.). Other sculptural variability is encountered in the European H, tubercu - lata. Mediterranean populations contain both entirely smooth (Figure 4-4) and highly lamellous (Figure 4-5) specimens. The elevation o f the spire can also differ markedly (Figures 4-12, 4-13). Such differences do not warrant formal recognition at any taxo nomic level. Figures 103 and 106 show an intriguing specimen of unknown identity. It is one of three rather similar specimens in the MNHN ranging in size between 52 and 68 mm, According to the lable they were collected in Shang Hai in 1854. The specimens show a unique combination o f characters: strong cords with some scales, and irregularly placed and deep folds. These specimens certainly do not represent any o f the known species in the area: H. discus Reeve, 1846, H. diversicolor, H. gigantea Gmelin, 1791, H. madaka Habe, 1977. The closest affinity is to H. rubra. However, the cording in H. rubra is usually much finer, specimens with folds are unknown to me, and H. rubra is a species endemic to southern Australia. The Shang Hai locality has to be treated with caution, because erroneous locality data abound particularly with 19th century specunens and publications. To mention but some of the most egregious cases, H. unilateralis Lamarck, 1822, has a type locality in Australia, but is restricted to East Afiica; H. squamosa Gray, 1826, also has its type locality in Australia, but is endemic to south Madagascar; and H. zealandica Reeve, 1846, described firom Kew Zealand is actually the Mediterranean H. tuberculata. A pathological condition o f the Shang Hai specimens can be ruled out, because the three specunens are very similar to one another and do not show any known signs o f accidents or disease, such as laterally shifted tow o f holes, holes closed out o f sequence, strong growth mark, abrupt change m sculpture, thickened shell, blister pearls, or downward growth o f aperture. P. Bouchet (pers. comm.) has suggested the possiblil- ity that these specimens may be representative o f a now ectinct species. Marine extmc- 187 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tion is an overlooked topic (of. Tegner et cd^ 1996; Davis e t al.^ 1996) and additional data is required to clarify this case. A particular effort to locate photographs o f living animals has bom fruits as shown in Figures 4-163-192. Many o f the species are illustrated here for the very first time including. Juvenile H. asinina (Figure 4-166), K roberti (Figures 4-167,4-168), K uni - lateralis (Figures 4-169,4-171), H. sp. nov. Owen et a/., in review. (Figure 4-170), H. coccoradiata (Figure 4-186). H. sem iplicata (Figure 4-187), H. hargravesi (Figure 4- 189), K squamata (Figure 4-191), and H. elegans (Figure 4-192). Biogeography and the fossil record The area cladogram as well as the taxon cladogram approach both place the origin o f the abalone in a generalized Indo-Pacific region. How does this find compare to the fossil record? The earliest known abalone fossils stem from the uppermost Cretaceous (Maastrichian). H aliotis lomaensis Andctsan, 1902, was found at Point Loma, Califor nia. H aliotis antillesensis Sohl, 1992, on the other hand was described from Puerto lUco and Jamaica (see Geiger & Groves, 1999, for review). Neither locality represents the Indo-Pacific by any stretch o f the imagination. A look at the putative sistergroups o f Haliotidae will shed some light on this apparent discrepang. Pleurotomariidae and the Trochidae have been proposed as sister groups o f Haliotidae. Haliotidae have also been placed m a basal polytomy o f Vetigastropoda with an apomorphic Pleurotomariidae - Trochidae clade, as a sistergroup to Fissurellidae, or in a polytomy with all Vetigas tropoda except for its sistergroup Pleurotomariidae (Tillier et al,, 1994; Ponder & Lind- berg, 1997; Harasewych e t aA, 1997 for review). All these families have a long standing fossil record reaching back into the Triassic (Knight e ta L , 1960; Tracey e t a l., 1993; Bandel, 1998), and Trochidae may have origmated as early as the Permian or Ordovi- 18 8 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cian. as discussed by Hickman & McLean (1990), althougk Tracey et aL (1993) ques tioned any Permian records. The exact date o f the split between these lineages will remain uncertain for the foreseeable future. Even molecular clock studies (e.g., Felsen- stein, 1985; Springer, 1995) will not be able to add any information due to the extremely variable speed o f the molecular clock (see Mindell & Thacker, 1996; Hillis et aL, 1996, for reviews). Accordingly, Haliotidae had on the order o f200 million years to alter their distribution pattern firom their area o f origin. Hence, the incompleteness of the fossil record accounts for the apparent discrepancy between the analytical approach and the fossil evidence. On top of this major hiatus, a negative taphonomic feedback due to the habitat o f abalone and their shell mineralogy casts further questions as to whether even the two finds firom the Maastrichian accurately represent the distribution o f abalone at that time; see Geiger & Groves (1999) for a more in-depth discussion. DISTRIBUTIONAL DATA AND MAPS M editerranean and West Afiican Species C^ignres 4-4-14,4-146,4-147,4-149,4- 150,4-170,4-172,4-174) Halioti s marmorat a Linnaeus, 1758 (Figures 4-14,4-146,4-149) Azores: Guadalupe, Praia das Conchas (MNHN no #, 2). Guadalupe, Praia Mouro Peixe (MNHN no #, 2). Spain: Canary Isl., Cape Palmas (ANSP 50527,1). MAURITA NIA: Port Etienne (MNHN no #, 1; MNHN no #, 1). Senegal: Cap Vert (LACM 76-43, 3; MNHN no #, 13). Presque lie de Cap Vert, Almadies (MNHN no #, 8). Cap Vert, N’Gor (AMNH 220228,13; MNHN NO #, 2). Dakar (MNHN no #, 1 ; MNHN no #, 2). Dakar, M’Gor (DLG AAB 14g, 2). Dakar, Anse Bernard (MHNG 21132,5). Dakar, 189 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 4-4-13. Shells o f the European-Mediterranean species o f H aliotis spp. Figures 4- 4-9. H aliotis tuberculata Linnaeus, 1758. Figures 4-4,4-7. DLG AAB 14af. Cala St. Maria, Montecristo, Italy. Between rocks and on sand, 3 - 18 m, 28.VI. - 20.VII. 1988. 40 mm. Very flat form. Figures 4-5,4-8. DLG AAB 14bl. Preveza, Greece. Leg. Van- glis Tzimas, 28. IV. 1995.41 mm. Highly lamellose form. Figures 4-6,4-9. DLG 14 am. Erquy, Brittany, France. 93 mm. Atlantic form o f H f. tuberculata. Figures 4-10- II. H aliotis tuberculata coccinea Reeve, 1846. DLG AAB 14 bg. Azores, Portugal. 60 mm. Note more elongated shape and the distinct spiral grooves. Figures 4-12-13. DLG AAB I4b. Naxos, Sicily, Italy. In rock canyons and open rock fields, 3 - 6 m, IX 1985. Appertural view showing variation in spire elevation. 190 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis tuberculata coccinea •Haliotis t tuberculata H aliotis marmorata H aliotis midae H aliotis rugosa Haliotis parva H aliotis mariae Haliotis queketti H squamosa H aliotis spadicea H aliotis pustulata H aliotis unilateralis H aliotis speciosa Figure 4-14. Distributioa o f European-Mediterranean, west African, and south African species o f H aliotis spp. Localities stemming from collection specimens are indicated with a solid circle, those from the literature with a solid square. For data see texL 191 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Almadines (MNHN no #, 30; SBMNH WW2472&2444,6). Point Almadies (SBMNH 6152,2). O ff Dakar, île de Corée (LACM 76-45,3; BMNH no #, 3). Petite Coast, Joal River (LACM 76-47,1). Gambia: Bathurst (BMNH 2341, I). Sierra Leone: Freetown (DMNH 012829, I), ghana: (BMNH no #, 2). Accra (AMNH 103367, I; NHB I705e, 1). O ff Busua, small isl. (AMNH 220229, 2). Liberia: Monrovia (ANSP 193228, 2; BMNH no #, 3). gabon: (AMNH 19706, 5; MNHN no #, 1). Libreville, Cap Esterias (MNHN no #, I). SÂo TomÉ: (MNHN no #, 6, MNHN no #, 2). Sâo Tomé and Prince (AMNH 238663, 5). Praia Emilia (MNHN no #, 2). Spanish Guinea, Santa Isabel (USNM 653538, 3). île de Prince (MNHN no #, 3). île du Prince, Hot Caroço (MNHN no #, 2). EQUATORIAL GUINEA: Fernando Poo (= Bioko), San Carlos Bay (AMNH 150778, 1). Ivory coast: (MNHN no #, 1). LTTERATURE, LOCALITIES: Ghana, Takoradi Harbour (Ubaldi, 1987). LITERATURE, RANGE: Gabon (Bernard, 1984). West Africa (Abbott & Dance, 1983: as rosacea), Haliotis st omat i aef ormi s Reeve, 1846 (Figures 4-14,4-147,4-150) ITALY: Sicily: (MNHN no #, 1). Sicily, Palermo (MNHN no #, 1). Giardini (HUJ 6313b, 3). Lampedusa (HUJ 6313a, 1). Malta: (DLG AAB 51a, 1; LACM A .2777,2; DMNH 097371,1; BMNH 1829,2). Halwtis L t uberculata Linnaeus, 1758 fig u res 4-4-9,4-12-14,4-172,4-174) IRELAND: Rosslare Strand (BMNH 193I.1.12.I, I). Great Britain: Channel Isl., Sark (DMNH OllOlO, 3; USNM 179300,4). Guernsey Isl. (LACM A.1463,4; LACM A J2 a, 2). Guernsey Isl., Lihou (BMNH 1838,4). Jersey (LACM 45894,1; LACM 2777,2). Channel IsL Herm (AMNH 259795, 1; NMBE Schuttleworth, 3). The Mihqueries, off 192 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Channel Isles (ANSP 303052, 2). Henn (USNM 179298, 2; USNM 179291, 8 ). W > France (Atlantic): Dunkerque (MNHN no #, I). Le Havre (LACM 28817, 3). Nor mandie, Die Chaussey (MNHN no #, I). Cherbourg (ANSP 50169,2; MNHN no #, 2). Mont Saint Michel (NMBE 132724, 3). St. Malo (MNHN no #, I; MNHN no #, 1). Brittany, Erquy (DLG AAB 14ag, 1; DLG AAB 14am, I). Baie de St. Brieuc (MNHN no #, 2). Binic, C. du Nord (MNHN no #, 5). Bréhat (MNHN no #, I). Paimpol (MNHN no #, 1). lie d’Er (MNHN no #, 9). Brittany, He de Batz (NHB 1705d, 2). Lannion (MNHN no #, 2). Roscoff (AMNH 91645, 1; MNHN no #, 2). Brittany, Brest (LACM 45899,1; MNHN no #, 1). Baie de Brest, Finistère (MNHN no #, 2). Brittany, Finistère, Camaret (NMBE 1327.540, 1). Trébeurdon, Cap du N (MNHN no #, 17; AMNH 101811, 1). lie de Yeu (MNHN no #, 1). Lorient (AMNH 239006, 3). Concarneau (MNHN no #, 1). lie de Glénans (MNHN no #, 1). Hendaye (ANSP 50498,2). SL Jean- de-Luz (NMBE 1327,6; MNHN no #, 20). Guethary (MNHN no #, 30; MNHN no #, 7). Nantes (USNM 73160, 1). Portugal: Algarve (NMBE 1327.985, 1). Algrave, Sagres (DMNH 059569, 7). Spain: Vigo, Bayona (SBMNH 36331, 2). Ruinas de Bolonia (USNM 739330, 1). Costa Brava, Beach o f Aiguabiava (NMBE 1327.611,2; SBMNH 24997, 1). Barcelona (ANSP 211557, 1). Valencia (AMNH 111953, 1; USNM 781971, 3). Balearic Isl., Menorca, lUa del Aire (DLG AAB 14at, 17). Balearic Isl., Menorca, Fomells (DLG AAB 14au, 50). Balearic IsL, Menorca, S’Amandar (DLG AAB 14av, 24). Balearic Isl., Menorca, Iliade Portos and CalaTorta (DLG AAB 14aw, 59). Balearic Isl., Menorca, Hla d*en Colom (DLG AAB 14ax, 64). Balearic Isl., Menorca, Cala Patenta (DLG AAB 14az, 3). Balearic IsL, hMlorca, Hla Dragonera and Cabrera (DLG AAB 14ay, 11). Balearic Isl., Mallorca, Santa Ponsa(DLG AAB 14d, 1). Balearic Isl., Mallorca, Colonia de Samt Jordi (DMNH 108794,1). Balearic Isl., Ibizza (AMNH Edi son A9853, 1; USNM 424773, 3). Balearic Isl., Ibizza, Calla Bassa, Santaria (DLG 193 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. AAB I4n, 6). Balearic EsI., NE o f Fonnentera (DLG AAB I4bi, 13). Balearic Isl, Fer mentera, 10 km N o f Es Pujols (DLG AAB I4ar, 15). Alicante, Javea (LACM 81-60, 3). Granada, La Herradura (MNHN no #, 3). Malaga (AMNH 111952, 1; USNM 781966,2). Malaga, Benalmadena-Costa (MNHN no #, 4). Mal%a, Estepona (MNHN no #, 1). Malaga, Calahonda (MNHN no #, 15; MHNG 21131, 5). Malaga E*rovince, E Marbella, Torre de Calahonda (LACM 94-20,4). Malaga, Tbrremolmos (MHNG 21165, 1). Cadiz, Tarifa (DMNH 073587,4; DMNH 073583,7). Cadiz, Tarifa, Torre de la Pena (MNHN no #, 3). Cadiz, Getares, Punta Camero (MNHN no #, 1; MNHN no #, 2). Cadiz, Sotogrande, Torre Guadiaro (MNHN no #, 1). Gerona, Soin Felui de Guinels (DMNH 179563, 5). Gerona, Puerto de la Selva (SBMNH 6173, 1). Almeria (MNHN no #, 10). Cadaques (AMNH 259794,3; MNHN no #, 15). CoUohando (ANSP 246246, I). Gibraltar. (DMNH 011033,2; DMNH 129548,1). CeutaN, Benzu(MNHN no #, I; MNHN no #, 3). Ceuta N, Pta del Saudino (MNHN no #, 1). Ceuta S, El Pineo (MNHN no #, 2). Ceuta S, Anse Almadraba (MNHN no #, 7). S-France (Mediterranean): Banyuls sur mer (LACM 45884, 6 ; LACM 80-51, 7 + 3 complete). Pyrenes, Peyrefîte near Banyuls (NMBE 1327.15,3; NMBE 1327.4,12). Port Vendres (ANSP 50514,1; USNM 857643,4). Roussillon Province, Paulilles (LACM 80-40, 1; NMBE 1327.10,3). Cap d’Agde (NMBE 1327.402,4). Côte d’Agde (MNHN no #, 3). Sète (MNHN no #, 3). Frontignan (NMBE 1327.405,1). Marseille ^M N H 1929.10.24.97-101,5; MNHN no #, 3). Ivbrseille, Endoume (MNHN no #, I). Bendor (DLG AAB I4as, 10; DLG AAB I4bd, 7). Provence, lies Embiez (MNHN no #, 1). Sanary (MNHN no #, 1). Toulon (MNHN no #, I, MHNG 13241,2). La Seyne (MNHN no #, 1). Hyères, La tour fondue (DLG AAB 14a, 3). Porquerolles (MNHN no #, 2; MNHN no #, 2). Valras (DLG AAB I4J, 4). Ramatuelle, La. Bastide Blanche ^4MBE 1327.21,1; NMBE 1327.5,26). St. Tropez ^4NHN no #, 2). S t Tropez, Plage de TEscalet (NMBE 1327.13,7; NMBE 194 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1327.6,2). St. Tropez, Salins (TSiM B E 1327.11, 3). Near S t Tropez (ANSP 280643,99; ANSP 280644,104). S t Raphael (MNHN no #, 2). Antibes (MHNG 17573, 10; MHNG 21228, 2). Nice (NMBE Schuttleworth, 4). Villefranche (MHNG 13242, 2; MHNG 13244,3). Mentou (MNHN no #, I). Provence, Cap Ferrat (MNHN no #, I). SE-Corse, Cap la Ciappa (DLG AAB I4ah, 19). Corse (DLG AAB I4h, I). Corse, Cap Corse (MNHN no #, 3). Corse, Potto Polio (BMNH no #, 3: complete). Corse, Cargèse (DLG AAB 14bm, 6). Corse, Viviano (MNHN no #, I). Corse, Ajaccio (MNHN no #, 1; MNHN no #, 4). G ulfof Ajaccis (BMNH 1563,21). Corse, Sanguinaires Isl. (MNHN no #, I). Cannes (AMNH 47284, 3; MNHN no #, I). Roussillon, La Franqui (MNHN no #, 1; MHNG 17574, 28). Roussillon, Collioure (MNHN no #, 4). Italy: Lovrana (MNHN no #, I). Trieste (DMNH 119952,2; SBMNH 28360, 1). Gulf of Trieste, Sis- tiana Bay (AMNH 198685,1). Grignano mare, near Trieste (DLG AAB I4bh, 4). Rim ini, CattoUca (NMBE 1327.25, I). SenigalHa (MHNG 16240). Ancona (LACM 21741, 2; MHNG 17827, I). Apulia, T.S. Gennara (DLG AAB I4w, 3). Apulia, Gargano (DMNH 079612,1). Apulia, Terre Colomela (DLG AAB I4x, 12). Tuscany, N ofPiom- bino, Populonia (DMNH 044178,5). Elba (BMNH 2258, 1). Montecristo, Cala Scoglio (DLG AAB I4ac, I). Montecristo, Cala Corfu (DLG AAB I4ad, I). Montecristo, Cala Maestra (DLG AAB I4ae, I). Montecristo, Cala S t Maria (DLG AAB I4af, 17). Naples (LACM 60-16, I; DMNH 175196, I). Naples, Zoological Station (AMNH 104, 5; NMBE 1327.19, I). Naples, NapIesM (DMNH 130953, I). Naples, PosiHpo (BMNH 1617, 2). W Naples, Golfo di Pozuoli, E Punta di Pennata (LACM 92-107,15). Capri (SBMNH no #, 2). Cape Miseno (SBMNH 240,5). Ischia IsL (SBMNH 2 0 6 ,1). Sicily, Giardini-Naxos (DLG AAB I4b, 31). E Siciliy (LACM 116523, 2). Sicily, Palermo (BMNH no #, 2; MNHN no #, I). Sicily, Catania (AMNH 240643,1). Sicily, Aci Trezza (AMNH 220199,4). Sicily, Messina (MHNG 13190,2). Sicfly SE, T. Vendicari (NMBE 19 5 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1327.16, 7; MHNG 13194, 6). Sicily, MondeUo (NMBE 1327.17,2; USNM 179289, 1). Sicily, Meiilli Marina (NMBE 1327.18, 1). Sicily, Bay Carini (USNM 710646, 1). Campania, Palinuro (AMNH 230255,2). Taranto, San Vito (AMNH 249112,5). Taranto, Marino di Pulsano (AMNH 249111, 3). Porto Cesare (AMNH 239005, 6). Tyrrhenian Sea, Panarea Isl. (AMNH 215140, 1). Portofino, Genova (ANSP 290332, 2). W Brin disi (NMBE 1327.7, 1). GaUipoU (USNM 42098,4). Sardinia, Porto Ferro (SBMNH WW209,4). Lampedusa (MHNG 13198,2). Malta: (LACM H-2687,1; LACM A2777, 2). Delimara Point (BMNH no #, 1). Yugoslavia: G ulf o f Piran (USNM 754029, 1: complete; USNM 754013,3). (stria, Umag (DLG AAB 14e, 1). Porec (DLG AAB 14m, 2). Rovinj (MHNG 13200, 10). Istria, S coast Katarina Isl. off Rovinj (ANSP 332522, 2). Adriatic, Bay of Bakar, Gulf o f Kvaraer (MHNG 13189,1). Krk Isl. (USNM 753776, 1). Novi (USNM 753821, 1). Privic Isl. (USNM 753853, 1). VruIJa Cove (USNM 753751, 1: complete). Cres IsL, Osor (DLG AAB 14t, 1). Losinj (DLG AAB I4aj, 1). Mali Losinj, Baldarka (DLG AAB 14u, 2). Dalmatia, Zara (= Zadar) (USNM 179288, 1). Zadar, Zaton (DLG AAB 14v, 3). Makarska35 miles S o f Zadar (LACM A.E.62-45, 8). Pag (DLG AAB 14y, 2). 5 miles N o f Prunosten, near Split (BMNH 2341, 6). Split (DMNH 029675, 4). Dalmatia, Korcula Isl., Osjak Isl. (LACM A.7759, 2; LACM A.7759, I). Promontaiy near Usarica, S of Kariobag (BMNH 2341, 6). Makarska (AMNH 108104,20). Opatija Beach (ANSP 318018,1). Dubrovnik (MHNG 21230,3: MHNG 13201, 6). Dubrovnik, Lorkum (MHNG 21129,1). Greece: Corfu (MNHN no #, 2; SBMNH WW2061,1). Eubôa, Limni (DLG AAB 14aa, 1). Crete, Stoa (DLG AAB 14ba, 3). Crete, Sonda Bay, Marthi (NMBE 1327.8, 1). W coast, Preveza (DLG AAB I4bl, 55). Near Athens (BMNH no #, 3). Kefallina IsL (DMNH 095476,1; DMNH 095504, 1). Saronikos Gulf (USNM 835593,1; SBMNH 6174,1). Poros Isl. (DMNH 095482,5). Spetsai Isl. (SBMNH 12249,2). Lenmos (SBMNH WW2040,2; MHNG 196 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 21164, I). Lesvos [= Lesbos ?I (AMNH 151988, 1). Samos IsL (AMNH 151989,1; AMNH 45733,2). Rhodos (USNM 669298, 1), Kos IsL, Bay o f Kalimnos (DLG AAB 14bo, 2). TURKEY: Bodrum (AMNH 239004, 5). Gulf o f Atalya (BMNH 2258,2). Foca (Med. Inlet.), Aegean Sea (USNM 709157, 1). CYPRUS: Ayia Napa (AMNH 275997,1). SW, Paphos (NMBE 1327.1905,5; NMBE 1327.1906,1). N Akamas Penin sula, Bath o f Aphrodite (MNHN no #, 1). SYRIA: Tartous (MNHN no #, 3). LEBANON: Beyrout (MNHN no #, 3; USNM 669101,1). Israel: Haifa, Shymona (DLG AAB 14f, 1). EGYPT: Alexandria (USNM 74545, 3). TUNESIA: Tabarka (USNM 740191, 1). 3 km W o f Bechateur (USNM 740264, I; USNM 740167, I). Bizerte (DMNH 127665,1). G ulfof Gabes, Bou Grara (MNHN no #, 3). Tunis, Gulf o f Gabes, Gerba/Jerba Isl. (BMNH 2258,1; BMNH 2258,2). lie Djerba, Sidi-^/brès near Houmt- Souk Pyrenes, Peyrefîte (NMBE 1327.799, 6). Algeria: (LACM 45951, 8). Alger (MNHN no #, 3; MNHN no #, 1). Oran (MNHN no #, 1; MNHN no #, 4). MOROCCO: Meinia (LACM 45795, 3; DMNH 011012,3). MeliUa, Cala Charranes (ANSP 215755, 2). MeliUa, Lengua de Tierra (ANSP 209639,3). MeliUa, MueUe Norte (ANSP 209640, 1). Tangier (MNHN no #, 4; MNHN no #, 2). Tangier Province, Mdiq (LACM 94-23, 1; MNHN no #, 7). Casablanca (MNHN no #, 1). Tetouan (MNHN no #, 1). Asilah (MNHN no #, I). Mogador (MNHN no #, 2; MNHN no #, 1). Agadir (MNHN no #, 2; MHNG 21810,1). 9 km N o f AgadË (NMBE 1327.1,1). Senegal: C ^ Vert, Ponte des Almadies (LACM 76-48,1 with dried bo<fy). Dakar (USNM 781776,1). LITERATURE, LOCALITIES: Messina, Taormina (Bolognari, 1953). Spain, Vigo; Italy, Beach o f Catanzaro; Tunesia, Susa (= Sousse) (Ubaldi, 1987). Akhziv; Bat Gal- lim; ‘Atlit; Dor; Netanya; "^Akko; Haifa (Barash & Danin, 1992). Punta Barcarello (Palermo); Ognina (Shacusa); Isola di Favignana; Taranto; Portofîno (Genova); G ulfof N icies; Cala Junco (Panarea Isl.); Orano (Algeria), Jerba (Tunesia) (Gianuzzi-Savelli et 197 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ed,, 1994). Capo Mbrtoia, Ventmiiglia; Bordighera; Varazze; Bergeggi ; Punta Manaia, Riva Trigoso; Baia del Silenzio, Sestri Levante; Arenzano; Vesima; Bonassola; Elba, Marina di Campo; Elba, Scoglio della Triglia; Piombino, Secca del Cimitero; Baratti; Golfo di Baratti, Piombino; S. Vincenzo; Calafuria; Tarquinia; Isola Palmarola, Arcipelago Pontino; Sorrento; Capo di Sorrento; Seiano; Isola di Vivara; Procida; Punta Pennata; Punta Pioppeto; Miseno; Posillipo; Porto Infieschi; Marina di Ascea; Pioppi; Punta Licosa; Agropoli; Salerno; Marina di Maratea; Capo Vaticano; Santa Domenica di Ricadi; S. Irene, Briatico; Punta Prosciutto; Lido Manduria; Taranto; Torre Pali, S. Maria di Leuca; Marina di Novaglie; Mattinata; Rodi Garganico; Marina di San Vito; Lido Riccio, Ortona Foro; Portonovo; near lake Chioggia; Costa Veneziana, Cavallino; Aurisina Filtri; Isola delle Femmine; Capo Gallo; Milazzo; Isole Lipari; Lipari, Can- neto; Aci Trezza; Porto Palo; Vendicari; Siracusa; Augusta, Castelluccio near Brucoli; Pantelleria; Isola Favignana; Favignana; Isola di Marettimo; San Antioco, Spiaggia Canisoni; Torre Cala Regina; Carloforte; Cagliari; Cagliari, Foxi; Villasimius, Isola Cavoli; Villasimius, Secca S. Caterina; Villasimius, Capo Boi; Villasimius, Punta Molentis; Villasimius, Isola Serpentaria; Portoscuso; San Caterina di Pittinuri, is Are nas; San Caterina Pittinuri, S’ Archittu; Torre Argentina; Capo Marargiu; Golfo di Arzachena; Isuledda; Cannigione Isuledda; Capo Caccia, Cala della Calcina; Alghero, Scogiiera Lazzaietto; Argentiera; Isola dei Gabbiani, Palau; Isola della Maddalena; Isola Santa Maria, Bocche di Boni&cio; Isola Razzali, Poloni ^edu lli et ai., 1982). LTTERATURE, RANGE: Channel Isl. - Senegal, Mediterranean (Lindner, 1975). SW Europe to Channel M ., Mediterranean (Poppe & Goto, 1991: as tuberculata form lam el - losay Mediterranean coast o f Israel, Egypt, Leganon, Syria, Cyprus, Aegean Sea, Adri atic Sea, Western Basin, Alboran Sea; Atlantik: Spain, Portugal Morocco, Canary Isl. (Barash & Danin, 1992). 198 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. HaUotis tuberculata coccinea Reeve, 1846 (Figures 4-10,4-11,4-14) Morocco: Cala Charranes, Melilla (ANSP 215756, 2). Spain, Malaga, S o f Torre Molino, Toroque Brada (ANSP 265699, 10), Canary IsL: Lanzarote (LACM 45966,2; LACM 45915, 3). Lanzarote, Arrecife (AMNH 151987, I; AMNH 170644, 1). Lan zarote, Orzola (MNHN no #, 5). Lanzarote, Las Coloradas (MNHN no #, 1). Lanzarote, Porto del Carmen (MHNG 21933, I). Gran Canaria, Las Palmas (ANSP 209637, 1), Gran Canaria, Las Palmas, Las Canteras Beach (LACM 62831,2). Gran Canaria, Puerto Rico (DLG AAB 14ao, 7). Gran Canaria, Sardina (DMNH 18200,4). Tenerife (DMNH 155574,2). Tenerife, Los Christianos (DMNH 179567,3). Tenerifa, Czuzruy IsL, Bajo Piedros (DLG AAB I4z, 15). Tenerife, Santa Cruz (AMNH 220245, 3; MNHN no #, 8). Tenerifa, Puerto de la Cruz (ANSP 202695,2; MNBE 1508.651, II). Tenerife, Punta Hilgado (MNHN no #, 4). Tenerife, La Tejita (MNHN no #, I; MNHN no #, 1). Tener ife, Las Galletas (MNHN no #, 2). Tenerife, San Andres (MNHN no #, I). Tenerife, Palm Nfer (MNHN no #, I). Tenerife de Gran Canarias (USNM 636758, 3: complete). La Palma, Santa Cruz de la Palma (MNHN no #, I). Fuerteventura (DLG AAB 14c, 3). MADEIRA: Santa Cruz (ANSP 88744, 9; DMNH 133467, 4). S-Madeira, Garajau (DLG AAB Waq, 2). Salvane IsL, Petite Salvage, S Side (BMNH 2324, 5). cape verde isL: (AMNH 19715, 1). Santa Luzia IsL, Curral (AMNH 241649, 2). AZORES: Payai (AMNH 19680, 3; USNM 31708, 7). Faial, Horta (MNHN no #, 1). Fayal, Castelo Branco (MNHN no #, 3). Graciosa IsL (BMNH 2341,12). Graciosa IsL, Hheu da Praia (MNHN no #, 1). San Miguel, Ponta Galera (MNHN no #, 5; MNHN no #, 1). San Miguel, nheu de Vila Franca do Campo (MNHN no #, 1; MNHN no #, 1). San Miguel, N (MNHN no #, 1). San W guel, S, Roia da Praia ^4NHN no #, 18). Santa Maria, NW, Anjos (MNHN no #, 3). Santa Marm, SE, Maia (MNHN no #, 1). Santa Maria, Praia Formosa (MNHN no #, 2). Santa Maria, Ponta do Marvao (MNHN no #, 1). Santa 199 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Maria, Vfla de Porto (MNHN no #, l). Flores (BMNH no #, 55). Flores, Ponta Delgada (MNHN no #, 2). LmERATURE, LOCALITIES: Cartagena, Spain. (Gianuzzi-Savelli et aL, 1994). LITERATURE, RANGE: Canary Isl. (Poppe & Goto, 1991). South African Species (Figures 4-14,4-142-145,4-148,4-151-156,4-159-162) ffaliotis mî dae Linnaeus, 1758 (Figures 4-14,4-142-145,4-148) ANGOLA: Lobito Bay (AMNH 45976, 3; AMNH 46340, I : doubflil, not mapped). South Africa: Saldanha Bay (AMNH 107426, l). False Bay (AMNH 186964, 11; AMNH 272141, 2). False Bay, Simons Town (AMNH 87538,3; AMNH 186831, 5). False Bay, Muizenberg (USNM 617743,4). Cape Town (DMNH 101956, 5; AMNH 45954.2). Strand (LACM S.2047, 1; LACM 22228, 1). Cape Peninsula, Kommetjie Beach (AMNH 186985, 1). Cape Province, Table Bay (USNM 636759, 7). Kleinmond (LACM A.8487.65,3; BMNH no #, 3). Cape Province, Onrust (LACM 10736,1; ANSP 211516,4). Hermanns (AMNH 205925,1; NMBE 1330.1169, 1). Hermanns, Abalone factory (BMNH 2341, 1). Hermanns, Gansbaai (BMNH 2341, 1). Cape Province, Bnf- fels Bay (AMNH 240634,1; AMNH Edison A9853, 15). Cape o f Good Hope (ANSP 50215,1; ANSP nncataloged, 1). Jefi&eys Bay (DMNH 049718,4; LACM 91284,14). Algoa Bay (DMNH 063297, 1; LACM 11667, 1). Aigoa Bay, Cape Recife (LACM 11709,1; LACM 45913,2). Gouritz River Month (ANSP 211525, 1). Gonritz River Month, outer ledges (ANSP 217651, 1). Mossel Bay (LACM A.2777a, 1; AMNH 101550.2). Knysna (MHNG 21234,1). Port Elisabeth (BMNH 2341,1; MNHN no #, 22). Port Elisabeth, Snmmerstrand (ANSP 276815,27). Port Alfred (DMNH 012307,6; DMNH 011028,1). Port Alfred, Cape Colony (USNM 249895,1). Port Alfred, Shelly 200 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Beach. (AMNH 186962, 2). Port Alfîed, near Grahamstown (USNM 187118,1; USNM 186886, 3). East London (NMBE 1330.8, 1). East London, Bonza Bay (LACM 28821, 1). London, Shelly Beach (NMBE ex 1510.343, 1). E Cape, Gonubie (AMNH 232367, 2; SBMNH 6186,5). Ciskei, Haga Haga (AMNH 1869663,3; AMNH 272140,2). Mor gans Bay, E coast (AMNH ex 200604, 2). Kei Mouth (DMNH 012210, 3). Port Shep- stone (LACM A.2777b, 1). Durban (NMBE 1330294,1). Capeland, Bushman's River Mouth [= Tugela, at Newark] (USNM 622256, 5). LITERATURE, LOCALITIES: Stony Point; Dassen IsL; Dawidskraal (Newman, 1968). Coffee Bay (Jacks, 1983). Saldanha Bay, Marcus IsL; Mas Bay near Cape Hangk- lip (Barkai & GrifBths, 1986). LITERATURE, RANGE: Table Bay - Port Alfted (Barnard, 1963). Saldanha Bay - Gonubie (Richards, 1987). Table Bay - Natal (Abbott & Dance, 1983). Saldanha Bay - Gonubie (Jacks, 1983). SL Helena Bay - W Transkei (Muller, 1984b). Haliotis parva Lmnaeus, 1758 (Figures 4-14,4-154-156) Namibia: Hottentot, Kleinward Strand (DMNH 012309,7). South Africa: Cape Town (AMNH 260669,1; AMNH 207957,1). Cape Town, Seapomt (DLG AAB 18a, 1). Cape o f Good Hope (DMNH 130284, 1; DMNH 133560, 1 > . False Bay (DMNH 010371, 1; AMNH 276889, 1). False Bay, Muizenberg (USNM 617742,4). Strand (LACM 45786, 5; LACM 45921,2). Shnonstown (AMNH 87540,3). Simon's Bay (ANSP 211515,1). Cape Province, Betty's Point beaches (AMNH Edison A9853, 8). Buffels Bay and Betty's Point (AMNH Edison A9853,12). Lialguhas [= Agulhas ?] (DMNH 015438,3; AMNH 272143,4). Cape L'AIguhas (ANSP 211522,2). Mosselbaai (DMNH 012311, 8). Jef&eys Bay (DMNH 124169,3; LACM 45792,2). Knynsa Cape (SBMNH 38831, 1). Algoa Bay g)MNH 131504,3; BMNH no #, 1). Port Elisabeth (DLG AAB 18f, 1). 201 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Port Alfred (LACM 45856, I). Port Alfred, near Grahamstown. (USNM 187116, 2; USNM 187117,1). BCleinmond (LACM A.8487.65,3). LTTERATURE, LOCALITIES: Table Bay; Still Bay (Barnard, 1963). LITERATURE, RANGE: False Bay - East London (Barnard, 1963). False Bay - Gonu bie (Richards, 1987). Table Bay; False Bay; Still Bay - Port Elisabeth - Port Alfred - Gonubie (Jacks, 1983). HaUo t i s queket U Smith, 1910 (F io res 4-14,4-161,4-162) South Africa: Transkei, Coffee Bay (DLG AAB 52a, 1; AMNH 145831, 1). Transkei, Xora (AMNH 232440, 1). Pondoland (NMW, 1; DMNH 12312, 4). East London (SBMNH 6654,1). Natal, Palm Beach (AMNH 272144,1). LITERATURE, LOCALITIES: Inhaca Isl. (Macnae & Kalk, 1958). Natal, Isezela; Port Alfred; Natal, Zuluiand, off O’ Neill Peak; Mozambique, Delagoa Bay (= Bahia de Lourenco Marques) (Barnard, 1963). Northeastern Somalia (Geiger & Pisor, 1999: not mapped). LITERATURE, RANGE: Port Alfred - Natal (Richards, 1987). Natal (Abbott & Dance, 1983). Transkei - S Natal (Mtiller, 1984b). Transkei - KwaZulu (Muller, 1986). Hal i ot U spadi cea Donovan, 1808 (Figures 4-14,4-151-153) South Africa: Robben Isl. (MNHN no #, 2). Table Bay (DMNH 136950,2). Cape Town (LACM 45798,1; ANSP 62750,1). W C ^ Town, near Dehoop Nature Reserve, o ff Witsands (AMNH 220030,2). False Bay (LACM 105825,2; AMNH 141073,2). False Bay, Saint James (LACM A.2777a,- 3). False Bay, Muizenberg (USNM, 603229, 26; USNM 67738,3). Strand (LACM 22197,1; LACM 45852,3). Kleinmond (DMNH 012314,1). Mffler Point (AMNH 186881,3). Shnonstown (AMNH 141083,1; AMNH 202 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 87539» I). Simon's Bay, E Cape Peninsula (ANSP 2II5I3, 3; BMNH 59.11J0.13,1). Mossel Beach (LACM 34261, 2; LACM A J777, 2). Cape o f Good Hope (LACM 59339,2; LACM 65-112, 1). Cape UAIguIhas (ANSP 211517,2; BMNH 1926.10.4.1- 3,3). Plettenberg Bay (BMNH no #, 1; ANSP 189719,1). Cape Province, Storm River Mouth (BMNH 2341,6). Jef&eys Bay (LACM A.8589,2; DMNH 111610,2). Port Elis abeth (LACM A.5678, 1). Knysa, Port Elisabeth (MNHN no #, >50). Algoa Bay (DMNH 049714,3; ANSP uncataloged, 2). Port Alfted (LACM 45905,1; LACM 45914, 2). Port Alfied, near Grahamstown (USNM 186885, 1). East London (LACM 10728,1; AMNH 96645, 8). East London, Glen Eden (LACM 10706,2). East Cape, Cmtza Reef (AMNH 232366, 4). Gonubie (AMNH 221705, 1). Morgans Bay, E Coast (AMNH 200604,4). Haga Haga (AMNH 182066, 1; AMNH 272153, 2). Kei Mouth (DMNH 101372,2). Transkei, Coffee Bay (AMNH 272512, 1). Transkei, Coffee Bay, 1 mile S Umtata River (ANSP 211523, 1; ANSP 211520,1). Umgazi River (ANSP 211519, 1). Port Shepstone (LACM A J777b, 1). Bushmans River (= Tugela, at Newark) (DMNH 131510,3; USNM 622255,6). LITERATURE, RANGE: Table Bay - Port Alfred, Durban; Tongaat, Natal (Barnard, 1963). False Bay - Natal (Richards, 1987). Table Bay - Natal (Abbott & Dance, 1983). Table Bay; False Bay - Durban and Tongaat, Natal (Jacks, 1983). Cape Peninsula, Par tridge Point - N Natal, Tongaat (Muller, 1986). Haliotis speci osa Reeve, 1846 (Figures 4-14,4-159,4-160) Senegal: Goree (ANSP 50187, 2: doubtful, not mapped), south africa: Table Bay (DMNH 009591,4). East London, Gonubie (LACM 29624, 1; AMNH Edison A9853, 1). Jef&eys Bay (AMNH 220128,1). Port Alfred. (ANSP 193234,1). Algoa Bay (NMW, 2). Pondoland (NMW, 1; DMNH 12316,1). 203 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE, LOCALITIES: East London; Kowie (= Port Alfied) (Jacks» 1983). LITERATURE, RANGE: Port Alfied - Natal (Talmadge, 1958b). Pondoland - Kowie - Port Alfied; Natal (Barnard, 1963). Gonubie - Natal (Richards, 1987). Port Alfied - Natal (Jacks, 1983). Port Alfied - W Transkei (Muller, 1984b). East African Species (Figures 4-14-23,4-130,4-133,4-IS7,4-158,4-169,4-171) Halioti s mari ae Gray, 1826 (Figures 4-14-16) Comoros: (BMNH no #, 2: doubtful). tanania: Zanzibar (BMNH 215,b, 1: doubtful). Oman: (DLG AAB 25d, 2 complete + 2 bodies). Muscat (LACM 45821, 3; AMNH 127217, 5). Qurm near Muscat (BMNH Smythe collection, 2). Dhofar province, near Sudh [= Sadh ?], Wadi Haart (LACM 111475,3; BMNH no #, 7). MasiralsL (BMNH 2258,4; BMNH Smythe collection, 4). Taqah (BMNH Smythe collection, 1). Near Ras el Hadd (BMNH no #, 2). Near Salalah, Rysut Power Station (BMNH 2375,1). Saudi Arabia: Jedda (DMNH 120252, 9: doubt ful). Aden (AMNH 104844,6: doubtful). LITERATURE, RANGE: Throughout Oman (Bosch & Bosch, 1981). HaUot i s pustulaùt Reeve, 1846 (Flignres 4-14,4-17-20) Jordan: Aquaba (BMNH 2258,2). Israel: Elat (AMNH 125441,2; AMNH 92656,2). Elat, Angelica Beacb (DLG AAB 09e, 1). Elat, Coral Beach (DLG AAB 09d, 1). Gulf o f Fara^um [= Faraun ?], Gulf o f Elat (USNM 769972,1: complete; USNM 769990,2: complete). Gulf o f Elat (AMNH 221666,2; AMNH 220214,2). E ^ p t: Smai, Dahab, Oasis (DLG 09a, 3; SMF 311910/4,3). Sinai, Dahab (SMF 311911/3,3; SMF 311909/1, I). Sinai, Dahab, Mangrove o f Nabq (SMF 311908/1,1). Sinai, DevSshead (DLG AAB 204 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ■ B Figures 4-15-23. Shells of East African abalone. Figures 4-15-16. H aliotis mariae Gray, 1826. LACM 45821. Muscat, Oman. Leg. D. T. Bosch Feb. 1963, ex. coll. Burch. 101 mm. Figures 4-17-20 H aliotis pustulata Reeve, 1846. Figure 4-17. DLG AAB 09r. Elat, Israel. 35 mm. Red variation. Figures 4-18-19. DLG AAB 09w. Pemba, N. Mozam bique. 41 mm. Dark specimen with deep spfral ridges sim ilar to those in H rugosa. Figure 4-20. DLG AAB 09m. Elat, Israel. 30 mm. dorsal. Figures 4-21-22. H aliotis rugosa Lamarck, 1822. DLG AAB 32a. SL Denis, Reunion Island. 40 mm. Figure 4-23. H aliotis unilateralis Lamarck, 1822. RP. Na’Ama Bay, E ^p L 21 mm. Specimen with more arched dorsal surface and ledge less pronounced. Compare to Figures 4-157,4- 158 for other specimens. 205 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 09c, 3). Sinai, Na^ama Nasra aal Et (DMNH 073898,4). Sinai, Na’ama, Masra al Et (AMNH 174080, 5). Sinai, Shann el Sheik, Ras um Sid (DLG AAB 09p, 4). Sinai, Shann el-Sheüc, Ras Mohammed, profile 3 (SMF 311907/1, 1: complete). Sinai, N o f Nuweiba (MHNG uncataloged, 4). S Tiran (DLG AAB 09q, 1). Ras Benas (USNM 604098, I). NW Gulf o f Suez, Wadi el Dom (LACM 67-86, 6). 21 km N o f Quseir (LACM 67-88, 4). Al-Ghardaqa (=Hurghada) (LACM 8980-68, I: complete). Hurghagda, Biological Station (EN no #, 1). Umm el (= al) Kiman (AMNH 179941,1). Saudi Arabia: Near Yanbu (BMNH 2363,1). Jedda (DMNH 121107,5; DMNH 051062, 4). N of Jedda (AMNH 151133; AMNH 148961,1): Jedda, 14 km N of (DMNH 120252, 9). Abu Latt Isl. (MNHN no #, 1). Aden (BMNH no #, I). Sudan: Port Sudan (AMNH 220221,1; DMMF lIE-8218,3 :2 complete). Port Sudan Harbor (BMNH 2341,1). S o f Port Sudan (UR 272, 1). 16 km S o f Port Sudan, N Towartit complex, Harvey Reef (BMNH 2341, 3). Sanganeb Atoll, S anchorage (SMF 311906/2,2: complete). San- ganeb Atoll, N anchorage (SMF 311905/3,2: complete). 20 km off Muhammed Qol, Shaab Baraia (BMNH 2341,4; BMNH 2341,4). DJIBOUTI: Djibouti (MNHN no #, 3). YEMEN: Socotra Isl., Hadibo (UR no #, 3). oman: Humer (BMNH Smythe collec tion, 1). Qurm near Muscat (BMNH Smythe collection, 1). Muscat (AMNH 127234,1). KenYa: Malindi, Gedi, Watamu Beach (AMNH 220209,1). Malindi (USNM 595039, 1). Shimoni (DLG AAB 09j, 4). Shimoni, Wasin Village (BMNH 2241,1). Mtwapa (NMBE ex 1507.955, I). Mombassa (BMNH no #, I; BMNH no #, 2). Faubourg de Mombasa ^dNHN no #, 3). Near Mombassa (AMNH 220222,2; AMNH Edison A9853, 2). Tanzania: Zanzibar, Kiwenga (DMNH 045981,4; ANSP 212431, 69). Zanzibar, Je, Boamo, 5 miles S o f Pîge (ANSP 213154,16). Zanzibar; Ras Nungwe (ANSP 212799, 23). Zanzibar, Mangapwani (ANSP 212982,1). Zanzibar, Kizmiakazi (ANSP 214118, 4). Zanzibar, Fumba (DMNH 045982, 1). Zanzibar, BCiwengwa (USNM 597238,1). 206 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Zanzibar, Uzoi (AMNH 220218,5). Zanzibar, Chango (BMNH 1811,1). S o f Mafia M. (LACM 45869,2). Dar es Salam (LACM 45848,1; DMNH 110695,10). Dar es Salam, Sinda IsL (USNM 704049, 2). Dar es Salam, Ras Kankadya (USNM 704020, 2). Mozambique: Pemba (DLG AAB 09w, 5). Puerto Amelia (LACM 45853, 3; LACM 111045,4). Inhaca IsL, Maputo (DLG AAB 09v, 10). Bazaïuta Isl. (DMNH 011048,2). Nacala Bay, Femâo Veloso (NMBE 1505.107, 1; NHBE ex 1507.89, 4). Mozambique Port (AMNH 80450, 3). Mozambique City (AMNH 114807, 6; ANSP 234109, 1). sey chelles: (BMNH no #, 1). Comoros: Grande Comore, Mitsamiouli (LACM 45935,4). Mayotte (BMNH no #, 1). Madagascar: Nosy Be (AMNH 220220,2). Nosy Bé, Andi- lana (DMNH 045923,2). Nosy Be, Ambka (AMNH Edison A9853,2). W Nossi Be, N shore o f Nosy Saktia (ANSP 257888,5). W Nossi Be, Nosy N’Tangam (ANSP 258698, 14). NW Nosy Be, at Andilanda (LACM 89-53, 6: 3 with dried bodies). 13" 3 1 'S 48* 32’ E (USNM 719376, 1). Tulear (LACM 144277,2; MNHN no #, 1). Tulear, Infati (SBMNH no #, 2; DLG AAB 09u, 2). N Tulear, Mora Mora (LACM 144355,1; LACM 89-50, 1: complete). Anakao (DLG AAB 09s, 3). Between Fort Dauphin and Monan- tenîna (DLG AAB 09y, 13). Androka (MNHN no #, 1). Reunion: (LACM A.2777,3; BMNH 1829,2). Cap Homard (DMNH 179566,4). 2 km S o f St. Leu (LACM 89-60, 2: complete). St. Giller (DLG AAB 09k, 2). Mauritius: (LACM S.1051, 2; LACM 22227.1). 0 2 5 miles N o f Black River Bay (DMNH 045913,7). 1 miles W o f Black River (ANSP 273941,1). S side Tombeau Bay (ANSP 272160,1). Port Louis (USNM 149075.1). LITERATURE, LOCALITIES: Madagascar, Ampalaza, Nosy Manitsa; Foulpointe; Tamatave ^autzenberg, 1929). S o f Aqaba(Mergner, 1979). Akhziv, Shiqmona (Tal- m a d g g, 1971; Fainzilber, 1984; Barash & Danm, 1992). RSA, N Kwa Zulu, Kosi Bay f u lle r ; 1984b). Lybia, Tobruk (CKanuzzi-Saveili ef 1994). 207 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LTTERATURE, RANGE: Mozambique; Madagascar; Zanzibar; Kenya; Red Sea; Cey lon - S India; Mauritius (Talmadge, 1956). Persian Gulf; E Africa; Somaliland; Red Sea (Talmadge, 1974). Red Sea(Sharabati, 1984: as varia). G ulf o f Suez, NW Indian Ocean (Abbott & Dance, 1983: as ancile). Gulf o f Suez and Elat; Red Sea; Madagascar; Sey chelles; Mauritius (Barash & Danin, 1992). Haliotis rugosa Lamarck, 1822 (Figures 4-14,4-21,4-22) Mozambique: (Dig aab 32b, 1). Reunion: 2 km S of S t Leu (LACM 89-60, 3, 1 with dried body). N o f S t GiUes (LACM 89-59, I; LACM 89-58, 1). St Denis (DLG AAB 32a, I). Saint Paul (ANSP 301680,4). S t Pierre (MNHN no #, I). Mauritius: (AMNH 220217, 1; ANSP 50202, I). Mauritius, S side Tombeau Bay, W (ANSP 399932, 1). South Africa: Jeffr^s Bay (AMNH 220224,1: not mapped). LTTERATURE, RANGE: Maurice, Reunion (Drivas & Jay, 1988: as pustulata). Haliotis s quamos a Gray, 1826 (Figures 4-14,4-130,4-133) Madagascar: Ambovomby (AMNH 226476, 1). Ambovomby, South M’car (DLG AAB 33b, I). Fort Dauphin (SBMNH no #, 2). Between Fort Dauphm and Monanten- ina(DLGAAB 33c, 1). LTTERATURE, LOCALITIES: Madagascar, Ikouko near Ambovombé; Faux-Cap ^Jautzenberg, 1932). H aliotis mut at eraUs L am arck, 1822 (F% ures 4-14,4-23,4-157,4-158,4-169,4-171) Israel: Elat (Neotype MHNG 18020,. 1; DLG 48a, I). Elat, Coral Beach (ANSP 350490,2). Off Elat (KAS no #: 6). EGYPT: Zafriana (MNHN no #, I). 30 km South o f Elat (SBS, 2). Sinai, Lagoon o f Dahab (SBS, 1). Smai, Dahab (SBS, 1). Sinai, Na’ama 208 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Bay (SBS, 1). Tîran, Jackson Reef (SBS, 1). Safega (RP, 1). SAUDI ARABIA: Abu Latt Isl. (MNHN no #, 2). Bab-el-Mandeb, ME5 stat. 236Ku (SMF 311920/1, 1). Djibuiti: (MNHN no #, 5; MNHN no #, 6). Ille des 7 frères (Stewart: 1; BMNH, 1). Ouaramous Isl. (KAS, I). Gulf o f Tadjoura, Moucha Isl. (AMNH 164632,1 [?]). sudan: 20 km off Muhammed Qol, Shaab Baraia (BMNH 2341,1). Port Sudan, Wingate Reef, S. Point (SMF 311905/1, I: complete). Sanganeb Atoll (SMF 311915/1, I). Sanganeb Atoll, N side lagoon (SMF 311904/1, I). Sanganeb Atoll, N anchorage (SMF ex 311905/3, 2: complete). UNITED ARAB EMIRATES: 500-800 nautical miles N o f Lulayyah Port, 25“ 24’ N 56“ 22’ E (SMF 311903/1,1 : complete). G ulf o f Oman, Shaqa, Hkor Fakkan, Martini Rocks, stat. 22,25“ 20’ 73” N 56“ 22’ 16” E (SMF 311922/4,4). kenya: Shimoni, Ras Rashid (BMNH 2241, 1). Aldabra: lie Picard, Passé Dubois (USNM 836532, I). Mozambique: Channel: Bassas Da India (KAS no #, I). Nacala Bay, Femâo Veloso (NMBE ex 1505.107, 26). Porto Amelia (AMNH ex 220215, 1). Zanzibar: (DMNH 001358). Pinna (CASIZ 1594, 1). Uzoi (AMNH 220218, 3). Mada gascar: Tulear (ND4HN no #, 1; MNHN no #, 1). Mauritius (CASIZ 1532, 3; BMNH 2305, 2; BMNH no #, 3). Pointe Fayette (ANSP 273696, 1). NW side o f Tamarin Bay (ANSP 273545, 6). Vcoas Point, 3 miles S Mahebourg (ANSP 274167, 1). Matson Point, Grand Bay (ANSP 273031,1). SE side Tamarin Bay (ANSP 273478,17). LITERATURE, RANGE: Red Sea (Sharabati, 1984: as cf. pustulata). Maurice, Reunion ODrivas & Jay, 1988: as varia). Ihdo-Pacifîc and C entral PachSc Species fig u res 4-24-44,4-63,4-66,4-116,4-117, 4-119,4-120,4-122-129,.4-136-141,4-163-166,4-175,4-176) 209 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Halîotâ asùtma Lnmaens, 1758 (F lo res 4-35,4-36,4-43,4-165,4-166) Andamanes: (DLG AAB Olh, 5; BMNH). Port Blair (BMNH 1838, l). Thailand: Koh Phuket, Ac Rawaî (USNM 661782, I). Phuket, E Kaew Yai Isl. (LACM 85-5, l: com plete). Phuket Isl., Airport Beach (ANSP 286645,1). Phuket Isl., Laam Ka, Rawai Beach (ANSP 286053, 1). Butang Group, Pulau Tanga (USNM 661644, 3). Goh Sindarar Nua (Chance Isl.) (USNM 661341, I). Singapore: (ANSP 50241,2; MNHN no #, 2). Viet nam: (AMNH 110486, l).N haTrang Harbor, A hnTrelsi. (AMNH 156199,1; SBMNH 035I I , 2). Nha Trang (MNHN no #, 2). Paracels (MNHN no #, I), china: (AMNH 19723, 18; AMNH 19724, 3). Japan: Honshu, Wakayama, Shirahama, Kii (AMNH 220141, I; LACM 45971, 2). Kyushu, Nagasaki, Enoshima (BMNH 1838, I). Oki nawa, Machinoto (DMNH 140587,3). Okinawa, Bolo Pomt (LACM 28828,12; AMNH 156001 ,5). Okinawa, off centre of Kadina Circle sea wail (AMNH Jackson 1992 A9873, 14). Okinawa, Hyakuna Reef (AMNH 220140, 2). Okinawa, Shioya, Shanawan Bay (USNM 589031, 1). Okinawa, off Oginmi (USNM 488037, 1). Okinawa, Onna Village, Horseshoe Cliffs (USNM 841169, 1). Okinawa, Naha Airport (SBMNH no #, 3). Oki nawa, Kadena R eef (SBMNH 25867, 1). Okinawa, Chinen Tide flat (SBMNH Albert #0-304,2). Okinawa, Moon Beach (SBMNH Hanselman #747-749,3; SBMNH Hansel- man #2767,2768, 2773,2774,4). Okinawa, Zampa-misaki reef (SBMNH no #, 1). Iri- mote (SBMNH Albert #0-304,1). Philippines: Luzon, Batangas (DMNH 187792, 1). Luzon, Batangas, Calatagan (DMNH 004401,2). Luzon, W Quezon (DMNH 188659, 1). Luzon, E (Quezon (DMNH 189644,1). Luzon, Quezon, Bondoc Penmsula (DMNH 193733,1). Luzon, Bataan Province, Luson (LACM 152166,1). Luzon, Zambales, Iba (ANSP 229239,1). Luzon, Lingayen Gulf, Pangasinan (DMNH 193445, 1). Luzon, Lingayen Gulf, Pangasinan, Nara Isl. QDMNH 193621,4). Luzon, Sorsogon, Magal- lanes (AMNH 80267,9; ANSP 223930,12). Luzon, Sorsogon, Badang, near Gubat 210 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Haliotis clathrata H aliotis squamata Haliotis diversicolor H aliotis dissona Haliotis nibiginosa 2 ^^?^ H aliotis jacnensis H aliotis nibiginosa Figure 4-24. Distributioa o f Indo-PactBc species o f H aliotis sp. (I). Localities stem ming &om collection specimens are indicated with, a solid circle, those firom the lit erature with a solid square. For data see text. 211 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-25-42. Shells o f Indo-Pacific abalone. Figures 4-25,28. H aliotis crebrisctdpta Sowerby, 1914. BMNH 1919.12J1.19. “New Caledonia”. 30 mm. Lectotype selected by Stewart & G eiger (1999). Figures 4-26,27,29,30. H aliotis glabra Gmelin, 1791. Figures 4-26, 29. DLG AAB 19b. BCushinoto, Wakayama Preference, Japan. 40 mm. Typical color pattern. Figure 4-27. DLG AAB 19b. Kushinoto, Wakayama Preference, Japan. H 1987.40 mm. Unkommon dark color pattern. Figure 4-30. DLG AAB 19c. Kushinoto, Wakayama Preference, Japan. 40 mm. Light color pattern. Figures 4-31-33. H aliotis ovina Gmelin, 1791. Figure 4-31. JK. Site 88, Astrolabe Reefi Fiji. 39 mm. Ventral view. Figure 4-32. DLG AAB 26h. Phuket, Thailand. 56 mm. Typical green variation. Figure 4-33. DLG AAB 26a. Helengeli, Maldives. 41 mm. Figures 4-34, 37. H aliotis varia Linnaeus, 1758. DLG AAB 15f. No localiQr. 44 mm. Typical specimen o f this extremely variable species. Figures 4-35,36. H aliotis asim na Linnaeus, 1758. DLG AAB Olo. No localiQr. 66 mm. Typical green specimen. Figures 4-38-39. H aliotis p u l - cherrima Gmelin, 1791. DLG AAB 34a. Anaa, Takapoto, Nîaa atoll, Tuamotu. 25 mm. 40-42. H aliotis diversicolor Reeve, 1846. Figures 4-40,41. DLG AAB 04a. J^ a n . 68 mm. Green finely mottled spedmeiL Figure 4-42. DLG AAB 29 a. No locality. 74 mm. Red, coarsely mottled specüneiL 212 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 213 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (ANSP 224376,6). Luzon, Sorsogon, Matnog (LACM 83718,2). Luzon, Albay, Tabaco Bay (LACM 31800,2). Luzon, Albay, Libon (LACM 83799,1). Samar (DMNH 192521, 7; MHNG 21817,1). E Samar, Borogan Village (ANSP 223337, 1). Mindoro, Calapan (AMNH 80446, 4; AMNH 121973,4). Mindoro, Calapan, Baco Isl. (DMNH 049140, 5). Mindoro, Puerto Galero, Little Balateros Cove (DMNH 192764,3). Mindoro, Cabra Isl. (LACM A.2777a, 2). Mindoro, Dalaovan (LACM 120218, 1). Mindoro, Pola (AMNH 98957, 1). Mindoro, Mansalay Bay (USNM 245532, 3). Mindoro, off Point Calavite (USNM 632562, 1: complete). Marinduque, Boac (DMNH 187195, 1). Min danao, off Zamboanga, Bilau-Bilau. Isl. (USNM 245539,1). Mindanao, off Zamboanga, Bilan Bilau Isl., o ff Moros (USNM 245507, 18). Mindanao, off Zamboanga, Panga- puyan Isl. (USNM 245540, 2). Mindanao, Zamboanga (LACM AHF Acc. 1205, 8 ; AMNH 220142,18). Mindanao, Zamboanga, Bilan-bilan Isl. (DMNH 043910,3). Min danao, Zamboanga, Santa Cruz Isl. (LACM 81-6, 1). Mindanao, Davao Bay (DMNH 118539.3). Mindanao, Davao, Gulf o f Mindanao, Davao City (USNM 747831,3). Min danao, Surigao Province (LACM 74720, 1). Mindanao, Misamis Occ., Calamba (AMNH 184049, 1). Panay, Iloilo, Bagacy (DMNH 192020, 1). Panay, Antique, San Royal (DMNH 192334,1). Cebu (LACM A. 2777c, 4; AMNH 80430,1). Cebu, Olango Isl. (DMNH 002169, 11; AMNH 80210,4). Cebu, off Olango, Cawhagen Isl. (ANSP 231846,1). E Cebu, 2 km S o f Olango IsL Salpa Isl. (ANSP 230484,2). E Cebu, off Mactan, Hilutungan IsL (LACM 81-4,1). Cebu, Bantayan Isl. (LACM 45938,1; DMNH 122436.3). Cebu, Malaboyoc (ANSP 104363,1). Cebu, Mad., Kaongrad (ANSP uncat aloged, 3). Cebu, Cebu City (USNM 631426, 4). Cebu, Cebu City, Camotes Sea (SBMNH 2719, 12). Mactan (USNM 245515, 3). Masbate Isl. (DMNH 176115, 3). Leyte Isl. (DMNH 194172,2). Leyte IsL, Vülaba (AMNH Edison A 9853,2). Leyte,, v niaka (AMNH 240626,8). Bohol (DLG Oli, 3). Bohol, Mantacas Isl. (USNM 245521, 214 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3). NW end o f Bohol, E side JagoUao IsL (ANSP 231453,1). N Palawan (LACM 90283, 2; 74725, 2). Palawan, Cuyo Isl. (DMNH 002253, 6 ; ANSP 223517, 5). Palawan, Calamian Isl., Lusong Isl., Bnsuanga (DMNH 178901,2; DMNH 178163,1). Palawan, Calamfan Group, Manglet Isl. (LACM 74702,1). Palawan, Culion (SBMNH no #, 2). Palawan, Ulugan Bay (USNM 245496, 2). S Palawan, Balbac (LACM 74740, 17). N Basilan Isl. (USNM 245554,1). Sulu Archipelago, Siasi (USNM 589461,4). Sulu Arch ipelago, Siasi and Bongao Isl. (USNM 313982, 1). Sulu Archipelago, SW o f Siasi Isl. (DMNH 028209, 2). Sulu Archipelago, SW of Siasi Isl., W o f Caluman Isl. (ANSP 318826, 11). Sulu Archipelago, Siasi Isl., Jolo Isl. (DMNH 049412,5; AMNH 105908, 4). Sulu Archipelago, Jolo (LACM 90417, 2). Off Sulu Archipelago, Jolo Isl., Muobo Beach (ANSP 318191,6). Sulu Archipelago, Bongao Channel, SW end Sanga Sanga Isl. (ANSP 230776, 7). Sulu Archipelago, Tabawan Isl. (ANSP 314975, 7). Sulu Archi pelago, SW o f Tabawan Isl., Nusa Isl. (ANSP 318273, 1). Sulu Archipelago, Dammai Isl. (USNM 245550, 1). Sulu Archipelago, Tawi Tawi Group, Papatag Isl. (USNM 245523, 1). Sulu Archipelago, Tawi Tawi Group, Simonor Isl. (= Simunul) (USNM 245544,1). Negros (LACM 45959,1). Negros, Near Giligaon, N o f hfeloh 0 9 ® 06’ 30” N 122“ 55’ 24” E (USNM 828722,1: complete). Siquijor Isl., W o f Little Point (USNM 828682, 1: complete). Catanduanes, Gigmoto (ANSP 223081, 1). Palau (= Belau): (AMNH 220138). Rattakadokoru Isl. (DMNH 069888, I). SW Rattakadokoru Isl. (ANSP 201667,3). Babelthuap Isl. (DMNH 046177, 1). S Babelthuap Isl., W Passage (ANSP 200961,2; ANSP 200898,2). E Babelthuap Isl., Reef I mile S o f Namelakl Pass (ANSP 203162,1). Babelthuap IsL Bridge Jetty (SBMNH no #, 2). Reef S o f W Babelthuap IsL, Karamando Bay (ANSP 200982,1). Malakal Harbor (ANSP 202838, 1). Koror to MalaicaL shore by causeway (ANSP 283702, 1). Koror Isl., Near Ferry Dock (ANSP 202962,1). Reef N o f Garokottan IsL (ANSP 20I231,1). Ngargersiul IsL 215 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (USNM 620863,1). Malaysia: Borneo, Sabah, Banggi IsL (NMW). Borneo, Sabah, Jes- selton (DMNH 059708,2). Borneo, Sabah, Jesselton, Tamjong Am (ANSP 275048,10; USNM 658296, 6). Borneo, Sabah, Sempora (BMNH 2277, 1). Borneo, Sabah, Sem- poma reef (ANSP 295384,3). Borneo, Kudat District (ANSP 255709,1; ANSP 255709, 1). Borneo, Mandi Darrah Isl. (AMNH 106758,1; USNM 633464,3). Indonesia: Suma tra, Batavia/Jakarta (MNHN no #, I). O ff Sumatra, Ptdau Bai, Batu Group (USNM 654482, 6). W Bali (LACM 45897,2). Bali, Sanur (DLG Old, I; DLG Olg, 6). Bali, Nusa Dua (LACM 86-164,2: complete). Bali, Denpasar (USNM 660546, 1). Sulawesi, Manado (DLG Ole, 3). Ceram, Pirn Bay, Tg Tutuhuhur (USNM 746703, 1: complete). Aroe and Tenimber Isl. (NHB 1707c, 7). Tanimbar, Seiaru, Tg Tuwaum (USNM 747661, 2). Tanimbar, central N shore Larat Isl. (USNM 747525,2: complete). Lompok (DLG no #, 1: complete). NW Lompok, S Ajer (= Gili Air) (LACM 88-62, 1). Lompok, Seng- gigi Beach, Mangait Point (SBMNH 5618,2). N o f Java, Thousand Isl. (AMNH 187261, 1). MoUuccas (= Maluku) (NMW). Moiluccas, Ambon (BMNH 2525, I; NMBE 1517.394, 3). MoUuccas, Nuhu Jaan, Elat Bay, W Nuhu Tjut, Kai Isl. (USNM 747138, 1: complete). MoUuccas, N Toba Isl., N o f Am (USNM 747881, 3: complete). Irian Jaya, Sarong (NMBE 1517.381, 2). Irian Jaya, Biak IsL (AMNH 99225, 2; USNM 600642, 10). Irian Jaya, Biak Isl., NR, KLM, Resthouse, 1 mile E o f Dock (ANSP 206481.5). Wan Jaya, Biak dock-reef about 1.5 mUes SW o f (ANSP 206359,2). Wan Jaya, Biak, Noesi Isl. (SBMNH no #, 1). Wan Barat, Noesi Isle, Mios Woendi Atoll, Reef E o f Padaido IsL (ANSP 205038, 1). Wan Jaya, Rani IsL, 20 miles W o f Biak Isl., R eef and Isl. 5 miles NW o f (ANSP 206704, 7). Wan Jaya, Yapen, W Samberbaba, Cape Tekopi (ANSP 205167,2). Wan Jaya, near HoUandia (USNM 611881,2). New Brittain IsU Rabaul Harbor (ANSP 274806, 5). New Brittain Isl., Rabaul (ANSP 274806.5). New Brittain EsI., 5 miles from Rabaul, Matapit Isl. (ANSP 304650,4). 216 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Barat Java, S Noekori Isl., E Padaido Isl. (ANSP 206178, l). Irian Barat, Abroeki Isl., Maransabadî Isl., Aoeri Isl., Geelvink Bay (ANSP 208693, I). PNG: Kapa Kapa (DMNH 047218,3). Aitape, Seleo Isl., NW reef (USNM 616056,1). Trobriand Group, Kiriwina Isl., Wawela (SBMNH 2322, 1). Port Moresby, Ela Beach (LACM 81-22,1: complete). Bougainville (LACM 45878, 4). Bougainville, Buin (LACM 45895, l). Admirality Isl., Manus (AMNH 220139, l; USNM 6II713, l). Solomon Isl.: BougainviUe, Buka Isl. (USNM 835589, I). Malaita, N Malaita Isl. (DMNH 050869, 4). Malaita, Auki (DMNH 046839, I). M alaita Isl., Ataa (AMNH 118203, 2; ANSP 289634,3). Santa Cruz Isl. (LACM 45958,1). Santa Cruz Group, Reef Isl., Pigeon Isl., lagoon (SBMNH Phillips# B3378, 2; SBMNH 1218, 5). Shortland Isl. (ANSP 310065, 1). Bungana Isl. (BMNH 2144, 1). Nggela Isl. (USNM 770325, 1). Guadalcanal, Marui/Marau Sound, Tavanipupu Isl. (SBMNH 3504, 1; SBMNH 1407,4). New Cale donia: Loyality Isl., Relais de We, Chauteubraind (DMNH 116414,1). Fiji: Viti Levu (LACM 45980, 2). QLD.: Torres Straits (LACM 22189, 3). Torres Strait, Moa Isl. (BMNH 2341,1). GBR, Hardy Reef (AMNH 104584,2; AMNH 249918,1). Lizard Isl. (USNM 795188, 1: complete). Low Isl. (LACM 45892,2). Green Isl. (DMNH 010960, 4; ANSP 234368,1). Off Caims (MHNG 21813,1). Cairns, reef (DMNH 031902,25). Cairns, Double Isl. (ANSP uncataloged, 4). GBR, Fitzroy Lagoon (AMNH 144301,1). GBR, Holboume Isl. (ANSP 134536,1). Bowen (DMNH 127389,2; LACM 63170,1). E o f Whitsunday Passage, Reef (AMNH 104760,1). Whitsunday Isl., Lindeman Isl. (LACM P.465.62,3). GBR, Capricorn Group, Tryon Isl. (LACM A .8589,1; LACM 25763,4). GBR, C ^ rico m Group, Heron Isl. ^A C M 45933, 1; AMNH 95698, 1). GBR, Capricorn Group, NW Is l (LACM 45941, 5; ANSP 236300,1). GBR, Capricorn Group, One Tree Isl. (ANSP 236317,1). Capricorn Isl., Wreck LsL (SBMNH WW2331, I). Yeppon (SBMNH 191,7). ïeppon, Keppel Bay, Keppel E sL (SBMNH 181,7). Kep- 217 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. pel Isl. (AMNH 208300,2), Near Gladstone (USNM 595680,1). Gladstone, Tryon IsL (SBMNH 23915,2). Lady EUiott Isl. (ANSP 219616, I). ShaUow Bay (LACM 45968, 3). NT: Damley Isl. (LACM A.2777b, 2; LACM 110388, I). Yirrkala, Arnhem Land (AMNH 105237, 3). WA: NE of Onslow, Passage Isl. (LACM 95-64,2 + 1 complete). Roeboume (LACM 34284,1). Port Headland (AMNH 121841,1). King Sound (AMNH 199648, 2). King Sound, Sunday Isl. (AMNH 199649,3). Scott Reef (AMNH 272131, 1; USNM 681666,2). Barrow IsL, S end Harcourt Bay (USNM 691911,1: complete). NSW: Sydney, Port Jackson (BMNH, 2: complete). SA: St. Vincents G ulf (LACM 45962, 12). LITERATURE, LOCALITIES: WA, Ningaloo Marine Park (Wüson, 1993). WA, North West Islet; NT, Murray Isl. (Cotton, 1943). Komodo (Dauphin & Denis, 1995). MoUuccas, Halmahera (Oostingh, 1925). Banda Neira (Adam & Leloup, 1938). Fiji, Viti Levu, Suva; Loyality Isl., Lifu; Guam (Talmadge, 1963a). N o f Penang (8* 30’ N 98* 00’ E) (Talmadge, 1974). Kepulauan Seribu (Stewart, 1986). Sulawesi, Gunung Tangkoko/Dtia Saudara Park (KitteL 1993). Indonesia, Batang Isl. (Baer & Baer, 1993). LITERATURE, RANGE: NSW (Alien, 1959). S-most isl. o f Japan and S (Kira, 1962). Ryukyu; Philippines; Indonesia; New Guinea; Solomon Isl.; New Caledonia; Fiji; Loy ality Isl.; Marianas; Caroline Isl.; GBR; N Australia; Japan (Talmadge, 1963a). Indone sia; N and E Australia; Marianas; Philippines; Chma; Ryultyu; Amami IsL; Indian Ocean (Habe & Kosuge, 1964). PNG ^Uhton, 1972). North West Cape - QLD (Wells & Bryce, 1985). Philippines (Springsteen & Leobrera, 1986). Indonesia (Dharma, 1988). North West Cape - Capricorn and Bunker Group (Wilson, 1993). Thailand, Malaysia, Aus tralia, Indonesia, Borneo, PhOippmes, Japan, Okinawa, Taiwan, New Caledonia, Belau, Solomon Isl. (Gosliner et aLy 1996). 218 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. HaUot i s clat hrat a Reeve, 1846 (non Lichtenstem, 1794) (Figures 4-24,4-138,4-141, 4-175,4-176) kenya: (BMNH 241, I). Madagascar: Tulear (MNHN no #, 2). 25 km N o f Tulear, Mora Mora Village (KAS no #, 2), Nosi Be, N Nosy Komba, Pointe Ambarionaombt (ANSP 258629,1). SWNosi Be, betweenAmbatoloakaand Madirokely (ANSP 259131, 2). Comoros: (DLG no #, 1: complete). Sandy IsL, Mayotte (NM J9665, 1). Anjouan: (BMNH no #, 1). Mayotte, Benthedi, N Pamanzi IsL, 12' 45’ 01” S 45' 17’ 09” E (MNHN sta. S 32,10). Mauritius: (DMNH ex 011016,3). BeUe Mare (CASIZ 044963, 2: complete). Trou aux Biches, Lagon (MNHN no #, 1). Aldabra Atoll: Picard Isl. (USNM 836531,1). seychelles: (BMNH no #, 1). Rodrigues Isl.: (BMNH 76.5.1.72,1). P t Mathwesi (DLG 44g, 1; KAS no #, 1). Chagos Archipelago: Ile du Coin, Peros Ban- hos (BMNH, 1). Peros Banhos, Ile Poule, seaward reef slope (BMNH 2307, 1). Mal- dive Isl.: GanAddu Atoll, south reef (BMNH, 1). Ari Atoll, NE o f Feridu IsL, Islet 5.5 km NE o f Feridu Isl. (ANSP 303927, 1: complete). Helengeli [= Helengüi] (A. Faucci collection, 23). Andaman Isl.: 80 km E o f S Andaman Isl., N end o f Invisible Bank (ANSP 292648, 2). Singapore: (BMNH; ANSP 196380, 1: complete). Raffles Light (ANSP 245726,3). Sentosa (KAS no #, 1). SW of Keppel Harbor, Palau Hantu (USNM 660824). South China SeA: Macclesfield Bank (BMNH 92.9 JO . 159.160, 2). Japan: Honshu, Hachijo Isl. (ANSP 240168,1). Okmawa, 1 km WNW o f Onna Village (LACM 78-29,1; LACM 79-75, 1). Okinawa, Onna Village, Horseshoe cliffe (USNM 838612, 1). Okinawa, 5 km ESE o f Zampamisaki, Bolo Point (LACM 78-25,1; LACM 78-100, 1). Okinawa, Serigaki Beach ^ L G 44k, 6 ; AMNH 276888,3). Okinawa, 1 km S o f Kuwae Hospital (USNM 838483, 1). Okinawa, I km S o f Kuwae Hospital near old Hamby Yacht Club (USNM 838483,1). Philippines: Bohol (NMW, I). Siquijor Isl., San Juan, San Juan Reef (USNM 802093,1). Luzon, Batangas (CASIZ 081039, 1). 219 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Luzon, Batangas, Devil’s Point (CASIZ, I). Luzon, Batangas, Calatagan (DMNH 205368.1). Luzon, Bataan Province (LACM 74724,1). Palawan, San Pedro Cove, Lina- pacan IsL (LACM 88-285^1, 1). Palawan, Calamian Group, Batunan Isl. (LACM 74701.1). Mindanao, Zamboanga, Yellow Beach. (KAS no #, I). Mindanao, Silino Isl. (USNM 245500, 1). Mindoro, Aro Point (KAS no #, I). Lubang (KAS no #, 1). Mari- caibo. Devil Point (KAS no I: complete). Palan (= Belau): Koror, Malakal Harbor (ANSP 203083, I). Arakbesan Isl. (ANSP 204544, 2). Indonesia: Java, Thousand Isl., Palau Pelangi (KAS no #, I). Java, Pulau-Pulau Seribu Isl., off Jakarta Roya, Pelangi and Putri Isl. (LACM 86-163, 8). Schouten Isl., Biak Isl. (USNM 600559,3). Sulawesi, off Menado, S Banuken and SOaden Isl. (LACM 88-56, 3: 1 complete). Bali (RP no #, 2). Bali, Lovma Beach, N coast (KAS no #, 6). Dual Tual (RP no #, I). Lesser Sunda Isl., Komodo Isl., Station JEM 87-4 (CASIZ 081123, I). Ambon, Nus Laut Isl. (DLG 44e, I: specimen from Baer, 1994). Malaysia: Borneo, Sabah, Sipidan (KAS no #, I). Borneo, Sabah, Sipidan, Sapi Isl. (KAS no #, I: complete). New Guinea: N coast near Madang, Pig Isl. (= Tab IsL) (CASIZ 086544,1). 2.5 km SW o f Biak, Dock-Reef (ANSP 206392, 2: complete). New Britain (RP, I). New Brittain, Rabaul (RP no #, 3). NW Nusa Perida, Toya Pakeh (DLG 44h, I; KAS no #, I). WA: Roebuck Bay (AMNH 220132.1). NE comer o f Seringapatam Reef (DLG 44i, I). Ashmore Reef (DLG 44f I; KAS no #, I). Freemantle, Cochbum Sound, Woodman Point (ANSP 358590,1). QLD.: Lizard Isl. (LACM 79-53, I; LACM 79-55, 1). Capricorn Isl. (CASIZ I0257I, 3). Swain’s Reef (CASIZ 102570,2). Middle Keppel Isl. (CASIZ 102572,3). Great Kep pel Isl. (KAS no #, I). South Keppel IsL, o ff'Ÿèpoon (CASIZ 102936). Keppel Group, Conical IsL (CASIZ 102937). Keppel Bay, hCddle Isl. (KAS no #, 3). Keppel Bay, Kep pel IsL (SBMNH, I; ANMH 220249,2). Keppel Bay (SBMNH, I). Keppel Bay, Pump kin Isl. (KAS no #, I). Humpy feL ^CAS no #, 3). Gladstone (RP no #, I). Moreton Bay 220 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (KAS no #, 1), Stanage Bay (KAS no #, 4; SBMNH, I), Great Barrier Reef, Grub Reef (LACM 83-44, 1). NSW: Sydney, Botany Bay (SBMNH 209,2). Lord Howe IsL, Neds Beacb (AMS ex 328700,1: complete). MarianaS: W Saipan (DLG 44j, 3; KAS no #, I). Guam (USNM 887465, 1; AMNH 220127,2). Guam, Orote Cliffs (AMNH 220528, 1). Guam, Tweed’s Cave (USNM 852034, I). Guam, o ff Gun Beach, Tumon Bay (USNM 698004, I). Solomon IsL: NE side Vanganu IsL, Marovo Lagoon, Kokuana Passage, Matui Isl. (LACM 89-77, 1). Honiara (CASIZ, I). Bunana Isl. (BMNH 2144, 2). New Caledonia: Central N side, Bogota Reefs (USNM 693386, I). Grand Reef o f Koumac (MNHN sta. 1316,3; MNHN sta. 551,1). Passe de Koumac (MNHN sta. 1310, 1). Sector o f Beiep (MNHN sta. 1217, 1; MNHN sta. 1128, 1). Touaourou (USNM 795269, 1). Cook’s Reef (CASIZ 102574). He des Pins (KAS no #, 2). lie des Pins, Kuto Beach (CASIZ 102494). Noumea, Ilot Charron, Baulari Bay (ANSP 275419,1). Noumea, Baie de Citron (ANSP 237557, 1). Noumea, Baie Ouemo (DMNH 19675,1; ANSP 271204, 1). Noumea, Touho, Koe Reef (DMNH 69885, I). Noumea, Touho (ANSP 238033, 1). Sector o f Touho (MNHN sta. 1264,1; MNHN sta. 1271,1). Bouraü (AMNH 107198,1). Ouen Isl., Prony Bay (MNHN sta. 2 3 2 ,1). Sector o f Yaté (MNHN sta. 735,1). N. O. “Afis” Campagne SMIB 5,23“ 25’ S 168“ 05’ E (MNHN sta. DW99, 1). Federate States o f Nficronesia: Upper Mortlocks, Losap Isl. (DMNH 205366,1). Kapingamarangi AtoU (UCMP loc. # 13107,1). Eniwetok AtoU (AMNH 92485,1). Eniwetok, Rigili Isl. (USNM 580735, 1). Kwajalein (KAS no #, 1). Kwajalein, West Reef ^ L G 44c, 2; KAS no #, 4). Kwajalein, Carlson Isl. (KAS no #, 3; KAS no #, 1). Rongerik, Bock Isl. (USNM 586290, 1). Tonga: Ha’apai Group, Cornfield and Camp bell (CASIZ 102573,1: complete). Vava’u Group, SW Vava’u Isl., S end Pangahnota Isl., cliff at W end o f Maugaui (LACM 86-220,1). Vava’u Group, N side Nuapupa Isl., outside o f lagoon ^A C M 85-89,1). Vava’u Group, between Longitau Isl. and Vaka’eftu 221 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. IsL (LACM 85-90.1). Eua IsL, A a'a Luma Beach (KAS no #, 1). WESTERN SAMOA: Savaii Isl., Mataatu Harbor, Eastern Reef (AMNH 178945,1). American Samoa: Fagan Bay (KAS no #, 2). LITERATURE, LOCALITIES: Indonesia, Bali, Temukus Roads (Boone, 1938). QLD., Hayman Isl. (Anon., 1975). QLD., Michaelmas Cay; WA, NW Cape (Anon., 1982). Kepulauan Seribu (Stewart, 1986). WA, Ningaloo Marine Park (Wilson, 1993). LITERATURE, RANGE: New Caledonia; tropical Australia; Hachijo-Jima; Singa pore (Talmadge, 1963a). PNG (Hinton, 1972: as crebrisculpta). E coast (Hinton, 1978: as crebrisculpta). As far S as Keppel Bay, Guadalcanal, Japan (Anon., 1982). Japan through N Australia (Abbott & Dance, 1983). Lesser Sunda Isl. (Dharma, 1988: as ere - brisculpta). New Caledonia (Salvat et a i, 1988: as coccoradiata). Scott Reef - N QLD (Wilson, 1993). Madagascar, New Guinea, Borneo, Philippines (Gosliner et a l, 1996). Madagascar, Rodriguez Isl., Chagos Archipelago, Maldives, Okinawa, Tonga (Stewart & Geiger, 1999). Haliotis crebri scul pt a Sowerby, 1914 (Figures 4-24,4-25,4-28) New Caledonia: (BMNH 1919.1231.19,1: lectotype). HaUot i s diversi col or Reeve, 1846 CF^ures 4-24,4-46-42) JAPAN: Honshu, Wakayama, Nada (MHNG 21818,1). Honshu, Wakayama, near Seto Marine Laboratory, Bansho-zaki (LACM 82-19, > 50). Honshu, Wakayama, Shi rahama, Kii (AMNH 260667,4; AMNH Edison A9853, 1). Honshu, Wakayama, Goza, Sime (ANSP 219612,1). Honshu, Wakayama, Nada-cho, Kusui (AMNH 220246,2; AMNH 241420,5). Honshu, Awa and Shhahama, Kii coast (DMNH 049707,3; AMNH 198684,2). Honshu, Ishikawa, Noto Peninsula, Lighthouse at AkasaJd (LACM 82-7, 222 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1). Honshu, Ishikawa, Noto Peninsula, Monzenrfukami (LACM 82-5). Honshu, Niig- ota, Sado IsL, Aikawa Macfai, Samuto Saki, Seki (ANSP 399942, 3). Honshu, Kana- gawa, Sagami Bay (DMNH 010974, 2; AMNH 112368, 10). Honshu, Kanagawa, Sagami Bay, Akiya (LACM H-792, 2; ANSP 209152,2). Honshu, Kanagawa, Sagami Bay, near Kanagawa (ANSP uncataloged, 4). Honshu, Kanagawa, Sagami, Hayama, near Kamakura (ANSP 86090, 14). Honshu, off Hondo, Sagami Bay, Misaki (USNM 206245, 2). Honshu, Kanagawa, Yokohama (USNM 43067a, 3). Honshu, Kanagawa, 0-shim a (AMNH 19699,2; MHNG 13180, I). ISO km S of Tolq^o, Miyake Jima Isl. (AMNH 186262, I). S o f Tokyo Bay, Hachijo Isl. (AMNH 198677, I; ANSP 252616, 5). Iblqro (MHNG 13168, 3). Honshu, Chiba, Station Boshu (LACM 120225, I). Hon shu, Akite, Fukura Awaji (ANSP 76329, 2). Honshu, Shizuoka, Toga (ANSP 252585, 4). Honshu, Mie (ANSP 252584, 10). Honshu, Mie, Ise (AMNH 80436, 2). Honshu, Mie, Anori (ANSP 252586,6). Honshu, Mie, Toba (AMNH 220192,1; DMNH 186156, 2). Honshu, Aishi, Mikawa (USNM 664049,1). Nagai Reef (LACM 91229,3; LACM 91274, I). Shikoku, Ehime, Misaki (ANSP uncataloged, 1). Shikoku, Kochi, Tosa (LACM P222.53, 2; AMNH 220247, I). Shikoku, Kochi, Tosa (DMNH 008691, 4, DMNH 008705, 4). Kyushu, nd Isl. (NHB 3835-b). Kyushu, Enoshima (DMNH 061282,2). Kyushu, Kagoshima (ANSP uncataloged. I; USNM 226854, 1). Kyushu, Kagoshima, Satsuma (ANSP 76324, I). Kyushu, Nagasaki (NHB 5438-a, 3; USNM 226858,5). Kyushu, Nagasaki, Hhado (AMNH 19686,2). Kyushu, Nagasaki, Krado Uzen (ANSP 80368,8). Kyushu, Nagasaki, Hhado Hizen (ANSP 262740,2; ANSP 49765, 4). Kyushu, Nagasaki, Matsushima Bay (ANSP uncataloged. I). Kyushu, Nagasaki, Mogi (USNM 226860, 8). Okinawa (USNM 671030, 2). Okinawa, Awa (ANSP 119841,15). KOREA: Cheju-do (= Quelpart Isl.) (LACM 71-256,1). China: (LACM 45940,1; AMNH 220174, 1). Hong Kong (LACM 45957,6 ; LACM 45831, 223 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1). Hong Kong, Port Shelter, Dock Rocks (ANS? 222671, 2). Fokine, Santu (NHB 3835.a, 9). Fikine, Foochow (Fuchau) (DMNH 011038, 2). Taiwan (= Formosa): (AMNH 164450,1; AMNH 182979, 6 ). Taipei (USNM 708574, 8). philippines: Min doro (AMNH 220172, l). Minodoro, Quadras (USNM 303330, l). Indonesia: Bali, Beach Hotel (ANS? 319655, 1). WA: E of Albany, near Point Valliant, Two Peoples Bay (LACM 87-96, 1). Exmouth Gulf, Exmouth (SBMNH 6185,1). Tonga: Tongatapu IsL, N o f Government Fisheries (LACM 81-56, 1). Ha'apai Group, Limu Isl. (LACM 72-153,3). LITERATURE, LOCALITIES: Yedo [= Tokyo] (Pilsbry, 1895). LITERATURE, RANGE: N Hokkaido and S (Kira, 1962: as supertexta). N Australia; Indonesia; Philippines; China; Ryulgm; Amami Isl.; Bonin Isl.; Kyushu Isl.; Shikoku Isl.; S Honshu - Boso Peninsula (Habe & Kosuge, 1964). Shirahama, Kii Peninsula and S (Habe, 1964). Shirahama, Kii Peninsula (Japan) and S (Lindner, 1975). Japan, Korea (Abbott & Dance, 1983: as aquatilis). HaUot a di ssona (Iredale^ 1929) (Figures 4-24,4-139,4-140) Marianas: (MNHN no #, 2). Federate States o f micronesia: (AMNH no #, 1). New Caledonia: (AMNH 44374,3; AMNH 44370,1). Noumea (DMNH116937,1). Noumea, Baie Citron (BMNH2341,1). Passe de Koumac (MNHN sta. I311,2; MNHN sta. 1319, 2). VANUATU: (MNHN no #, 1). FIJI: (MHNG 13171, I). S o f Koro Levu Isl., Tave- uni (USNM 695560,1 [?]). Kandavu, E o f Yauravu Pass (USNM 696660,1). Viti Levu, Suva Mam Reef Rat Tail Passage (USNM 835591, 1: complete). Astrolabe Reeve (JK no #, 1; JK no #, 6 with dried bodies). Tonga: Tongatapu, Sopu (USNM 672192,2). T ongat^u IsL, N o f Government Fisheries (LACM 81-56,1; BMNH no #, 2). Ha’apai G roiç, Lhnu Isl. (LACM 72-153,3). 224 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE, LOCALITIES: Michaelmas Cay (Iredale, 1929: type locality). Haiiotis dohmi ana Dunker, 1863 (Figures 4-24,4-120,4-123) Indonesia: Java, Jakarta Raya, Pulau-Pulau Seribu IsL, Pelangi and Putry Isl. (LACM 46-162, 1: complete). PHILIPPINES: Tatri Taui, Tataam Isl. (USNM 245559a, I). Mar shall Isl.: Jaluit Isl., Jabor Isl. (SBMNH 24051, I). Solomon Isl.: Santa Cruz, Pigeon Isl. (SBMNH 1217,1). QLD.: Wreck Bay, Martha Ridgway Reef (DMNH 31783, I). NEW CAIedonia: Noumea, Amadee Lighthouse (DMNH 116937, I). Passe Koumac (MNHN sta. 1323, 2 [?]). Unia near Yate (USNM 795141, 1 [?]). Vanuatu: Efate (AMNH 221448, 2). SW Efate (USNM 793717, 1: complete). Efate, Point d’Arbel (USNM 787480, 1: complete; USNM 787378, I). SE coast o f Santos Isl. (USNM 787248,1; USNM 787211, 2). Reef S o f Utja, Aneityum (= Anatom) (USNM 692098, 2). Reef near TJipthav, Aneityum (= Anatom) (USNM 692374,6). N Port Aneityum, Aneityum (= Anatom) (USNM 692069,2). TONGA: Tongatapu, Nuku’alofa (SBMNH no #, 5). LrrERATURE, LOCALITIES: Dutch East Indies; Formosa (= Taiwan) (Tal- madge, 1963a). Kepulauan Seribu; Bali (Stewart, 1986). LITERATURE, RANGE: New Caledonia (Abbott & Dance, 1983). Haliods fa tid Geiger, 1999 (Figures 4-43,4-116,4-117) MARIANAS: Mang Island, N Marianasm (USNM 486708, 1: holotype). Agrigan Island, (USNM 487953,1: paratype). NEW BRITAIN: Kumbun Island (ANSP 284325, I: paratype). SOLOMONS: Santa Ana Island (AMNH 81878, I: paratype; AMNH 81899,1: paratype). VANUATU: (MHNG21226,1: paratype). TONGA: (USNM no #, I paratype). NE side o f Tofiia bland, Ha^apai Group (USNM 702975,7: paratypes). 225 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. - Haiiotis ovina ’ . • * Haiiotis . ' * glabra % ^ - Haiiotis planata . ■ ■ - • — '# Haiiotis ûïmma . . Haiiotis varia • * T . Haiiotis n. sp. Geiger • . . . * • # Figure 4-43. Distribution o f Indo-Pacific species o f Haiiotis spp. (II). Localities stem ming firom collection specmiens are indicated with a solid circle, those from the litera ture with a solid square. For data see tex t 226 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reef off Tofùa Island (USNM 702512,9: paratypes). Vavau (USNM no # [ex. San Diego Society o f Natural History No. 32719], 2: paratypes). HaUotis glabra Gmelm, 1791 (Figures 4-26,4-27,4-29,4-30,4-43) Japan: Honshu, Wakayama, Kushinoto (DLG AAB 19b, 10; DLG AAB 19c, 4). Hon shu, Wakayama, Awa and Shirahama, Kii coasts (AMNH 220173, 1). Kyushu, Kagoshima (DMNH 186155, 1). Philippines: Luzon, Bataan (DMNH 181156, 1). Luzon, Bataan, Cabcaben (DMNH 002732, 6). Luzon, Bataan, Corregidor Isl. (LACM 74727,2; ANSP 95395, 1). Luzon, Bataan, W tip o f Corregidor Isl. (ANSP 230117,2). Luzon, Bataan, Luson (LACM 152164, 1). Luzon, Bataan, Lusang (ANSP 246218, 2). Luzon, Bataan, Maviveles (ANSP 230399,4; USNM 283263, 1). Luzon, Bataan, E end o f Sisiman Bay (ANSP 230038,9). Luzon, SE Bataan, at Cabcaben (ANSP 229787, 29). Luzon, SW Battan, Lusong Cove (ANSP 230060, 1). Luzon, Bantangas, Mari- caiban Isl. (USNM 245502,21). Luzon, Bantangas, E o f Maricaban Isl., between Cule- bra Isl. and Malajibomanoc Isl. (LACM 84-162, 1). Luzon, Batangas, Pagaspas Bay (LACM 74729, 2). Luzon, Zambales, Iba (DMNH 002763, 7; ANSP 229243, 14). Luzon, Manila Bay, Cavite (DMNH 122844,2; DMNH 003203,5). Luzon, Manila Bay, Caballo Isl. (ANSP 229670,1). Luzon, Quezon, Baler (AMNH 103697,1). Luzon, Sor- sogon, Sorsogon (LACM 74728,2). Luzon, SE Soisogon, Magallanes (DMNH 002447, 2; LACM A .1930, 4). Luzon, Tayabas Province, Dingalan Bay, N Otpegon Point (SBMNH 12242,1). Marinduque (DMNH 187194, I). Marinduque, Quadras, Laylay, Baie de Boac (USNM 306, 1). Mindoro (DMNH 010979, 3; LACM S.11I6, 3). Min doro, Caiman (DMNH 049711,9; DMNH 049411,17). Mindoro, Calîçan, Hoec^Oato Isl. (ANSP 249511,14; ANSP 242232,4). Mindoro, Calapan, Nazareto, Silonay Isl. (BMNH 1962891, 5; SBMNH 04884,8). Mindoro, Lubang (LACM A.2777a, 4). Min 227 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. doro, Silonay Isl. (NHB 1716b, 5). Mindoro, Silonay Isl., Nazareto Calapan (USNM 656741, 7). Negros (DMNH 010981, I). Bohol (DMNH 186029, 3). Bohol, Jagna (DMNH 186157, 8). NGndanao, Surigao del Sur, Marihatag (DMNH 176017,1). W n- danao, Davao (DMNH 187932, 1). Mindanao, Zamboanga (AMNH 220187,4; USNM 310209, 6). Mindanao, San Ramon Farm, near Zamboanga (USNM 245531, 2). Min danao, near Zamboanga, Ayala (USNM 245561, 35). Mindanao, Dapitan (USNM 303634,1). N o f Mindanao, Camiguin Isl., Mambajas (ANSP 223742,5). Cebu (BMNH 2341, 4). SE Masbate, off Talajit Isl. (USNM 234209, 1). Dalanganem Isl. (USNM 245494,9). Palawan (MHNG 21235, 1). LITERATURE, LOCALITIES: Molluccas, Bachan (Oostingh, 1925). Bali, beach between Boegboeg and Boeitan (Adam & Leloup, 1938). Java (Dauphin & Denis, 1995). LITERATURE, RANGE: N Australia; Indonesia; Philippines; Reunion Isl. (Habe & Kosuge, 1964). Philippines (Talmadge, 1963a; Springsteen & Leobrera, 1986). Philip pines - N Australia (Abbott & Dance, 1983). Moluccas and Lesser Sunda Isl. (Dharma, 1988). Haiioti s jacnensis Reeve, 1846 (Figures 4-24,4-124-129) INDONESIA: Java, Jakarta, near Thousand Isl. (DLG no #, 1). JAPan: (LACM 91304, 2). Honshu, Wakayama, Shirahama, Kii (AMNH 276887, I; AMNH 279268, I). Shikoku, Kochi, Tosa Bay (LACM 22230,2). Loochoo (= Ryukyu) (LACM 91219,2; ANSP 252455,6). Okinawa, off Sunabea wall, 4 km SSW o f Kadena Circle (LACM 78-28,1). Okinawa (LACM S.4628, 1; LACM 45886,1). Okinawa, ESE Zampa-mis- aki, Bolo Pomt ^A C M 78-25,1). Okinawa Gunto, Kerama Retto, SW side o f Gishifu Sbima Q JICM 77-72,1). Okinawa, Madimoto (AMNH ex 187058,1). Okinawa, Imbu (SBMNH no #, I). Philippies: N Samar (DLG AAB 45c, 2; DLG AAB 45e, 15). Mari 228 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. anas: Saipan (DMNH 009712,3; DMNH 115477,3). Guam (SBMNH 26856,1). Guam, Tweed’s Cave dive area (USNM 852057,1; USNM 851181,1). Federation o f Microne sia: Upper Mortlocks Losaap Isl. (DMNH 205367, 8). MARSHALL ISL,: Eniwetok Atoll (AMNH 182982, 2; SBMNH no #, 2). Eniwetok A toll, Aniyaanii Isl. (LACM 114416,1: with dried body). Eniwetok Atoll, between Parry and Eniwetok Isl. (LACM 113210, 2: with dried bodies). Enewetok (Fred), tagooa (SBMNH no #, 1). Eniwetok, Ragoa and Buiri Isl. (USNM 655738, 1). Eniwetok, Rigili Isl. (USNM 581238, 1; USNM 607247, 1). Bikini, Namu Isl. (USNM 580699, 6 ; USNM 580124, 3). Bikini, Bikini Isl. (USNM 579750, 1; USNM 579387, 1): Bikini, Okini Isl. (USNM 579207, 3). Bikini, Yomyaran Isl. (USNM 580299, 1). Bikini, Enyu Isl. (USNM 580961, 1). Rongelap, Kobelle/Kabelle Isl. (USNM 585759, 1; USNM 582105, 33). Rongeiap, Kieskiechi Isl. (USNM 585415, 2). Rongelap, Erapwotan (USNM 584113, 3). Ronge lap (USNM 610131, 1). Rongelap, Burok Isl. (USNM 585859,2: complete). Rongerik, Latobach Isl. (USNM 582952, 1). Rongerik, Enyvertik (USNM 583456, 1: complete). Taka Atoll (USNM 615515, 4). Ailuk Atoll, Ailuk Isl. (USNM 615154, 3; USNM 615155,1). Ujelang Atoll (USNM 614614,1). Wbtho Atoll, Wotho Isl. (USNM 614442, 1; USNM 614437,4). Ujae Atoll, Wbtya IsL (USNM 607363, 6). Ujae AtoU (USNM 614535, 3; SBMNH no # 6). Lae Atoll (USNM 615020, 2; USNM 615030, 2). Lae Atoll, Lae Isl. (USNM 614914,12). Majuro Atoll, Laura (SBMNH 6131,1). Kwajalein Atoll (USNM 486159,2). S Burok IsL (USNM 585859, 1: complete), kmbati: Gübert Isl., Onotoa Atoll (USNM 622624,1). PNG: New Britain (RP no #, 1). New Caledonia: (LACM 45882,4; MNHN no #, 2). Noumea, Baie Ouemo, near radio station (ANSP 399938, 1). Grand Reef o f Koumac ^/B4HN no #, I; MNHN sta. 1323,2). Loyality Isl., Lifou (USNM 423136,2). Tonga: Nlve (= Niue) (USNM 360742,1; SBMNH no #, 4). Tongatapu Group, 3.5 km o f S-most tip Eua Isl., Ha’a Luma (LACM 81-55, 6 ; 229 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. AMNH 222563, 3). Eua Isl. (BMNH uo #, 2). Eua Isl., Aa’A Luma beach (USNM 851407,3; SBMNH 6169,4). Fiji: Astrolabe Reef, site 121 (JK uo #, I). ILES WAL LIS: Futuna, W Anse de Sigave (USNM 676583,1). LITERATURE, LOCALITIES: New Caledonia, lie Nou; Philippines, Bohol, Jacna; Okinawa; Cebu; Guam; Palau; Truk; Ponape; Eniwetok; Bikini; Rongelap; Kwajalein; Suva; New Caledonia, Lifu; New Hebrides (= Vanuatu), Espiritu Santo; Rabaul (Tal madge, 1963a). LITERATURE, RANGE: Ryulqai; Philippines; Maraina IsL; Carolines; Marshall Isl.; Fiji; Loyality Isl.; New Hebrides; Bismarck Archipelago (Talmadge, 1963a). Amami Isl. and S (Habe, 1964). Philippines; Ryulqm Isl.; Amami Isl. (Habe & Kosuge, 1964). Philippines (Abbott & Dance, 1983). HaUotis ovina Gmelin, 1791 (Figures 4-31-33,4-43) Mauritius: ^M N H 010990, 3; NMBE Schuttleworth.401,1). Balaclava (MNHN no #, I). Maldives: Addu Atoll, Feddu lagoon (BMNH, 6 : complete). Addu AtoU, seaward reef (BMNH no #, 1). Addu AtoU, W o f S tip o f WUingUis (ANSP 305183, 1). N Male AtoU, N o f Male, Dunidu Isl. (ANSP 305219, 1). SE side o f N Male AtoU, Imma Isl. (ANSP 305562, I). N Male AtoU, N Kagilsland (ANSP 303991,2). N Malosmadulu AtoU, N and NW o f Ongu Isl. (ANSP 304430,1). Rasdu AtoU (ANSP 399939,2). Tilad- ummati AtoU, NW o f Filadu Isl., Faro Islet (ANSP 304839, 1). Helengili (DLG AAB 26a, I), andamanes: (BMNH no #, 1). Thailand: Phuket (DLG 26h, I). Phuket, SW end o f Bon M . (LACM 85-3,1). O ff Phuket, E Mai Thon IsL (LACM 85-8,2). Phuket, E Kaew Yai Isl. (LACM 85-5,12:1 with cMed body). Phuket, SW Bon Isl. (LACM 85-3, I Ixxfy, I: complete). Gulf o f Siam, Ko Tao (USNM 405685,2). Te Vèga, Goh Sindarar Nua (= Chance Isl.) (USNM 661337,1). Malaysia: Peninsula, Perhentian Isl. (BMNH 230 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2264, I). Pulau Pangkor (BMNH 2264, I). N Borneo, Kudat, Airport Trench (ANSP 262759, I: subfbssil). N Borneo, Tigafai Isl., Malawali channel (USNM 666858, 1). N Borneo, Sempoma, Bahaydulong Isl. (USNM 666729, 1). Off Mersing Coast, Pulau Babi Besar (ANSP 399937, 1). Borneo, Sabah, Jesselton (NMW). N Borneo, Maliangin Isl. (ANSP 298206, 1). New Guinea, New Britain, 5 miles from Rabaul, Matapit Isl. (ANSP 304687, 1). New Guinea, Manus Isl., Pere village (AMNH 215141, 1). Indone sia: Java, Pangandaran (DLG 26], 3). Java, off Jakarta, Raya, Pulau-Pulau Seribu Isl., Pelangi and Putri Isl. (LACM 86-162, 1 + 1 complete; LACM 86-163, I). Java, Batavia/Jakarta (MNHN no #, I). Java, Jakarta Bay, Thousand Isl. (DLG no #, 10). Ambon (NMBE Schuttleworth #402, 2). Bali, Lovina Beach (DLG 26c, I). Irian Jay a, Biak, near KLM Resthouse, 1 mile E of Dock (ANSP 206482,2). Schouten Isl., Biak (USNM 600567,1). Sumatra, N o f Sipora, Pulau Siburu (USNM 654703,2). Te Vega, Pulau Bai, Batu Group, off Sumatra (USNM 654480,1: complete). Vietnam: Nha Trang (MNHN no #, I; SBMNH 03515, 1). Nha Tmag, Hon Leo Isl. (DLG 26m, 6 ; LACM 69-111, I). South China Sea in vicinity o f Nha Trang (LACM 109718,2). Cam Ranh Ba, Gran Passo Die (AMNH 198787,1). Vanfong Bay, Hon Lon (LACM 67-187, 1). Paracels (MNHN no #, 3). Japan: Okinawa (LACM A .8589,1; LACM AHF Acc. 1063, 2). NE Okinawa (ANSP 325440, 1). Okinawa, Ukibaru Shima (DMNH 120273, 3). Okinawa, Bolo Point (AMNH Jackson 1992 A9873,32; AMNH Jackson 1992 A9873, 3). Okinawa, off Sunabe sea wall, 4 km SSW o f Kadena Circle (LACM 78-28,2). Oki nawa, WNW o f Onna Village (LACM 78-27,1). Okinawa, Onna Village, Horseshoe Cliffs (USNM 828892, 1). Okinawa, Kadena Yacht Club, Channel (USNM 821388,1; USNM 821127,1). Okinawa, Yakata, Katabaru, Onna Village (USNM 809879,1). Oki nawa, Ada (SBMNH no #, I). Okinawa, Sobe Pomt (SBMNH2766,1). Okmawa, NNW Oku (LACM 77-61,1). Okmawa, 1.5 miles S o f Henna (ANSP 302731, 1). Okinawa, 231 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Niakama, Catholic Camp (AMNH 248812, I). Okinawa, Serigaki Beach (AMNH 276892,2). Okinawa, Shioya (USNM 600811, 3). Okinawa, Shioya, Shanawan Bay (USNM 489109, I). Okinawa, o ff Ogimi (USNM 488118, I). Okinawa, Chinen reef flat (SBMNH no #, 5). Okinawa, Henza reef (SBMNH no #, 2). Okinawa, Tsuke reef flat (SBMNH no #, I). Kyushu, Osumi, Osima (USNM 343830,5; USNM 273338, 3). Philippines: Luzon, Bataan (DMNH 181157,2). Luzon, Sorsogon, Magallanes (ANSP 223931.1). Luzon, SW Corregidor Isl. (USNM 235293, 1). Luzon, 13“ 43’ 59” N 120“ 49’ 11” E (USNM 801679, 1: complete). Minodro, Calapan (DMNH 049717, 3). Min doro, Puerto Galera (LACM 74730, 1; LACM 74732, 1). Mindoro, Daiaovan (LACM 74737.1). Bohol, Panglao (DLG 26d, 6). Cebu IsL, Cebu City (SBMNH 3502, 1). Leyte (DMNH 194171, 1). Sulu Archipelago, Sanga Sanga Isl. (DMNH 049193, 1). Sulu Archipelago, Bet Siai & Bongao Isl. (USNM 313983, 1). Sulu Archipelago, Tawi Tawi Group, Tataan Isl. (USNM 245559, 1). Sulu Archipelago, Jolo, Bufauau Isl. (USNM 245548,2). Sulu Archipelago, Jolo, Marongasi Isl. (USNM 245535,3; USNM 245546, 1). Sulu Archipelago, Dammai Isl. (USNM 24556,3). Sulu Sea (LACM HH-3740, 1). Mindanao (LACM 45791, 1). Mindanao, Zamboanga province, Zamboanga Bay (LACM 74731, 1). Palawan, Linapacan Isl., San Pedro Cove (LACM 88-285.16, 1). (Quezon, Polillo Group, Palasan (LACM 74734,1). (Quezon, Baler (AMNH 132075, 1). Marinduque, Boac (AMNH 132109,1). S Negros Oriental, Siaton, Tambobo (USNM 808480.1). Negros Isl., Maloh (S tip o f Isl.) (USNM 821756,1). Negros Oriental, Gui- hulhga (SBMNH 28938, 1). marianas: Near Guam (AMNH Edison A9853,2). Guam (MNHN no #, 1). W Saipan, Lagoon (USNM 595505, I). Palau (= Belau): (AMNH 220206.1). Koror IsL (DMNH 046176,2). Near Ngatpeal Pass (DMNH 069889,3). Babelthuap (AMNH 121795, 1; USNM 62093, 3). E Babelthuap, Reef 1 mile S o f NamelakI Pass (ANSP 203153,8). Babelthuap, 1 mile S o f W Passage, 07“ 33’ N 134“ 232 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 29’ E (ANSP 200987, 2). Kotor IsL, Near Koror (ANSP 202455,2). Korukigiiri IsL, Malakal Harbour (ANSP 202046,1). Federate States o f Micronesia: Truk (LACM C- 133,2). Truk, Out o f Moen, Paradise Reef (SBMNH 3037, 1). Yap IsL (LACM 91228, I; ANSP 250072,2). Upper Mortlocks, Losaap IsL (DMNH 037623,3). Ponape (AMNH 284463, I; AMNH 284465). Pohnpei (SBMNH no #, I; SBMNH no #, I). Pohnpei, Green Reef, Jokaj Passage (SBMNH no #. I). Round Rock, Helen Channel, Helen Reef (ANSP 208680, 1). Marshall isl.: Kwajalein AtoU (SBMNH 6135, 1). Eniwetok, Jeit- eiripucchi Isl. (USNM 581709, 1; USNM 584690, 1). Enewetok, Enewetok (SBMNH no #, 1). Eniwetok, RigiU Isl. (USNM 581274,10; USNM 581595, I: complete). Eni wetok, Bokonarappu Isl. (USNM 655777, 3). Eniwetok, Rujoru Isl. (USNM 581515, 9). Eniwetok, Rajoa Isl. (USNM 655723, 1). Eniwetok, Jiriiniea Isl. (USNM 581672, 2). Eniwetok, Mui Isl. (USNM 581624, 1). Enewetok AtoU, Bubble Butt Piimacle (USNM 809786, 1). Ujalang AtoU (USNM 614617,2). WA: Port Headland (DLG 26k, 2). Perth (LACM 45896,1). QLD.: SE Lizard Isl. (LACM 79-55,1). Whitsunday Group, Hook Isl. (DMNH 114349,2). W GBR, Hawk Reef, E of Whitsunday Passage (AMNH 104761.1). Hyman Isl. (AMNH 187150,2). GBR, Cairns, Scott Reef (AMNH 178750, 1). Curtis Isl. (AMNH 80424,2). GBR, off TownsviUe, John Brewer Reef (BMNH 2341.1). Friday Isl., Tonoabatu (BMNH). Wreck Bay, Martha Ridgway Reef (DMNH 031805, 1). GBR, Hardy Reef (AMNH 104585, 3; AMNH 249919,3). GBR, Beaver Isl., Beaver Ree( Knchinbrook (ANSP 399930,1). Capricorn Group, Tryon Isl. (USNM 617150.1). Port Douglas (SBMNH 202,1). PNG: Madan& Madang (Channel (SBMNH 2594,2). Solomon EsL: Malaita ^M N H 050876, 1). Malaita, Roncador Reef (LACM 78-55.25, 1). BougainviUe (ANSP 327759,1). BougainvUle, Buin (LACM 45788, 1). Santa Ana Isl. (AMNH 81900,1). Reef Group, Ngakando E sL (SBMNH 2704,1). Santa Cruz Group, Pigeon Isl. (SBMNH 1216,6). New Caledonia: (AMNH Edison A9853, 233 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2). Bouraü (AMNH 107201, 3). Bouraü, 2 miles NW o f Pass (ANSP 269963, 3). Bourail, Plage de Poe (ANSP 237088,2). Touho Bay (DMNH 069890,1; ANSP 238028, 3). ne Ain 3 miles ENE o f Touho NW (ANSP 270255, I). Voh, Barrier Reef (DMNH 069891, l). Gatope IsL, 5 miles WSW o f Voh, SW (ANSP 269724,2). Noumea (USNM 796586,1: complete). Noumea, Amadee Light House (DMNH 116991,1 +1 complete). Noumea, E end o f Vata Bay (ANSP 275417, 1). Dumbea Pass, 1.5 miles NNW of off Noumea (ANSP 270454, 2). lies aux Goélands, about 7 miles SE o f Dumbea Pass off Noumea (ANSP 270532,1). Grand Reef of Koumac (MNHN sta. 1316,2; MNHN 1319, 3). Yate (ANSP 238321, 1). La Foa (DLG 26i, 2). Brony (DLG 261, 7). Port Koué (MNHN no #, 2). Loyality Isl., Relais de We (DMNH 116516, 1). Loyality IsU Lifou, Relais de We, Bay Chateaubriand (DMNH 116453,2+1 complete). Loyality Isl., Uvaa Atoll, Guetie Isl. (USNM 692693, I), fiji: (AMNH 220205, 3). Yanu-yanu (= I-Ioma Isl.), Astrolabe R eef (LACM 71-205, 1; LACM 71-203, 1). Vanua Levu (DMNH 010989, 1), W Vanua Levu, Rambi Isl., Georgia Cove (USNM 695337, 1). Vanua Levu, NW Rambi Isl., Albert Cove (USNM 695312, 1: complete). Vanua Levu, NE Yandua Isl. (USNM 694701, 1). Viti Levu (LACM 45912, 2; LACM 67-93). Viti Levu, Waya Isl. (DLG 26o, 1). Vhi Levu, Korolevu (LACM 79-42,1). VM Levu, Nanu-i-Ra (USNM 638512, 1). Viti Levu, Suva Harbor (USNM 532329, 3; USNM 532378, 3). Viti Levu, S t Annes-on-Sea, Nadroga (USNM 835597,2). O ff Viti Levu, Malolo Group, Cast away Isl. (SBMNH 6132,2). Yasawa Group, Nacula Isl. (LACM 81-38, 1). Koro, Smoothwater Reefs (USNM 695816, I). Koro, Reefs W o f Stoneaxe Road (USNM 695749,2). Koro, SW o f Tulane Harbour (USNM 695910,1). Taveuni (USNM 695744, I). Taveuni, N ofVuna Point (USNM 695685,1). Tavauni, W o f Koro Levu Isl. (USNM 695606, 2). Kandavu, E o f One Isl. (USNM 696061,1). Kandavu, S o f Nagaloa Pass (USNM 696454, I). Kandavu, S o f Waya Isl. (USNM 696335,1). Kandavu, NW o f 234 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Solondamu, W Coast (USNM 69710, I). Kandavu, Reef E o f Yauravu Pass (USNM 696487, I). Kandavu, NE side o f Nagaloa Pass (USNM 696422, 2). Kandavu, E of Korelevu Bay (USNM 696138,2). Matuku, off SW Point (USNM 686411,1). Vambina (=Vambia) (ANSP 193214,2). TONGA: Ha’apai group, Luangahu Isl. (LACM 72-151, I). Vava’u Goup, Vava’u Isl., Kapa IsL W Luaofa Isl. (LACM 86-231,1). Vava’u Goup, Vava’u Isl., Kapa Isl., W Sisia Isl. (LACM 86-225, 1). Tongatapu Isl. (AMNH 249110, 1; ANSP 305800, 2). Tongatapu Isl., Sopee Reef (AMNH 240645, I). Tongatapu, N Makaha’a Isl., Nuku’alofa Bay (AMNH 220204,2). Samoa: Navigators Isl. (= Samoa) (ANSP 50233,1; USNM 89084,2). Tutuila Isl. (USNM 488747,1). Tuamotus: Makatea Isl. (DMNH 072530,7). LITERATURE, LOCALITIES: Tongatabu, Nukualofa; Monu Reef (Ostergaard, 1935). Lady Elliot Isl. (Cotton, 1943). Swain’ s Reef (Talmadge, 1961). Samar; Cebu; Schouten Isl.; New Caledonia, Cook’s Reef (Talmadge, 1963a). Gulf o f Siam; C^obba, WA (Talmadge, 1974). Bali, Sanur (Baer & Caillez, 1985). Fiji, Matagi Isl. (Baer, 1986). Maldwes, Male, Kurumba; Maldives, Rasdu, Kuramathi (Baer, 1989). Barrow Isl. (Wil son, 1993). Sulawesi, Toigna Isl.; Bira; Pulau Hari (Anon., 1995). LITERATURE, RANGE: QLD - NW Australia (Allen, 1959). SW isl. o f Japan (Kira, 1962). Ryukyu Isl.; Philippines; Dutch New Guinea; Solomon Isl.; New Caledonia; Loyality Isl.; Fiji; Indonesia; Carolines; Marianas (Talmadge, 1963a). Kyushu Isl.; Shikoku Isl.; S Honshu - Kii Peninsula; Indian Ocean (Habe & Kosuge, 1964). PNG (Hinton, 1972). S Japan through N Australia (Abbott & Dance, 1983). WA, Point (Juobba - QLD (Wells & Bryce, 1985). Philippines (Springsteen & Leobrera, 1986). Indonesia (Dharma, 1988). New Caledonia (Salvat et a i, 1988). Indo-West Pacific; WA, Carnarvon - Capricorn and Bunker Group (Wilson, 1993). 235 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. HaUot i s planata Sowerbyv 1882 (Figures 4-43,4-119,4-122) Sri Lanka: (ANSP 50192,3; USNM 89090,1). Thailand: Phuket, Patang (DLG AAB 3 la, I). S Phuket, Ko Racha Noi (DLG AAB 31 b, 5). INDONESIA: Bali, Nusa Dua (LACM 86-164, I + I complete). W Lombok, N o f Sengiggi Beach, Santigi Point (LACM 88-65, 1). New Guinea, Seleo IsL, NW o f reef, Aitape (USNM 616058,1). Kai IsL Tajundu IsL (= Tayandu IsL), E Ree Isl. (USNM ex 746732, 1: complete). Japan: Kyushu, Kagoshima, Kikai, Osumi (DMNH 186025, I; ANSP 86239,2). Okinawa, I km NNW Oku (LACM 77-61,1). Okinawa, Bolo Point Reef (LACM 27416,2). Oki nawa, Imbu Reef (AMNH I4I006,2). Okinawa, Imbu Bay (AMNH 163495, I). Oki nawa, Tancha Reef (SBMNH no # I). Okinawa, Tsuken reef (SBMNH no #, 4). Okinawa, Onna reef (SBMNH no #, 2). philippines: (NMW, 2). Panay (USNM 256594, I). E end o f Negros Isl., Admiraiity Isl. (USNM 488924, I). Apo Isl. (USNM 828711, I: complete). MARIANAS: Agrigan Isl. (USNM 487952, 4). WA: Port Headland (LACM 45846, 3 [?]; LACM 45790, I [?]). SOLOMON IsL: Bougainville (LACM 45845, I). Santa Ana Isl. (AMNH 81889, 1). Choiseul, ChoiseuI Bay (AMNH 81888, I). Guadalcanal (NMW, I). Fiji: (DMNH 130282, 1; AMNH 132598, I). Viti Levu Isl. (LACM 67-93,4; ANSP 50190, I). Viti Levu IsL Suva (LACM 72-147, 2). Viti Levu Isl., Mbengga Isl., Vanga Bay (LACM 71-202,1). Vhi Levu, Makuluva (USNM 531869, I). Yasawa Isl. (USNM 617856,1). Vanua Levu, W o f Ngaratoka Pass (USNM 695049, I). TONGA: Nuku’alofa district, Makaha’a Motu (LACM 72-150, I). Tongatapu (BMNH no # ,2 ). LITERATURE, LOCALITIES: Okinawa; N Coast o f Luzon; Malaita; Fiji, Suva; S Java, Wynkoops Bay (= Pelabuhaniatu, Teluk) (Tahnadge, 1963a). Kagoshima, Amami Isl. (Aral et a i, 1988). 236 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LrrERATURE, RANGE: From Yakushima S (Habe, 1964). Ryukyu IsL; Philippines, Solomon IsL; Fiji; Indonesia (Talmadge, 1963a). Andamanes, Ceylon (Talmadge, 1974). Philippines (Springsteen & Leobrera, 1986). Indonesia (Dharma, 1988). Indian Ocean, Rowley Shoals, Cartier Isl. (NT) (Wilson, 1993). Haiiotis pul cherri ma Gmelm, 1791 (Figures 4-38,4-39,4-44) Kiribati (= Marquesas Isl): Fanning Isl. (= Tabuaeran) (ANSP 185728, 1). Lord Hood Isl. (= Fatu Huku) (ANSP 199729,2; AMNH 19781,11). Tuamotus: Makatea Isl. (ANSP 199337, 12). W Makatea Isl. (DMNH 072616, 1). Rangiroa AtoU, Kai Ora Hotel (DMNH 115875,2). Rangiroa, Ocean Side o f Avatoru (LACM 96-10, 2). Rangiroa, Ayatoru Pass (USNM 782744, 1). Anaa (USNM 775845, 16). Anaa, Takapoto, Niau AtoU (DLG AAB 34a, 4). Takapoto AtoU (DMNH 186717, 3; MNHN no #, 10). Takapotu Isl. (LACM 117636, 1). Takaroa AtoU (DMNH 129590, 3; USNM 617399, 2). Takaroa Isl. (ANSP 185003,10; ANSP 199320,56). Marutea AtoU (DMNH 130747, 1). Nig[e]ri (= Nihiru) (LACM 59325,7). Mahini (DMNH 156569, 1; AMNH 263365, 1). Mahini, King George Isl. (SBMNH 6187,1). Manihi, E end o f Main Motu (USNM 798134,1). Raroia Isl. (ANSP 199095,9; ANSP 250551,2). Raroia Isl., Ngrumaea Isl. (USNM 698332,2; 698529,2). Raroia Isl., Kahongi Isl. (USNM 698576,2). Raroia Isl., Oneroa Isl. (USNM 720725, 2). Raroia Isl., S Opakea Isl. (USNM 722376, 1; USNM 723121,1). Raroia IsU Matira Isl. (USNM 698609, 16). Raroia IsL, Garumaoa Isl. (USNM 722122, 7). Raroia Isl., N Garumaoa Isl. (USNM 720196, 2; USNM 723462,24). Raroia IsL, Gavarivari Isl. (USNM 723276, 1). Raroia, Tekatikati Isl. (USNM 721286, I). Vahitahi (AMNH .107583, 5; USNM 613279, 15). Toau AtoU, Marangnta felet (ANSP 155933,6). Toau Atoll, Ocean Beach (ANSP 156000, 6). Hua- nine (MNHN no #, 4). Nengonengo Isl. (USNM 638322,1). Anuanu Rare, W o f NE 237 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Haiiotis pulcherrima Figure 4-44. Distributfon of the central Pacific (Tuamotus) Haiiotis pulcherrima. Local ities stemming from collection specimens are indicated with a solid circle, those from the literature with a solid square. For data see text. 238 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. IsL (USNM 725320,9). Tatakoto AtoU (USNM 782721,3). Tureia IsL (USNM 638297, 2). Ravaka Isl. (= Raraka) (ANSP 264346,4). Temoe Isl. (= Timoe Isl.) (USNM 671841, 1). Society Isl.: (LACM A.6186.52,4). Tahiti (LACM A.6I86.52a, 2; USNM 835592, 1). Moorea Isl. (ANSP 185142, 2; MNHN no #, 5). GAMBIER ISL.: Motu Tepapuri (USNM 726394, 1). Henderson Isl.: (USNM 732017, 24; BMNH 1913.7.28.7-9, 3). PITCAIRN ISL.: (USNM 789557, 1). Ducie AtoU (USNM 732115, 5). Oeno (USNM 731587,22; USNM 789609,6). LITERATURE, LOCALITIES: Tahiti; Makatea; Aratika; Motutunga; Taenga; Marutea; Fakahina; Amanu; Vahitahi; Pinaki; S Marutea; Taraouroa (= Taraura Roa ?) (Dautzenberg & Bouge, 1933). Raraka, Raroia (Talmadge, 1963a). LITERATURE, RANGE: New Caledonia (not on map); Taumotu, Lord Hoods Isl. (Habe & Kosuge, 1964). Tuamotu Archipelago; Gambler Isl. (Talmadge, 1963a). E Polynesia (Abbott & Dance, 1983). New Caledonia (not on map) (Salvat et a i, 1988). Hiüwti s e ubi gmos a Reeve, 1846 (Figures 4-24,4-136,4-137) NSW: Lord Howe Isl. (BMNH 81.2.20.19-22,4; USNM 851408,2). NSW, Lord Howe Isl., Middle Beach (USNM 7914422,6). Lord Howe IsU Neds Beach (AMS C328700, 1: complete; AMS C.328698, 1: complete). QLD., North-West Isl., Capricorn Group (MNHN no #, 1). Tonga: Nive (=Nhie) (USNM 360748,1). LITERATURE, RANGE: Lord Howe Isl. (Talmadge, 1963a; Hinton, 1978: as howen - sis). E Australia, Lord Howe Isl. (Abbott & Dance, 1983: as hawensis). 239 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Kaliotis squamat a Reeve, 1846 (F lares 4-24,4-63,4-66) nicobar: (BMNH 1829, 1). VIETNAM: Pulo Condor (9“ 20" N 106“ 40" E) (MNHN no #, I) Indonesia: Bali, Nusa Dua (LACM 86-164,1: complete). SW Java, Sukabumi, Queens Bay (AMNH 103486,2). Java, Batavia/Jakarta (MNHN no #, I). Java, Drini Beach, near Jogjakarta (DLG AABOg, 2). QLD.: GBR, off Caims, Arlington Reef (AMNH 178752,2). Near Gladstone (USNM 595682,1). NSW: Sydney (MNHN no #, 2). NT: Taipang/Taypang Bay [=Trepang ?] (AMNH 114549,1; MNHN no #, I). Nhu- lunby, Gove Peninsula (BMNH 2341, I). Yirrkala, Arnhem Land (USNM 602298, 14). Point Charles (ANSP 267918,2). WA: Cape Leveque (USNM 618426, 3). King Sound (LACM A 2777,2; AMNH 111949,1). Broome (LACM 45877,5; AMNH 169622a, 1). Broome, Gantheaume Point (LACM 45979,3; DMNH 124207,1). Broome, Quondong (AMNH 110146, 1). Broome, Roebuck Bay (AMNH 199647, 4; AMNH 182972, 4). Port Headland (LACM 45802,1; LACM A .6548,1). Rosemary Isl. (DLG AAB I3e, 2). Nicol Bay (LACM 45887, 1). Dampier (DLG AAB 13f, 4). Dampier Archipelago (SBMNH no #, 4). Point Samson (NHB 11 239-a, I; NHB II 244-a, 1). Barrow Isl. (USNM 692029,1). Barrow Isl., Airport Beach (USNM 691996,4: complete). Between Cape Dupuy and Cape Malouet (USNM 691874,4: complete; USNM 691827,13: com plete). NE o f Onslow, Montebello Isl., Trimouüle IsL (LACM 95-66,2). NE o f Onslow, Pasage E sL , Great Sanify IsL (LACM 95-63, 8 + 1 complete; LACM 95-64,1 + I with dried body). NE o f Onslow, Weld Isl. (LACM 95-60,2). Yanchey"s [= Yanchep ?] Beach (LACM S .I1I4, 1). Murat, Exmouth. (AMS C.328734, 3: complete). Learmouth, Exmouth G ulf (AMS C328735,2: complete). S. o f Exmouth townsite. Exmouth Gulf (AMS C 328738,3: complete), Bundegi Ree^ Exmouth (AMS C328732,3: complete). O ff Camarvan (AMS C.328740,2: complete). Port Gregory, N o f Geraldton (AMS C.328733,15: complete). O ff Perth (DMNH 012213,3). Swan River (BMNH no #, 5). 240 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Shark Bay ^M N H 2341,1). Vlaming Head (AMNH 96444,3). Cape Preston, Regnard Bay (SBMNH no #, 1). Cape Preston (AMNH 220131, I). SE o f False Cape Bossut, Lagrange Bay (ANSP 233326, 1). LITERATURE, LOCALITIES: Java, Karang Hawoe; Bali, beach between Boegboeg and Boeitan (Adam & Leloup, 1938). Sunda Straits; Prinz Eiland (= P. Panaitan); S coast o f Java, Wynkoops Bay (= Pelabuhanratu, Teluk) (Talmadge, 1963a). Ningaloo Marine Park (Wilson, 1993). LITERATURE, RANGE: W and NW Australia; Indonesia (Talmdge, 1963a). Shark Bay - NT (Wilson, 1993; Wells & Bryce, 1985). Bali, Lesser Sunda Isl. (Dharma, 1988). Ha Uot i s vari a Lmnaeus, 1758 (F^ures 4-34,4-37,4-43,4-163,4-164) Tanzania: Zanzibar, Pwani Mchangani (ANSP 399945, 6: doubtful). Mauritius: (DMNH 011016, 3; DMNH 129904, 3: doubtfiil). Maldive Isl.: Rasdu Atoll (ANSP 253542, 1: doubtful). India: Kilikkarai (DLG 15], 1). Hare Isl. (= Musal Trvu), 6 miles SW Mandapam (ANSP 301461, 1). Coromandel Coast (AMNH 220252,3). Shingle Isl., E of Krusadai Isl., SE Gulf o f Mannar (ANSP 301249,5). Waltair (USNM 622097, 1: complete). Sri Lanka: (BMNH 1838, 6; AMNH 48366, 1). Tricomale (DMNH 069887,3; BMNH 1838, 12). Tricomale, E shore Fort Frederick (ANSP 211220,10). Trincomale, Elisabeth PL (ANSP 210955,1). NUaveli, Pigeon Isl. (LACM 84-7,6). Nayinativu Isl. (ANSP 225034, 3). Weligama Bay (ANSP 210842,1). Andamanes: (DMNH 010980,3; BMNH 73.3.10.17,2). Port Blair (BMNH 1838.8; BMNH 1838, 3). thadand: PhukeL Airoport Beach (ANSP 286646,2). PhukeL Laam Seng, 1 mile S o f Laam Son (ANSP 285840,2). Vietnam: Nha Trang (MNHN no #, 1). China: (LACM 45867,1). Hong Kong (BMNH no #, I: complete). Hong Kong, E sick High IsU Port Shelter (ANSP 222679,14). Taiwan: (DMNH 118533,72; LACM 45854,3). Taipei Co, 241 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. E o f CM-Iung (= Keelung) (LACM 88-80, 3). W Lan Yü (= Hung-tou Hsu) (USNM 653768, 3). Japan: Honshu, Chiba, Yokahama (USNM 43067; USNM 43067,1). Hon shu, Kanagawa, O-shima, Amanago (ANSP 141829,1). Honshu, Kanagawa, Oshima, Osumi (USNM 343823,3). Honshu, Kanagawa, Oshima, Osumi, Kikai (USNM 304309, 4). Honshu, Wakayama, Kushimoto, Shiono-misaki (AMNH 239007, 4). Honshu, Wakayama, Shirahama, Kii (ANSP 252612, 3). Shikoku, Kochi, Tosa , Kashiwajima (ANSP 86241,1). Shikoku, Kochi, Tosa, Kashizima (USNM 273362,4; USNM 343832, 3). Kyushu, Kagoshima, Amami, O-shima (AMNH 241419, 5; ANSP uncataloged, 1). Ryukyu Isl. (= Loo Choo) (NMW, 1). Oku (LACM 77-61,1:87 mm). Mukojima, Bonin Isl. (= Ogasawara-shoto) (ANSP 252592,2; USNM 343833,3). Bonin Isl., Chichi Jima (USNM 621934, I; USNM 621866, 1). Okinawa, Taguchi (DMNH 140614,1). Oki nawa, SSW Kadena (LACM 77-55,1). Okinawa, Itoman Reef (LACM 72045,4; LACM 72156.2). Okinawa, Imho Beach (LACM 28830, 3). Okinawa, Metasaki Reef (LACM 29332,4). Okinawa, Oima Village, Horseshoe Cliffs (LACM 79-75, 1). Okinawa, 8 km E o f Kawata (LACM 77-79,2). Okinawa, Kadena Reef (AMNH 141067, 3). Okinawa, Sobe (AMNH 102963,2). Okinawa, hnbu Bay (AMNH 156010,5; AMNH 156010,5). Okinawa, Sobe (AMNH 102963a, 1). Okinawa, Madimato (AMNH 187058,7). Oki nawa, 5 miles N o f Kin (AMNH Jackson 1992 A 9873,7). Okmawa, Fuchaku (ANSP 302807.2). Okinawa, White beach, 1.5 miles S ofHenne (ANSP 302730,1). Okinawa, Onna reef (SBMNH no #, 4). Okmawa, Yakena reef (SBMNH no #, 2; SBMNH no #, 16). Okinawa, Zampa-misaki (SBMNH no #, 1). Okinawa, off Civil Air Terminal, Naha (SBMNH no #, I). Okinawa, Machinato Reef (SBMNH no #, 1). Okinawa, Kiushi IsL Shuri (USNM 362834, 1). Okinawa, Shima, Odonari (USNM 488158, 1). Okinawa, Shanawan Bay, Shioya (USNM 489109a, 1). Philippines: Luzon, Cape Engaho (DLG 15b, I). Luzon, Batangas, Verde Isl. (DMNH 049712,3). Luzon, Batangas, Calatagan 242 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (DMNH 004402, 8). Luzon, Batangas, between Culebra and Malajibomanoc IsL (LACM 84-162,2). Luzon, Bataan Province, Lusong Cove (LACM 74741,24). Luzon, Bataan Province, Luson (LACM 152163,2). Luzon, Bataan Province, S Corregidor Isl. (DMNH 003200, 1; LACM 74739,2). Luzon, SE Bataan, Cabcaben (ANSP 231301, 1). Luzon, Manila (LACM A.2023,5). Luzon, Manila Bay, Caballo Isl. (ANSP 229671, 2). Luzon, W tip of Corregidor Isl. (ANSP 230142, 10). Luzon, Sorsogon, Magallanes (LACM A.1930, I; ANSP 223923,6). Luzon, Sorsogon, Matnog (LACM 74735, 2). Luzon, Sorsogon, Barrio Lupi, Prieto Diaz (ANSP 224201,3). Luzon, Sorsogon, Albay Gulf, Tabaco Bay (LACM 23549,3). Luzon, Zambales, Iba (ANSP 229242,13). Luzon, Maricaban Isl. (USNM 245506, II; USNM 245511, 1). Luzon, Tayabas Province, Din galan Bay, N o f Otpegon Point (SBMNH 12243, 8). E Luzon, Butauanan Isl. (USNM 245501, 1). Luzon, W Palaui Isl. (USNM 245492, 2). Burias, Port Busin (USNM 245509,1). Minodro (DMNH 011020,2; BMNH, 1: complete). Mindoro, Lubang, Isl., Hulagaan Point (LACM 74742, 13). Minodro, Cabra Isl. (DMNH 049713, 7; LACM 45803, 1). Mindoro, Calipan (LACM 110395,5; SBMNH 21572,2). Mmdoro, Calapan (ANSP 242231, 7; ANSP 249501, 17). Mmdoro, Calapan, Hoecapilato (ANSP 399936, I). Mmdoro, Dmnai Point (AMNH 80448, 5; AMNH 80448a, 2). Mindoro, Ambil Isl. (AMNH 220188,6). Mindoro, Naujan (USNM 30337, 1). Mmdoro, Lubang, Port THiq (USNM 245541,4). Catanduanes, Gigmoto (DMNH 002110,16; AMNH 220254,3). Catanduanes, Bugas Point (LACM 74722,1 with dried body). Samar (DLG I5u, 1). Samar, Batag Isl. (USNM 245504, I). Samar, Borongon (DMNH 001906, 7; ANSP 223339, 15). Samar, Catbalogan (USNM 245526, 2; USNM 245514, 1). Palawan, Cafamian Group, Culion E sL (DMNH 140315,1; LACM 74736,1). Palawan, Calamian (jroup, Culion, Leper Colony (LACM 91279,2; AMNH 197954,1). Palawan, Calamian Group, Busuanga, Port Galton/Balton (USNM 245519,1). Palawan, Calamian Group, 243 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Busuanga IsL, Dînaran Isl. (LACM 74703, l). Palawan, Cuyo Isl. (ANSP 223512,1). Palawan, Observatory Isl., Malubutglubut Isl. (USNM 245491,4). Palawan, Ulugan Bay (USNM 245499, I). NW Palawan, Ulugan Bay (AMNH 132220, I). Palawan, Dalanganem Isl. (USNM 345393,4; USNM 245495,1). Palawan, Balabac Isl. (SBMNH no #, 5). Panay, Borokay Isl. (DMNH 122611, 1). Bohol, Panglao (DLG I5g, 3). Min danao, Camiguin Isl., Port Pio (USNM 245490, 2). Mindanao, Zamboanga (DMNH 061283, 1; AMNH 220255, 3). Mindanao, Santa Cruz Isl., o ff Zamboanga (USNM 245549, I). Mindanao, Davao (DMNH 136928, 2; DMNH 118538, 2). Mindanao, Davao, Pongit, Hijo (ANSP 133357, 1). Mindanao, Near Davao (ANSP 118912,10). Mindanao, Balut Isl., Darangani Isl. (USNM 245557,3). Mindanao, Lianga Bay (USNM 245558, I). Mindanao, Polloc (USNM 245516, 5). Mindanao, Panabutan (USNM 245542, 5; USNM 245508,2). Sulu Archipelago, Siasi Isl. (DMNH 028211 ,4; AMNH 279267, I). Sulu Archipelago, outside reef and barrier reef S o f Muso and Siasi Isl. (ANSP 318245, 39). Sulu Archipelago, Sanga Sanga Isl. (DMNH 003939, 4). Sulu Archipelago, SW Sanga Sanga Isl., Bongao Channel (DMNH 003018,6; ANSP 230772, 50). Sulu Archipelago, Jolo Isl., off Muobo Beach (ANSP 318344, 1). Sulu Archipel ago, Tabawan Isl. (ANSP 318295, 1). Sulu Archipelago, Jolo Isl. (USNM 245537, 1). Sulu Archipelago, Tawi Tawi group, Papatag Isl. (USNM 245512, 1). Cebu (LACM 34588, 1; ANSP 50185,2). Cebu, Cebu City (SBMNH no #, I). Cebu, Bantayan Isl. (LACM 45924,1). (Quezon, Polillo Group, Cabaloa (LACM 74721,1). Quezon, Polillo Group, E Ikon Isl. (LACM 74738, II). Quezon, Baler (AMNH 103709, 2). Masbate Province, U cao IsL, Taclagan Bay (LACM 74726, I). Masbate, Wumusung Point (USNM 245560, I). Capul (BMNH 1829,3). 2 km S o f Olango, E o f Cebu, Salpa (ANSP 230457,1). Negros OrientaL Guihulhga (SBMNH 28937,2). Guam: Alupat IsL Agana Bay, Guam Isl. (ANSP 197355,1). Palau (= Belau): SE shore Aurapushelaru 244 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Isl., N ^akal Harbor (ANSP 202501, 1). Airai, Babelthuap, reef oposite, 07* 22’ N 134“ 34’ E (ANSP 202687, 1). Federate States o f Micronesia: Round rock, Helen Channel, Helen Reef (ANSP 399941, 1). Indonesia: Ambon (DMNH 009556,2; NHB 5439d, 2). Ambon, Saparoea (MNHN no #, I). E o f Ambon, Haruku Strait, Pombo Isl. (USNM 746572,1: complete). N Sulawesi, off Manado, S Bunaken and Siladen Isl. (LACM 88- 5 6 ,1: complete). Java, off Jakarta Raya, Pulau-Pulau Seribu Isl., Pelangi and Putri Isl. (LACM 86-163, 1). Java, Batavia/Jakarta (MNHN no #, I). Java, Sukabumi (AMNH 220130, I). Timor (AMNH 221449, I). Sumatra, Banda (ANSP 281223,2). Lompok, Senggigi Beach, Mangait Point (SBMNH 5617, 2). SW Suinatra, N o f Sipora, Pulau Siburu (USNM 654889A I; USNM 654703A, 2). SW Sumatra, Mentawai Isl., Pulau Stupei (USNM 655162,2). Off Sumatra, Pulau Batu Group (USNM 654480A, 3). Flo res (NHB 4394.b, 6). Celebes Isl. (USNM 617400,3). Irian Jaya, Biak Isl., Biak (LACM 88-43,1). Wan Jaya, Biak Isl., W o f Biak (LACM 88-52,2). Irian Jaya, N Japen, Ambai Isl., Maroepi (ANSP 208567,1). Wan Java, Japen, W Samberbaba, Cape Tekopi (ANSP 205122.1). Wan Jaya, Pai Isl., Mios Woendi AtoU, Reef E o f Padaio Isl. (ANSP 205067, I). Wan Jaya, Noesi IsL Mios Woendi AtoU, Reef E o f Padaido Isl. (ANSP 205039,1). Wan Jaya, Abroeki Isle, Maransabadi, Isl., Aoeri Isl., Geelvmk Bay (ANSP 208636,2; ANSP 207547, 2). W Wan Jaya, n% r Sorong (AMNH 125348, 6). Wan Jaya, Sorong (NMBE 1507382, I). Aroe and Tenimber Isl. (NHB 5439-b, 3). N Am, N Toba Isl. (USNM 755376,1). Tanimbar, Jamdena Straits, 2 mUes N o f Tg Nuan (USNM 747571, I). Tanimbar, Jamdena Straits, W Mitak Isl. (USNM 747546,1). Kai Isl., N o f Du Rowa, N o f Nuhu Rowa (USNM 746993,1). Kai Isl., Tajundu Isl., E Ree Isl. (USNM 746732, I: complete). Kai IsL, TajtmduIsl., Warantneu (USNM 746772, 1). Ceram Isl. (USNM 837284.1). Halamahera Isl., Tbbelo District, Bobi (USNM 837126,2). Hahnahera Isl., JaUoIo District, Pasir Putih (USNM 837195, 18). Malaysia: Peninsular, Perhentian Isl. 245 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (BMNH 2264,1; BMNH 2264,1). O ff Mersing coast, Pulau Babi Besar (ANSP 319229, 4). Borneo, Sabah (DMNH 049710,2; AMNH 175467,1). Borneo, Sabah, Sapi (DMNH 201327,1), Borneo, Sabah, Jesselton, Sapi IsL (USNM 658276, I; USNM 658277, 4). Borneo, Sabah, Jesselton (ANSP 274999, 6 ; ANSP 275051, 7). Borneo, Sabah, Jessel ton, Sapangar Isl. (NMW). Borneo, Sabah, Jesselton, Gaya Isl. (USNM 658287, 6). Borneo, Sabah, Jesselton, Sabang (USNM 658293,6). Borneo, Sabah, Sepoma (BMNH 2277, 2). Borneo, Sabah, Malawali Isl. (NMW; USNM 887464, I). Borneo, Sabah, Malawali Isl., Permuan (NMW, I). Borneo, Kudat, Bak Bak (USNM 632352,3; USNM 633018, 1). N Borneo, Kudat (AMNH 220133,1). Borneo, Marudu Bay (ANSP 255847, I). Borneo, 6 miles N o f Kudata, W Marudu Bay (ANSP 255710,2). N Borneo, Mandi Darrah Isl. (AMNH 106759,3). S Gomumu Isl., S o f Obi Isl. (USNM 746446,2: com plete). Manipa Isl., N Tg Hapale (USNM 746475, 1: complete). WA: 35 miles N o f Broome, James Prince Point (ANSP 233549, 15). 17 miles N o f Broome, 0.5 mile S of WiUy Creek (ANSP 233507, 3). Broome (AMNH 272135, 2; AMNH 279269, 1). Broome, Port (BMNH; LACM 72012,4). Broome, Entrance Point (ANSP 233014,19). Broome, Cable Beach (BMNH 2341, 2). I mile NE o f Broome, Gantheaume Point (ANSP 232692,21). Broome, 2 miles SW o f Jetty (ANSP 233107, 2). Broome, Rid del’s Beach (SBMNH 6138, 5). Port Headland (LACM 45861, 3; LACM 45801,2). Perth (DMNH 0 I2 2 I4 ,3). Perth, Swan River ^M N H no #, I). Onslow, 2 miles W of Onslow Jetty, Ward R eef (ANSP 267421, 1). 130 miles from Onslow, Cape Preston (AMNH 163394, I). Barrow Isl., S Flacourt Bay (USNM 69I88I, I: complete). Dampter Archipelago (SBMNH no #, 2). ()uobba Pomt, 40 miles N o f Carnarvon (ANSP 238529,4). Monte Bello Isl. (BMNH no #, I). I mile SE o f False Cape Bossut, La Grange Bay (ANSP 233333,3). Cygnet Head (AMNH III9 5 0 ,2). Point Samson (NHB 5439e, I). 4 miles N o f Red Bluff (ANSP 268029,1). Christmas Isl. (BMNH no #, I). 246 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 80 Mile Beach (NHB 5439-f, I). Carmarvan, Cambridge Bay (BMNH I913.7J.I95, I). NT: Darwin (LACM 45850, 1; LACM 45880,2). Port Darwin (ANSP 193247,5). Point Charles (ANSP 267917,5). Port Essington (BMNH 1829, 3). NE end of Gulf of Carpenteria, Eyiandt (USNM 602260, I). QLD.: G ulf o f Carpenteria, Wallaby Isl. (SBMNH 31463, 5). Torres straits (BMNH; AMNH 285411,1). Torres Straits, Thurs day Isl., W Snodon (BMNH no #, 1). Lizard Isl., Casurina Beach (USNM 783328, 1). SE Lizard Isl. (LACM 79-55, 1; LACM 79-54, 1). SW Lizard Isl. (LACM 79-53, 3: complete). Hook Isl. via Prosperine (SBMNH 228, 6). Port Douglas (SBMNH 203,3). Low Isl. (SBMNH 226, 2; BMNH, 1: complete). Caims (DMNH 031897,4; AMNH 272156, 2). Caims, Hasting’s Reef (DMNH 179654, 3). GBR, o ff Caims, Arlington Reef (AMNH 178751,1; LACM 45947,8). GBR, Fitzroy Lagoon (AMNH 144302,6). Townsville (LACM 45800,2). Off Townsville, Davies Reef (BMNH 2341,1). Orpheus Isl., Fig Tree Bay (SBMNH 6137,2). Townsville, Palm Group, Esk Isl. (SBMNH 227, 3; SBMNH 2639, 1). Whitstmday Group, Black Isl. (ANSP uncataloged, 2). Whitsun day Group, Hook Isl., Mackerel Bay (BMNH 2341, 3). Whitsunday Group, Hook Isl., Langord Reef (AMNH 104533,2). Whitsunday Group, S MoUe Isl., Paddle Bay (LACM 29660,2). Whitsunday Group, Glochester Isl. (AMNH 220129,4). GBR, Lmdeman Isl. (LACM45943,12; ANSP 138604,1). Proserpine (AMNH 272149,1). Proserpine, Shute Harbour (LACM 45799,2). Bowen (AMNH 189533, 12). Mackay (SBMNH no #, 1). Yeppoon (AMNH 215146,2; SBMNH 213,3). Bamborough Isl. (AMNH Jackson 1992 A9873,2). Keppel Isl. (AMNH 272155,3; AMNH 163124,2: complete). Capricorn Isl. (AMNH 85706,1). Capricorn Gourp, North West Isl. (ANSP 236332,2; ANSP uncata loged, 3). Capricorn Group, One Tree Isl. (ANSP 236314,2). Capricom Group, Tryon Isl. (USNM 617151,4). Capricom Group, Wreck Isl. (SBMNH WW2332, 2). GBR, Holboume Isl. (ANSP 138767,2). Curtis Isl. (AMNH 80432,2). Hinchinbrook Isl., 247 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Palm IsL Groiq), Orpheus Isl. (AMNH 178912,2). Hinchinbrook, Beaver Reef (AMNH 121864,2; AMNH Edison A9853,2). Brook IsL (ANSP 134521,6). Heron Isl. (DMNH 094026,1; DMNH 186028,4). GBR, Pandora reef (ANSP 134667, 1). Queen’s Beach (LACM 45906,3). PNG: Kapa Kapa (DMNH 047217, 2). Madang (AMNH 215144, 2). Madang province. Astrolabe Bay, Nagada, Tab Isl. (LACM 80-24). Port Moresby Bay (ANSP 219621, 3; ANSP 247692, 2). Port Moresby (ANSP 242539, 2). New Britain, Rabaul Harbor (ANSP 274801, 6). New Brittain, Matapit Isl., 5 miles &om Rabaul (ANSP 304733, 3). Admiraiity Isl., Koruniat Isl., near Manus (ANSP 181932, 3). Trobriand Isl. (AMNH 215145, 1). Goodenough Isl. (AMNH 124564, 1). Conflict Group (BMNH). Louisade Archipelago (BMNH no #, 4). Solomon Isl.: Guadalcanal, NW (DMNH 053167,3). Guadalcanal, Honiara (DMNH 059188, 3; BMNH no #, 3). Guadalcanal, Honiara, Point Cruz (SBMNH 1997, 5). Guadalcanal, Marau Sound (AMNH 183045, 1). Guadalcanal, Marau Sound, S &/brapa Isl. (LACM 78-68.15, 1). Guadalcanal, Marau Sound, Tavanipupu Isl. (SBMNH 25669,3; SBMNH 3503, 14). Guadalcanal, Merow (AMNH 82286, 1). Gudalcanal, Lunga (AMNH 220186, 3). Tulaghi IsL (AMNH 81883, 1; AMNH 82253,1). Bunana (= Bunala) (LACM 45842,2; BMNH 2144,1). Malaita (LACM 45902,2). Malaita, Su’u (AMNH 82130,41; AMNH 82130, >50). Malaita, Ataa (ANSP 289619,2; AMNH 118202, 3). N Malaita (DMNH 050854,1). N Malaita, Auki (DMNH 046851,2; BMNH, 3: complete). Shortland Isl. (ANSP 310059,2). Shortland Isl., Bougainville (LACM 45864, 1; LACM 45859,2). Shortland Isl., Bougainville, Buin (LACZM 45873,4; ANSP 267125,4). Shortland Isl., Bougainvffle, Kieta (AMNH 82159,1; AMNH 82166,5). Shortland Isl., Faisi (AMNH 82124,3). Choiseul IsL, Bambatani (AMNH 82235,3; AMNH 82244,1). Choiseul Isl., Luti (AMNH 82254,7). Choiseul IsL, Choiseul Bay (AMNH 81887,2; AMNH 81886, 4). Santa Ana M . (AMNH 81877,1). San Cristoval (LACM 45872,1; ANSP 167545, 248 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1). New Georgia (AMNH 220185, 4). New Georgia, Mimda (SBMNH 1602, 1). Santa Cruz Group, Reef IsL, Pigeon Isl. (SBMNH no #, 1). New Caledonia: (AMNH 220216, 2). Koumac (MNHN sta. 1303, 1). Vanuatu: Efate (ANS? 222846, 1). Efate, Undine Bay (DMNH 053602, 1, 1: complete; DMNH 052472, 3). Efate, Mele Bay (BMNH 2341,2; SBMNH 25677,3). Efate, N Mele Bay (SBMNH 0091,2). Efate, Mele Bay, N o f Port Vila (BMNH 2341,5; SBMNH 3173,2). Efate, Vila, Le Lagoon (SBMNH 2625, 11:2 dried bodies; SBMNH 40081, 7: 4 dried bodies). Efate, off Port Havaimah (SBMNH no #, 4; SBMNH 25963, 3). Efate, Baie d’Erakor (SBMNH 25645,3). Efate, N Moso Isl., Shortland Isl. (BMNH 2341, 1). Efate Isl., E Ennis Point, Erakor Reef (LACM 77-37, 2). Lamap, MaHcoIo Isl. (ANSP 243153, 1). Fiji: (BMNH; LACM A.1463, 1). Vanua Levu, Savusavu Bay (AMNH 141409,1). Tonga, Vava’u (DLG 15n, 2). Tongatapu (SBMNH no#, 1: with dried body). Tongatapu, Havannah IsL, Nuku’alofa (SBMNH no #, 5). Cook Isl.: Aitutaki (SBMNH 2732,2). LITERATURE, LOCALITIES: Lompok, Labuan Pandan; BCarakelang (= Karakelong) IsU Bee; Waigeo, Wunoh Bay; W Kur [= KoorJ Isl., Kilsuin; S Rot[t]i Isl., Buka Bay (Schepman, 1909). MoIIuccas, Halmahera; Bachan; Tidore (= Soasiu); Misol; SB o f Ceram, Kur [= Koorj IsL; N o f Timor, Kisser [= KlsarJ; Solor; Flores, Larentuka; Java; S Java, Palabuan [= Palabuhanratu ?]; N Celebes; N Togean Isl.; Sangir Isl.; Taiaut (= Talaud) IsL (Oostingh, 1925). Bangaan Isl. ^redale, 1929). Bali, beach between Boeg- boeg and Boeitan; Celebes, G ulf o f Paloe (= Pain); Between Banda Neira and Goeno- eng Api; Banda N eka (= Bandanaira); N o f Misoôl, Weim Isl.; Port o f Soembawa (= Sumbawa); Pisang Isl.; Banda Archipelago (= Bandanaira), between Lonto Isl. and Goe- noengApi (Adam& Leloup, 1938). Gulf o f Carpenteria, Grocte Eyiandt (Cotton, 1943). India, Pamban area (Satyamurti, 1952). L ^ e ; Kwajalein; Schouten Isl.; Trobriand Isl.; Bougainville; Lkh; Fiji, Suva (Talmadge, 1963a). Capricorn Group; Keppel Isl.; Bowen; 249 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Townsville; Magnetic Isl.; Caims; Darwin; Broome; Biak (Talmadge, 1974). NW Aus tralia, Cockatoo Isl. (Anon., 1975). Norfolk Isl. (Wilson, 1993). LITERATURE, RANGE: N Australia (Allen, 1959). S Honshu and S (Kura, 1962). Ryukyu; Philippines; Marianas; Caroline Isl.; Marshall isl.; Dutch New Guinea; Solomon Isl.; Papua New Guinea; Loyality Isl.; Fiji; Friendly Isl. (Talmadge, 1963a). N, E and W Australia; Indonesia; Philippines; China; Ryukyu; Amami Isl.; Kyushu Isl.; Shikoku Isl.; S Honshu - Kii Peninsula; Malaya; Nicobar; Ceylon; India; Mauritius; Red Sea; E coast o f Africa (Habe & Kosuge, 1964). PNG (Hinton, 1972). Point Quobba, WA - QLD (Wells & Bryce, 1985). Indonesia (Dharma, 1988). Indo-West Pacific; Abrol- hos,WA- S QLD (Wilson, 1993). Australian Endemic Species (Figures 4-45-67,4-131,4-132,4-134,4-135,4-183- 192) Haüot i s brazKfi Angas, 1869 (F%ures 4-45,4-48,4-67) NSW: The Entrance (AMNH 220152,2). Solitary Isl., Coffe Iferbcr (SBMNH 6642, I). Split Solitary Isl. (AMS C.149015,1: complete; AMS CJ28770, 1: complete). Bro ken Bay (USNM 162181,2). Sydney, Vaucluse Point (ANSP 50231, 1; USNM 515804, I). Sydney, Collaroy Beach (LACM 45960,3; LACM 45964,4). Port Jackson (DMNH 010965, 1). Port Jackson, ofiT Shark Bay (BMNH no #, I). Port Jackson, Bottles and Glass Rock (ANSP 50227,1). CronuUa (LACM45875,2). Shellharfaor (AMNH 179676, 4). Jervfe Bay (SBMNH 52707,1). O ff Currarong, near J«vis Bay (AMS C327855,1: complete). O ff Bowen Isl., Jervis Bay (AMS C.328703, I: complete). Whale Point, Ctockhaven Bight, 3 5 ® 02’ S 150“ 53’ E (AMS C J28702,1: complete). Bass Pomt, SE of Shell Harbour, 3 4 ® 36’ S 1 5 0 ® 35 E (WAM C328704,1: complete). 250 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-45-60. Shells o f abalone endemic to Australia I. Figures 4-45,48. H aliotis brazieri Angas, 1869. DLG AAB 43a. Cofifs Harbor, Solitary Island. 30 mm. Figures 4- 46-49. H aliotis hargavesi Cox, 1869. DLG AAB 42b. Cofifs Harbor; Solitary Island. 31 mm. Figures 4-47,50. H aliotis coccoradiata Reeve, 1846. BMNH type lo t Type local ity : “?*. 57 mm. This is the largest o f three species and not the one marked “type”. Fig ures 4-51-52. H aliotis cyclobates Péron & Lesueur, 1816. DLG AAB 35a. Port Lincoln, South Australia. 57 mm. Figures 4-53-54. H aliotis elegans Philippi, 1844. DLG AAB 24b. Albany, Western Australia. 72 mm. Figures 4-55, 58. H aliotis s. scalaris Leach, 1814. DLG AAB 20a. Western Australia. 6 6 mm. Figures 4-56, 59. H aliotis scalaris emmae Reeve, 1846. DLG AAB 06b. South Australia. 58 mm. Figures 4-57,60. Halio - tis laevigata Donovan, 1898. LACM H-791. Phillip Isl., Victoria. Earl Hufhnan coll. 115 mm. 251 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 252 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE, LOCALITIES: NSW, Lake Macquarie; Port Jackson, Watson’s Bay (Angas, 1871). SA, Amo Bay; QLD., Caloundra (Cotton, 1943). Port Jackson (Anon., 1975). Sydney (Wilson, 1993). LITERATURE, RANGE: S QLD. - Jervis Bay (Wilson, 1993). H aliotis coccoradiata Reeve, 1846 (Figures 4-47,4-50,4-67,4-186) QLD.: Lizard Isl. (USNM 766895, I: complete). Moreton Bay (DLG AAB 23e, I). Brisbane, Balmoral Beach (AMNH 220156, 1). Stanage Bay (DLG AAB 23f. I). Bowen, Brampton Reef (AMNH 220154,4). Proserpine (AMNH ex 272149,1). Moss- man (AMNH 96443,2). Cumberland Group, on reefs (BMNH 1829, 3). GBR, Linde- man Isl. (LACM 45942, > 50). NSW: Montague IsU off Narooma (AMS C.328732, I: complete). Ballina (LACM S.2926,1). The Entrance (AMNH 220155,3). 25 miles S o f Newcastle (SBMNH no #, 3). S shore Broken Bay near The Entrance (ANSP 137975, 8). Solitary Isl., Coffs Harbor (DLG AAB 23d, 1). 15 km N o f Forster, Black Head (BMNH 2341, 10). 30 km S o f Forster, Seal Rocks (BMNH 2341, I). Port Jackson (LACM 45988, 2; LACM 45974,3). Manly Beach (USNM 842776, 5). Sydney, Long Bay (DMNH 118535,2). Sydney, Long Reef (AMNH 183923, 13; AMNH 220158, 8). Sydney, Long Reef, Collaroy (BMNH Smythe collection. I). Sydney, Long Beach, Col laroy Beach (LACM 45879,2; ANSP 228191,3). Sydney, Botany Bay (AMNH 279266, I). Sydney, Botany Bay, Solander Monument (ANSP 399935,7). Sydney, Botany Bay, Bare IsL (SBMNH 222,2). Sydney, Kumell (AMNH 179794,4; ANSP uncataloged, 2). Sydn^, Long Reef (LACM 79-58,3). Syrhiey, Middle Harbor (LACM 79-59,1). Syd- n ^ Harbor, Bottle and Glass Reef OLACM A 6548,1). BoIIaroy, Long Reef (LACM H- 3338,2). Jervis Bay, Green Point (lA C M 80-63, I). Broulee-Batesman Bay (NHB II 250-a, 2). South. Durras, Batesman B ^ (NHB 11250-b, 1). McMasters Beach (AMNH 253 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 220157,6). Angourie (ANSP 2I96I5, I). Rous County, Tweed Heads (ANSP 138447, 2). VIC: Melbourne, Port Philip Bay, Ricket Point (LACM 79-49, 1). TAS: Hobart (MNHN no #, I). LITERATURE, LOCALITIES: VIC, TuIIaburga Isl. (Cotton, 1943). Botany Bay (Wil son, 1993). LITERATURE, RANGE: NSW - E VIC (Allen, 1959; Wilson, 1993). SE Australia (Hinton, 1978; Abbott & Dance, 1983). Halioti s q^cl obat es Pérou & Lesueur, 1816 (Figure 4-Sl, 4^52,4-67,4-185) QLD.: Capricorn Group (LACM 45891, I). SA: Venus Bay (NHB 10 906-b, 1). Spencer GuIL Moonta Bay (DMNH ex 008703,1; LACM 45952). Spencer Gulf (LACM A.2777a, 1). Encounter Bay (AMNH 80425,4). Port Willunga (AMNH 220168,4). NormanviUe (AMNH 220167,5). Edithburg (SBMNH 6161,2; SBMNH 6141,1). Port Gawler (SBMNH no #, 2). Adelaide (DMNH 049720, 6 ; LACM A.2777,2). Adelaide, Glenelg (DMNH 009567, 2; LACM 34580, 1). Adelaide, Brighton (LACM A.52, 5). Adelaide, Henley Beach (DMNH 179565, 2). Adelaide, Outer Harbor, Section Bank (LACM 45987, 1). Adelaide, Outer Harbour (DMNH 049720, 6 ; DMNH 136901, 3). Adelaide, Seacliff Reef (AMNH 252365, 1). Adelaide, Semaphore Beach (NHB 10.906a, 2). Adelaide, West Beach (AMNH 121848,2). Adelaide, Largs Bay (SBMNH no #, 3; SBMNH 185,2). Port Adelaide (LACM 45889, 1). St. Vincent’s Gulf (LACM 45939,2; AMNH 220166, 1). St. Vincent Gulf, Marina (ANSP 219620,1). Point Vin cent (LACM S.584,4; LACM 22187,3). Wallaroo (ANSP uncataloged, 1). Tumby Bay (LACM 28820,1). Port Lincoln (ANSP-193224, 1; AMNH 252700,3). Kangaroo Isl. (AMNH 11945, 3; AMNH 141077,1). Eyre Peninsula, Peake Bay (AMNH Edison 254 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. A9853, 3). TAS: Rocky Bay (NHB 11 225-b, 1). WA: Perth. (LACM 45937,1). Swan River (AMNH 23099,1; USNM 16958, 1). LITERATURE, LOCALITIES: Port Philip (Cotton & Godfrey, 1933). SA, Corny Point (Cotton, 1943). VIC, Portsea (Macpherson & Gabriel, 1962). Tiapara Isl.; West Isl. (Shepherd & Laws, 1974). Garden Isl. (Wilson, 1993). LITERATURE, RANGE: Streaky Bay - Spencer Gulf (Cotton & Godfrey, 1933). SA and VIC (Allen, 1959). Esperance - South East Cape (Shepherd, 1973). Esperance - SA (Wells & Bryce, 1985). Esperance - W VIC (Ludbrook & Gowlett-Holmes, 1989). Recherche Archipelago - SA (Wilson, 1993). Haüotis el egans Philippi, 1844 (Figures 4-53,4-54,4-67,4-192) NSW: Sydney, Mosman River (MNHN no #, 1). SA: Adelaide (BMNH 2351,1). WA: Ledge Paint (AMNH 220181,1). Albany (DLG AAB 24b, 2). Jurien Bay (AMNH Jack son 1992 A9873,4). Augusta (WAM S1003, 1: complete; WAM S1004, 1: complete). Green Isl. (AMNH 276881, 1; AMNH 220182, 1). Margaret River (LACM 45881, 3). Cowaramup Bay (AMNH 120839,4). Rockingham, P t Peron (LACM 45965, 1). Swan River (BMNH no #, 1). Perth (SBMNH 5142, 1). Freemantle (USNM 877162, 2; SBMNH no #, 3). Rottnest Isl. (LACM 117638, 1; LACM 87-91,1). Horrocks Beach, via North Hampton (SBMNH 212, 2). Shark Bay (SBMNH W4567, 1). Broome (AMNH 169622,2). LTTERArURE, RANGE: Jurien Bay - Esperance (Wells & Bryce, 1985). Geraldton - Esperance (Wilson, 1993). 255 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Haüot i s hargravesi Cox, 1869 figures 4-46,4-49,4-67,4-189) NSW: La Perouse, Botany Bay (ANSP 399934, I). O ff Currarong, near Jervis Bay (AMS C J27855,1: complete). Solitary IsL, Cofife Harbor (SBMNH no #, 10). W. side, off Cofife Harbour (AMS C.327856, I: complete; AMS C J27857, 1 body). Split Soli tary Isl. (AMS C.149015, 1: complete). Cook IsL, Tweed River Head (AMS C.327854, I: complete). WA: Quaboba (= Quobba) (BMNH no #, 2). LITERATURE, LOCALITIES: NSW, Broken Bay Heads (Angas, 1871). Jervis Bay (Wilson, 1993). LITERATURE, RANGE: SE Australia (Hinton; 1978). NE Australia; E Australia (Abbott & Dance, 1983: including ethologus), S qld, N NSW (Wilson, 1993). Hal i ot i s l aevi gat a Donovan, 1808 (Figures 4-57,4-60,4-67,4-184) VIC: Torquay (DMNH 127475,1; DMNH 127654,1). E o f Geelong, Point Londsdale (NMBE 1478.95,1). PhiUip Isl. (DMNH 186161,1). Westemport Bay, W Head, Phillip Isl. (LACM H-791, 2). San Remo near Melbourne (ANSP 98914, 7). Cambertwell (MNHN no #, 1). Portsea (LACM 45806, 1). Portland Bay (AMNH 105235,2). Lakes Entrance (SBMNH 236, 5). SA: Yorke Peninsula, Stansbury G-ACM 45813,2; USNM 890100, 1). Yorke Peninsula, Sultana Bay (LACM 45815,2). Yorke Peninsula, Wool Bay (LACM A .3664,2; A .2777,1). NormanviUe (AMNH 220198,2). Kangaroo Isl. (ANSP 219617,1). Ocean Beach, FImders (AMNH 111935, 1). Edithburg, Hickey’ s Point (SBMNH 671, 2). Hardwicke Bay (LACM 45794, 1). Corny Point (AMNH 121847,1). Tumby Bay (SBMNH no #, 1). Port Lincoln (SBMNH 241, 3; SBMNH 25536, 1). 25 miles S o f Port Lincoln, Thorny Passage (AMNH 1666125,1). Wallaroo (ANSP uncataloged, 1). Port Victoria (AMNH 100811, 5). TAS: (LACM 17619, I; = Van Diemen’s Land: BMNH 37.7.15.299, 1). FUnders (LACM 79-47, I; BMNH 256 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 933.2.451, I). NWTAS (LACM A.7759, I). Montagu (AMNH 199047, 2). Stanley (BMNH 2341,3). WA: E o f Albany, Two Peoples Bay, near PL Vaillant (LACM 87-96, 1). Albany (DMNH 012211,3; LACM 127440,1). Albany, Nanarup (AMNH 110345, 1). Near Albany, King George Sound, Frenchman Bay (LACM 87-97,1). E o f Albany, Cheyne Beach (BMNH 2237,2). Swan River (BMNH no #, 2). Margaret River (LACM A .5787,1). Geralton (LACM 116190,2; ANSP uncataloged, 2). Port Augusta (AMNH 152476, 9). Augusta (AMNH 272138, 2; AMNH 220197, I). Augusta, Ring Bolt Bay (AMNH 169587,2). Esperance, Esperance Bay (SBMNH 242,3). LITERATURE, LOCALITIES: B C in g George Sound (Cotton & Godfrey, 1933). Murchison; Reevesby Isl. (Cotton, 1959). Sorrento; QueenscliS; VIC, Portsea (Macpherson & Gabriel, 1962). SA, West Isl.; Tiapara Reef (Shepherd & Hearn, 1983). SE Tip o f Eyre Peninsula, Taylor Isl.; Owen Isl. (Shepherd e t a f, 1992a). SA, Sceale Bay; McLaren Point; Taylor Isl.; Yanerbie; Anxious Bay; Waterloo Bay (Shepherd et al„ 1992b). LITERATURE, RANGE: S t Francis Isl. - Glenelg (Cotton & Godfiey, 1933). Cape Naturaliste - TAS, Bumie (Shepherd, 1973). Cape Naturaliste - VIC, TAS (Wells & Bryce, 1985; ^RTIson, 1993). Cape Naturaliste - Esperance; Amo Bay - W o f Head o f Bight, S Coast Kangaroo Isl., SW Eyre Peninsula - Wallaroo (Mace, 1986). S WA - E Bass Strait (Ludbrook & Gowlett-Holmes, 1989). HaUot i s n e t Gray* 1826 ^% ures 4-61,4-64,4-67,4-188) SA: Aldinga (AMNH 220225,2). Kangaroo Isl. (USNM 347085,2). Kangaroo Isl., Hog Bay (LACM A. 8589,1). Yorke Peninsula (AMNH ex 220176, 1). Giles Point (LACM 22216, 1; LACM A 3 7 7 7 ,2). Adelaide, Glenelg (AMNH 105233, 3). Port Willunga (SBMNH 21047,1). G ulf o f ^%cente (ANSP uncataloged, 2). Eye’ ^ s Penin- 257 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-61-66. Shells o f abalone endemic to Australia H. Figures 4-6 U 64. H aliotis roei G r^ , 1826. DLG AAB 17a. Freemantel, Western Australia. 81 mm. Figures 4-62, 65. H aliotis rubra Leach, 1814. LACM 45950. S t Vincent Gulf, South Australia. 79 mm. Figures 4-63,66. Haliotis squamata Reeve, 1846. DLG AAB 13c. Port Headland, Western Australia. 50 mm. 258 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. sula, EUistoa (SBMNH 19805,4). WA: Shark Bay (DLG AAB 17b, I). Kalbairi (NHB 1708-b, 10). 6 miles S of Geralton, Imile S o f Greenough River mouth (LACM 45916, I; SBMNH 0001, 8). Geralton, Moore Point, Geralton Lighthouse (LACM 95-58,4). Yanchep Beach (LACM 22218, 1). N o f Perth, Trigg IsI. (BMNH 2341, 13; USNM 790684.2). N o f Perth, Marmion Beach (AMNH 241175,1). Perth (DMNH 011037,2; DMNH 175704, 1). Perth, Watermans Bay (USNM 847083, 2). Swan River (ANSP 193249, 1; BMNH 38.7.222627, 2). Cottesloe (LACM 45827). Freemantle (DMNH 012212, 3; AMNH 80447, 17). Freemantle, Rottnest Isl. (LACM 28832, 1; USNM 794728, 1: complete). Rottnest Isl., Cape Vlamingh (USNM 878898,2). Freemantle, Trigg Isl. (ANSP 263203, 1; AMNH 111944, 1). Bunbury (BMNH 2351, 1). Rocking ham (LACM 45817,3). S o f Cape Clairault, Moses Rock (LACM 95-53,25). Leeuwin- Naturaliste National Park, Cowaramup Bay (LACM 45907,3; LACM 87-100,11). Cape Leeuwin (BMNH 2351,2). N Cape Mentelle (LACM 95-55, 1 + 1 complete). Augusta (SBMNH 273,2). Near Augusta, E Cape Leeuwin (LACM 87-98,2). Yallingup (AMNH 80428, 2; AMNH 80442, 3). E o f Cape Naturaliste, Bunker Bay (LACM 95-57, 1). Cape Naturaliste, Bunkers Bay (LACM 87-93,14; USNM 691539,3: complete). Cape Naturaliste (BMNH 2351,6). Cape Naturaliste, N o f Lighthouse (USNM 291502, 1: complete). Albany (BMNH 2351, 1). E o f Albany, Two Peoples Bay (LACM 87-96, 23). Near Albanay, King George Sound, Frenchman Bay (LACM 87-97, 5; LACM 87- 9 4 .2 ). King George Sound, Mistaken Isl. (USNM 858568,2). King George Sound, Pricess Royal Harbour (USNM 867220,2). King George Sound, Badter Bay, Quaran tine (USNM 867074,6). Peaceful Bay (SBMNH 184,7). LITERATURE, LOCALITIES: Yorke Peninsula, Corny Point; Guichen Bay; Encounter Bay; Port Willunga; Edhhburgh; Amo Bay; S t Francis Isl.; Beard Bay (Cot ton & Godfiey, 1933). King George Sound; Ellenbrook; Guichen Bay (Cotton, 1959). 259 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. VIC, Portland (Macpherson & Gabriel, 1962). SA, Hapara Reef; West Isl. (Shepherd & Laws, 1974). Greenough (Anon., 1987). WA, N o f Pert, Waterman Bay (Wells & Keesing, 1989). WA, ^farmion Marine Park; SA, Point Brown (Wilson, 1993). LITERATURE, RANGE: Shark Bay - Cape Otway (Anon., 1987). Shark Bay - South East Cape (Shepherd, 1973). Shark Bay - W VIC (Wells & Bryce, 1985; Ludbrook & Gowlett-Hohnes, 1989; Wilson, 1993). Geraldton - Cape Leeuwin (Mace, 1986). Haltotb rubra Leach, 1814 (Figures 4-62,4-65,4-67,4-183) NT: Port Darwin (DMNH 010998,3). QLD.: 40 miles N o f Port Douglas near Gura- dine (AMNH 121839, 2). GBR, Lindeman Isl. (LACM 45826, 49; LACM 45830, I). TownsviUe (SBMNH no # 2). NSW: Ballina, Lennex Head (LACM A.2777, 1). 15 km N of Forster, Black Head (BMNH 2341, 4). Port Stephens (DMNH 179568, 1). Sol dier’ s Point (AMNH 267915,1). Newcastle (USNM 101771,2). 25 miles S of Newcas tle (SBMNH no #, I). The Entrance, Blue Bay (AMNH 100809, 2). Swansea Head (LACM 45936, I). Sydney, Newport (USNM 835596,2). Sydn^, Long Reef (MNHN no #, 1; MNHN no #, I). Sydney, Narrabeen (DMNH 036875, I). Sydney, Newport Reef (LACM 63152,2). Sydney, Mossman (AMNH ex 96443,1). Sydney, N o f Manly, Long Reef (ANSP I34141,3; AMNH 157802,1). Sydney, Manly, Fairy Bower (USNM 842816,1; USNM 842817,3). Sydney, Coogee Bay (ANSP 136952,82). Sydney, Bondy Bay (BMNH 44.12.6205-6,2). Sydney, Collaroy (USNM 784028,1). Botany Bay, La Perouse (ANSP 134399, 4; AMNH 220230, 2). Botany Bay, Solander Monument (ANSP 149674,2). Port Jackson (LACM A 3664a, 4; LACM 105829,1). Vicmity o f Sydney, Palm Beach (AMNH 220232,2>. Nowra (NHB II 225-^ I). Woononga Beach (LACM 45929,1). Bellambie Point (AMNH 152500,2). Jervis Bay, Callala Bay (DLG AAB 22^ I). Jervis Bay, Iluka (LACM 63331, 1). S side o f Jervis Bay (LACM 79-43, 260 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 5). Jervis Bay, Huckissoa (LACM 80-64,1). Cape Jervis (AMNH Edison A 9853,2). UUadulla (SBMNH 223,17). 17 mües S of UUaduUa, Pebbly Beach (SBMNH 225,7). Narooma (LACM 30502, I). Batesman Bay, Batehaven, Corrigan’s Beach (BMNH 2341, 4; BMNH 2341, 4). Broule Batesman Bay (NHB II 225-a, 2). Eden (AMNH 276883, I; ANSP uncataloged, 2). Eden, Two Fold Bay (NHB II 225-d, I; NHB II 250-c, I). VIC: Mallacoota (NMBE 1477.19,1). Lakes Entrance (DMNH 011000,2). Momington (LACM S.4635, 1). Melbourne (LACM 59331,2; ANSP 209463,2). Mel bourne, Port Phillip Bay, Rickett Point (LACM 79-49, 8). Richmond near Melbourne (ANSP 94152,1). San Remo near Melbourne (ANSP 98915,3; ANSP 98037,10). Gee long West (LACM A.8487.68,1; LACM 45828,1). Torquay (DMNH 127548,1; DMNH I85341,2). Sorrento (BMNH Smythe collection. I). Port Phillip, Sandringham (ANSP I349I5, I). Phillip Isl. (LACM H -797,4; ANSP 2I96I9, I). PhiUip Isl. Kitty MUler Beach (LACM 28834, 7). PhiUp Isl., Western Port (AMNH 173497, 4). PhiUUp Isl., SW o f Ventnda (BMNH 2341, 14). Flinders and Philipp Isl., near Melbourne (BMNH 2036, 8). Lome (LACM S.504I, 6 ; LACM 23652,4). Otways, Skene’s Creek (BMNH 2341.1). 9 mUes ESE o f Port CampbeU, near Castle Rock, Gibson Beach (BMNH no #, I). Warmambool (ANSP 2I96I8, I). Portland Bay (AMNH 105236, 3). Portland (BMNH 2351,3). Port Fairy (ANSP 134967, 6). Portsea (LACM 45833, 3). Lome, Point Grey (USNM 767396, I). SA: Port MacDonneU (SBMNH 25566, 3; AMNH 175334, 4). Port Noarlunga (DMNH 010999, 2). South Semaphore near Adelaide (ANSP 219627,3). GUes Point (USNM 347083,4). HaUett’s Cove (LACM 45973,2). Point WUIunga (LACM 20217,3; LACM 45976,2). Pott WUIunga (LACM A2777c, I; LAC3MI52I68,1). Aldmga Bay (MNHN no #, 2). Aldinga (AMNH 240646,1; BMNH 2351.1). Cape Jervis (AMNH 240647,2). Kangaroo IsL (AMNH 80453,2). Kangaroo Isl., Hog Bay (LACM A2777a, I). St. Vincent Gulf (LACM 45950,2; MNHN no #, I). 2 6 1 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Moonta Bay (LACM 45920,2). Wallaroo (ANSP uncataloged, 2). Ybrke Pola [= Ybrke- town ?) (DMNH 008694, I). Corny Point (SBMNH no #, 3). Spencer GuIL Port Pieri (USNM 381723, 5). Westemport Bay, West Head, Flinders (LACM 79-47, I). Near Edithburgh (USNM 877521, 2). Eyre’s Peninsula (SBMNH 50069, I). Port Lincoln (SBMNH no #, 3; SBMNH 190, 4). Flinders (LACM 71876, 10; USNM 365732, I). Stansbury (AMNH ex 220178, I). Hardwicke Bay (AMNH 220231,4). Eye’s Penin sula, Elliston (SBMNH 18863, 2). Streaky Bay (USNM 835595, I). TAS: (ANSP 185897, 1; = Van Diemen’s Land: BMNH, no #, 4). Stanley (BMNH 2341, 3). Hobart, Battery Point (LACM 20390, 1; LACM 45910, 10). S TAS, Saltwater River (LACM S.1113, 3; LACM 22217, 3). Adventure Bay, Bruny IsI. (ANSP 312839, I). Middleton Channel (LACM 45819, 1). WA: Eyre (DLG AAB 22c, 1). Esperance (DLG AAB 21b, 2). Albany (LACM 127441,4; AMNH 105234,5). Cape Naturaliste (BMNH 2237,1). Cape Leeuwin (LACM 127442, 3). Cowaramup (AMNH 169687, 1; AMNH 152482, 1). Geographe Bay (USNM 877522, 5). Perth (LACM 45835, 1; LACM 45981, 1). Safety Bay, Penguin Isl. (USNM 847082,1: complete). LITERATURE, LOCALITIES: SA, Guichen Bay; Beachport; Kangaroo Isl.; Middle ton; St. Francis Isl; Marino Rocks; WA, Ellenbrook (Cotton, 1943). SA, Port Elliston, Swan River; King George Sound; Robe (Cotton, 1959). Flinders Isl.; VIC, Shoreham; Frankston, Mordialloc, Sorrento, Port Philip (Macpherson & Gabriel, 1962: some as conicopora). Bastion Point; Sandpatch Point; TuIIaburga Isl. ^ c S h a n e et aLy 1988). TAS, Nmepm Point (Prince et aL, 1987). TAS, Port Esperance, Blubber Head (Prince et al.y 1988). SA, Tiapara Reef (Shepherd & Heam, 1983). VIC, Gabo Isl.; Petrel Pt.; Petrel Rock; Cape Schanck; Apollo Bay; SA, West Is!.; Waterloo Bay, Pearson Isl.; WA, Cape Arid (Brown, 1991). TAS, Hogan Isl.; Kent IsL; Babel IsL; Cape Barren Isl.; Waterhouse IsL (Nash, e t oL, 1994). 262 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LETERATURE, RANGE: St. Francis Isl. - SA, Glenelg; Yorke Peninsula (Cotton & Godftey, 1933). Rottnest tsl. - Cofis Harbour (Shepherd, 1973). Freemantle - VIC (Wells & Bryce, 1985: as conicopora). TAS, Flinder^s Isl., King Isl., Kangaroo Isl. (not in Gulf of Vincente) - Broken Bay, SW Eyre Pen. - Wallaroo, Amo Bay > Cape Wondorma (Mace, 1986). Freemantle - VIC; SA - NSW, TAS (Wilson, 1993: part as conicopora), Hal i ot i s scalaris Leach, 1814 (Figures 4-55,4-56,4-58,4-59,4-67,4-190) NSW: Sydney, Long Reef (LACM H-338a, I). VIC: Flinder’s Isl. (LACM 28835,2). Philip Isl., Westemport Bay, Sandy Point (LACM 79-50, 1). Philip Isl., Cat Bay (SBMNH 204, 3). Westem Port (ANSP 132937, I). TAS: (AMNH 182974, 2; = Van Diemens Land: NMBE Schuttleworth, 2). Blyth (LACM 45927, I). SA: Tumby Bay (AMNH Edison A9853, 1; SBMNH 205, I). Port Lincoln (SBMNH 224, 1; SBMNH 25216,8). Point Lincoln (WAM S1007,1: complete). Adelaide (LACM 45986,1). Ade laide, Glenelg, Broadway Reef (LACM 91288, 1). Adelaide, Glenelg, Kemps Ground (AMNH 252366, I). Marino (MNHN no #, 4; MNHN no #, 2). Pt WiUunga (LACM 120215,2; LACM H-794, 5). S o f Adelaide, near Rapid Bay, Second Bay (LACM 76- 42). Spencer Gulf, Moonta Bay (LACM 45954,2; DMNH 008703,2). WaUaroo (ANSP uncataloged, 2). Stansbury (AMNH 220178, 1). Yorke Peninsula (AMNH 220176,6). NormanviUe (AMNH 220175, I; AMNH 220239, 2). Cape Jervis (AMNH 80426,2). St. Vincent G ulf (AMNH Edison A9853,1). Kangaroo Isl., Hog Bay (LACM 22219,1; LACM S.1I12,2). Port McDonneU (AMNH 175334,1). WA: Geralton (LACM 45855, 1). Jurien Bay (AMNH Jackson 1992 A9873,10). N o f Perth, Marmion (AMNH272147, 2). Perth (LACM 45849, 1). Perth, 2 mile Beach o ff Sorrento, o ff Limestone Reef (AMNH 220240,2). Perth, Bums Beach (USNM 711213,1). Swan River (ANSP 50223,. 2; BMNH 52.7.2837-38,2). Freemantle (DMNH 010200,2; AMNH 199100,2). Free- 263 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. mantle, Rottnest Isl. (LACM 87-91, 4; AMNH 203252, 3). Perth, Cockbom Sound, Woodmans Point (USNM 672361, 3). Bunbury (AMNH 240638, 3; BMNH no #, 7). Geography Bay (AMNH 252370, 1). Cape Naturaliste (DMNH 011004, 2; USNM 691501,2: complete). Cape Naturaliste, Eagle Bay (BMNH 2237,2). Cape Naturaliste - Cape Leeuwin National Park, Gracetown, Cowaramup Bay (LACM 45834, 1; LACM 95-56, 1: complete). Cape Naturaliste, Bunker Bay (USNM 691540,2: complete). S o f Cape Clairault (LACM 95-53,2). Margaret River QLACM 45919,7; LACM 45945, 11). Albany (SBMNH 36202, 1, MHNG 21812, 1). Albany, Frenchman’ s Bay (DMNH 011007, 1; LACM 30387, 1). Albany, Freemantle Bay (LACM 29948, 1). NT: Darwin (BMNH Smythe collection, 1). Indonesia: Java (AMNH 23110,1). LITERATURE, LOCALITIES: Esperance; Ellenbrook; Edithburgh; St. Francis Isl.; Point Sinclair; Kingston; Guichen Bay (Cotton & Godfrey, 1933). Corny Point; Gulf o f Vincente (Cotton, 1943). Port Willunga (Cotton, 1959). VIC, Sorrento (Macpherson & Gabriel, 1962: as emmae). SA, West Isl. (Shepherd & Godoy, 1989). WA, Marmion Nferme Park (Wilson, 1993). LITERATURE, RANGE: Geraldton - central TAS (Shepherd, 1973). Jurien Bay - VIC and TAS (Wells & Bryce, 1985). Geraldton - SA (Ludbrook & Gowlett-Holmes, 1989). Geraldton - W SA; VIC, TAS - W SA (Wilson, 1993: as emmae). f faUoit s sem^Ucaia Menke, 1843 (Figures 4-67,4-131,4-132,4-134,4-135,4-187) NSW: Port Jackson (NMBE Schuttleworth 397,1). Wbononga Beach [= Woonana ?] (LACM 120209,1 [?]). VIC: Sorrento (AMNH 272148,1). WA: Esperance (DLG AAB 11c, 3). Albany, Goode Beach (LACM 29630,1). Busselton (LACM 127439,1). Bun bury (AMNH 220242,1). Freemantle (DMNH 012825,2; WAM S1005, 1: complete). Garden Isl. (DLG AAB 11a, 1). Freemantle Rottnest Isl. (SBMNH no #, 1). O fl Free- 264 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. L - ........ H aliotis roei Î f f .' M Î s H aliotis coccoradiata • ? H aliotis cyclobates . . V X ’ . ' - * • ■ • ? ? . Haliotis elegans • ? b ' H aliotis hargravesi ■ -f. • : H aliotis laevigata * * ? > . 7 H aliotis nibra ^ ■ s t V Haliotis scalaris 4 . 7 f c H aliotis sem iplicata V . / V G H aliotis australis C / ' Haliotis iris % ^ H aliotis virginea Figure 4-8. Distribtrtion o f endemic Australian and New Zealand species o f H aliotis spp. Localities stemming from collection specimens are indicated with, a solid circle, those from the literature with a solid square. For data see tex t 265 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. mantle, Camac IsL (SBMNH 6168, 2). Cowaramup Bay (AMNH 152485, 2; AMNH 169641 ,2). Cape Mentelle (WAM S1006,1: complete). LITERATURE, LOCALITIES: WA, Marmion Marine Park; Cottesloe (Wilson, 1993). LITERATURE, RANGE: Freemantle - Esperance (Wells & Bryce, 1985). Geralton - Esperance (Wilson, 1993). New Zealand Species (Figures 4-67-79,4-177-182) Haliotis australi s Gmelin, I791(Figures 4-67,4-68,4-71,4-177,4-178) New Zealand: N Isl., Whangaroa, Stephenson’ s Isl. (DMNH 131736,2). N Isl., Doubt less Bay (AMNH 116153,1). N Isl., Northland, Bay of Isl. (DMNH 010963,4; LACM 2777c, 1). N Isl., Northland, Bay o f Isl., Kerikeri, Opito Bay (LACM 112167,1). N Isl., Northland, Bay of Isl., Russell (LACM 110349,2). N Isl., Great Barrier Isl., Tryphena (AMNH 80451, 2; AMNH 141080, I). N Isl., Northland, Whangarei Head (ANSP 219626, 2). N Isl., Northland, Whangarei Bay (LACM 45874,2). N Isl., Northland, W hangarei Pat’ s Beach (LACM 28822,1). N Isl., Whangamata (SBMNH 31773,2). N IsL, N Auckland (LACM S.1050, 1; LACM 22214,1). N Isl., N Auchland, Whale Bay (AMNH 91998,2; AMNH 220147,3). N Isl., Bay of Plenty (AMNH 199438,4; USNM 889966, 1). N IsL, Auckland, Marfelts Beach (DMNH 155567,1). N Isl., off Alderman Isl. (DMNH 040906,1). N Isl., Cape Colville, Fletcher’s Bay (DMNH 130278,1; ANSP 167580, 1). N IsL Hauraki Gulf (LACM I1046I, I). N Isl., Mahia Peninsula (LACM 6 3015,1). N Isl., Mahia, Hawke’s Point (SBMNH 13080,2). N Isl., Kuaotunu Beach (LACM 91285,2). N Isl., Castlepoint (LACM 25764,1). N Isl., Otaki (BMNH 2113, 5). N M ., Wellmgton (LACM 2777a, I; LACM 22221,1). N Isl., Wellington, Makara Beach (MNHN no #, I). N Isl., Wellington, E Isl. Bay (LACM 87-87,2). N IsL, Welling- 266 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-68-79. Shells o f New Zealand abalone. Figures 4-68, 71. H aliotis australis Gmelm, 1791. DLG AAB 40b. New Zealand. 53 mm. Figures 4-69,70,72,73. H aliotis virginea Reeve, 1846. Figures 4-69, 72. DLG AAB 30a. Marlborough. 41 mm. Dorsal and ventral view o f green variation. Figure 4-70. DLG AAB 30a. 41 mm. Marlborough. Dorsal. Figure 4-73. DLG AAB 30b. Taupiri Bay, Northland, North Island. 3 1 mm. Dorsal view o f red crispata variation. Note undulating folds and coarser cords. Figures 4-74-79. H aliotis iris Gmelin, I79I. Figures 4-74, 77. DLG AAB 07a. No locaHty. 102 mm. Adult with characteristic muscle scar. Figures 4-75,78. DLG 07c. KaDcoura, South Island. 41 mm Subadult without muscle scar, but with indication o f flared columella. Figures 4-76,79. DLG 07f. Nelson, South Island. 26 mm. Juvenile without muscle scar and no flared columella. Note simüarily to adult H virginea, 267 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ton. Breaker Bay (LACM 112168, 3). N IsL, Wellington, Lyall Bay (LACM 45969,4; AMNH 276878,2). N Isl., Wellington Harbor (AMNH 199051,4). N Isl., Wellington, Isl. Bay (AMNH 84995,2). N Isl., Wellington Heads (AMNH 80429,1). N Isl., Welling ton, Sinclair Head (USNM 637952,2). N Isl., Leigh (LACM 87-82,1; USNM 680555, 1). N Isl., Leigh, Okahari Point (LACM 87-83,1). S Isl., Pakawau (LACM 2777b, 1). S Isl., Cook Strait, Picton (AMNH 109103,3). S Isl., Nelson (DMNH 011853,61). S Isl., W Nelson, Paturau River Mouth (ANSP 283647, 3). S Isl., Kaikoura (AMNH 182267, 5; MNHN no #, I). S Isl., Bluff (AMNH 220150,3). Chatham Isl. (ANSP 399933, 1; SBMNH 33445, 1). Otago, Kartigi Beach (USNM 680557, 3). Chatham Isl., Owenga Beach (LACM 62958, 4; AMNH 276880, 1). Stewart Isl. (LACM 45975, 1; AMNH 92403, 2). Stewart Isl., Paterson, Inlet (AMNH 105536, 3; ANSP 284733,2). Stewart Isl., Native Isl. (AMNH 126329, 4). Stewart Isl., TraiU’s Bay (AMNH 126331, 3; 126330,3). LITERATURE, LOCALITIES: Lyall Bay. (Lindner, 1975). N Isl., Mount Maunganui (Ubaldi, 1986). LITERATURE, RANGE: New Zealand, Chatham, Snares, Auckland (Suter, 1913). North and South Isl., Snares, Stewart Isl., Chatham Isl. (Ubaldi, 1986). Haliotis iris Gmelin, 1791 (Figures 4-67,4-74-79,4-179,4-180) New Zealand: N Isl, Gisborne (AMNH 100806,19). N Isl, Opononi, Hokianga Har bor (USNM 680553, 2). N Is l, Whangaroa, Mahinepua (SBMNH no #, 1). N Isl, Ahipara Bay, Reef Point (AMNH 116079, 5; AMNH 116100,1). N Is l, Maunganui (LACM 18926,2). N Is l, Northland, Bay of Is l (LACM A.2777c, 1; AMNH 272137, 1). N b l , Northland, Bay of Is l, Whangarei Heads ^A C M A 3948,3; LACM A.2777b, 2). N b l., Northland, Bay of b l., Russell QACM 110354,3). N b l., NW Pomt on Motu- 268 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. tapu FsL (ANSP 234041,4). N IsL, Great Barrier Isl. (LACM 45948, 1; 45977,2). N IsL Great Barrier Isl., Port Abercrombie (LACM A.2777, I). N Isl., Great Barrier IsL, Midland Beacb (SBMNH no #, 2). N Isl., Waiheke Bay (LACM 45796, 1; ANSP 219623, 3). N Isl., Auckland, Whiüanga (USNM 749321, 1). N Isl., Gulf o f Hauraki, Takatu Passage, Kawau Isl. (USNM 617409, 2). N Isl., Auckland (BMNH 1894,4). N Isl., N Auckland, Little Manly (USNM 680556,4). N Isl., Tarakani (= Mount Egmont), Cape Lighthouse (DMNH 033841,2). N IsL Paekakarüd (DMNH 155558, l; DMNH 155902, 1). N Isl., Uriü Beach (LACM 25770, 1). N Isl., Leigh (LACM 127462, 7). N Isl., Leigh Marine Laboratory (LACM 87-82, 1). N Isl., W o f Leigh Marine Laborato ries, Okahari Point (LACM 87-83,2). N Isl., Kuaotumo Reef (LACM 91772,1; LACM 91283,2). N Isl., Turanga Bay (LACM 91305, 1). N IsL Mercury Bay (USNM 749320, 2; USNM 749241, 1). N Isl., Whangamata (SBMNH 31774, 3). N Isl., Bay o f Plenty, Mt. Maunganui (USNM 680554,1; SBMNH 2662,4). N Isl., Hawke’s Bay, Cape B C id - nappers (LACM 88-175). N Isl., Castlepoint (MNHN no #, 1). N Isl., Makara Beach (MNHN no #, 2). N Isl., Wellington, Lower Hutt (DLG AAB 07d, 2). N Isl., Welling ton, E Isl. Bay (LACM 87-87,2). N Isl., Wellington, Lyall Bay (AMNH 84994,2). N Isl., Wellington, Reef Islet (USNM 637951, 2). N Isl., Wellington, Breaker Point (SBMNH no #, 2). N Isl., Wellington, Rækaihau Point (DLG no #, 11: complete). N Isl., Cook Strait (MNHN no #, 2). S Isl., Chetwode IsL (AMNH 220190,2). S Isl., Cook Strait, Picton (AMNH 109101,1; AMNH 190111,2). S Isl., Queen Charlotte Sound, Canîbal Cove, N edge (DLG AAB 07e, 1). S Isl., Cable Bay (AMNH 116197,5). S Isl., Nelson (DMNH 118541,1). S Isl., Nelson, Fairwell Spit (DMNH 049716,2). S Isl., Nelson, Golden Bay, Tata Point (AMNH Edison A9853, 1). S Isl., W Nelson, Kaihaka (AMNH Edison A9853,1). S Isl., Kaikoura (DMNH 038954,2; ANSP 219624, l). S IsL, Thnaru (AMNH 80427). S Isl., B. St. Otago, Tautuhu (SBMNH 28350, 1). S Isl., 269 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Christchurch (NHB 1709-b. 4). S IsL, Christchurch Beach (LACM 28672, 2). S IsL, Shag Point (LACM 23455, 3; AMNH 111943, 1). Stewart IsL (DMNH 049715, 2; LACM 23454, 1). Stewart IsL, Ruggedy Point (LACM 140020, 1). Stewart Isl., Half Moon Bay (LACM 45811, 3). Stewart Isl., Ringa Ringa (AMNH 126327, 2; AMNH 220189, 2). Stewart Isl., Harrold’s Bay (AMNH 126326, 3). Stewart Isl., Horseshoe Bay (AMNH 126472, 1). Stewart Isl., Native IsL (AMNH 126325, 1). Stewart Isl., Mason Bay (AMNH 238752,1). Stewart Isl., Traill’s Bay (AMNH 126324,3). Chatham Isl. (SBMNH no #, 1). LITERATURE, LOCALITIES: Bay of Plenty (Ubaldi, 1986). Chailqr Inlet; Omaru; The Nuggets; Cape Runaway; Cape Egmont; Anchor Isl.; Bench Isl.; The Neck; Cape Jackson; Cape Young; Coal River; Codfish Isl.; Coromandel; E Ruggedy; Ruggedy W; Hapuku Rocks; BCakanui Point; Moeraki; New Plymouth; Palmer Head; Rangitoto b l; Ruapuke Isl.; Sandy Point; Stony Creek; Te Kakako Isl.; Three leg woodhen; Tinui Isl; Tongue Point; Tory Channel; Tupari Bay; Cape Turikirae; Waikawa; Waituna; Waitui Bay; W o f Cave Point; Christmas Isl.; Billhook Bay; Black Beach; Bushby Bay; Clay Point; Smolqr Cape; Fighting Bay; Karoti Light; Cape Koamaru; Lawa’s Point; Nichol son Bay; Perano Head; Robertson Point; Rollers Beach; Saddle Point; SE Harbour; Staircase; Steep Head (McShane et al., 1994). LITERATURE, RANGE: New Zealand, Chatham, Auckland (Suter, 1913). N and S Isl.; Stewart Isl.; Chatham IsL (Ubaldi, 1986). Haliotis vi rgi nea Gmelin, 1791 (Figures 4-67,4-69,4-70,4-72,4-73,4-181,4-182) New Zealand: N Isl., Taipa Beach (AMNH 115954,2). N Isl., Doubtless Bay (AMNH 116163, 2). N Isl., Northland, Taupiri Bay (SBMNH 6182, 1). N Isl., Northland, W hangarei (LACM 28818,2; AMNH 85756,4). N IsL, Near Whangarei, Taiharuru 270 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (AMNH Edison A9853,2). N Isl., Northland, Hauraki Gul^ Leigh, Goat IsL (AMNH 220259, 1; ANSP 250827, 2). N IsL, Northland, Hauraki Gulf, near Cape Rodney, Cat Isl. (AMNH 220260, 2). N Isl., Northland, Hauraki G ulf (DLG AAB 30c, 1). N Isl., Northland, Bay o f Isl. (DMNH 027831,33). N Isl., Northland, Bay of Isl., Tapeka Point (LACM H-3637). N Isl., Northland, Bay o f Isl., Russell (LACM 152307, 1; SBMNH no #, 2). N Isl., Bay o f Plenty (DMNH 027554,2). N IsL, Auckland (DMNH 155568, 1). N Isl., Auckland, Waiheke Isl. (LACM 45866). N Isl., Slipper Isl. (DMNH 073974, 2). N Isl., Wellington (LACM 45925,1; LACM S.133,1). N Isl., Wellington, Lyall Bay (LACM 45903,2; AMNH 167717,3). N isi., Kuaotunu Beach (LACM 91287,1; LACM 91277, 1). N Isl., Mercury Bay, WhManga (AMNH 220261, 3). N Isl., Mercury Bay, Simpson’s Beach (ANSP 231956, 2). N Isl., Gisborne (AMNH 220137, 2). N Isl., Hawkes Bay, Mahia (SBMNH 13079, 3). N Isl., Great Barrier Isl. (LACM 45865,3; SBMNH 31794, 1). N Isl., 1 km W o f Leigh Marine Laboratory, near Okahari Point (LACM 87-83,2; 87-84,3). N Isl., Leigh (LACM 127461,1). S Isl., Marlborough (DLG AAB 30a, 4). S Isl., Marlborough Sounds, Arapawa Isl. (AMNH 252349, I). S Isl., Cape Campbell (AMNH 122292, I). S Isl., Southland (LACM 112162, 3). S Isl., Dunedin (LACM A J1 4 7 ,2). S Isl., Dunedin, Dowling Bay (LACM H-2848,2). S Isl., Awarau Bay (LACM 45923, 3). S Isl., Kai Koures Coast (DMNH 039249,1). S Isl., Kaikoura (AMNH 182268,6; SBMNH 26233,9). S Isl., Oamani (DMNH 054637,2; DMNH 133524, I). S Isl., Mahinepoa (DMNH 133525, I). S Isl., Nelson (SBMNH 31775,2). S Isl., Timaru (AMNH 199425,5; ANSP 219622,3). S Isl., Thnaru, Caroline Bay (AMNH 220257,2). S Isl., Foveaux Strait (AMNH 169585,3; AMNH 87449,4). Stewart Isl. (LACM 45863,1; ANSP 263374,2). Stewart Isl., Ocean Beach (DMNH 027820, 4; AMNH 141079,1). Stewart Isl., Patterson Inlet (AMNH 105535,2). Stew art Isl., Harrold’s Bay, Beach Isl. (AMNH 126333, 1; AMNH 126332). Stewart Isl., 271 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Ringa Ringa (AMNH 126334, 1). Stewart IsL, Horseshoue Bay (ANSP 231941, I). Stewart IsL, Ocean Beach (ANSP 232279,5). Chatham IsL (AMNH 183966, l; AMNH 23094,4). Chatham Isl., Kaingaroa (ANSP 250828,2). Chatham IsL, Waitangi (SBMNH 6180,1). Chatham IsL, Red Bay (DMNH 026452, 2). Campbell Isl. Monument Har bour (DMNH 027398,4). Campbell Isl., Monument Harbor (AMNH 183965,1; AMNH 84996, 1). Campbell Isl., Perseverance Harbor (ANSP 235404, 3). Auckland Isl. (BMNH 1905.11.28.228, 1). Auckland Isl., Enderby Isl., Derry Castle Reef (SBMNH no #, 6). LITERATURE, LOCALITIES: N Isl., Tutukaka; Mercury Bay; Takapuna; Matapouri; Chatham Isl., Waitangi; Owenga (Ubaldi, 1986). LITERATURE, RANGE: New Zealand, Chatham, Stewart, Kermadec IsL Campbell Isl. (Suter, 1913). E coast of North Isl. to Hauraki Gulf; Cook Strait Area; S most South Isl. - Stewart Isl.; Auckland Isl.; Campbell Isl.; Chatham Isl. (Ubaldi, 1986). Northwest Pacific Species (Figures 4-80-86,4-118,4-121) Hal i ot i s discus Reeve, 1846 figures 4-80,4-83,4-86) USSR: Kurilen, Moneron Isl. (DLG AAB 03c, 2). CHINA: Tsingtao Sung (USNM 404389, 3). JAPAN: Hokkaido, Teshio Prov., Yagishiri Isl. (USNM 471854, 15). Hokkaido, Montebetsu (DLG AAB 03b, 1). Hokkaido, Hiyama, Tomari Village, Nakayama Said (LACM 78-14,7). Hokkaido, Hqrama, Tomari (LA(3M 78-9,2; LACM 78-8,2). Hokkaido, Nemuro, Nemuri (AMNH 80433,1). Hokkaido, Geshio, Mashike (AMNH 19789, 5). Hokkaido, Geshio, Mashike, Teshiwo (ANSP 80391, 1; USNM 273417,2). Hokkaido, Oshhna, Hakodate (ANSP 219611,1; MNHN no #, 1). Hokkaido, Abashiri (ANSP 252615, 6; ANSP uncataloged, 1). Hokkaido, Ishikari, Otaru (USNM 272 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Ï Figures 4-80-85. Shells o f the northwest Pacific abalone. Figures 4-80,83. H aliotis dis - eus Reeve, 1846. DLG AAB 03b. Montebetsu, Hokkaido, Japan. 82 mm. Figures 4-81. 84. H aliotis gigantea Gmelin, 1791. DLG AAB 27c. Nada, Wakayama Preference, Japan. 75 mm. Figures 4-82, 85. H aliotis madaka Habe, 1977. LACM 31925. Japan. Cate coll. 163 mm. 273 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H aliotis discus - " -- • H aliotis g^gantea y • H aliotis madaka H aliotis exigua • 1- • • Figure 4-86. Distributioa o f northwest Pacific species o f H aliotis spp. Localities stem ming from collection specimens are indicated with a solid circle, those fiom the iitera- ture with a solid square. For data see texL 274 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 22683, 4). Honshu, Wakayama, Mio (AMNH 239003,2; ANSP uncataloged. I). Hon shu, Wakayama, Seto Marine Laboratory, Cape Bansho-zaki (LACM 82-19, 3). Hon shu, Wakayama, Shirahama, B C Ii (LACM 110919, 2; AMNH 260666). Honshu, Wakayama, Shirahama, B C ii, Kii Channel (AMNH Edison A9853,2). Honshu, Niigota, Sado Isl., Aikawa Machi, Samuto Said, Seki (ANSP 325476,3). Honshu, Niigota, Sado IsL, Aikawa, Riuzen (USNM 205829, 1). Honshu, Noto (AMNH 80443, 1; AMNH 141071.1). Honshu, Noto, Monzen-flikami (LACM 82-5). Honshu, Amori (AMNH 173245,1; MNHN no #, 2). Honshu, Amori, Cape Shiriyazaki (LACM 78-15,2; LACM 82-17, 3). Honshu, Amori, Asamushi Marine Biological Station, Mutsu Bay (LACM 82-16, 4; LACM 87-12, 2 + 3 with dried bodies). Honshu, Akita, Funagawa [= Funakawa ?] (USNM 226857, 4). Honshu, Iwate, RBcuzen (LACM 59342, 1; USNM 343822.2). Honshu, Iwate, Fukura, Awaji (ANSP 76327, 1). Honshu, Miyagi (ANSP 241102.2). Honshu, Kanagawa, Sagami Bay (AMNH Edison A9853,2; SBMNH no #, 2). Honshu, Kanagawa, Yokohama (USNM 43065,4). Honshu, Ishika, Mikawa (AMNH Edison A9853, 1). Honshu, Chiba, Station Boshu (LACM 120226, I). Honshu, Chiba, Boso Peninsula, Katsu-ura (USNM 753692,2). Honshu, Shizoaka, Matsuzaki (ANSP 252589, 6). Honshu, Mie (ANSP 252583,2). Shikoku, Ehime, Misaki (ANSP 119840, 13). Shikoku, Kochi, Tosa (LACM 91286,2; LACM 91275,1). O ff Shikoku, Dd-suido (SBMNH 6172, 1). Kyushu, Ka(g)oshima (AMNH 115319,1). Kyushu, Kumamoto, Amakusa Marine Laboratory (LACM 82-27,11). Kyushu, Kumamoto, Amakusa Marine Laboratory, Tsigushima IsL (LACM 82-25,2). Kyushu, Kumamoto, Amakusa Marine Laboratory, S Tamioka Peninsula (LACM 82-26, 1). Kyushy, Nagasaki, Sasebo, Shra- hama Beach Cove (AMNH 272136,1). Kyushu, Nagasaki, Kuchinotsu (AMNH 241418, 6). Kyushu, Nagasaki, Nagasaki (NHB 1711-b, 1). Kyushu, Nagasaki, Matsushima Bay (ANSP uncataloged. I). Kyushu, Nagasaki, Enoshima (AMNH 46574, 1). Kyushu, 275 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Saga, Takashima (USNM 226852, 1). Bonin Isl. (= Ogasawara-shoto) (USNM 226853, I). Korea: Korea Strait, Cheju-do (= Quelpart Isl.) (ANSP 298275, 3; LACM 71-256). Pusan (DMNH 011046, 5; AMNH 198669, I). Masan, Kyongsong Namdo (USNM 623631, 1). Off Inchon, Paenr Myong Do (USNM 673371,2). LITERATURE: Localities: S[h]imoda; Hakodate; Goto Isl. (Pilsbry, 1895). Ibaragi, Otsu, 180 miles N o f Tokyo (Ino & Harada, 1 961). Honshu, Akita, Oga; Kyushu, Ehime, Uwajima; Kyushu, Tokushima, Anan (Arai et aL, 1982). LITERATURE: range: N Japan; Honshu through S Kyushu (Kira, 1962). Halioti s exi gua Dunker, 1877 (Figures 4-87,4-118,4-121) Japan: Hokkaido, Oshima, Hakodadi [= Hakodate ?] Bay (USNM 1652, I). Okinawa (LACM 45851, I; LACM 45847,1). Okinawa, Camp Kue, W Nakagami-Gun (USNM 670615,1). Okinawa, Onna Village (USNM 821324,2). Okinawa, Onna Village, Horse shoe Cliffs (USNM 838974, 16; USNM 838847, I). Okinawa, Kadena Yacht Club, sea wall at channel (USNM 821168, 3; USNM 821128, I). Okinawa, Machinato Reef (SBMNH no #, 1; SBMNH no #, 2). Okinawa, Kouri-jima (SBMNH no #, 3). Okinawa, Imbu (SBMNH no #, 2; SBMNH no #, 5). VIETNAM: Nha Trang Bay, Hon Lo Isl. (DLG AAB 54a, 3). LITERATURE, LOCALITIES: Kagoshima, Amami Isl.; Kagoshima, Amami Isl., Setouchi; Kagoshuna, Kakeroma Isl., Shodon Bay (Arai et aL, 1988: as K planata). LITERATURE, RANGE: Okinawa (Talmadge, 1963a). Haliotis g^antea Gmelm, 1791 fig u re s 4-81,4-84,86) Russia: Sakahalin Isl. (LACM 31490,1). Japan: Honshu, Wakayama, Shirahama, Kii (LACM 110815,2; DMNH 049414,1). Honshu, Wakayama, Minabe (AMNH 239002, 276 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 4). Honshu, Wakayama, near Seto Marine Laboratory, Cape Bansho Zaki (LACM 82- 19,2). Honshu, Miura Pensinsula, Misaki (Entrance to Tokyo Harbor) (USNM 753693, 2). Honshu, Kanagawa, Sagami Bay (AMNH 220241, I; SBMNH no #, I). Sagami (USNM 228275, 1). Honshu, Kanagawa, between Tokyo and Sagami Bay, Misaki (AMNH 120582,1). Honshu, Chiba, Station Boshu (LACM 129224,1). Honshu, Chiba, Katsuura(LACM A.8487.68,1; SBMNH no #, I). Honshu, Shizuoka, Matsuzaki (ANSP 252588,4), Honshu, Mie (ANSP uncataloged. I). Honshu, Hiroshima (SBMNH no #, I). Shikoku, Kochi, Tosa (ANSP 192975, 2). Shikoku, Kochi (= Tosa), Oinoshima (DMNH 011026, 1; DMNH 008698, 4). Kyushu, Nagasaki, Nagasaki (NHB 1711-b). Kyushu, Nagasaki, E[n)oshima (LACM 120220, 2; LACM A ^777a, 2). Kyushu, Nagasaki, E[n]oshima, Market (LACM 120228, 1). Okinawa, Ishinone, Izu (ANSP 219613,1). Palau (= BELAU): (USNM 589521,1: doubtful). LITERATURE, LOCALITIES: Yedo (= Tokyo); Hakodate (Pilsbry, 1895). LITERATURE, RANGE: Honshu - S Kyushu (Kira, 1962: as sieboldn), Kyushu Isl.; Shikoku Isl.; S Honshu - Boso Peninsula; S Korea (Habe & Kosuge, 1964). Haliotis madaka Habe, 1977 (Figures 4 - 8 2 ^ 4-85,4-86) Japan: Honshu, Chiba, Station Bonshu (LACM 112927,1+3 dried radulae). Honshu, Kanagawa, Sagami (LACM 119068, 1; USNM ex 228275, 1). Honshu, Shizuoka, A/bt- suzakim (ANSP 399931,1). Honshu, Kurihama, Miura Peninsula, entrance of Tokyo Bay, Kii (USNM 753694,1). Shikoku, Ehime, Misaki (ANSP 252573,2). Shikoku, Okinnshima, Tosa (DMNH 011021,2; DMNH 008706, 1). Kyushu, Nagasaki, Market o f Enoshima (LACM 120227,3). Kyushu, Kumamoto, Amakusa Marine Laboratory (LACM 82-27,9). 277 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE: range: Honshu - S Kyushu (Kira, 1962: as gigantea). Japan, Korea (Abbott & Dance, 1983: as gigantea). East Pacific Species (Figures 4-87-102,4-104-107,4-109,4-110,4-112,4-113,4- 167,4-168,4-173) ffaliotis corrugat a Wood, 1828 (Figures 4-88,4-91,4-100) CANADA: British Columbia, Victoria, Breakwater (AMNH 249114,1 [?]). Califer- nia: San Francisco County, San Francisco, Farailon Isl. (DMNH I I I I 64, I; DMNH 111165.1). San Francisco County, Roclg Point (AMNH 198682,1). Monterey County, Monterey Peninsula (SBMNH 1919, I). San Luis Obispo County, Moro Bay (DMNH 066221, I; DMNH 095516, I). San Luis Obispo County, Cambria (SBMNH 229, 2). Santa Barbara County, Santa Barbara (DMNH 118926, 1; 136735, I). Santa Barbara County, Point Conception (LACM 63-39.8,3). Santa Barbara County, Coal Oil Point, Goleta (SBMNH 23446, 5). Santa Barbara County, Coal Oil Point, Isla Vista (SBMNH 25528.1). S Anacapa Isl. (LACM 64-28.6,1). Anacapa Isl. (SBMNH 44473,2; SBMNH 44470N, 2). Anacapa Isl., ocean side (SBMNH 44471,2). Santa Barbara County, Santa Rosa Isl. (SBMNH 170,3). Santa Barbara County, Santa Cruz Isl. (LACM 69-32.7,2; LACM 40-163.4,1). Santa Barbara County, W Santa Cruz Isl. (SBMNH 6153,1). Santa Barbara County, N Santa Cruz Isl. (LACM 63-5.6,1; AMNH I9877I, I). Santa Bar bara County, Santa Cruz Isl., Smuggler’ s Cove (LACM 48-42.11, 3; SBMNH 44469, 2). Santa Barbara County, S Santa Cruz Isl., N Gull Isl. (LACZM 73-11.9, 4). San Clemente M ., near Pyramid Cove, HoresCove ^A C M 39-129.5,1). San Clemente Isl. (LACM 146745, I; LACM 23453,2). SE San Clemente Isl. (LACM 20951, I). San Clemente Isl., Wilson Cove (LACM 66-51.6,2). SE Santa Clemente Isl. (LACM 69- 278 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ■ jf Haliotis k. kamtschatkana Figure 4-87. Distribvrdott o f east Pacific H aliotis L kam tschatkana. Localities stem ming fiom collection specimens are mdicated with, a solid cficle, those fiom the litera ture with a solid square. For data see texL 279 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-88-99. Shells o f the northeast Pacific abalone I. Figures 4-88,4-91. H aliotis corrugata Wood, 1828. LACM 23453. San Clemente Island, California. Leg. Dameoun Burford, XII. 1953. 151 mm. Figures 4-89, 4-92. H aliotis cracherodii Leach, 1814. DLG AAB 37c. Channel Islands, California. 82 mm. Figures 4-90,4-93. H aliotis fu l - gens Philippi, 1845. DLG AAB 05b. San Diego, California. 168 mm. Figures 4-94,4- 97. H aliotis k. kamtschatkana Jonas, 1845. LACM 117633. Vancouver Island, British Columbia. 88 mm. Figures 4-95, 4-98. H kamtschatkana assim ilis DaU, 1878. DLG AAB 36b. California^ 83 mm. Dorsal and ventral view o f specimen with somewhat uncommon orange (dark) band near row o f holes. To enhance the contrast o f the orange band m black and white, the blue alpha-channel o f the RGB file was used. Figures 4- 96,4-99. H aliotis nifescens Swainson, 1822. DLG AAB 46<L Point Conception, Cali fornia. Ex col. K L Stewart, leg. Thomas 1976.209 mm. 280 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 281 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 29.12,8). Santa Barbara County, Santa Barbara (BMNH1822,1). Santa Barbara County, Santa Barbara Isl. (LACM 72-97.8, 2). Santa Barbara County, 10 miles off Santa Bar bara Isl. (SBMNH 18825a, 2). Santa Barbara County, San Miguel Isl. (USNM 778933, I). Channel Isl., Cortes Bank, near Bishop Rock (LACM 71-2063,1; LACM 69-34,4: complete). Ventura County, Port Hueme (AMNH 111142,1). Ventura County, Marina Oxnard, Channel Isl. (SBMNH 178,1). Ventura County, N San Nicolas Isl. (LACM 72- 101.8,2; LACM 68-35.6, 1). Ventura County, NE San Nicolas Isl. (LACM 69-15.7,1). Ventura County, S San Nicolas Isl. (LACM 69-14.5, 1). Los Angeles County, Santa Catalina Isl., Isthmus Cove (LACM 71-99.12,3). Los Angeles County, Santa Catalina Isl., Farnsworth Bank (LACM 65-32.4,2). Los Angeles County, Santa Catalina Isl., N Avalon (LACM 62-5.7,1). Los Angeles County, W Santa Catalina Isl. (LACM 64-26.8, 4; LACM 68-25.5,2). Los Angeles County, W Santa Catalina, Ironbound Cove (LACM 69-31.7,1). Los Angeles County, Santa Catalina Isl. (DMNH 066025, 2). Los Angeles County, Santa Catalina Isl., USC lab (USNM 784077,2). Los Angeles County, S Santa Catalina Isl., E o f Isthmus (LACM 67-11.6, 2). Los Angeles County, Point Dume (AMNH 198676, 1). Los Angeles County, N Malibu (AMNH 240630, 1). Los Angeles County, Malibu (SBMNH no #, 1). Los Angeles County, Redondo Beach (LACM 146743, 5). Los Angeles County, Los Angeles, Palos Verdes (DMNH 111166,1). Los Angeles County, Palos Verdes, Whites Point (LACM 59328, 1; AMNH 111940,1). Los Angeles County, Palos Verdes, Portuguese Bend (LACM 146748, 1; LACM 146738, 1). Los Angeles County, Paradise Cove (LACM 66-58.5,1; AMNH 93406, 1). Los Angeles County, Los Angeles, San Pedro (DMNH 136735, I; DMNH 066013, 1). Orange County, Laguna Beach (MHNG 21808, 1). Orange County, S Laguna Beach (AMNH Edison A9853, 2). Orange County, between Newport Beach and Laguna (USNM 524678,2). San Diego County, Point Loma (LACM 72-111.6,1; AMNH Jack 282 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. son 1992 A9873,2). San Diego County, La Jolla (DMNH 111972, 1; LACM 67-24.5, 1). San Diego County, San Diego, Mission Bay (LACM 62-3.6,1; LACM 71-1012,1). San Diego County, San Diego (AMNH 100818, 1). Mexico: Sonora, Guyamas (DMNH 101967, 4; AMNH 123807, 2). NW Guadalupe, Poloto Roca (LACM 72-121.4, I). Guadalupe, Inner IsL, Melpomene Cove (LACM 49-155.6, 1). SW Guadalupe, Five Fathom Bank and Basalt Arches, Isla Afuera, Ha Adentro (LACM 72-120.4, 7). W Guadalupe (LACM 65-412,2). S Guadalupe (LACM 65-42.6, 3). Mexico, Baja Cali fornia, Pacific Side: Middle Coronado Isl. (LACM 63-41.10,6). Coronado Isl. (LACM 71-98.5,1). S Punta Pequena (LACM 71-1792,1; LACM 71-181.5,1). Punta Pequena (LACM 71-6.2,1). Todas Santos Bay (AMNH 47262,4; ANSP 50144, 3). Todos San tos Bay, N Punta Banda, Between Black Rock Blufflf and Red Earth Bluff (SBMNH 13344, 1). Punta Abreojos (LACM 71-3.8, 5). Punta San Eugenia (LACM 41-40.10,1: complete). S Puerto Santo Tomas (LACM 66-1.4,1). Santa Tomas (SBMNH 5672,2). S Punta Banda (LACM 67-53.1, 1 with dried body; LACM 63-42.6, 4 bodies). S o f Todos Santos, Punta Banta (DMNH 053918, 1; LACM 146751,2). NE to E Isla Todos Santos (LACM 67-45.8,1+2 with dried bodies). SW Isla Natividad (LACM 72-117.8, 5; LACM 71-165.5,4). S Isla Natividad (LACM 72-116.4,3). San Martin Isl. (USNM 601642, 1). N Isla San Geronimo (LACM 67-62.5, 8). S Isla San Geronimo, Sacra mento Reef (LACM 71-91.8). Islas San Benito, San Benito del Geste (LACM 146753, 1; 72-118.9,2). Tres Benitos Isl. (AMNH 183129, 1; AMNH 254127, 1). Cedros Isl. (USNM 264495,1). E Isla Cedros, Pueblo de Cedros (LACM 71-92.7,1). Off SE Isla Cedro, Five Fathom Bank (LACM 71-93.7, 2). SW Isla Cedros, S Piedra Colorada (LACM 72-115.6,2). NE Isla Cedros (lA C M 72-113.6,5; LACM 71-151.9,14). Punta Rompiente ^A CM 71-161.6,3). Thurloe Head (LACM 71-170.1, 3). Bahia San Bar tolomé (= Turtel Bay) (LACM 146750, 10; AMNH 77712,2). Isla Asuncion (LACM 283 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 67-66.10, 8). Punta San Pablo (LACM 71-178.12,3). Bahia Magdalena, NW Punta Bercher, Isla Santa Magdalena (LACM 67-70.2,14). Bahia Magdalena, Man o f War Cove (LACM 71-183.8, I). N Bahia Magdalena, Sail Rock, E Punta Entrada (LACM 71-14.14,1). LITERATURE, LOCALITIES: All Channel Islands (Cox, I960). Bahia Tortugas (Munoz-Lopez, 1976). LITERATURE, RANGE: Montery - San Quentin Bay (Oldroyd, 1927; Bonnot, 1948). Monterey (California) - Lower California (Abbott, 1954). Point Conception - Turtle Bay (Cox, 1960). Point Conception - Central Baja California (Lindner, 1975). Point Conception - Ascuncion Isl. (McLean, 1978). Point Conception - Santa Maria Bay; Channel Islands; Guadalupe Isl. (Haaker et uL, 1986). Haliotis cracherodi i Leach, 1814 (Figures 4-89,4-92,4-100) California: Del Norte County, Crescent City (USNM 104142a, 3). Mendocmo County, near Albion, Cormorant Cove (LACM 49-4.7,1). San Francisco County, San Francisco (USNM 4266, 1). San Francisco County, San Purissima (USNM 98321, I). Monterey County, Pacific Grove (USNM 664043,1). M onter^ County, Pacific Grove, near Lucas Point, DMNNH 035477, 2). Monterey County, Pacific Grove, Asilomar State Park (DMNH 120828, 11). Pacific Grove, 17 miles Fan Shell Beach (SBMNH 27121,1). Monterey County, Monterey (DMNH 049704,1; ANSP 50163,2). Monterey County, Monterey, Sand Dollar Beach (DMNH 112596,1). Monterey County, Monterey, Pacific Grove (DMNH IIII6 9 , 6 ; LACM 59-12.7,13). Monterey County, Carmel (DMNH 122144,1; ANSP 155145,1). M onter^ County, Carmel Point ^A C M 79-1142,3; USNM 812885,1: complete). Monterey County, Carmel, Mission Pomt (SBMNH no #, 3). Monterey County, N Point Sur Lighthouse (LACM 69-38.1,1). Sonoma County, 284 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Russian River (LACM 48-47.1, l)> Marin County, Moro Bay, Horseshoe Cove (LACM 146754.1). San Luis Obispo County, San Simeon Bay (USNM 212719, 4). San Luis Obispo County, Cambria (AMNH 85473,4; SBMNH 219,7). San Luis Obispo County, W Cayucos Creek (LACM 77-112.3, 1). San Luis Obispo County, Cayucos (LACM 29567, I; LACM 146765, I). San Luis Obispo County, Cayucos, Garcia Ranch (SBMNH 46551, 1). San Luis Obispo County, Avila Beach (LACM 68-37,7, 6). San Luis Obispo County, N Shell Beach (LACM 77-111.2,2). San Luis Obispo County, San Luis Obispo (USNM 4258, 1). San Luis Obispo County, S Shell Beach, Dinosaur Cave (LACM 68-36.5, I). San Luis Obispo County, Pismo Beach (ANSP 187052, 1). San Luis Obispo County, N Piedros Blancas Point (DMNH 090273,1; LACM 76-12 J , 2: complete). San Luis Obispo County, Moro Bay (DMNH 095738,1; DMNH 095519,1). San Mateo County, Franklin Point (LACM 62-8.5,1). Santa Cruz County, Grayhound Rock to Ano Neuvo Point, Waddell Beach (LACM 67-95.8, 5). Santa Barbara County, S Point Sal State Beach (LACM 77-312,1). Santa Barbara County, E Carpentaria State Beach (LACM 77-30.4, 1). Santa Barbara County, Point Conception (LACM 110443, 2; LACM 88-131). Santa Barbara County, Jamala (SBMNH 46519, 1). Santa Barbara County, Goleta, Coal Oil Point (SBMNH 232, 1; SBMNH 23445,4). Santa Barbara Cotmty, Santa Cruz Isl. (DMNH 090277,1; LACM 40-163.5,3). Santa Barbara County, S Santa Cruz Isl., Willow anchorage (LACM 40-167.8, 1 body; LACM 40-167.9, 1). Santa Barbara County, Santa Cruz Isl., Foumeys Cove (LACM 69-11.3, 3; SBMNH 19137.1). Santa Barbara County, Santa Cruz Isl., SW Smugglers Cove (LACM 48- 42.12.1). Santa Barbara County Santa Cruz IsL, Pelican Bay (USNM 682092,3). Santa Barbara County, Santa Cruz Isl., Arch Rock (SBMNH 28304,2). Santa Barbara County, Santa Barbara Isl. (ANSP 155137,1; 50538,3). Santa Barbara County, San Miguel Isl. (LACM 67-38.7,4; ANSP 239564,2). Santa Barbara County, San Miguel IsL, Tyler 285 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Bight ^A C M 38-15LI, L ; LACM 38-151.1,1). Santa Barbara County, Santa Rosa Isl. (AMNH 198673,1; USNM 60296,3). Ventura County, Point Mugu (LACM 146777,1; LACM 146783,1). Ventura County, Oxnard Marina (SBMNH 221,5). Ventura County, Point Hueme (SBMNH 44392M, 1). Ventura County, Point Dume (DMNH 035329, 3). Ventura County, San Nicolas Isl. (DMNH 174760, 11; LACM 59322, 5). Ventura County, San. Nicolas IsL, Honey Moon Cove (LACM 146781, 3). Ventura County, San Nicolas Isl., Dutch Harbor (SBMNH no #, 2). Los Angeles County, Santa Catalina Isl. (LACM 146766, 1: complete). Los Angeles County, Santa Catalina Isl., Fisherman’s Cove (LACM 146782, 5). Los Angeles County, W Santa Catalina Isl. (LACM 64-26.9, I; LACM 68-25.6,1). Los Angeles County, Santa Catalina Isl., NW Fouth of July Cove (LACM 52-143,1: complete). Los Angeles County, SW Santa Catalina Isl., Ironbound Cove (LACM 69-31.8, I). Los Angeles County, Santa Catalina Isl., Cherry Cove, W Isthmus Cove (LACM 48-15.5,2). Los Angeles County, Santa Catalina Isl., Isthmus Cove (LACM 71-213.2,2; LACM 71-99.13, 4). Los Angeles County, Santa Catalina IsU Avalon (SBMNH 48,4). Anacapa Isl. (LACM 146774,2; USNM 778883,2). San Clemente Isl. (LACM 39-125.2,2; AMNH 93906, 1). San Clemente Isl., China Point (LACM 39-2.2,1). San Clemente IsUWilson Cove (LACM 66-51.7,1). San Clemente Isl., NW Harbor (LACM 39-1253,5). San Clemente Isl., Pyramid Cove (LACM 39- 1 2 8 3 ,1 : complete). San Clemente Isl., near Pyramid Cove, Horse Cove (LACM 39- 129.6,1; LACM 39-129.6,5: complete). Los Angeles Coimty, Leo Carillo State Park, Sequit Point (LACM 62-233, 1). Los Angeles County, Malibu (LACM 61972, 4; LACM 54786,3: complete). Los Angeles County, Topanga Creek (LACM 68-192.4,1). Los Angeles County, Redondo (USNM 199890,1; SBMNH no #, 2). Los Angeles County, Palos Verdes Peninsula: Torrence, Flat Point ÇDLG AAB 37e, 1). Torrence, Algal Cove (DLG AAB 37g, 1). Lunada Bay (LACM 60-13.7,1). B luff 286 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Cove (LACM 46-32.8, I: complete; LACM 146760, 2). Point Vincente (LACM 61- 10.4, 6 ; LACM 146764,1). Portuguese Bend (LACM 65-80.5, 6; LACM 54781, I: complete). Flatrock Point (LACM 10911,5). Whitens Point (DMNH 121423,20; DMNH 008702,3). Malaga Cove (DMNH 113801,8; ANSP 263045,2). Clayden Drive (DMNH 074540,2). 2 miles N o f Marine Land (DMNH 108687,2). Reef Point (AMNH 183127, 1). Los Angeles County, Point ^^cente (USNM 120131,2). Los Angeles County, San Pedro (LACM 18748, 1; LACM 111094, 7). Los Angeles County, Los Angeles, San Pedro, Royal Palms Beach (DMNH 0421296, 5; LACM 72-96.5, 3). Los Angeles County, San Pedro Bay (ANSP 50539,9). Los Angeles County, Los Angeles, Point Fer- min (DMNH 066023,3; LACM 61812,2). Orange County, Anaheim Landing (LACM 146767, 17). Orange County, Laguna Beach (DMNH 008699, 1; DMNH 136499, 8). Orange County, Newport Beach (DMNH 088024,3, AMNH 103822,2). Orange County, Laguna Beach (ANSP 324965,1). Orange County, Corona del Mar (DMNH 112133,3; BMNH 2258,7). Orange County, Corona del Mar, Newport (LACM 69-72, 7; LACM 40-142.9,1: complete). Point Loma (AMNH 84651, 1, AMNH 100827, 1). San Diego County, La Jolla (DMNH 008683,2; AMNH 240632, 1). San Diego County, La X oU a, Devil^s Slide (SBMNH no #, 1). San Diego County, La Jolla, Bird Rock (SBMNH no #, 3). San Diego County, San Diego (ANSP 50534,5; USNM 3406,1). San Diego County, San Diego, Mission Bay ^A C M 146761, 1; LACM 68-26.4,1). Mexico, guadalupe Isl.: (DMNH 010968, 2; LACM 146799. 1). Seal Beach (DMNH 111608, 1). NW Anchorage (AMNH 198671, 1). NE anchorage (LACM 10349, 3). NW Guadalupe Isl. (LACM 19653,2). S Bluff (LACM 49-159.6,6). Melepomeme Cove (LACM 146789, 2). S Guadalupe IsL (LACM 31930,1; LACM 18045,1). Leeward Isl. (AMNH 236683, 1). N Lee Cove (SBMNH no #, 2). mexico, Baja California, Pacific coast: 25 miles S o f Tiajuana (AMNH 114751, >50). N Coronado IsL (SBMNH 13327,1). S Punta Banda, 287 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Arbolito (LACM 68-40.5, 1; LACM 67-53.2, 2). Punta Banda (LACM 63-43.1, II; SBMNH 0888,4). N Punta Banda (LACM 63-54.5, I). Punta Banda, Kennedy Cove (SBMNH no #, 5). NE Punta Banda, Punta Vanda Beach (SBMNH 14626, I). N Punta Banda, between black Rock Bluff and Red Earth Bluff (SBMNH 13346, 4). Punta Banda, LaBufadora (SBMNH 1771, 10; SBMNH 40242, 16). Punta Clara, S o f Rio Santo Tomas (SBMNH I7I72, I). O ff San Jose Point (AMNH I2I798, 1). San Quintin (DMNH 126410,1; USNM 34496,1). San Quintin, W o f Kenton Hill (SBMNH 16119, 1). Isla San Martin, Hassier Cove (LACM 68-31.4, 3; LACM 73-2 J , 3). Isla San Mar tin, Hassler Cove, Boulder Rock (SBMNH 13428, 1). Guerrero Negro (DLG AAB 37b, 4). Cedros Isl. (USNM 4317,1; USNM 264497,6). SW Isla Cedros, S Piedra Colorada (LACM 72-115,7,1). N Isla Cedros (LACM 51-41.4, 1). NE Isla Cedros (LACM 71- 151.10.2). Isla Cedros, S Bay (LACM 34-162.1,1). Punta Asuncion (LACM 50-26.10, 1). Puerto Santo Tomas (LACM 64128, 1; LACM 67-2.4, 8). Punta Santo Tomas (AMNH 183128, 1; AMNH 183128, 1). N Baja, Ensenada (AMNH 284464, 1; AMNH 254126.2). O ff Ensenada ML (AMNH 175667,1). SE Isla San Geronimo (LACM 67- 63.1,6; SBMNH 13523,5). San Geronimo Isl. (SBMNH 13522, 1). N o f Ensenada at K-lOO (LACM 64-32.4, 1; LACM 63-53.4, 1). N o f Ensenada, El Sauzal (LACM 67- 94.1.2). Ensenada (USNM 664047,2; USNM 664048,2). Isla Todos Santos (LACM 67-45.9, 1). Isla San Benito (LACM 146787,1; USNM 265598, 1). Isla E San Benito (LACM 50-233,4). Isla W San Benito (LACM 50-223,1; LACM 59333, 1). Middle San Benito Isl. (AMNH 85676,3). Punta Colnett (LACM 46-14.1, 2). S Natividad Isl. (SBMNH 14095,2). Turtle Bay (USNM 2759,2; SBMNH 18246,6). San Bartholome Bay (USNM 265141,1; USNM 554377, 3). S o f Tijuana, El Mono and Los Gaviotas (LACM 6-183, 1). O ff Soledad B ^ (AMNH 121800,1). Thurloe Head (LACM 71- 1 7 1 3 ,2 ; LACM 31926, 1). Punta Rompiente (LACM 71-161.7,1; LACM 71-1623, 288 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 7). N Magdalena Bay (SBMNH 23249, 1). Mexico: Nayarit, Tres M aria Isl. (DMNH 112686.1). Guyamas (AMNH Edison A9853,2; AMNH Edison A9853, 8). Socorro Isl. (AMNH 88826,2). LITERATURE, LOCALITIES: California, Point Arena (Bonnot, 1930). Point Lobos State Park, Cypress Cove (Bonnot, 1948). California, Montana de Oro State Park (Minchin, 1975). California, Point Arguello (Haaker et nA, 1995). LITERATURE, RANGE: Coos Bay, Oregon - Santa Rosalia, Baja California (Oldroyd, 1927; Bonnot, 1948). Part o f Oregon; San Francisco - San Quentin Bay; all off-shore Isl. (Bartsch, 1940). Coos Bay, Oregon - Lower California (Abbott, 1954). Coos Bay, Oregon - Cape San Lucas, Baja California (Cox, 1960). Mendocino County - S Baja California (McLean, 1978). Mendocino County - S Baja California (Haaker et aL, 1986). H aliotis dt üU Henderson, 1915 ^% ures 4-107,4-110,4-113) equador, Galapagos: Santa Cruz, Academy Bay (AMNH 267218, 2; AMNH Edison A9853, 1). Santa Cruz, Bahia Academy, Eocacro (LACM 146876, 1). Santa Cruz, 2 miles E o f Academy Bay (LACM 72-200.7, 1). Santa Cruz, off Academy Bay (AMNH 220170, 1; SBMNH no #, 1). Santa Cruz (DMNH 106715, 1; LACM 152364, 1). S Santa Cruz (AMNH 145835, 1: in alcohol with some tissue; USNM 812778, 1). Isla Floreana, Isla Onslow (LACM 30144,1 body). Isla San Salvador, ex p isce (AMNH 139043.1). San Salvador, Bahia James (LACM 34-273.7,1; SBMNH no #, 1). San Sal vador; 1 mile W o f BaMa James (LACM 66-210.14,1). Baltra IsL (AMNH 156380,1). Femandina, Punta Espmosa (LACM 152363,3; LACM 72-197.1,2). Espagnola, Bahia Gardner (LACM 34-78, 10). Isabella, Punta Albemarle ^A C M 11732,1). Santa Maria, 0.5 miles S o f Playa Negro ÇACM 34-3003,1). Santa Maria, NW o f Bahia Post OfiBce 289 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (LACM 3 4 -7 2 2 ,1). Isabella, Caleta Tabus (LACM 34-20.12, 1; LACM 34-21.9, 1). colombia; N o f Isla Gorgona (LACM 38-119.5,1 + 1: complete) LTTER A rU R E, LOCALITIES: Charles Isl. (DaU, 1889: as Kpotartalesii ?). LITERATURE, RANGE: Gorgona Isl., Colombia; Galapagos Isl. (McLean, 1971). Galapagos, W Colombia (Abbott & Dance, 1983). Gorgona Isl. - Galapagos (Finet, 1993). Haliotis fulgens Philippi, 1845 (Figures 4-90,93,100,173) Washington: Straits of Juan de Fuca (AMNH Edison A9853, 1 [?]). California: Saa Luis Obispo County, Moro Bay (DMNH 095515, 1). San Luis Obispo County, Oceano (DMNH 095520, 1). San Luis Obipo County, near Cambria, LefEingweil Landing (LACM 47-72,1). Santa Barbara County, N Point Conception (AMNH Edison A9853, 4). Santa Barbara County, Santa Barbara (LACM 146806, 1). Santa Barbara County, Goleta (DMNH 090340, 1; AMNH Edison A9853, 1). Santa Barbara County, Coal Oil Point (SBMNH 23448, 5). Santa Barbara County, Santa Cruz Esi. (LACM 40-163.6, 1: complete; AMNH Edison A 9853,1). Santa Barbara County, N Santa Cruz Isl. (SBMNH 12589, 1). Santa Barbara County, Santa Cruz Isl., Smuggler’ s Cove (SBMNH 28302, 1). Santa Barbara County, Santa Rosa Isl. (ANSP 50151,1; USNM 73589,2). Ventura County, Point Mugu NA..S (AMNH 254136,5). Los Angeles County, Santa Catalina Isl., Isthmus Cove (LACM 71-99.14,3). Los Angeles County, Santa Catalina Isl., Iron- bound Cove (LACM 69-31.9, 3). Los Angeles County, Santa Catalina Isl., N Avalon (LACM 62-5.8,2). Los Angeles County, Santa Catalina Isl., Catalina Harbor (LACM 146805,2). Los Angeles County, W Santa Catalina Isl. (LACM 68-25.7, 3; LACM 146812,1). San Clemente Isl. (AMNH 100803,1; AMNH220184,2). SE San Clemente, Pyramid Point (LACM69-29.13,1; LACM 39-129.8,1). SW San Qemente Isl. (AMNH 290 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 100814, U USNM 684686, I). Los Angeles County, M dibu (SBMNH 28175, 1). Los Angeles County, 2 miles NE Palos Verdes (DMNH 034839, 1). Los Angeles County, Palos Verdes, Portuguese Bend (LACM 146835, 4; LACM 65-80.6, 1). Los Angeles County, off Palos Verdes Estate (DMNH 111167,1). Los Angeles County, Palos Verdes, Flatrock Point (LACM 60-29.1, I). Los Angeles County, Palos Verdes, Bluff Cove (LACM 31927,1;AMNH 198679,1). Los Angeles County, San Pedro (DMNH 066015, 1; LACM 59332,7). Los Angeles County, Palos Verdes, Lunada Bay (LACM 60-13.8, 6 ; LACM 64-25.5, 2). Los Angeles County, S Redondo Beach. (LACM 41-217.2,1: with dried body; AMNH 100822,4). Orange County, San Onofre (LACM 69-363,1: with dried body). Orange County, Laguna Beach (DMNH 119863, I; LACM 146809, 6). Orange County, Anaheim Landing (LACM 146798, 1). Orange County, Corona del Mar, S Newport Jetty (LACM 69-73,1). Orange County, Newport (DMNH 066026,1; AMNH 111941,1). San Diego County, Pointe Loma (AMNH Jackson 1992 A9873,3; SBMNH no #, 2). San Diego County, La JoUa (DMNH 136744, 1; LACM 54785, 1: complete). San Diego County, La Jolla, Boomer Beach. (LACM 146807, 1). San Diego County, La JoUa, Devil’s Slide (SBMNH 29017,1). San Diego County, La Jolla, Bird Rock (SBMNH no #, 3; SBMNH no #, 4). San Diego County, San Diego (AMNH 100828, 6 ; ANSP 50511, 3). Mexico, Baja California, Pacific Coast: N Coronado Isl. (SBMNH 15227, 1). SE Isla San Geronimo (LACM 67-632,3; SBMNH 13524, 1). E Isla San Martin, Hassler Cove (LACM 68-313,2). San (Quentin Bay (USNM 34497, 1). N Baja, S of San C^umtin, El Consuelo (AMNH 254131,1). Todas Santos Btty (ANSP 50512,2; USNM 740978,1). Punta Colnett (LACM 46-142, 3). Punta Banta (DMNH 053919, 1; SBMNH 43471,1). S Punta Banda, Arbolito (LACM 67-44.5,1). Punta Banda, Punta Vanda Beach (SBMNH 14627,1). Punta Conception (SBMNH 140220, 1). 10 miles N of Ensenada (AMNH 85560,1). Ensenada (LACM 146814, 1, AMNH 291 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 183124, I). N Ensenada at K-IOO (LACM 64-32.5, I). Turtle Bay (= Bahia San Barolome) (LACM 146813,3; SBMNH 18250,2). Bahia Thurloe (LACM 146822, 6). Santa M aria Bay (USNM 2642111, 1). Punta Hughes, N Bahia Santa Maria (LACM 67-69.3, 12). E Punta Hughes, Bahia Santa Maria (LACM 67-68.2,2). Magdalena Bay (ANSP 151204,1; ANSP 152214,2). Magdalena Bay, Punta Entrada (LACM 71-184.7, 2). Magdalena Bay, Magadalena Isl., S side o f Hughes Point (AMNH 77554, I). îvfag- dalena Bay, Magdalena Isl., NE Punta Belcher (LACM 67-72.4, 1; LACM 6 7 -7 0 3 ,1). Magdalena Bay, Man of War Cove (LACM 71-183.9,4). N entrance Bahia Magdalena, Sail Rock (LACM 71-14.15,7). S Punta Pequefia (LACM 71-181.6,3; LACM 79-253, 2: with bodies). NE Punta Abreojos (LACM 50-323, 1). Punta Abreojos (LACM 71- 3.9, 1). Islas San Benito, San Benito del Este (LACM 50-23.4, 1). 20 miles E Punta Eugenia, Hancock Cove (LACM 51-45.2, 1). Islas San Benito, W Benito del Centro (LACM 50-89.3, 5). W San Benito Isl. (USNM 264398,4; USNM 265600, 1). Near Santa Rosalia (LACM 146816,2). Isla Cedros, S bay landing (LACM 41-35.1, 3). NE Isla Cedros (LACM 71-151.11,4). Islas Cedros, N Pueblo de Cedros (LACM 67-65.9, 1). E Isla Cedros, Viscaino Bay (LACM 146818,1). Cerros Isl. (ANSP 50530,2; USNM 6209, 1). E Isla Asuncion (LACM 67-66.11,1). San Juanico (SBMNH 17154, 1). La Paz (USNM 533690,3; USNM 60430,1). Punta Rompiente (LACM 71-161.8,1). Mex ico other: Guadalupe Isl. (DMNH 066029, 1; LACM 146820, I). N Guadalupe Isl. (SBMNH 17589,5). S Guadalupe Isl. (AMNH Edison A9853, 1). Guadalupe Isl., Five Fathom Bank, Isla Adentro, Isla Afuera (LACM 72-120.5, 1). Guadalupe Isl., S Tip, Mono Sur (USNM 660565,1). Guadalupe Isl., N Anchorage (SBMNH 38687,2). Gtqra- mas (AMNH Edison A9853, 4). Acapulco (NMBE ex 1329.1420, I: doubtfW, not mapped). 292 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE, LOCALITIES : GuH^ La Paz (Bartsch, 1940). Bahia Tortugas; Isla Natividad (Munoz-Lopez, 1976). LITERATURE, RANGE: Monterey Bay - Lower California (Oldroyd, 1927). Farai lon Isl. - G ulf o f California (Bonnot, 1948). Farailon Isl. - Gulf o f California (Abbott, 1954). Point Conception - Magdalena Bay (Cox, 1960; McLean, 1978; Haaker ef aL, 1986). Haliotis k, kamt schat kana Jonas, 1845 (Figures 4-87,94,97) Alaska: Ketchikan County, S Revillagigedo Isl., Ketchikan (LACM 62-200.7, 2; LACM 11737, 3). Ketchikan (AMNH 220196, 2; MNHN no #, I). 20 miles N o f Ketchikan, Behm Canal, S Cedar Isl. (AMNH 198688, 5). Ketcuikan (AMNH 173857, 1). Prince o f Wales County, Craig (LACM 146827,4; LACM 34585,1). Prince o f Wales County, Prince o f Wales Isl. (LACM 146831,2). Prince o f Wales County, W Prince o f Wales Isl. (AMNH 198693,22). Kenai-Cook Inlet County, Kenai Peninsula, Cook Inlet, Kachemak Bay, W Ksitsna Bay, Kenai-Cook Inlet County, 59" 29’ N 151® 37’ W (USNM 894178, 1). McDonald Spit (LACM 83-104.19, 4). Fort Wrangell (USNM 777785,5; USNM 132950, 1). Sitka County, Sitka (DMNH 112684, 1; LACM 110444, 2). Sitka County, Sitka, Berry Isl. Reef (LACM 10529, 5). Sitka County, Alexander Archipelago, Sitka Sound, Japonski IsL (LACM 63-85.1, 1). Sitka County, Sitka Sound (LACM 146823,2; LACM 146825, 1). Sitka County, Sitka Sound, Biorka Isl. (USNM 160556, 3; SBMNH 14166, 7). Sitka County, Baranoff Isl., Sitka Sound, Pirate Isl. (LACM 73-15.6,6). Sitka County, Baranoff Isl. (LACM 146826,1). Sitka County, W o f Sitka, Kruzof IsL (SBMNH 14172,5), Noyes Isl. (AMNH 198692,2; ANSP 252561, 1). Gravina IsL (DMNH 112633,2; DMNH 098712, 1). Annette IsL 0)MNH 131453, I; ANSP 182730,2). Annette Isl., Metlakatka (DMNH 131567,2; AMNH 100829,4). 293 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Kodiak Isl. (AMNH 121675, 1). Fox Isl., Unalaska IsL, Dutch Harbor (AMNH 86802, 3). Forrester Isl. (USNM 216402, 2; SBMNH 3404, 3). Dali IsL, Rose Inlet (USNM 603181, 4). Canada, British Columbia: Prince Rupert (SBMNH 14173, 2; SBMNH 14169, 8). ()ueen Charlotte Isl. (LACM 34278,2; LACM 146833, 1). Queen Charlotte IsL, S end o f Moresby Isl., Dolomite Narrows (LACM 66-67.2, 1). Queen Charlotte Isl., S end Moresby Isl., Rose Harbour (LACM 69-54.2, 2; LACM 69-54.1, 1 body). Queen Charlotte Isl., Kunghit Isl., Rose Harbour (LACM 146824, 2; USNM 58845, 12). Queen Charlotte Sound (SBMNH 29525, I). Queen Charlotte Isl., Graham Isl. (ANSP 152379, 2). Queen Charlotte Isl., Langara Isl. (A N SF152367, 1). Queen Char lotte Isl., N Lnagara Isl. (LACM 60-240.1, 1). Vancouver Isl., Victoria Breakwater (DMNH 087871,2; AMNH 249113,1). Vancouver Isl., SE Victoria Breakwater (LACM 66-63.2, 3; SBMNH no #, 3). Vancouver Isl., near Victoria, McNeill Bay (LACM 73- 35.4,4). \%ncouver Isl., Victoria (LACM 10341,15; SBMNH 1159, 1). Vancouver Isl., Victoria harbor (DMNH 100778,2). Vancouver Isl., Johnstone Strait side, Seymour Narrows (LACM 73-39.2, 1). Vancouver Isl., Barkley Sound (DMNH 111993, 1; DMNH 111992, 1). Vancouver Isl., Barkley Sound, off Bauke Isl. (DMNH 111996, 1). Vancouver Isl., Barkley Sound, Broken Group, Effingham Isl. (LACM 34807,2). Van couver Isl., Ucluelet (AMNH 160605, 3; SBMNH no #, 1). Vancouver Isl., Nootka Sound (BMNH 553.4.131,1). Vancouver, Nootka district, Nuchatlitz Inlet, Louise Bay entrance ^A C M 73-383,14+2 dried bodies). Vancouver, Nootka district, S Nuchatlitz Inlet, Louise l%oon (LACM 60-25930,3). Vancouver Isl., Ruppert District, Hope Isl., entrance Bull Harbor ^ C M 63-2834,15; LACM 63-283, 3 bodies). Vancouver Isl., Rupert District, Kyuquot Sound, IJnfon laL, W White Cliff Head (LACM 60-253.11,1). W Vancouver Isl., Alyquot [= Clayoquot?] (USNM 60293,8; USNM 777784,4). Ocean Fall District, Birdswell IsL, Campbell IsL, New Bella Bella (LACM 82-44.1,3). Calvert 294 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Isl. (AMNH 100831, 1; NHB 6593c, 3). Calvert Isl., Mouth o f River’ s Inlet (AMNH 198687,5). Fitz Hugh Sound (AMNH 104177,4; AMNH 198693,6). M UI Bank, Price Isl. (AMNH 198691, 7). Washington: San Juan Isl. (AMNH 240633, 1; USNM 85343, 2). San Juan Isl., W end Cactus Isl. (AMNH 88075, 8; AMNH 88075a, 1). San Juan IsL, Lonesume Cove (AMNH 108026, 1). Olympic Peninsula, Clallam County, Neah Bay (DMNH 112624, I; NHB 6593d, 1). California: Humboldt County, Point Delgada (LACM 7I-I07.1, 1). Mendocino County, S Shelter Cove (60 miles S o f Eureka), Tolo Bank (LACM 71-106.1,4). San Francisco County, SE Farailon Isl. (LACM 62-9.5,16). Monterey County, Monterey Bay, off Del Monte (LACM 60-22.6, 2; LACM 64-232.1, 1: with preserved epipodium and radula). Monterey County, S Monterey (LACM 146837, 5; LACM 146838,3). Monterey County, N San Jose Creek Beach, Carmel Submarine Canyon (LACM 60-24.13, > 50; LACM 60-24.14, 2 bodies). Monterey County, Monterey, Coastguard Breakwater (LACM 63-6.2, 3 bodies; LACM 68-30.1, 8). Monterey County, Pacific Grove (LACM 54409, 16). Santa Cruz County, Gray- hotmd Rock to Ano Nuevo Point, Waddell Beach (LACM 67-95.9, 2). Santa Cruz County, Santa Cruz (ANSP 219590,2). San Luis Obispo County, between Point Buchon and Point San Luis (DMNH 095737,2; DMNH 112620,1). San Luis Obispo County, Moro Bay, off Point Estero (DMNH 11977,1). San Luis Obispo County, off Cayucos (ANSP 183595,1). LITERATURE, LOCALITIES: Point Estero (Cox, I960), (^ueen Charlotte Isl., Hippa Isl.; Louise IsL; Lyell Isl.; Juan Perez Sound; Vancouver Isl., Clayoquot Sound; Nanoose Bay; Alert Bay; Main Coast, Johnstone Strait; Goose Isl. Group; Aristazabal Isl.; Caa- mano Sound; Estevan Group; Banks Isl.; Porcher Isl. (Sloan & Breen, 1988). LITERATURE, RANGE: Sitka - Redondo (Oldroyd, 1927). S Alaska through Point Conception (Abbott, 1954). Sitka - Point Conception (Bonnot. 1948; Cox, 1960). Sitka 295 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. - Monterey (Haaker et al.y 1986). Icy Strait, N tip o f Sitka - Baja California (Sloan & Breen, 1988). Halioti s kamt schat kana assimili s DaU, 1878 (F^ures 4-95,4-99,4-100) Washington: San Juan County, Cactus Isl. (DLG AAB 36c, 2). California: Montery County, Monterey (USNM 60297, 3; USNM 56470, 1). Montery County, S o f Mon terey (SBMNH no #, 1). Montery County, Carmel Bay (SBMNH 10348, 1). Monterey County, Hollister, off Coxo Point (AMNH 220145, 3). San Luis Obispo County, Moro Bay (DMNH 066027,2; DMNH 095518, 1). San Luis Obispo County, 10 miles N of Moro Bay (SBMNH 22903,2). San Luis Obispo County, off Moro Bay (LACM 45816, 1). San Luis Obispo County, Point Estero (LACM 146842, 4; LACM 146846,4). San Luis Obispo County, Cayucos (LACM 146839,2; AMNH 198695,18). San Luis Obispo County, o ff San Simeon (LACM 146740,2). San Luis Obispo County, Point Buchon (AMNH 115300, 2). San Luis Obispo County, S Point Buchon (LACM 146847, 5; LACM 146843, 3). San Luis Obispo Coimty, Cambria (LACM 146849,2). San Luis Obispo County, off Port San Luis Obispo (USNM 535850,2). Santa Barbara County, Pt. Arguella, Rocky Pomt (DMNH 124214,1). Santa Barbara County, off Jalama State Beach, between Pomt Conception and Point Arguella (SBMNH no #, 1). Santa Barbara County, 5 miles N o f Point Conception (AMNH 198690, 1). Santa Barbara County, off Point Conception ^M N H 095623, 1; DMNH 112617,1). Santa Barbara County, Near Point Conception, Ghost Reef (AMNH 186255, 1). Santa Barbara County, near Point Conception, Coho Anchorage (AMNH 220143,3; ANSP uncataloged, 2). Santa Bar bara County, between Coal Oil Point and Naples (SBMNH 22177,1). Santa Barbara County, Goleta, Coal Oil Point (SBMNH 28323,2). Santa Barbara County, Santa Bar bara, Goleta (AMNH Edison A9853,3). Santa Barbara County, N Santa Barbara (LACM 296 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 146848.1). Santa Barbara County, Santa Rosa Isl. (AMNH 198693,4; USNM 877517, 5). San Clemente Isl. (AMNH 220144, 7; SBMNH no #, 1). O ff San Clemente Isl. (AMNH 100820,1). Santa Barbara County, Santa Cruz Isl. (USNM 98329, 1; SBMNH 18903, 1). Los Angeles County, Redondo Beach (DMNH 111982, 1; LACM 146855, 1). Los Angeles County, Hermosa Beach (LACM 146850,1). Los Angeles County, Par adise Cove (LACM 66-58.6,2). Los Angeles County, Palos Verdes Peninsula, Whites Point (LACM 146853, 4; LACM 35602,6). Los Angeles County, San Pedro (AMNH 254140, 2; USNM 181130, 1). Los Angeles County, San Pedro, off White’ s Point (SBMNH 23985, I). Los Angeles County, Portugese Bend (SBMNH 24709, 1). Los Angeles County, Santa Catalina Isl. (AMNH Edison A9853,2). Los Angeles County, S Santa Catalina IsL, Farnsworth Bank (LACM 65-183, I with dried body; LACM 67- 163, 1 with dried body). Orange County, Newport Beach (AMNH Edison A9853, 1; SBMNH 23131,2). San Diego County, San Diego (LACM 70228,2; AMNH 117368, 1). San Diego County, San Diego, 1 mile off Bird Rock (LACM 146856,2). San Diego County, San Diego, Point Loma (LACM 23456, 2; LACM 147291, 1). San Diego County, San Diego, La Jolla (LACM 72-109.6,4; LACM 46-16.4,1: complete). San Diego County, San Diego, Mission Bay (AMNH Edison A9853,2). Mexico, Baja cali fornia: Pacific Coast, S Punta Banda, Arbolito (LACM 67-52.1,5; LACM 63-42.7,2). BC del Norte, Punta Banda, La Bufadora (SBMNH 40237,2). Pacific Coast, S Puerto Santo Tomas (LACM 67-49.1,2; LACM 67-1.10,9). Santa Tomas Bay (SBMNH no #, 1). Bahia Tortuga (LACM 11271, 13). Turtle Bay (= Bahia San Barolome) (AMNH 198697,1; AMNH 254134,1). Pacific Coast, 4 miles off Isla San Martin, Six-and-a- half Fathom Bank (LACM 71-1483,1). Islas Coronado, Coronado del Norte (LACM 146817.1). Mexico: Mazatlan (AMNH Edison A9853,1). 297 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE, LOCALITIES: Farallones; S o f Carmel, Yankee Point; Seaside (Smith & Gordon, 1948). Point Loma (Cox, I960; MuIIiner, 1984). LITERATURE, RANGE: Faralloa Isl. - San Diego (Oldroyd, 1927). Monterey County - San. Diego (Bartsch, 1940). Farailon Isl. - San Diego (Bonnot, 1948). Farailon Isl. - San Diego (Abbott, 1954). Point Conception - Turtle Bay (= Bahia San Bartholome) (Cox, 1960). San Luis Obispo County - Bahia Tortugas (Haaker et aL, 1986). ffaliotis robert i McLean, 1970 (Figures 4-107,4-109,4-112,4-167,4-168) Cocos Isl.: (DLG AAB 41a, 1). Near Isla Bautista, off Bahia Inutil, SE o f Bahia Igle sias (LACM 151184, 2). Bahia de Chatham (AMNH 276890, 1; AMNH 232184, 5: with (fried bodies). LITERATURE, RANGE: Cocos Isl. (McLean, 1971). ffa lio ta ruf escens Swainson, 1822 (F%ures 4-96,4-99,4-100) California: Humboldt County, Shelter Cove (LACM 70-70.4,4). Mendocino County, Cape Mendocmo (AMNH 254135,1). Mendocino County, 60 miles S o f Eureka, Shel ter Cove (AMNH Jackson 1995 A9873, 1). Mendocmo County, Fort Bragg (DMNH 182043,1; AMNH 183027,3). Mendocino County, Mendocmo (ANSP 7942,4). Men docino County, Punta Arena (LACM 49-5.3,42). Mendocino County, near Albion, Whitesboro Cove (LACM 49-7.3,3). Mendocmo County, near Albion, Cormorant Cove (LACM 49-4.4, 8). Mendocino County, Gualale River (USNM 543615,3). Mendocmo County, Gualala (USNM 98326,2; USNM 98327a, I). Mendocmo County, Van Damme Beach State Park, N Little River Cove (LACM 49-8.1, I; LACM 64-8, I). Marin County, Point Reyes (LACM 17835,2). Sonoma County, Fort Ross, Cementary Cove (LACM 50-11.2,1). Sonoma County, N o f Russian River, E Beach ^A C M 50-62,1; 298 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Haliotîs fiilgens -5 H àliotis cracherodîi H aliotis corrugata : H aliotis kamtschatkana assim ilis H aliotis rufescens Haliotis sorenserti Figure 4-100. Distribution o f east Pacific species o f H aliotis spp. (I). Localities stem ming from collection specimens are indicated with a solid circle, those from the litera ture with a solid square. For data see text 299 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. % LACM 4 8 -4 7 2 ,1). Sonoma County, Bodega Head (LACM 48-5.6,1; LACM 49-113.1, 1). Sonoma County, Bodega Bay, Horseshoe Cove (LACM 49-108.1,4; LACM 49- 38.1, 3). Sonoma County, Shell Beach (LACM 50-842,7). Sonoma County, Stewart’ s Point (LACM 50-10.3, 8; LACM 50-7.3,21). San Mateo County, Pillar Point (AMNH 154937, 8). San Mateo County, Moss Beach (LACM 63-1.4,2; SBMNH 5555, 6). San Mateo County, Moss Beach, Kelp Cove (AMNH 89182,1). San Mateo County, Franklin Point (LACM 62-8.6, 1). San Francisco County, Farailon Isl. (DMNH 111168, 1). San Francisco County, San Francisco (AMNH 19735, I). San Francisco County, Pigeon Point (AMNH 99375, 1). San Francisco County, Purissima (USNM 98313, 1). Santa Cruz County, Grayhound Rock to Ano Neuvo Point, Waddell Beach (LACM 67-95.10, 3). Santa Cruz County, Beach near Santa Cruz (ANSP 170249, 2). Monterey County, Big Sur, S Lighthouse Rock (LACM 47-16.10,1: complete). Monterey County, Pacific Grove (LACM 59340, 5; LACM 59-12.8,2). Monterey County, Pacific Grove, Point Joe (LACM 146861,1). Monterey County, Pacffic Grove, Asilomar State Park (DMNH 120827, 4). Monterey County, Monterey (USNM 60433, 2; USNM 31267, 3). Mon terey County, Monterey Bay, o ff Cannery Row and Hopkins Marine Station (LACM 60-14.4,15). Monterey County, Carmel (ANSP 155146,3). Monterey County, Carmel, N side o f Mission Point (DMNH 035655, 1). Monterey County, N o f San Jose Creek Beach, Carmel Submarme Canyon (LACM 60-24.15, 1). Monterey County, Carmel, N side o f Mission Point, o ff Carmel River State Beach (DMNH 114896, 6; DMNH 114914, 8). Monterey County, Monterty^ (LACM 110621,3; LACM 146863, 1). Mon terey County, Monterey, Coast Guard Breakwater (LACM 54784,3: complete; LACM 68-30.2,3). Monterey County, Monterey Peninsula, Pomt Pinos ÇDMNH 036487, 1). Monterey Cotmty, Soberanes Point (LACM 64-12.1, 2). San Luis Obispo County, Piedras Blancas (DMNH 119434,2; AMNH 220233,4). San Simeon (AMNH 100799, 300 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2). San Luis Obispo County, Cambria (AMNH 85472, 1; SBMNH 230, 1). San Luis Obispo County, Cambria Pines (SBMNH 23722, 2). San Luis Obispo County, N Cayu- cos (LACM 46-35.5,1; LACM 77-112.4,2). San Luis Obispo County, Cayucos, Garcia Ranch (SBMNH 46553, 1). San Luis Obispo County, Moro Bay (DMNH 010955, 4; DMNH 066020,2). San Luis Obispo County, Moro Bay, Point Estero (AMNH 276884, 1). San Luis Obispo County, S Shell Beach (LACM 68-36.6,1). Santa Barbara County, Point Conception (LACM 62364,7; SBMNH 28305, 1). Santa Barbara County, 5 miles SE o f Point Conception (LACM 6339.10,2). Santa Barbara County, Point Conception, Coho Anchorage (AMNH 198678, 1; AMNH 175668, 1). Santa Barbara County, Santa Barbara (SBMNH no #, 5). Santa Barbara County, Santa Barbara Isl. (LACM 23451,1; ANSP 50155,1). Santa Barbara County, Santa Barbara, Goleta (AMNH Edison A9853, 2). Santa Barbara County, Goleta, Coal Oil Point (SBMNH 231,3; SBMNH 23447,1). Santa Barbara County, Jalama (SBMNH 28310, 1). Santa Barbara County, off Jalama (SBMNH 28306, 1). Santa Barbara County, Tajiguas (LACM 63-36.7, 1; SBMNH 235, 1). Santa Barbara County, Santa Rosa Isl. (SBMNH 12590,2). Santa Barbara County, S Santa Cruz Isl., W Yellowbanks (LACM 69-32.8, 1). Santa Barbara County, off San Miguel Isl., Wilson Rock (LACM 82-59.9,2: complete). Santa Barbara County, San Miguel IsL SE o f Bay Point (LACM 67-38.8,2; LACM 67-38.8, 1 radula + some tis sue). Santa Barbara County, NW San Miguel IsL (LACM 82-58.5,4). Ventura County, San Nicolas Isl. (DMNH 119114,1). Ventura County, S San Nicolas IsL (LACM 69- 14.6). Ventura County, Oxnard, Channel Isl. Marina (SBMNH 179,1). San Clemente Isl. (AMNH 220234, 2). San Clemente Isl. (AMNH 83987, I). O ff Anacapa Isl. (SBMNH no #, 6). Los Angeles County, N Malibu (AMNH 240637,2). Los Angeles County, o ff San Pedro (LACM 18156, I). Los Angeles County, San Pedro (AMNH 100800, 1 ; ANSP 50507,2). Los Angeles County, Pt. Fermia Lighthouse (LACM 55- 301 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 24 .5 .1). Los Angeles County, Catalina Isl. (AMNH 71334,3). Los Angeles County, off Catalina Isl. (SBMNH 28151,1). Los Angeles County, W Santa Catalina Isl. (LACM 146841,2). Los Angeles County, Paradise Cove (LACM 66-58.7, 1). W hite’s Point (SBMNH 177,1). Orange County, Newport Bay (AMNH 121687, 1; AMNH 198675, 2). Orange County, Corona del Mar (SBMNH 13689,1). Orange County, Laguna Beach (MHNG 21807,2). San Diego County, Del Mar (AMNH Jackson 1995 A9873,2). San Diego County, La Jolla (LACM 72-109.7, 2; AMNH Jackson, 1995 A9873, 4). San Diego County, La Jolla, Bird Rock (SBMNH no #, 5). San Diego County, Point Loma (SBMNH no #, 1; SBMNH no #, 2). San Diego County, San Diego (ANSP 50506, 2; USNM 522562, 2). Mexico, Baja California: N Baja, Ensenada (AMNH 284471,2; 284470, 3). Pacific Coast, Punta Banda (LACM 63-42.8,2; SBMNH no #, 2). Pacific Coast, S o f Punta Banda, Arbolito (LACM 67-47.6,5; LACM 67-53.4,3). Punta Banda, Kennedy Cove (SBMNH no #, 1). Punta Santa Tomas (SBMNH no #, 2). Pacific Coast, 18.5 miles W o f Santo Tomas, Puerto Santo Tomas (LACM 67-46.9, 1 + 1 with dried body; LACM 67-1.11, 2). E o f Cedros Isl. (AMNH 97182, 1). Pacific Coast S o f Isla San Geronimo (LACM 71-91.9,2). Pacific Coast, E Isla San Geronimo (LACM 67- 61.9.1). LITERATURE, LOCALITIES: California, Crescent City (Bonnot, 1930). Between Piedras Blancas and San Simeon; S White Rock No. 1; S Harlan Rock; N Carmel Bay; S Point Conception; NE and SW San Miguel Isl.; N P t San Luis; 5 miles N Point Estero (Bonnot, 1940). Point Hueme (Friedman e t a/., 1993). Santa Cruz Isl. (Davis et aL, 1996). LTTERATURE, RANGE: Bodega B ay- La Paz (Oldroyd, 1927). Mendocino County and S (Bartsch, 1940). N CA, Point Saint George - La Paz (Bonnot, 1948). Oregon, 302 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Sunset Bay - Turtle Bay (= Bahia San Bartholome) (Cox, I960). Oregon, Sunset Bay - Bahia Tortugas (Haaker et al., 1986). H aliotis sorenseni Bartsch, 1940 (Figures 4-100,4-101,4-104) california: San Luis Obispo County, Moro Bay (LACM 146852,10; DMNH 095517, 1). Santa Barbara County, off Point Conception (LACM 67901, I). Santa Barbara County, N Point Conception (DMNH 034815, I). Santa Barbara County, S o f Point Conception (LACM H-3418, I). 18 miles S of Point Conception (LACM 146867, 1). Santa Barbara County, Santa Barbara Isl. (LACM 17834,1). Santa Barbara County, off Santa Barbara Isl. (SBMNH 16192, 1; SBMNH 18826, 1). Santa Barbara County, E Santa Barbara-Isl. (LACM 72-97.9,5). Santa Barbara County, Santa Cruz Isl., off Smug glers Cove (AMNH 175672,1; SBMNH 28303, 1). Santa Barbara County, S Santa Cruz IsL W o f Yellowbanks (LACM 69-32.9,5). Santa Barbara County, off Santa Cruz Isl. (SBMNH 18902, 1). Santa Barbara County, Santa Cruz Isl., Pmnacle reef (SBMNH no #, I). Santa Barbara County, Santa Rosa Isl. (SBMNH 166, 1). Ventura County, Point Hueme, Harbor (AMNH 240640, 1). Los Angeles County, off Point Dume (LACM 146868, 1). Los Angeles County, Palos Verdes, off Malaga Cove (DMNH 118516,1). Los Angeles County, Tanner Bank (LACM 69-35.4,2). Los Angeles County, off San Pedro (LACM 17823,2). San Clemente Isl. (LACM 146865,1; LACM 21843,1). S San Clemente Isl. (AMNH 198785, 1; MNHN no #, 1). SE tip o f San Clemente Isl., Pyramid Point (LACM 69-29.14,1). Anacapa Isl. (AMNH 198774,2). S Anacapa Isl. (LACM 64-28.7, 1). Los Angeles County, Santa Catalina Isl., Emerald Bay (LACM 146866.1). Los Angeles County, SW Santa Catalma Isl., Eronbound Cove (LACM 69- 31.10, 1). Los Angeles County, S Santa Catalina Isl., Isthmus, Bird Rock (AMNH 198786.1). Los Angeles County, W end Catalina M . (LACM 68-25.8,2; LACM 68- 303 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-101-106. Shells of Northeastern Pacific abalone I, and unidentified species. Figures 4-101,4-104. H aliotis sorenseni Bartsch, 1940. LACM 146869. S. o f Point Conception, California. 24.1.1940. Ex. Huffinan. 159 mm. Figures 4-102,4-105. H ati - Otis w alallensis Steams, 1899. LACM 62-15. Albion, Mendocino County, California. Leg. J. H. McLean 11. XI. 1962. Intertidal. 110 mm. Figures 4-103,4-106. H aliotis sp. MNHN no #. “Envoi de M. de Montigny, Consul de France à Shang Hai, Mars 1854.” 52 mm One o f three very similar specimens o f an cmidentified species with affinities to the Australian H rtibra^ but with strong folds in shell and Shang Hai locality data. 304 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 25.8.1). Los Angeles County, off S Catalina IsL, Farnsworth Bank (LACM 70-27.4,1; LACM 67-16.4,4). Channel Isl., Cortes Bank, S o f Bishop Rock (LACM 69-34.8, 2). Orange County, Newport Beach (ANSP 252572, I; AMNH 240639, I). San Diego County, Point Loma (SBMNH no #, I). Mexico, BAja California, PACIfic Coast: S o f San Diego, Isla Coronados (LACM 71-98.6, 1). S o f Ensenada (LACM 146870, I). Punta Banda, La Bufadora (SBMNH 6190,1). S Punta Banda (LACM 67-53.5, 8; 67- 4 4 .1 ). SW Isla Natividad (LACM 72-117.9, 1). Isla Natividad, 1 ml. SW o f Twelve Fathom Bank (LACM 71-165.6,9). Bahia San Bartolomé, near Kelp Point (LACM 54- 13.4, 3). N end o f lia San Geronimo (LACM 67-62.6, 1). Punta San Pablo (LACM 71- 178.13, 8). E Isla San Martin, Hassler Cove (LACM 68-31.6,2). Isla San Martin, South Anchorage (LACM 67-60.7, 1). Isla Asuncion, E anchorage (LACM 67-66, 3). Isla Cedros (LACM 71-156.6, 1; AMNH 254137, 1). Thurloe Head (LACM 71-170.8, 2). Punta Rompiente (LACM 71-161.9, 5). San Felipe (DMNH 111162, 1). LITERATURE, LOCALITIES: N Point Conception; San Diego; Santa Cruz Isl. (Cox, 1960). Isla Guadalupe (Chace, 1966). LITERATURE, RANGE: Point Conception - Astmcion Isl. (McLean, 1978). Point Conception - Bahia Tortugas; Channel Isl. (Haaker et oL, 1986). Haliotis wal aUe ns i s Stearns* 1898 (Figures 4-100,4-102,4-105) Washington: Abalone Point, near Westport (AMNH 182797, 1; SBMNH 2943, 1). Oregon: Coos County, Coos Bay (LACM 42-34.2,1). California: Del Norte Cotmty, Crecent City (USNM 104142,5). Humboldt Cotmty, Eureka (DMNH 012826,1; AMNH 220266.1). Humboldt County, Shelter Cove (LACM 70-70, 1). Sonoma County, Fort Ross (SBMNH 43470,2). Sonoma Cotmty, 0 5 miles S o f Fort Ross (LACM 63-573, 2). Sonoma Cotmty, Bodega Bay, Stewart’ s Point (LA(]M 50-7.9,7; 112722,5). Sonoma 305 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. County, out o f Bodega Bay (DMNH 11246, 1). Sonoma County, N Russian River, E beach (LACM 4 8 -4 9 2 ,1). Sonoma County, Salt Point Ranch (LACM 64-6 3 ,5; LACM 64-203, 6 ). Mendocino County, Fort Bragg (DMNH 112644, 1; ANSP 257076, 2). Mencocino County, S entrance Noyo Anchorage (LACM 71-140.1, 1). Mendocino County, Albion (LACM 62-15,4; SBMNH 10331,3). Mendocino County, near Albion, Cormorant Cove (LACM 49-4.8,2). Mendocino County, near Albion, Witesboro Cove (LACM 49-7.4,3). Mendocino County, Ptmta Arena (LACM 146873,1; LACM 49-5.4, 3). Mendocino County, Gualala (AMNH 100830, 1; USNM 635437,1). Mendocino County, Mendocino, Agate Beach (SBMNH 43472,3). Mendocino County, Little River, Van Damne State Park (LACM 64-9.1,2; LACM 64-8.4,1). Monterey County, 1 1 miles S o f Pacific Grove, off Nankee Point (AMNH 140003,2). Monterey County, off Carmel River state beach (DMNH 074501, 5). San Francisco County, SE Farailon Isl. (LACM 62-9.6, 3). San Mateo County, Moss Beach (DMNH 034748, 1). San Luis Obispo County, N Point Buchon, Montana de Oro state park (DMNH 112654, 1). San Luis Obispo County, San Simeon (AMNH 198686, 1). Santa Barbara County, Arguello, Jalama Park (AMNH 220265,1). Santa Barbara County, off Point Conception (DMNH 112154,1). Santa Barbara County, Coho Anchorage, Point Conception (DMNH 111980, 2). Santa Barbara County, Santa Cruz Isl. (SBMNH 18904,1). Santa Barbara County, Santa Rosa Isl., W end (USNM 877515,1). Santa Barbara County, Santa Rosa IsU Ford Point (SBMNH 6140, 1). Santa Barbara County, Santa Rosa IsL, Hohnson’s Lee (AMNH 198691, 1). Santa Barbara County, San Miguel Isl. (SBMNH 23853,2). Ven tura County, San Nicolas M . (AMNH 254138,2). San Clemente Isl. (AMNH 220263, 2). Orange County (MHNG ex 13166, 1). San Diego County, San Diego (AMNH 220264, 1). San Diego County, La Jolla (SBMNH 23676,1). San Diego County, San 306 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Diego» Mission Bay (AMNH Edison A9853, 1; AMNH 240644,1). Mexico: Baja Cali fornia, Babin Todos Santa (DLG AAB 38a, I). LITERATURE, LOCALITIES: Abalone Point near Westport (Bonnot, 1930). Hop kins Marine Station, Point Cabrillo (Lowry & Pearse, 1973). LITERATURE, RANGE: Westport - Point Conception (Bonnot, 1948; Abbott, 1954). British Columbia - S California (Abbott & Dance, 1983). British Columbia - San Diego (Haaker et al., 1986). Western Atlantic Species (Figures 4-107,4-108,4-111,4-114,4-115) HaUot i s aurantium Simone, 1998 fig u res 4-107, 4-108,4-111) LITERATURE, LOCALITIES: Sâo Paulo, off Ubatuba, 24“ 07’ S 44“ 06’ W; Amapà, off Cabo Orange, Brazil; off Vitôria, Espirito Santo, Brazil; Espirito Santo, E of Ponta do Ubù, 21“ 05’ S 41“ 19’ W; Rio de Janeiro, 21“ 56’ 05” S 40“ 07’ 00” W; Rio de Janeiro, off Cabo Frio, 23“ 05” S 40“ 05’ W; Rio de Janeiro, off Campos Bay, 22“ 06’ 06” S 40“ 08’ 38” W (type locality); Rio de Janeiro, off Cabo de Sâo Tomé, 22“ 27’ 06” S 40“ 30’ W; Rio Grande do Sul; off Conceiçâo (Simone, 1998). Between Vitoria and Salvador, Baja, 20“ 10’ 0” S 39“ 49’ 3” W (Silva & Guerra, 1968: fid e Klappenbach, 1968: as barbouri). Off Vitôria, E. S.; O ff Santos (Rios, 1975). (Rios, 1975,1985,1994) San Pablo, 24“ 18’ S 44“ 50’ W; 24“ 20’ S 44“ 40’ W (Bdappenbach, 1968: as pourtale - sit). HaUot â pourtalesü Dali, 1881 (F%ures 4-107,4-114,4-US) USA: Texas, 185 km SSW of Galveston, West Flower Garden Bank (LACM 71-359.1, 2). Florida, Petersburg (AMNH 100599,1). Florida, Anna Maria Isl. (AMNH 244314, 307 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. % H aliotis w atattem is Haliotis roberti H aliotis dalli H aliotis pourtalesü Haliotis aurantium Figure 4-107: Distribution o f east Pacific (II), tropical Pacific, and western Atlantic species o f H aliotis spp. Localities stemming from collection specimens are indicated with a solid circle, those fiom the literature with a solid square. For data see tex t 308 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3), Florida, Lee County, W Fort Myers (DMNH 103228,2). Florida, Lee County, 90 miles W Fort Myers, 90 fins (AMNH 107598). Florida, SW o f Egmont Key (420 m) (DMNH 103239,2; AMNH 194257,1). Florida, Key Largo, 5.5 miles SE ‘ P a rk er B” at Elbow, 9-100 fins (AMNH 125538, 1). Florida, Sombrero Light (AMNH Edison A9853, 1). Florida, o ff Sand Key, S by E 85 fins, Eoüs Station 338 (USNM 333715,2). 25“ 16’ 55” N 83" 37’ 4 7 ’ W (USNM 833627, 1). LITERATURE, LOCALITIES: Key Largo, 3 miles o f Sand Key, approximately 90 fins (Foster, 1946). Florida, 6 miles E of Sombrero (Aguayo & Jaume, 1947). NE Yuc- tan Peninsula, 21" 21’ N 8 6" 30’ W (Harry, 1966). Cuba, Havana (Sarasua, 1968). 24" 57’ N 83" 40’ W (Guice, 1968). E o f Mississippi Delta, 29" 20’ N 87" 46’ W (Jung, 1968). O ff Suriname, T 35’ N 56" 52’ W (Nijssen-Meyer, 1969). Brazil, OffMaranhao; West Flower Garden Bank, 27" 52’ N 83" 40’ W; off SW Florida, 27" 47’ N 83" 40’ N; 28 Fathom Bank, 27" 53’ N 93" 27’ W; Little Sister Bank, 27" 52’ N 94" 15’ W; Sidner Bank, 27" 57’ N 92" 22’ W; Alderdice Bank, 28" 04’ N 91" 59’ W; Elvers Bank, 27" 50’N 92" 53’ W; Rezak Bank, 27" 55’ N 92" 23’ W (Titgen & Bright, 1985). Venezuela, N State o f Anzoâtegui, S Isla Tortuga (Martinez & Ruiz, 1994). LITERATURE, RANGE: Off Lower Florida Keys (Abbott, 1954). SE United States - Brazil (Abbott & Dance, 1983). 309 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-108, 111, H aliotis aurantium Simone, 1998. MZUSP28.39I, paratype. “À leste da ponta do Umbù, ES [Espiritu Santo: = province] +/- 48 m, mergulho autônomo, sobre Laminaria abussalis, Eurico Cabral de Oliveira Fo. col. l4/x/l986.’ * 16 mm. Fig ures 4-109,112. H aliotis roberti McLean, 1970. DLG AAB 41a. Cocos Island. I l mm. Figures 4-110, 113. H aliotis dalli Henderson, 1915. LACM 72-197.7. Punta Espinosa, Isla Femandina (Narborough), Galapagos. Leg. Richard M. Lease, RV SEARCHER. 2 5 .1. 1972. Intertidal, lava tidepools. 27 mm. Figures 4-114,115. H aliotis pourtalesü Dali, 1881. LACM 71-359.1. West Flower Garden Bank, 185 km SSW o f Galveston, Tœcas. 10 mm. Figures 4-116,117. Haliotis fa tu i. USNM 487953. Tonga. 30 mm. Fig ures 4-118, 121. H aliotis exigua Dunker, 1877. NMW 1955.158.2133. Japan. 21 mm. Figures 4-119, 122. H aliotis planata Sowerby, 1882. Figure 4-119. NMW 1955.158.2124. Philippmes. 39 mm. Figure 4-122. NMW 1955.158.2125. GuadalcanaL Solomon Islands. 33 mm. Figures 4-120, 123. H aliotis dohmiana Dunker, 1863. DLG AAB 49a. Brony, New Caledonia. 43 mm. 310 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. lO S g 311 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-124 - 129. H aliotisjacnenesis Reeve, 1846. Note scales from suture but not entirely reaching row o f tremata with width o f bare space being variable. The coloration o f the shell is extremely variable. Figures 4-124, 127. RP no #. No locality. 12 mm. Dorsal and ventral. Figure 4-125. DLG AAB 45L 1,000 Islands group. Bay o f Jakarta, near Java, Indonesia. 28 mm. Dorsal view o f elongated specimen. Figure 4-126. USNM 423136. Lifu. 15 mm. Dorsal with wide bare space between scales and row o f tremata. Figure 4-128. DLG AAB 45e. N. Samar, Philippines. 19 mm. Dorsal view o f round specimen. Figure 4-129. RP no #. New BritaiiL 20 mm. Dorsal. Figures 4-130, 133. H aliotis squamosa Gray, 1826. DLG AAB 33c. Between Fort Dauphm and Monanten- ina, hW agascan 83 mm. Leg. Marc Girona (Tahiti). Col. Carmen & Michel Blanchard (1960 - 1965). Figures 4-131, 132, 134, 135. H aliotis sem iplicata Menke, 1843. This species displays a wide array o f colors. Figures 4-131,134. DLG AAB 11a. Garden Island, Western Australia. 38 mm. Red color form, dorsal and ventral. Figure 4-132. DLG AAB Ilk . Rottnest Island, Western Australia. 35 mm. Yellow color form. Figure 4-135. DLG AAB 1 Ig. No locality. 50 mm. Green color form. Figures 4-136,137. Hali - Otis rubiginosa Reeve, 1846. BMNH type collection. Type locality: “?*. 43 mm. Fig ures 4-138,141. H aliotis clathrata Reeve, 1846. RP. South New Caledonia. 29mm. Figures 4-139, 140. H aliotis dissona (Iredale, 1929). JK. She 142, Astrolabe Reef, Fiji. 17 mm. 312 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 313 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-142 - 145, 148. H aliotîs midae Linnaeus, 1758. Figures 4-142, 143. DLG AAB 8b. Simons Town, False Bay, South AMca. 95 mm. Dorsal and ventral view of white west coast variation. Figure 4-144. DLG AAB 08d. Port Alfred, South Africa. Dorsal view o f dark east coast variation. Figures 4-145, 148. DLG AAB 08c. No local ly data. Dorsal and ventral o f Juvenile; note marked contrast in sculpure and red tinge in nacre near spindle. Figures 4-146,149. H aliotis marmorata Lmnaeus, 1758. NMW. “Cape o f Good Hope” [doubtful locality]. 44 mm. Dorsal and ventral view. Figures 4- 147,150. H aliotis stomatiaeform is Reeve, 1846. DLG AAB 51a. Malta Island. Dorsal and ventral view. Figures 4-151-153. H aliotis spadicea Donovan, 1808. Figure 4-151. DLG AAB 12d. No locality data. 21 mm. Dorsal view of juvenile specimeiL Figures 4- 152, 153. DLG AAB 12c. Muizenberg, False Bay, South Africa. 6 6 mm. Dorsal and ventral view o f adult Figures 4-154- 156. H aliotis parva Linnaeus, 1758. Figures 4- 154,155. DLG AAB 18g. Tanskei, South Africa. 32 mm. Dorsal and ventral view of typically mottled specunen. Figure 4-156. DLG AAB 18. Cape Town, South Africa. 19 mm. Dorsal view orange color variation. Figures 4-157, 158. H aliotis unilateralis Lamarck, 1822. MHNG 18020: neotype. E lat Smai, Israel. 33 mm. Dorsal and ventral view. Specimen with flat dorsal surfrtce and pronounced ledge. Compare to Figure 4-23 for an other specimen. Figure 4-159. H aliotis speciosa Reeve, 1846. NMW. Algoa Bay. 30 mm. Dorsal. 314 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 315 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 4-160. Haliotis speciosa Reeve, 1846. NMW. Algoa Bay, South Africa. 30 mm. Ventral. Figures 4-161,162. H aliotis queketti Smith, 1910. DLG AAB 52a. Coffee Bay, Transkei, South Africa. 29 mm. Figures 4-163, 164. H aliotis varia Lmnaeus, 1758. Lompok, Indonesia. IV 1988. Photographs and © Henry Chaney. Figure 4-163. Dorsal. Figure 4-164. Ventral. Figure 4-165,166. H aliotis asinina Linnaeus, 1758. Figure 4- 165. Ashmore Reef, Timor Sea. Photograph and © Clay Bryce. Adult specunen. Figure 4-166. Heron Island, Queensland, Australia. June 1983. Photo #100560/61. Photograph and © Bill Rudman. Juvenile specimeiL Figures 4-167, 168. H aliotis roberti McLean, 1970. Bahia Wafer, Isla del Coco, Costa Rica. H 1991. Photographs and © Henry Chaney. Figure 4-167. Lateral view of spawning specimen. Figure 4-168. Dorsal view, on coralline algae. 316 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 317 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 4-169. H aliotîs unilateralis Lamarck, 1822. Hurghada, Egypt 3-4 m under stone at the foot o f a coral reef. Photograph and © Urs W iiest Figure 4-170. H aliotis sp. nov. (Owen et of., in prep.). Greece. Photograph and © Buzz Owen. Figure 4-171. Haliotis unilateralis Lamarck, 1822. E lat Israel. Neotype MHNG 18020. Photograph and © Solly Singer. Figure 4-172. H aliotis tubermlataSAsxaaAxxs, 1758. Ilia d’en Colom, Menorca, Balearic Islands, Spain. 11m . VH-Vm 1989. Slide #370. Photograph and © Daniel L. Geiger. Figure 4-173. H aliotisfulgens Philippi, 1845. Adult specimen in situ, o ff Monterey, California. Photograph and © Henry Chaney. Figure 4-174. H aliotis tuberculata Linnaeus, 1758. Banyuls-sur Mer, France. Slide #87. Photograph and © Daniel L. Geiger. Righting behavior. Figures 4-175, 176. H aliotis clathrata Reeve, 1846. Tulamben, Bali, Indonesia. Under a rock. Photographs and © Karen Gowlett- Holmes. Two specimens shown. 318 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 319 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figures 4-177,178. H aliotis australis Gmeim, 1791. Stewart Island, New Zealand. Pho tographs and © Karen Gowlett-Holmes. Figure 4-177. Under rock, dorsal. Figure 4- 178. Ventral, showing the white sole o f the foot. Figures 4-179, 180. H aliotis iris Gmelin, 1791. Photographs and © Karen Gowlett-Holmes. Figure 4-179. Wellington, North Island, New Zealand. Dorsal. Figure 4-180. Stewart Island, New Zealand. Ven tral, showing the black sole o f the foot Figures 4-181,182. H aliotis virginea Gmelin, 1791. Photographs and © Karen Gowlett-Holmes. Figure 4-181. Potnua, North Island, New Zealand. Two specimens dorsal, under rock. Figure 4-182. Mil&rd Sound, South Island, New Zealand. Ventral, showing pale white sole o f fixjt Figure 4-183. H aliotis rubra Leach, 1814. Lagoon Bay, Tasmania, Australia, dorsal under a rock. Photograph and © Karen Gowlett-Holmes. Figure 4-184. H aliotis laevigata Donovan, 1808. Duke o f Orleans Bay, Western Australia, Australia. Under a rock. Photograph and © Karen Gowlett-Holmes. 320 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 321 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 4-185. H aliotis cyclobates Péron & Lesueur, 1816. Edithburgh, South Australia, Australia. On Pirma bicolor Gmelin, 1791, its usual habitat. Photograph and © Karen Gowlett-Holmes. Figure 4-186. H aliotis coccoradiata Reeve, 1846. Malabar Bay, New South Wales, Australia. Under a rock. Photograph and © Karen Gowlett-Holmes. Fig ure 4-187. H aliotis sem iplicata Menke, 1843. Cockbum Sound, Western Australia, Aus tralia. Under a rock. Photograph and © Karen Gowlett-Holmes. Figure 4-188. H aliotis roei Gray, 1826. Albany, Western Australia. Photograph and © Clay Bryce. Figure 4- 189. H aliotis hargravesL Split Solitary Island, northern New South Wales, March 1986. AMS C150035. Dorsal, photo #025409. Photograph and © Bill Rudman. Figure 4-190. H aliotis scalaris Leach, 1814. Duke o f Orleans Bay, Western Australia, Australia. Under a rock. Photograph and © Karen Gowlett-Holmes. Figure 4-191. H aliotis squam ata Reeve, 1846. Dampier Archipelago, Western Australia. Photogr^h and © Clay Bryce. Figure 4-192. H aliotis elegans Philippi, 1844. Rottnest Island, Western Australia. Pho tograph and © Clay Bryce. 322 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 323 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter 5: A Total Evidence Cladistic Analysis of the World-Wide Haliotidae. INTRODUCTION Two studies concerning the phytogeny o f the Haliotidae have been published to date. Brown (1993) took a phenetic approach and used allozytne frequencies for 17 nominal species to infer the phytogeny with Nei’s genetic distance. Lee & Vacquier (1995) used the l 6kD protein called Tysin’ to infer the phytogeny o f 22 abalone species employing a parsimony-based approach. The taxonomic overlap between the two stud ies is 13 taxa. The two studies are in broad general agreement with one another (Figures 5-1 and 5-2). First, they both show a north Pacific clade, comprised of the west Ameri can species and the large Japanese species {H. discus, K gigantea) to the exclusion o f the smaller Japanese species such as H. diversicolor. The second second shared clade comprises the taxa endemic to Australia, to the exclusion o f wide-spread taxa also occur ring in Australia (e.g., H. ovina, H. varia). For the remainder, the tow taxonomic sam pling density precludes any further general statements. For instance, o f the five species endemic to South Afiica (H. midae, H. parva, H. queketti, H. spadicea, H. speciosa) only H. midae was included in these studies. Both p^)ers have a serious deficiency, in that the type species o f the genus, H. asinina^ was not included. Hence, the tentative taxonomic recommendation on the genus level made in Lee & Vhcquier (1995) are lack ing a sound base (see also Geiger, 1996,1998a, for discussion). 324 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. MATERIALS AND METHODS Source o f specimens Table I gives the sources of all data and specimens used in this study. No animals could be located for the following taxa considered valid (cf. chapter l)iH, crebrisculpta, K cyclobates, H. exigua, K pulcherrima, H. sorenseni, and H. squamosa. These taxa are not listed in the morphological data matrices. Lee & Vacquier (1995) had some tis sue o f H, sorenseni available, hence, only the other 5 o f the total o f 56 taxa will not be included in any o f the data analyses o f the family. SEM preparation Preparation and viewing of radulae as well as desiccation o f epipodia followed the protocols o f Geiger (1996) and Stewart & Geiger (1999). Cladistic methodology My basic understanding of the cladistic method has been presented in detail in Chap ter 3 (= Geiger & Fitzhugh, m review). For the analyses presented here, the following parameters were used. All characters were unordered and equally weighted. No differential transition/transversion o f gap- cost were used, and GAPMODE = NEWSTATE was set. Due to the number o f taxa analyzed, heuristic search options o f PAUP 3.1 (Swofford, 1991) were used, with ran dom taxon addition in effect and random seed number entered manually. Between 10 and 1 ,0 0 0 heuristic searches were performed depending on how quickly the number o f equally parshnonious trees remained stable. Subsequent searches with Tree Bisection and Reconnection (TBR) on non-minimal length trees to escape possible local minima were performed. The maximum length o f the submfnimal trees was dictated by hnple- 325 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 5-1: Source o f specimens and data. no data available. Abbreviation for source o f specimens furnishing morphological characters (Morphologr column): ANSP: Acad emy o f Natural Sciences, Philadelphia. BMNH: British Museum o f Natural History. BO: Buzz Owen Collection, Gualala, California. CASIZ: California Academy of Sci ences, Invertebrate Zoology, San Francisco, California. DLG: Daniel L. Geiger collec tion, Los Angeles. JK: Joan Koven collection. Astrolabe hic., Washington (DC). LACM: The Natural History Museum o f Los Angeles County. S (1998): Simone (1998). SAM: South Australian Museum, Sydney. SBMNH: Santa Barbara Museum o f Natural His tory. USNM: United States National Museum, Washington, (DC). WAM: Western Aus tralian Museum, Perth. Other abbreviation : W (unpubl.): Wray, unpublished data. L&V (1995): Lee & Vac quier, 1995. B (1993): Brown, 1993. 326 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 5-1. Taxoa asînina aurantium australis èrazieri clathrata coccoradîata corrugata cracherodii cyclobates dalli discus dissona dîversicolor dohm iana elegans fa tu i Julgens gigantea glabra hargravesi iris jacnensis kamtschatkana laevigata maemka mariae marmorata midae avfna parva planata pourtalesii pulcherrim a pustulata quekett i Morphology 16S mtDNA DLG LACM 85-2 W (unpubl.) S (1998) DLG W (unpubl.) SAMC32701-05 SAMC149014 DLG USNM 795269 BMNH I8872.9.l68-168a SAMC328730-32 - DLG ANSP 50503 W (unpubl.) LACM 54780 LACM 79-23 fysin allo^m e ANSP 315293 LACM 82-12 CASIZ 44960 JK LACM 82-20 DLG USNM 360940 WAM SI003-4 USNM 486708 LACM 146819 BMNH 1878326.2 - BMNH 18542.19.117 W (unpubl.) W (unpubl.) W (unpubl.) W (unpubl.) ANSP 230038 SAMC327854-57 SAMCI49015 DLG LACM 113210 CASIZ 21339 LACM 87-97 BMNH DLG BMNH no # DLG CASIZ 20810 LACM 85-3 DLG DLG USNM 856447 DLG DLG W (unpubl.) W (unpubL) W (unpubl.) W (unpubl.) W (unpubl.) W (unpubl.) W (unpubl.) W (unpubl.) L&V (1995) B (1993) B (1993) L&V (1995) B (1993) L&V (1995) B (1993) L&V (1995) B (1993) L&V (1995) B (1993) L&V (1995) - L&V (1995) B (1993) L&V (1995) B (1993) L&V (1995) 1 3 (1993) L&V (1995) B (1993) L&V (1995) - L&V (1995) 1 3 (1993) L&V (1995) - L&V (1995) 327 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 5-1: continued roberti roei rubigpiosa rubra nifescens rugosa scalaris sem iplicata sorenseni spadicea speciosa squamata stomatiaeform is tuberculata unilateralis varia vtrginea walallensis SBMNH no# W (unpubL) — — LACM SAMC328696-700 W OmpubL) L&V (1995) B (1993) SAMC352383 - - — LACM 71-495 W (unpubl.) L&V (1995) B (1993) LACM 1813-49 - L&V (1995) B (1993) DLG LACM 87-100 W (unpubl.) — WAM SI 007-8 W (unpubl.) L&V (1995) B (1993) WAM S1005-6 - - - - W (unpubl.)- L&V (1995) - DLG - - - DLG - - - SBMNH no# - - - SAMC328733-41 - - - BO - — - DLG W (unpubl.) L&V (1995) - DLG DLG LACM 85-5 W (unpubl.) L&V (1995) - CASIZ 44954 - • B (1993) LACM 1813-49 W (unpubl.) L&V (1995) - 328 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. mentatîoa limits in PAUP 3.1, not memory (RANQ limitations. PAUP 3.1 can not store more than approximately 30,000 trees internally; the precise number o f maximally stor able trees depends on the number o f taxa as well as the number of characters. In cases of multiple most parsimonious resolutions, a strict consensus tree is shown. Tree statis tics refer to any o f the fully resolved topologies, not the strict consensus tree. Abbreviations used are Cl (consistency index), RI (retention index), RC (rescaled consistency index), and MPR (maximum parsimonious resolution). Cladograms were prepared in MacClade 3.04 (Maddison & Maddison, 1992) and further modified in McDraw™ 111.1 (Claris, 1987). Outgroup comparison Characters were polarized through outgroup comparison where possible. Outgroups were chosen fiom all member o f Vetigastropoda, because the relationships amongst Vetigastropoda are currently unresolved (cf. Baten, 1975; Haszprunar, 1988b; U llier et al., 1994; Hickman, 1996; Harasewych et uA, 1997). The character states were derived firom both original observations (Pleurotomariidae: AMNH 187302, AMNH 3483, AMNH19855. Tfochidae: LACM 46-32.12. Lepetodrilidae: DLG. Fissurellidae: LACM 45-1.5) as well as fiom the literature (Hickman, 1984; Haszprunar, 1988c; McLean, 1984b, 1988; Franz, 1989; Hickman & McLean, 1990; Sasaki, 1998). If multiple taxa were available, the most basal taxon was chosen according to available phylogenetic hypotheses (Trochidae, Turbinidae: Hickman, 1996. Pleurotomariidae: Harasewych et a i, 1997). 329 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ALLOZYME FREQUENCIES The use o f the allo^rme frequency data provided by Brown (1993) provides a par ticular challenge, because the original frequency data had to be transformed into char acter-state data. In its most basic form character coding is a problem o f belief representation. This problem can be subdivided into three areas; - What is the cut-off level at which a low frequency occurrence o f an observation is still considered to represent a character state found in a species? - How are the electromorphs of proteins stained with the same stain treated? Do these different alleles belong to the same locus, or are they from di&rent loci? As differ ent character states in one character resulting in massively polymorphic taxa, or using presence / absence coding for each electromorph resulting in a clean, binary data matrix? - A third but minor problem is when are two electromophs given the same identifier? This is a similar problem to when we call two legs the same length. For the frrst problem the general statistical significance level of 0.05 can be used for the inclusion/exclusion argument. For sample sizes o f up to 20 individuals every elec tromorph found was mcluded in the binary matrix, whereas single instances o f particu lar electromorphs in sample sizes between 21 and 40, and double instances in sample sizes between 41 and 60 were excluded. No larger sample sizes were used by Brown (1993). Assuming that the specimens used were a representative sample from the entfre population, this treatment compares fovorably with practices in m orpholo^ and is more thorough than what ^ i c a l sample sizes in studies with sequence data allow. For mor phological studies usually far fewer than 2 0 specimens are studied, and for sequence 330 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. data rarely more than a single specimen is assessed. Hence, the 0.05 inciusion/exclu- sion argument is rather conservative when compared to other types o f data. The second problem is the most difBcuIt. Stains are considered to be specific for a particular en^m e, however, it is not neccessarily known whether that en^nne is pro duced by a single- or a multi-copy gene. Addtionally, epigenetic bands may have been identified as discrete allo^nxies; without the original gels, the nature of the band (epige netic, actual alio^one) can not be resolved. Consequently, the question o f homolo^r or orthology versus paralogy becomes central. A second problem concerns the the staining method, which is a substrate modification to produce a color in the presence of a partic ular en^rme. It is assumed that the substrate reaction is highly specific for the en^nne in question. One can certainly entertain the possibility that one substrate may be uti lized by more than one target protein, hence, staining protein may not be a reliable indi cator o f similarity. I have opted here to take the position o f the advocatus diaboli, meaning that I assumed multi-copy genes and multiple use o f substrates due to absence o f evidence to the contrary. As a necessary consequence, all electromorphs are coded in a presence^absence matrix. Such a practice may strike some as inappropriate, but such are my subjective b elief, which are open to reinterpretation, o f course, by subsequent workers. On the other hand I believe that bands o f the same electric motiliQr a id same stain ing properties provide good grounds to postulate their shnilarity and, hence, to explam them by means o f primary homology, further assuming that Brown (1993) did not iden tify epigenetic bands as discrete allo^m es. The similarity is based on topographical identity being the sam e motility and character-state identity being the same staining property. For a more detailed discussion o f belief formation and its inherent subjectivity see (Zhapter 3 (= Geiger & Fitzhugh, m review). 331 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table 4-2 gives the recoded ailo^rme frequency data o f the 16 abalone taxa used by Brown (1993). Note that H. conicopora has been synonymyzed under H. rubra by Geiger (1998a), therefore, the data from these two nominal taxa were combined. Data from multiple populations o f the same species were averaged. Outgroup comparison As the identification o f the electromorphs is relative to the other taxa studied, addi tion o f data to the published ones is virtually impossible without repeating most of the original study. Consequently, the data-set for the ingroup is limited to that o f the origi nal ingroup, and outgroup comparison is impossible. The cladistic analysis of that data set, hence, will show unrooted cladograms, or networks. These networks may be rooted anywhere within the tree, for which reason it is inappropriate to call groups o f taxa clades. I will, therefore, rather use the informal word ‘grouping’ to highlight some of the particular finds from the analysis of that data-set. Reanafysis of allozyme data with comparison to the results of Brown (1993) An initial search with TBR on minimal trees only produced eight equally most par simonious trees o f 117 steps (Cl = 0.479, RI = 0.555, RC = 0.266). A search with TBR on non-minimal length trees o f tq > to 120 steps found four additional topologies o f 117 steps. This case illustrates clearly the importance o f branch swapping on non-minimal length trees in heuristic searches. A branch-and-bound search with upper bound =117 steps fotmd the same 12 trees. Brown’ s (1993) tree is fully resolved (Figure 5-1: A), whereas the reanalysis o f the allozyme frequencies as character state data using parsimony K characterized by a basal polytomy (Figure 5-1: B). Was a lot o f information lost during the transformation o f 332 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-L Trees generated from alloqmae frequency data (Brown» 1993). A) Neighbor joining tree o f Brown (1993) redrawn, with geographic distribution added. B) Strict consensus tree of 12 equally most parsimonious resolutions from the allozyme data only. 56 informative characters, 117 steps. Cl = 0.479, RI = 0.555, RC = 0.266. NZ: New Zealand. RSA: Republic o f South Africa. H aliotis conicopora is synonymized under if. rubra. Note the North Pacific group, and the majority of endemic Australian species form one group. The three New Zealand species, however, are either dispersed over the tree (A), or are are united in a basal polytomy (B). 333 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I 0 5 Net’s Genetic Distance (D) B australis rubra rubra rubra conicopora laevigata laevigata laevigata scalaris roei roei cyclobates coccoradiata midae iris N virginea Z discus fidgens kamtschatkana rufescens corrugata cracherodii < s s 1 I z corrugata cracherodii discus fulgens rufescens kamtschatkana coccoradiata rubra cyclobates laevigaxa scalaris » roei ‘ midae ■ his “ australis ■ virginea z I û d Z 334 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. fiequenqr data to character state data? For this questioa a particularity o f distance meth ods such as neighbor joining using Nei’s genetic distance has far reaching consequences. The distance algorithm ignores other potential resolutions and is particularly sensitive to the initial, most similar taxa. Although the higher resolution o f the topologies is touted by advocates o f phenetic methologies as a clear advantage (see Wenzel, 1997, for discussion), the irrelevance o f the analytical procedure to evolutionary questions has been lost 6 om sight As a consequence, the patterns resulting &om the parsimony analy sis are the only meaningful ones, because they are based on the Darwinian notion of descent with modification, and hence also on a cause - effect relation o f spéciation and observed distributional patterns o f character states among taxa. Resolution is secondary, as polytomies may be real, the so called ‘hard’ polytomies (Wenzel, 1997). The distance topology and the recoded parsimony topology do not squarely contra dict one another, despite the relatively low resolution o f the parsimony tree. The com mon elements are as follows. The north Pacific species and most o f the species endemic to Australian are grouped together. The three New Zealand species as well as the single south Afiican species are united in the large basal polytomy, hence, no statement about them precise relationships can be made. To ^>pease advocates o f compatibility analysis prior to combination o f data-sets, the present analysis does not contra-indicate combi nation with the other data (see below). However, the compatibility argument is vacu ous, because the primary quest o f a phylogenetic analysis is explanation o f observations; inclusion or exclusion o f data in a total evidence analysis depends solely on the rele vance o f the data to the question addressed (see Chapter 3 = Geiger & Fitzhugh, in review). The presence or absence o f allozymes as properties o f organisms is relevant b ^ o n d any doubt 335 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. DNA CODING STRATEGIES FOR 16KD CDNA The DNA-sequence data on a sperm acrosomal protein (I6kD Lysin) published by Lee & Vacquier (1992, 1995) was incorporated as follows. All positions were included in the present analysis, though the original alignment and data representation was some what altered as will be explained below. Character number refers to the recoded matrix. The new coding strategies are those mtroduced in Chapter 3 (= Geiger & Fitzhugh, in review), hence, I will only briefly summarize the procedures here. For a detailed Justifi cation see Chapter 3. ‘ ^Data contraction* reduces a stretch o f bases or amino acids of questionable align ment to a single position. It uses a higher level o f generality for the observations at hand. It was used in both the translated region (= open reading fiame. ORF: positions 21,153) as well as the 3' untranslated (UTR: positions 5 ,6,20,75, 132) region. ‘ ^Stretch coding* introduces (aut-)^m orphic character states fi)r stretches o f ques tionably aligned sequence m a few taxa only. It is directly comparable to practices in morphology, where the introduction of new character states is used for observations that cannot be classified in existing states. It has been used in positions 45-46,101-104, and 111-114 in the UTR o f the lysin cDNA. Table 4-3 gives the recoded data from Lee & Vacquier (1995), with explicit notes on the coding strategies employed. In order to obtain the most mArmation from the data, additional recoding strategies were explored. It can be argued that the use o f the amino-acid sequence for coding regions o f genes such as the ORF of lysm offers more phylogenetic meaningful infer- mation, because problems due to the redundancy o f the genetic code and the limited number o f character states—6 ur bases as opposed to twenty amino acids—are avoided. This approach can be taken further, noting that coding regions form functional entities. 336 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Hence, it may not be the actual ammo acids, which show the greatest amnunt o f infor mation, but the functional groups. The amino-acid sequence was recoded according to the seven functional groups (Stryer, 1995) and re-analyzed. The strict consensus tree of 174 MPRs was characterized by a large polytomy (tree not shown). Accordingly, this approach was abandoned and the amino-acid sequence o f the lysin’s ORF was used for further analysis. Outgroup comparison Outgroup comparison is un&rtunately impossible. The lysin protein o f a trochid has been sequenced but is only 50% o f the length o f the abalone lysin (Hellberg, 1998). Alignment is impossible and even the question o f whether these functional proteins are homologous, or in molecular parlance *orthologous% arises (c f Hellberg & Vacquier, 1999). However, as lysis is under positive Darwinian selection (Lee & Vo:quier, 1992; Vacquier & Lee, 1993; Lee et al., 1995; Swanson & Vacquier, 1995), such high dissim ilarities o f putatively homologous gen% between families are not surprising, but do not help with outgroup comparison either. For other Vetigastropoda, no lysin data are avail able. Hence, I searched for unrooted trees. Reanafysis of the fysm data Lee & ^ ^ q u ie r (1995) partitioned the data into the ORF and the UTR. My initial treatment will follow them data partitioning to show that the effect o f recoding the data is (fistiact 6 om the effect o f combmmg the data-sets. For the same reason, the taxa were the same as used by Lee & Vacquier (1995). Differences between their study and the present study include the above mentioned alignment and coding variation, but also extend to the treatment o f gaps. Whereas Lee & Vacquier (1995) treated gaps as mfes- 337 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-2. Trees generated &om data o f Lee & Vacquier (1995). EM-RS: European- Mediterranean and Red Sea. IP: wide-spread Indo-Pacifîc. NZ: New Zealand. RSA: Republic of South Africa. A) ORF tree redrawn, from Lee & Vacquier (1995: fig. 4a) with geographic distribution superimposed. B) ORF strict consensus tree from recoded data. 316 steps» Cl = 0.560, RI = 0.673, RC = 0377. Q UTR tree redrawn from Lee & Vacquier (1995: fig. 5a) with geographic distribution superimposed. D). UTR strict consensus tree from recoded data. 423 steps. Cl = 0.709, RI = 0.858, RC = 0.609. E) Strict consensus tree o f ORF and UTR combmed. 781 steps. Cl = 0.647, RI = 0.787, R C = 0.509. 3 3 & Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. r u f e c e n s s o r e n s e n i k a m ts c h a tk a n a w a la lle n s is s i e b o l d i i d is c u s h a n n a i c r a c h e r o d ii c o r r u g a ta f i d g e n s r o e i s c a l a r is la e v ig a ta c y c l o b a t e s r u b r a m id a e p u s tu la ta tu b e r c u la ta I O Z RSA. c o c c m e a v a r ia o v in a d i v e r s i c o lo r a u s t r a li s i r i s I m g B I r u f e s c e n s I s o r e n s e n i ' w a la lle n s is ^ k a m ts c h a tk a n a ^ g ig a n te a d is c u s ic o r r u g a ta t c r a c h e r o d ii tf id g e n s a u s t r a li s i r i s Z ( S v e r s ic o lo r v a r i a o v in a C u c o c c m e a tu b e r c u la ta ^ p u s tu la ta ^ r u b r a a z & s c a l a r is g u te v tg a ta midae 339 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. r u f e c e n s s o r e n s e n i k a m ts c h a tk a n a i w a l a l l e n s i s § s i e b o l d i i a . d i s c u s h a n n a i T g c r a c h e r o d i i o c o r r u g a ta Z J u lg e n s i r i s N Z r o e t e i s c a l a r i s l a e v i g a t a 1 c y c l o b a t e s r u b r a < n u d a e R S A p u s tu la ta 1 1 tu b e r c u la ta c o c c in e a 1 “ ( v a r i a o v in a ûÿ ( V e r s i c o l o r 1 a u s t r a li s N Z D c r a c h e r o tu i w a la U e n s ts g t g a n te a c o r r u g a ta r u f e s c e n s j u l g e n s a u s t r a li s p u s t u l a t a tu b e r c u la ta ( V e r s i c o l o r r u b r a m id a e 340 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. E c c rufescens sorenseni kamtschatkana a walallensis § gigantea cu discus Tg corrugata O cracherodii Z Julgens iris australis roei scalaris laevigata rubra 1 cyclobates < i fi pustulata tuberculata s coccmea cu omna varia ^ dîversicolor m i dae R SA . 341 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. mg infennatioa (gapmode = missing), Geiger & Fitzhugh (in review = Chapter 3) argue for a fifth character state (gapmode = newstate), which is followed here. ORF (Figures 5-2: A and 5-2: B): a heuristic search with TBR on non-minimal length trees o f up to a tree length o f 318 using 1,000 replications revealed 120 equally most parsimonious trees of 316 steps (Cl = 0.560, RI = 0.673, RC = 0.377). Figure 5-2: B shows the strict consensus tree. When compared to the tree published by Lee & Vac quier (1995: fig. 4; Figure 5-2: A) the two correspond fairly well in that they both show a North Pacific group. In Lee & Vacquier (1995) the larger remaining taxa are joined in a large polytomy containing, however, groupings that unite the species endemic to Aus tralian, the widespread species o f the Indo-Pacific, as well as the European and Red Sea species. The two New Zealand species are separated, though. This rather surprising pat tern o f multiple colonizations o f New Zealand implied by this analysis agrees with finds fiom other groups o f organisms such as beeches (Motofagus), Afiican violets (Gesneri- aceae), a caddis fly genus {H ydrobiosella), and platycercine parrots discussed by Humphries & Parenti (1986). My re-analysis shows rather similar patterns in that the North Pacific species form one group (although less resolved), and the remaining taxa are joined in a large polytomy (Figure 5-2: B). Note that the two Japanese species [H. discus y K gigantea) are now located in this large polytomy, whereas they were previ ously fotmd nested withm the west American abalone species (£ T . fiilgens and up). It seems then that the position o f the two Japanese species in Lee & Vacquier's original analysis stems fiom arbitrary alignment. Within the latter polytomy, however, the species endemic to Australian are not in a single group, the European and Red Sea species are closely associated with one o f the species endemic to Australian, and one o f the New Zealand species (Ü iris) is nested withm the widespread Indo-Pacific group- tng It is clear that changes in coding does have a significant impact on the phylogenetic 342 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. hypotheses generated. Re-evaluatîoa particularly o f the homoplastic character state changes as implied from my coding strategies did not reveal any coding problems. Therefore, E interpret the inferred homoplasies that remained despite the careful recod ing carried out to be best explained by multiple changes at these sites, and not due to additional coding problems. UTR (Figures 5-2:C and 5-2:D): thirty-eight equally most parsimonious trees of length 423 were found with branch swapping on non-minimal length trees o f up to 425 steps and 1,000 replications (Cl - 0.709, RI = 0.858, RC = 0.609). This re-analysis also grouped the North Pacific species, the species endemic to Australian, the Red Sea - European species, and the wide-spread Endo-Pacific species, with the two New Zealand species {K australis, K iris) being separated by two nodes. The difference between my (Figure 5-2: D) and Lee & Vacquier’ s (1995: fig. 5a; Figure 5-2: C) analyses lies in the resolution within some o f these groups. The north Pacific group shows three nodes in Lee & Vacquier’s work, whereas two nodes separate H. julgens and H, nifescensy respec tively, from a large polytomy in the re-analysis. The position o f the Japanese species undergoes the same changes as discussed for the ORF above. Additionally, the position o f the two New Zealand species differs between the two analyses, though the New Zealand species represent individual spéciation events, and not a small radiation on this remote island group. ORF combined with UTR (Figure 5-2£): A heuristic search with TBR and 10 repli cations found two equally parshnonious topologies of 781 steps. Subsequent searches with TBR on non-minimal length trees (< 785 steps, 100 replications. Cl = 0.647, RI = 0.787, R C = 0.509) found the same two equally most parsimonious trees. A branch-and- bound search with t^p er bound o f 781 was aborted after almost 100 hours o f search on a 604/120MI& processor that did not get passed in approximately 22% o f all topologies 343 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. to be evaluated. Branch-and-bound searches become very problematic with more than 20 taxa, unless the data-matrhc is extraordinarily clean (D. Kizirian, pers. comm.). Mol ecular data matrices in general do not satisfy such a condition. The strict consensus tree o f these two trees is shown in Figure 5-2: E. As the consensus tree is unrooted, I will identify groups rather than clades as the rooting may be placed anywhere on the tree resulting in non-monophyly o f apparent clades. The first group, comprising the north Pacific species, contains the only two tri chotomies found. Note that the Japanese species are again nested within the west Amer ican species. The second group contains the species endemic to Australian. Third, a small wide-spread Indo-Pacific group can be identified. Note the dispersed occurrence o f the two of the three New Zealand taxa {K australis, K iris), which are separated by one node implying again separate colonization events. It is mteresting to note that the Red Sea H. pustulata forms a group with the Euro pean taxa H. tuberculata and H. t. coccinea. This pattern resounds with one o f three biogeographical hypotheses discussed by Geiger & Groves (1999 = Chapter 2) and Geiger (in press = Chapter 4), the Tethys origin. This model is based on published chro mosomal count data, which suggests an origm o f the abalone in the Tethys, with subse quent eastward dispersal to the Indo-Pacific, and then to the north Pacific. However, due to the limited taxon samplmg o f this reanalysis, I want to de-emphasize this find rather, than to use it as a comer stone o f the Tethys hypothesis. One may also note the apparent derived position o f the (pusttdata {tuberculata, coccinea)) group, which would be a counter indication to the required basal position in support o f the Tethys model. Additionally, the Red Sea is geologically speaking a relatively young rift-shaped depres sion o f approximately 20 My o f age (Reiss & Hottinger, 1984). 344 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. PARTIAL I 6S MTDNA (CHARLEY WRAY, UNPUBL. DATA) Twenty-six unpublished I 6S mtDNA sequences were kindly made available to me by Charley Wray (see Table 5-1), a former post-doctoral associate at the University o f California, Berkeley. The sequences o f540 bp were treated with my new coding strate gies; forty-six positions o f questionable alignment were excluded (positions 1-18, 257- 266, and 337-354). The sequences are not shown here, because they are unpublished data of Charley Wray. Figure 5-3 shows the strict consensus topology o f 15 MPRs (Length = 189; Cl = 0.434; RI = 0.698; RC = 0303). For outgroup comparison, Genbank was searched for alignable vetigastropod sequences, but none could be found. The network (Figure 5-3) shows the two familiar groups being the north Pacific species and the Australian endemic ones. The Indo- Pacific is polyphyletic in contrast to the combined analysis discussed below. The two New Zealand species do not form a group within this data-set either. The combination o f the lysin ORF and UTR, the allotyme frequencies, and the 16S mtDNA data yielded three MPRs (Figure 5-4). The features shared with the already dis cussed topologies are the North Pacific group mcludmg the tropical eastern Pacific H. roberti and the Caribbean K poitrtalesiiy and the species endemic to Australia to the exclusion o f the wide-spread taxa also occurring in tropical Australia (K ovma, ff. varia). The three New Zealand species (K australiSy H iriSy H. virginea) are distributed over the entue tree, which shows that either three diffèrent colonization events occurred for this remote group o f islands, or that the abalone fituna o f New Zealand is a relict frnma from an ancient stock, but is certamly not a discrete radiation on these islands. The wide-spread frido-Pacific taxa (E L diversicolary K ovina, H. varia) and K glabra with a more restricted distribution in the Indo-Malayan archipelago (see Chapter 4 = Geiger, m press) form another group to the exclusion o f the enigmatic type species o f 345 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. E asinina australis cyclobates laevigata scalaris roei rubra midae tuberculata corrugata kamtschatkana sorenseni walallensis pourtalesii fulgens discus gigantea rnadaka roberti glabra varia iris^ diversicolor ovina I pustulata rugosa IP NZ RSA EM I Z I ' NZ I- | 2 Figure 5-3. Strict consensus o f 15 MPRs topologr o f 16S mtDNA sequences (Wray, unpubl. data). EM: European-Mediteiranean. RS: Red-Sea. 10: Indian Ocean. IP: Ihdo- Pacîfîc. NZ: New Zealand. RSA: South Afiica. 346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. asinina australis cyclobates laevigata scalaris roei rubra IP N Z C corrugata cracherodii discus gigantea madaka roberti pourtalesii Kamtschatkana sorenseni rufescens rufescens walallensis julgens iris midae diversicolor glabra varia ovina tuberculata tuberculata 1 2 NZ RSA " C tuberculata coccinea pustulata rugosa L —virginea % I NZ Figure 5-4. Strict consensus tree o f 3 MPRs from ail allozyme (Brown, 1993), lysin (Lee & Vacquier, 1995), and 168 mtDNA (Wray, unpubl.) data combined. EM-RS: European-Mediterranean-Red-Sea. IP: Indo-Pacific. NZ: New Zealand. RSA: South. Africa. 347 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. the genus (for discussion see Chapter 1 = Geiger, 1998a). This shows that the very dis tinct shell morphology o f K asinina is also reflected in its position on the phylogenetic tree. MORPHOLOGICAL CHARACTERS The radnla in abalone The radula is one o f the unique structures o f mollusks and has played a very impor tant role in gastropod systematics and classification (e.g., Thiele, 1931). It is a chitinous band with attached chitinous teeth. A radular formula is commonly used, which reads number o f rows x marginal teeth + lateral teeth + rachidian (R) + lateral teeth + - mar ginal teeth. Warén (1990) has shown that during ontogeny the marginal teeth are derived fiom lateral teeth and that the distinction o f laterals and marginals is, therefore, artifi cial. However; because this distinction is so widely accepted, and has descriptive utili^, it is used here as welL The arrangement and fine structure o f the teeth has been hnpor- tant for the recognition o f taxa at virtually every level of gastropod classification. How ever, the u tili^ o f the radula to address species level problems m the family Haliotidae had been called into question (Talmadge, 1956; Barnard, 1963). These early claims have been disproved subsequently (Geiger, 1996; 1998a, 1999a; Simone, 1998; Stewart & Geiger, 1999). Table 5-2 gives the data matrix for the radular characters; the charac ter states are discussed below. Geiger (1996) introduced terminology to identify various parts o f the teeth o f abalone radula fig u re 5-5) and this terminology has been adopted fi)r this account. The radular structure and the radular formula change durmg early growth. The radula is seen first in the post-torsional veliger (Crofts, 1937; Dinamani & McRae, 1986). Competent larvae have at least three rows o f radular teeth (Tong & Moss, 1992). At 348 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 " O 3. 3 " CD CD ■ D O Q . C a O 3 " O o CD Q . P lew o to w riid ae Soigeweilidae (Scissureita) Iiepetodirllldae (Lqpetodrilue) FiPPiurelUdae (jpia«MJreii4) Vroohidoe {Tegula) Vuiixmidoe (x lp ^ a ) a&ininn tiwreuitim auatretlia braaieri olatturata oocoocadiata ooixuffata craobexvdii daUi diaaua diaaona diveraicolor aJaffana fa tu i tui^ana giffantaa glabtra hargravaai ix ia jaonanaia Jcamtaahatkana laevigata 0 0 0 0 0 0 0 0 mdaka 0 0 1 X X 0 0 0 0 0 0 1 1 1 1 1 1 0 0 1 0 0 muriae 0 0 0 X 0 X X 0 0 0 0 1 1 0 0 1 0 0 0 1 0 m m orata 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 «•* 1 0 0 0 0 0 midaa 0 0 I X 0 0 0 0 0 0 0 0 1 1 0 0 1 0 0 1 1 0 ovina 1 1 0 0 0 0 0 0 X 0 0 1 1 1 1 1 1 0 0 0 1 0 paxva 0 0 0 0 0 X X 0 0 0 0 p la m ta 1 1 X X 0 X 5 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 p o u rta lea ii 0 0 0 0 0 X 0 0 0 0 X 0 0 0 0 0 1 0 0 0 0 1 puatulata 1 5 0 X 0 X 5 0 0 0 Û 0 0 0 0 0 0 0 0 0 1 0 q u eka tti 1 0 0 0 0 X 0 0 X 0 0 1 0 0 0 1 0 0 0 0 0 0 ro b erti 1 0 X X 0 X 0 0 0 0 X 1 1 1 1 0 1 0 0 s 0 0 roei 1 0 0 0 0 0 X 0 X 0 0 1 0 1 0 0 0 1 0 0 X 0 ivbiginoaa 0 0 0 X X 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 0 xvbra 0 0 0 0 0 0 X 0 0 0 0 0 1 1 1 1 0 5 0 1 0 0 rufaaoana 0 0 X X X 0 0 0 X 0 0 1 0 1 1 0 1 0 0 0 0 1 xvgoaa 1 1 5 X 0 0 0 0 0 0 0 1 0 1 1 1 0 0 0 0 1 0 acalaria 0 0 0 0 0 0 5 0 X 0 0 1 1 1 1 0 1 0 0 0 0 0 aeaUplicata 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 apadicaa 1 1 X X X 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 apacioaa 1 0 0 0 0 X X 0 0 0 0 1 1 0 1 0 1 0 0 0 0 0 agumata 1 0 0 0 0 X 0 X X 0 0 0 0 1 1 1 0 1 0 1 1 0 a ta m tia efo m ia 1 0 X 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 tuberculata titberculata I 0 0 0 0 X X 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 tuberculata ccxxinea 0 0 0 0 0 0 ? ? 0 0 0 0 0 0 0 1 1 0 0 0 0 0 m d la ta ra lia 1 5 5 X 0 X 5 0 0 0 0 1 0 0 1 0 0 1 0 0 0 0 varia 1 1 X X 0 X 0 0 0 0 0 0 1 1 1 1 1 0 0 0 1 0 virginea 1 1 0 X X 0 0 0 0 X 0 0 1 1 1 1 0 0 0 1 0 0 walallanaia 1 0 X X X 0 X X X 0 0 0 0 0 1 0 0 1 0 1 0 0 " O CD ( / ) ( / ) ê Table 5-2, Character states for the radular characters, For description of character states see text. 5 = (0,1), - = inapplicable. CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 ■ D 3 . 3 " CD CD ■ D O Q . C a O 3 " O O CD Q . ■ D CD C/) C/) wnticteson primaiy sm oominm gr C llS p dentideft lîîîW .W W .I U l o Figurç 5-5. Diagram illustrating the terminology used for the radular teeth found in the family Haliotidae. The diagram is taken from Geiger (1996). metamorphosis, the radula formula is >6 - 8 x 3 + R + 3 (Tong, 1985; Dtnamani & McRae, 1986), and marginal teeth and additional rows are formed during subsequent growth (Dinamani & McRae, 1986). The radular formula for a three-week old juvenile is5 + 4 + R + 4 + 5 (Crofts, 1929). The rachidian tooth o f this stage is pluricusped and similar to those o f other small vetigastropods (Pleurotomariidae: Hickman, 1984a. Hali otidae: Garland et u/., 1985; Tong, 1985; Dinamani & McRae, 1986. Trochoideæ Hick man & McLean, 1990; Warén, 1990. Scissurellidae: Thiele, 1912; Marshall, 1993). The fully developed radular pattern, with its five lateral teeth, is found after two months (Crofts, 1929). The radula formula given by most authors for mature spechnens is oo + 5 + R + 5 + 0 0 (Fischer, 1885; Herbert, 1990 and references therein), where ‘oo’ signifies a large but unspecified number. However, Dai & Wu (1989), Wu & Huang (1989), and Wu (1991) specified the number o f marginal teeth between 55 and 65 for the various species studied. Outgroup comparison The radulae o f Haliotidae differ in various aspects from those o f other Vetigas- tropoda. The rows of the radular teeth are almost symmetrical in abalone, but are dis tinctly asymmetrical in pleurotomariids. The rachidian tooth is well-formed in Haliotidae, but reduced in Pleurotomariidae. The fine outer marginal teeth in Haliotidae show denticulate cusps (Wu & Huang; 1989; Herbert; 1990; Geiger, 1996; 1999a; Stew art & Geiger, 1999); m Pleurotomariidae, however, afim o f articulated bristles is found (Hfrkmmn, 1984a; Harasewych & Askew, 1993; Anseeuw & Goto, 1996). A comparison o f Pleurotomariidae and Haliotidae to Scissurellidae is not appropriate as discussed above. The independence o f radular morphology and foedmg ecology has to be ques tioned due to the extensive morphological plasticity o f the radula in response to the 351 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. feeding ecologr o f the respective animals. The radulae o f Pleurotomariidae and Halioti dae will not help to resolve their phylogenetic relationship* because the former is that o f a specialized spongivore, the latter that o f a strict macroalgal herbivore. The coding o f paedo- and peramorphic structures adds further problems, as in the case o f the radular characters o f Scissurellidae and Haliotidae. One could consider stage- specific structures, which would overstate the degree o f differentiation. In this case, the serrate rachidian o f mature Scissurellidae is coded differently fiom the rachdian with a smooth cutting edge in Haliotidae. Alternatively, heterochronic processes are taken into account, creating characters with inapplicable character states. In this second case, the rachidian o f Scissurellidae and Haliotidae < 5 mm is coded as serrate for both families, but the character state o f the rachidian for animals > 10 mm is inapplicable to Scis surellidae. The use o f the radula to resolve family level relationships within Vetigas- tropoda is, therefore, questionable (see also Haszprunar, 1993). 1 have coded the radulae o f the outgroups accordmg to ^p aren t shared similarities. It could also be argued that most character states should be coded as inapplicables, but 1 consider a clearly specified shared similarity statement—though admittedly tentative— more meaningful than the escape route o f m ^plicables. Characters and character states The characters and then states are illustrated in three figures. Figure 5-6 shows the modifications o f the central field (rachidian tooth and lateral tooth 1). Figure 5-7 shows modifications on the lateral teeth 3-5 and Figure 5-8 illustrates the variation in the mar ginal teeth. 1: Posterio-basal projection on rachidian tooth (Figure 5-6). Absent: 0; Present: 1. 352 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-6. nixistratioa o f radular character states for central field. Number on images refer to character number. A. Haliotis. stomatiaeformis. Scale bar = 100 pm. B. H aliotis glabra. Scale bar=200 pm. C. H aliotis elegans. Scale bar=200 pm. D. H aliotis roberti. Scale bar = 100 pm. 353 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-7. Illustration o f radular character states for lateral teeth 3-5. Note denticles on outer edge o f teeth in A » B, and D, in contrast to C, for wMcti the outer edge is smooth. A: H aliotispustulata, B: H aliotis tuberculata, C: H aliotis asinina. D. H aliotis varia. Scale b ar= 100 pm. 354 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5-8. Illustration o f radular character states for marginal teeth. The ^unmetrical condition (0) is shown in A. Condition (I) for character 7 is seen m E. Condition (I) for character 8 is shown in C. A: H aliotis pustulata. Scale bar = 100 ^im. B. Outer margin als o f Æ asinina. Scale bar = 10 fim. C. Inner marginals o f H. asinina. Scale bar = 100 pm. D. Lateral view o f middle marginals o f H. tuberculata. Scale bar = 10 pm. D. Inner margrnafs o f H. varia. Scale bar = 100 pm. 355 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2: Thickened cutting edge on rachidian tooth and lateral tooth 1 CFigure 5-6). Absent: 0; Present: 1. 3: Lateral tooth 1 with acute angle formed by cutting edge and primary ridge (Figure 5- 6). Absent: 0; Present: I. 4: Concave primary ridge on lateral tooth I (Figure 5-6). Absent: 0; Present: I. 5: Size o f primary ridge (Figure 5-6): < 10% the size o f the secondary ridge: 0; > 10% larger than secondary ridge: 1. 6 : Denticles on outer margin o f lateral teeth 3-5 (Figure 5-7). Absent: 0; present: I. 7: Denticles on inner mammal teeth at least by one denticle asymmetrical (Figure 5-8). No: 0; yes: 1. 8 : Denticles on inner marginal teeth at least by three denticles asymmetrical (Figure 5- 8). No: 0; yes: 1. 9: Rachidian tooth broader than lateral tooth 1, and lateral tooth 1 approximately same size as lateral tooth 3 (Figure 5-6). No: 0; Yes: 1. 10: Rachidian tooth posteriolateral wings with knobs/joints (Figure 5-6)? Absent: 0; present: I. 11: Lateral tooth 1 much narrower than rachidian tooth and lateral tooth 3 (Figure 5-6)? No: 0; Yes: 1. THEEPIPODIUM The epfpndrnm is a fleshy girdle with various projections situated on the dorsal por tion o f the foot below the margin of the shell (Crofts, 1929; Cox, 1962). It is best devel oped in HaKntidae, but is also recognized, m other vetigastropod taxa (Pleurotomariidae: Anseeuw & Goto, 1996; Scissurellidae: Haszprunar, 1988a, 1988c; Trochidae: Hick man & McLean, 1990). The epipodium is known to have a species-specific morphology 356 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CD ■ D O Q . C g Q . ■ D CD C/) C/) 8 ■ D CD 3. 3 " CD CD ■ D O Q . C a O 3 " O O CD Q . Pleurotcinariidae SoisBurellldae {ScHssurella) Lqpetodrilidae (Lepetodrilus) F issurallidoe {Fissmlh) TrookWoB (ategruia) Turbonidae (L iotia) aainina aurontiun auBtJraÜB taraxi&ri c iatb rata pcxM xwadiata carrugmta oraalw rçdii (iftXIi <Uaau9 diaaona (dried bocÿ) diyeraiooloe eXegana fatw i fuXgeaa gigmtea gXaiwa hargravaai iria jaonanaia (dried body) kamtaobatkana Xaevigata 0 0 0 0 0 0 0 1 0 1 0 mdetka 2 0 1 0 I 0 0 *— 0 0 1 0 0 0 0 5 5 0 0 0 0 0 0 m riae 0 1 0 0 0 1 0 0 0 - 0 0 0 0 0 5 5 0 0 0 0 0 0 mazmrata 1 0 1 1 1 0 0 - X 0 0 0 1 0 1 0 0 0 0 1 0 1 0 midaa 2 0 1 1 0 0 0 2 0 1 0 0 0 0 5 5 0 0 1 0 0 0 ovina 0 0 1 1 1 0 0 - 0 1 0 0 0 0 0 5 5 0 1 0 0 0 0 paxva 0 1 1 1 0 0 0 0 1 0 1 0 pourtaXeaii 2 0 1 0 1 0 0 - 1 0 X 0 1 0 1 1 1 0 0 2 1 0 0 pXonata 0 0 1 1 1 0 1 - 1 1 0 0 2 0 1 1 1 0 0 0 0 1 0 puatuXata 2 1 0 1 1 1 0 0 0 0 0 0 2 1 0 1 1 1 0 0 0 0 0 quekatti 2 0 1 1 1 0 1 - , 1 0 0 0 0 0 1 1 1 0 0 * * 1 0 1 0 robaxti 2 0 1 1 1 0 0 - 0 0 X 0 5 1 1 1 1 0 0 0 2 0 0 0 roei 2 0 0 1 1 1 0 1 0 0 0 0 0 1 1 1 0 0 — 0 1 1 0 xubiginoaa 0 0 1 1 0 0 0 - 1 1 X 0 2 0 1 0 1 0 0 1 1 1 0 rubra 2 0 1 1 1 1 0 - 0 1 X 0 2 0 1 0 1 1 1 0 0 0 0 rufaaoana 2 0 1 0 1 0 1 - 1 0 0 0 0 0 1 1 1 0 0 - 1 0 1 0 rugoaa 2 1 0 1 1 1 0 0 0 0 0 0 2 0 1 0 1 1 0 1 0 1 0 aoaXaria 1 0 1 1 1 0 0 - 1 1 0 0 ? ? ? ? ? ? ? ? ? ? ? Î aemipXioata 0 0 1 1 1 0 0 - 1 1 X 0 1 0 1 1 1 0 0 w 2 1 1 0 apadicea 1 0 1 1 0 0 1 - 1 0 0 0 1 0 0 1 1 1 0 2 1 1 0 apeoioaa 2 1 1 1 1 0 0 0 2 0 X 0 0 0 1 0 0 0 0 1 1 1 0 aguamata 1 0 1 1 1 0 0 - 2 0 X 0 2 0 1 0 1 0 0 2 0 0 0 atanatiaefoxmia 1 0 0 1 0 0 0 - 1 0 X X ? ? ? ? ? ? ? ? ? ? 7 ? tubarouXata tuberculata I 1 1 1 1 1 0 1 1 1 X X 0 0 1 1 1 0 0 1 0 1 0 tubarouXata oocoinea I 1 1 1 1 1 0 1 1 0 X X 0 0 1 1 0 0 0 1 0 1 0 uniXataraXia 0 1 1 1 1 0 0 1 1 0 0 X 2 0 1 0 1 0 0 0 0 0 1 varia 0 0 1 1 1 0 0 - 1 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? virginea 1 0 1 0 1 1 0 - 0 0 0 0 2 0 1 0 1 0 0 — 1 0 1 0 waXaXXenaia 2 0 1 1 1 1 0 - 1 0 0 0 0 0 1 1 1 0 0 - 2 0 0 0 ■ D CD C/) C/) w Table 5-3: Character states for epipodial characters. For description of character states see text. 5 * - (0, 1). - = inapplicable, H, ovina H. fa tu i H. elegans H. dathrata H. varia Figure 5-9. Sample epipodîa o f Haliotis spp. Semischematic reconstructions &om cam era lucida dnmings. H aliotis ovina (LAÇM 85-3). H aliotis spadicea ^DLCQ. H aliotis elegans (ÜSNM 360940). H aliotis fa tu i (USNM 486708); from Geiger (1999). Haliotis dathrata (DLG no if); ôom Stewart & Geiger (1999). H aliotis varia CDLG no if); fiom Geiger (1999). Dorsal and ventral tentacles in H spadicea. Undulating mldep^odial foldûlÆ fa tu i and& varia. CauMower-shapedprojections in H elegans. Palm-sh^ed projections în H dathrata. 358 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (Owen etaLy 1971), but has received little attentioa since then with some notable excep tions (Geiger, 1996, 1999a; Simone, 1998; Stewart & Geiger, 1999). The data matrix for the epipodial characters is given in Table 5-3 and Figure 5-9 shows some sample epipodîa with the various character states. Outgroup comparison The epipodium o f Haliotidae is particularly well developed and can be termed hyper trophied as compared to those of the remaining Vetigastropoda. The comparison to other vetigastropods proved to be difhcult, due to the extent to which the haliotid epipodium is hypertrophied. The postulation o f shared similarities for character states could not be satisfactorily resolved. Comparison between non-haliotid vetigastropods proved to be more conducive than the comparison o f haliotid and other vetigastropod epipodia. As a consequence, the establishment o f character-state polarity through outgroup comparison only succeeded at a rudimentary level. Characters and character states 1: Symmetry o f epipodium. Symmetrical: 0; a^mmetrical, dorsal side more developed: I; asymmetrical, ventral side more developed: 2 . 2: Midepipodial tentacles. Absent: 0; present: 1. 3: h/Gdepipodial fold. Present: 0; absent: 1. 4: Dorsal 6 mge tentacles. Absent: 0; present: 1. 5: Ventral ftinge tentacles. Absent: 0; present: 1. 6 : Cauliform projections (e. g., H. elegans). Absent: 0; present: 1. 7: Bare lateral 6 cmg surface (âce)? No: 0; Yes: 1. 8 : Arrangement o f midep^odial tentacles. Scattered: 0; m line: 1. 359 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 9: Palmate structures? Absent 0; present without secondary palms on surface: I; present with secondary pahns on surface: 2 . 10: Epipodial fold. Straight: 0; undulating: I. II: Tuberculate cones on face. Absent: 0; present: I. 12: Sheeth surrounding tentacles. Absent: 0; present: 1. Hypobranchial gland The hypobranchial gland is a highly folded tissue located in the roof o f the mantle cavity. The character states are given in Table 5-4. A sample illustration o f a hypo branchial gland is shown in Figure 5-10. The glandular tissue shows a few modifica tions between species o f abalone, which were coded as follows: I : A few, irregular side branches from main lamellae? No: 0; At the tip: I; Only in pos terior part: 2 . 2: Does posterior part contain lamellae, which intersect anterior lamellae? No :0; Yes: I. 3: Length o f hypobranchial gland. Less than six tunes as long as wide: 0; more than six times as long as wide: I. 4: Number o f lamellae on hypobranchial gland. Less than 20:0; More than 2 0 :1. 5: At tip many regularly spaced, secondary lamellae? Absent: 0; present I. 6 : Height o f lamellae on right side o f hypobranchial gland (m situ). As high as lamellae on left side: 0; higher than on left side: I. 360 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Z ] CD ■ D O Q . C s Q . ■ D CD % O 3 CD 8 ■ D C 5 - 3 CD C 3. 3- CD CD ■ D I C a O 3 ■ o S & o c C/) o ' 3 PI eufotom ari l()ae S o ls a u re llid a e ( SoiagureJIa) L e p e to d rilid a e (Lepatodrilua) F ia a u re llid a e (F ^aaw rella) Troohidae {Teçfula) Turbonidae (Turbo) t^ainiM aaaimilia aurantium a u s t r a l i s b r a s ie r i o Z atb rata o ooooradiata oorrusrata axraoharodii d a i i i (m issing) d isp u s diaeona (d rie d body) d iveraiaolor. eieg an s f a tu i fu ig en s gigrantea glabra hargravaai i r i s Jaonanaia (d rie d body) bamtaohatkana 0 1 0 1 0 0 la e v ig a ta 0 0 X X 0 0 ? ? 7 ? 7 7 madaka ? ? ? ? ? ? 0 0 0 1 0 0 mariaa 2 1 0 X 0 X _ — marmorata 2 X 0 X 0 X • • midaa 1 1 X X 0 0 0 0 7 7 0 0 ovina 2 1 0 5 0 X parva 0 1 0 X 0 X 1 1 I 1 1 0 p ia n a ta 2 X X X 0 X 0 1 0 5 I 0 pourtaleaii 0 0 0 X 0 X 0 0 0 0 0 0 puatulata 1 X X X 0 0 0 0 1 1 0 1 quekatti 0 X 0 X 0 0 0 0 0 1 0 0 roberti 0 0 0 X 0 0 0 0 0 1 0 1 r o e i 0 X 0 X 0 0 1 1 0 1 0 1 rubiginoaa 0 0 0 X 0 0 0 1 0 1 1 0 rubra 2 X 0 X 0 0 0 5 0 0 1 0 rufaaoana 0 X 0 5 X 0 ? ? 7 7 7 7 rugoaa 0 0 0 X 0 X 0 0 1 1 0 0 acalaria 2 X 0 X 0 0 ? ? 7 7 7 7 aemiplioata 2 0 X X X X 0 1 1 1 0 0 apadicea 0 X 0 0 5 0 0 0 1 1 0 1 apeoioaa ? 7 ? 7 7 7 2 1 1 1 0 1 aguamata 0 X X X 0 X 0 1 0 0 1 0 atomatiaeformia 0 0 X X 0 X 1 1 1 1 1 0 tuberculata tuberculata 1 0 X X 0 X 1 1 0 1 0 0 tuberculata coccinea 0 0 0 X 0 0 0 0 0 1 0 0 u n ilateralia ? 7 7 7 ? 7 1 1 1 1 1 0 varia 2 X 0 0 X 0 7 7 7 7 7 7 virginea 1 0 0 X 0 X 0 1 0 5 1 0 walallanaia 1 0 0 X 0 X Table 5-4. Character states for hypobranchial gland characters. For description o f character states see text. 5 = ( 1, 0), 3 inaplicable. at Figure 5-10. Example o f a hypobranchial gland o f K fia u i. Holotype USNM 486708. r = rectum. Scale bar = 5 mm. From Geiger (1999). 362 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Potential characters not used I have not used some characters and observable features o f abalone. The underlying reason for the exclusion o f this data is uncertain belief formation» i.e., I strongly doubt prim a facie that the observable shared similarities can be explained by common ances try. Such considerations are somewhat subjective as pointed out in Chapter 3, hence, I will offer arguments to support my position. Shell morphology Abalone shells are extremely plastic in virtually every aspect o f their morphology, which includes both qualitative and quantitative features. Striking examples for qualita tive characters include the following: lamellosity o f the shell (Figures 4-4,4-5); height o f the spire (Figures 4-12,4-13); rotundity o f the shell, which is under environmental influence (cf. Ino, 1952) and changes during ontogeny (Stewart & Geiger, 1999); shell coloration, although useful in other groups o f organisms (e.g., Westemeat, 1993; Swen son & Bremer, 1997), is extremely variable in abalone as discussed in Chapter 4 (= Geiger, in press). The most prominent quantitative character is the number o f open holes in abalone. I have discussed the variability o f this character already in Chapter 1 ( - Geiger, 1998a) and Chapter 2 (= Geiger & Groves, 1999). Shell mmeralogy Shell mineralogy was shown to differ between abalone species in three distinct pat terns (Dauphin e t a l., 1989; Dauphin & Denis, 1995). These results were already dis cussed in Chapter 2. Additional problems with mineralogical characters arise when evaluating studies regarding the mmeralogy o f Recent organisms into account The con nection between sea water temperature and mineralogical composition o f shells, com 363 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. bined with the observatfoa that the supposedly aragonite-specific Feigl stain also binds to high magnesium caicite (C. Hedegaard, pers. comm.), are particularly significant. The influence o f sea temperature on the magnesium content in calcium has been demon strated in various invertebrates (e.g., Weinbauer Velirimov, 1995; Hastings et a i, 1998). As Haliotidae occur over a substantial temperature range fiom below 10°C in the case o f the pinto abalone (HI kamtschatkana) to greater than 20® C for many tropical species, it is highly questionable that the shell mineralogy will harbor any phylogenetic signal. ANALYSES WITH MORPHOLOGICAL CHARACTERS An outgroup-rooted analysis o f all 32 ingroup taxa for which at least some molecu lar data was available, with morphological data and the six outgroups added, was per formed. The heuristic search was constrained so that the ingroup was monophyletic [constraints outgroups=((l-6)(7-58))], because the monophyly o f the femily Haliotidae is not in question (Figure 5-11). The analysis found 680 MPRs of length 1290 (Cl = 0.519, R I=0.699, RC = 0.363). The strict consensus tree showed hardly any resolution. The only topologies recovered were: (Scissurellidae (Lepetodrilidae, Turbinidae)); {{pustulata^ rugosa) virginea) asinina) {tuberculata tuberculata, tuberculata coccinea) australis); {{laevigata^ scalaris) roei); and (pourtalesiU w alallensis), I suspect that the large amoimt o f missing data m the she outgroups was largely responsible for the many MPRs found. The low information content o f the morphological data is also evident fiom the relationsh^s within the outgroup taxa. Although the between-family relation ships within Vetigastropoda is currently uncertain (cf. Baten, 1975; H as^runar, 1988b; Tillier e t a f, 1994; Hickman, 1996; Harasewych et al., 1997), the monophyly o f the Trochoidea (— Trochidae and Turbinidae) is undoubted (K ckm an & McLean, 1990; 364 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. é £ Lepetodrilidae Turbinidae Scissurellidae Pleurotomariidae Trochidae Hssurellidae asinina virginea pustulata rugosa corrugata assimilis sorenseni cracherodii discus walallensis pourtalesii Julgens gigantea kcantschatkana madaka rufescens iris midae diversicolor glabra varia ovina tuberculata coccinea t. tuberculata australis coccoradiata rubra cyclobates scalaris laevigata roei I § IP NZ IlO z NZ RSA | e m NZ Figure 5-11. Strict consensus tree o f 680 MPRs 6 om combmed analysis o f allo^rme (Brown, 1993), lysin O Lee & Vacquier, 1995), I6S mtDNACWiay, unpubl.), and mor phology (this study), with she vetigastropod outgroups. The low resolution is due to excessive missing data. 365 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Hickman, 1996). Figure 5-11, however, showed these two families separated by three nodes. An unrooted analysis for the same 32 ingroup taxa as above was carried out with ail available data, but without the outgroups. This analysis was performed in order to assess whether the large amount o f missing data in the outgroup taxa was responsible for the low resolution in Figure 5-11. The analysis found 38 MPRs o f length 1256 (Cl = 0.525, RI = 0.703, RC = 0369. Figure 5-12). This sharp drop in number o f equally parsimo nious trees clearly demonstrates that the amount of missing data in the outgroups causes the parsimony algorithm to find many equally parsimonious'topologies. The polytmy for the north-Pacific group, which was resolved in the previous analyses (see Figure5- 4) without the morphological data can not be explained by missing data alone. It seems that the morphological data actually reduced the resolution in that group. Note that the between-outgroups relationships are peculiar in that the two top shell families Trochi dae and Trubinidae, which are not doubted to be sistergroups, are separated by three nodes. This fact points to the questionable nature of the morphological data set, although one can also argue that these morphological clmacters are not useful at the betweeen families level, but may be well suitable to elucidate within-family relationships. An analysis without the allozyme data was performed to determine the relative con tribution to the number o f MPRs o f missing data compared to the low information con tent in the morphological data. The allozyme data-set is the one with the least taxa, hence, with most missing data. This analysis included 30 taxa. H aliotis coccoradiata and K virginea were omitted, because the only molecular data available were allo^m e frequencies. The analysis o f lysm, 16S mtDNA, and the morphological data produced 106 MPRs o f 1123 steps (Cl = 0.537, RI = 0.719, RC = 0386. Figure 5-13). The num ber o f MPRs was approximately 2.5 times as large in the analysis made without the 366 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. { E asinina walallensis pourtalesii assim ilis corrugata cracherodii discus Julgens gigantea kamtschatkana madaka rufescens sorenseni iris varia glabra ovina diversicolor rubra coccoradiata cyclobates roei scalaris laevigata midae australis tuberculata coccinea t. tuberculata virginea pustulata rugosa IP 2 NZ RSA NZ | e m NZ |io Figure 5-12: Strict consensus tree o f 32 ingroup taxa fîrom combined analysis o f allo^m e (Brown, 1993), lysm (Lee & Vacquier, 1995), 16S mtDNA (Wray, unpubL), and morphology (this stiufy), without ou% toi^. 367 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. asinina IP walallensis pourtalesii assimilis corrugata P cracherodii discus fulgens gigantea k^tschatkana madaka rufescens. sorenseni iiis NZ varia glabra ovina diversicolor cyclobates rubra roei scalaris laevigata midae RSA australis NZ tuberculata coccinea t. tuberculata pustulata rugosa | e m |io Figure 5-13, Strict consensus tree for combined analysis o f lysin. (Lee & Vacquier, 1995), 16S mtDNA (Wray, unpubl.), and morphology (thisstucfy),, without allozyme OBrown, 1993) data. 36S Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. allozyme data. The only area o f the tree that is affected is the north Pacific taxa. When including allozymes, the {pourtalesii, w alallensis) clade is recovered, byt without ailo^unes these two taxa are found in the overall polytomy o f the north Pacific taxa. Additionally a small rearrangement within the Australian endemic species is found, but in both cases the Australian endemic species are fully resolved. The consensus tree shows, that in some areas o f the tree the morphological data seem to remove resolution due to character conflict. This is seen by the large polytomy o f the north Pacific taxa.lin other areas o f the tree the morphological characters do not contradict the molecular data as shown by the fully resolved remainder o f the tree. In conclusion, the use o f morphological data m this particular study highlights two problem areas.^The first is that when a small morphological data-set is added to a larger data-set o f other characters, the amount o f missmg data may prove to be problematic in a total evidence cladistic analysis. Second, morphological data, like any other data set, may be useful in one area o f the tree, but may even obscure evolutionary relationships amongst taxa in other areas o f the tree. The latter is clearly shown with the north Pacific taxa and the outgroups. CLASSIFICATION The classification o f abalone has been unresolved. Fourteen genus-level taxa have been proposed, which were discussed in Chapter 1 (= Geiger, 1998a). Lee & Vacquier (1995) made some taxonomic recommendations, but these are fundamentally flawed in that the ^ p e species o f Hcdiotis s. s., ff, asinina, was not been included in their study. Accordingly, Geiger (1996,1998a) recommended use o f a smgle genus until a compre hensive phylogenetic hypothesis has been fbimed. The present study has advanced this goal it is appropriate to discuss the classification within Haliotidae. I will highlight 369 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. those Sidings that are consistent between the several different analyses and omit from the discussion all those potential groupings and taxa for which there is no sound basis fbr an in-depth appraisal. HaiiotiSj sensus stricto The type species H. asinina occupies an isolated position in the phylogenetic tree. It does not group with any other abalone species. There are two possible ways to deal with this conclusion. Either the genus H aliotis is monotypic, which is a little-favored option, particularly amongst phylogeneticists, because a monotypic genus is by defini tion monophyletic, i.e., it does not help in the understanding of shared similarities among species. The alternative is to treat H aliotis as the single genus for the family Haliotidae, i.e., to continue the current practice. To restrict the use o f H aliotis to ail the species that are not assigned to another genus-level taxon (see below) is not advocated, because it would make that taxon polyphyletic. Nordoüs The most prominent clade contains the north Pacific species on both sides o f the Pacific, i.e., the large Japanese species and all west American species, as well as the tropical American species on either side o f Panama. For these species either the name Nordotis or N otohaliotis has been employed (e.g., Kira, 1962). N otohaliotis has as its type species the Australian H. rubra (see below), therefore, N otohaliotis can not be applied fr)r the north Pacific clade. Two genus-level taxa have as type species members o f the north Pacific clade: Nordotis Habe & Kosuge, 1964 (type = K gigantea Gmelin, 1791), and U sahaliotis Habe & Kosuge, 1964 (type = H cracherodii Leach, 1814). 370 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Applymg the first reviser’s principle, I recommend the name Nordotis to be applied to this group. Nordotis is characterized by one rather weak radular synapomorphy, the concavity in the primary ridge on the lateral tooth 1. This character is also found in m some other abalone taxa, but is a rather consistent feature o f the north Pacific species. The epipodium is highly asymmetrical with the ventral side being more strongly developed than the dorsal side. In the UTR portion of the lysin gene all species contain two notice able gapstretches with an interspersed variable region in positions I to 10, an insert sequence in positions 26 to 39, and a consistent gapstretch in positions 145-153 (see Table 4-3). Notohaiiotis The second consistent group contains the endemic Australian species, to the exclu sion o f those warm water Australian species, which have a wide Indo-Pacific distribu tion, such as H. dathrata^ H ovmoy and K varia, A number of genus-level taxa could be applied to this group: E xohaliotis Cotton & Godfirey, 1933 (type: H, cyclobates)^ A/unnourrr Iredale, 1927 (type: Æ ôrozfm^Aéoto/iforis Cotton &Godfiey, 1933 (type: K scalaris), and Notohaliotis Cotton & Godfr^, 1933 (type: K rubra). The most fie- quently used name is Notohaliotis. Note that Padollus has often been used for some of the Australian species, particularly H. scalaris. The type o f Padollus, however, is the south Afoican H. parva, hence Padollus can not be applied to the clade o f Australian endemic species. No diagnostic syn^m orphies can be given for this weakly supported clade. The species are grotq>ed by rather diffuse characters, such as character state *d’ in position 75 of the lysin UTR. 371 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Sankaliotis The last consistent group is the Indo-Pacifîc species. Membership in this clade is not as well defined as in the other clades but it seems to contain at least K diversicolor, K glabra, H. ovina, and K varia. Ovinotis Cotton, 1943 (type: H. ovina) and Sanhali - Otis Iredale, 1929 (type: K varia) have been proposed as the generic name for species in this clade. Sankaliotis has priority over Ovinotis. No diagnostic synapomorphies can be given for this clade. The species are grouped by rather diffuse characters. It seems that a strong posteriobasal projection on the rachidian tooth and a strongly angulated primary ridge on lateral tooth one are found in some o f these taxa (Figure 5-6: B). These characters are also foimd in some other taxa such as H. facnensis, which I suspect to belong in this clade, but the present data do not justify a &m conclusion. Use of Genus-level taxa I recommend the use o f only the single genus-level taxon H aliotis, because the cladistic hypothesis is not firmly established and is based in some cases on too little data. Additionally, not all species can be assigned to any given genus-level taxon with any degree o f certainty. Those who wish to emphasize particular groupings may use subgenera, which are not mandated fbr unambiguous identification o f a species with a bmomen. It also avoids the use o f the polyphyletic taxon H aliotis s.s. 372 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Conclusions The conclusions o f this dissertation are summarized as follows: CHAPTER 1: RECENT TAXA There are 56 valid species with 10 valid subspecies out o f the ever 200 species- level taxa proposed (Geiger. 1998a. 1998b. 1999; Geiger & Stewart, 1998; Stewart & Geiger, 1999). The species count has been reduced from the previously accepted 75 by 20 species. One additional species was described. This demonstrates that a critical eval uation o f the ingroup taxa is a crucial first step in any phylogenetic analysis. CHAPTER 2: FOSSIL TAXA Abalone are rare in the fossil record. This scarcity is due to a combination o f the rocky habitat and the nacreous shell, which makes abalone little likely to be preserved in the fossil record. The first records finm the Upper Cretaceous (Maastrichian) does most likely not represent the root o f the lineage. The ecology of abalone seems not to have changed over the documented time span. No on-shore/off-shore pattern o f vertical distribution could be detected. CHAPTER 3: DNA SEQUENCE ALIGNMENT The critical evaluation o f the coding related aspects in cladistic analysis has revealed inconsistent, particularly in how DNA sequences are used. As I used a combination of various such, an evaluation is at the heart o f a cladistic analysis. A strict compari son o f the treatment o f morphological data with that o f sequence data, based on philos ophy o f science (jparticularly the nature o f observation, abductive inference, explanation, classification theory, and relevance) has led to the postulation o f a new methodology 373 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. for the treatment o f sequence data in phylogenetic systematics (Geiger & Fitzhugh, m review; Geiger, 1999b). First, DNA sequence alignment must be limited to the smallest identifiable fragment, what 1 call minimum fragm ent alignment (relevance issues). Sec ond, g ^ s must be treated as a fifth character state, because they are postulated during the classification act o f observations. Third, any differential weighting scheme is with out a basis, because observations are equivalent as such, and because such weighting schemes introduce an explanatory element into the observational phase. Fourth, differ ences in alignments are due to questionably aligned sequences. Recoding of such areas can only be accomplished with data reduction strategies such as exclusion and data con traction. Flexible coding strategies such as elision, polymorphic coding, missing data coding, presence/absence coding, and case sensitive coding, introduce internal conflicts and inconsistencies. For few but very dissimilar taxa, new coding strategies o f block coding and stretch coding h&ve been introduced, which have been applied in Chapters 4 and S. CHAPTER 4: BIOGEOGRAPHY The distribution o f all 56 species is documented, based on specimen data from museum collections (Geiger, in press). There are no abalone with a global distribution. The rather vague distributional indications for some species have been specified more exactly. The biogeographic analysis usmg Brooks parsimony revealed a general Indo- Pacific origin o f the family- The precise origin can, however, not be pinpomted more precisely with the data at hand. Two o f the biogeographical hypotheses (Tethys origin. Pacific Rim origin) are considered less, likely, but the Indo-Pacific hypothesis is not without inconsistencies, most likely due to the potentmlly long evolutionary history o f the fomily, reaching back at least to the middle Mesozoic. 374 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CHAPTER 5: TOTAL EVIDENCE CLADISTIC ANALYSIS All data analyses show the following patterns: - a north Pacific group including the Japanese as well as the west American species; - a group o f species endemic to Australia; - a group of widespread Indo-Pacific species; - the three New Zealand species are dispersed over the entire tree. Some molecular synapomorphies can be identified for some o f the above groups, whereas the morphological characters do not show clear evolutionary patterns. The combination o f molecular data sets contributes to the resolution of the combined topol ogy, despite some missing data. The addition o f morphological characters, although constituting a complete data-set, allows multiple character state optimizations leading to a large polytomy among the north Pacific taxa. Outgroup comparison is currently of little value, because o f the questionable simi- lariQf statements for the morphological characters, which are the only data available for the six vetigastropod outgroups. These sunHari^ statements lead to rather questionable resolutions among the outgroups. The large amount of missing data due to no molecular data being available for the outgroups accounts for the nearly unresolved strict consen sus tree. Only four o f the 17 proposed genus-level taxa may be used at this time. The only genus H aliotis, s. L, for all abalone is suggested. Additionally, four subgenera may be used: Haliotis^ j. s., for its Indo-Pacific type species K asmina; Nordotts^ for the north Pacific species; NotohaliotiSy for the species endemic to Australia; and SanhaliotiSy for some o f the wide-spread Indo-Pacific taxa. All other genus-level taxa do not warrant recognition at this time. 375 Reproduced with permission of the copyright owner. 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Induction o f polyploidy and embryonic development o f the abalone, H aliotis diversicolor, with temperatrue treatm ent American M alacologie Bulletin 14:139-147. Yaron, I. 1983. The Haliotidae of the Red Sea. Levantina 42:486-493. 423 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. U M I films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. 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Asset Metadata

Creator Geiger, Daniel Ludwig (author)

Core Title A total evidence cladistic analysis of the Haliotidae (Gastropoda: Vetigastropoda)

School Graduate School

Degree Doctor of Philosophy

Degree Program Biology

Degree Conferral Date 1999-12

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag biology, zoology,OAI-PMH Harvest

Language English

Contributor Digitized by ProQuest (provenance)

Advisor McLean, J.H. (committee chair), Zimmer, R.L. (committee chair), [illegible] (committee member)

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c17-573630

Unique identifier UC11350940

Identifier 9987597.pdf (filename),usctheses-c17-573630 (legacy record id)

Legacy Identifier 9987597

Dmrecord 573630

Document Type Dissertation

Rights Geiger, Daniel Ludwig

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA