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Pubblicazione Mensile Edita Dalla Unione Malacologica Italiana Distribution and Biogeography of the Recent Haliotidae (Gastropoda: Vetigastropoda) Worid-wide Daniel L. Geiger Autorizzazione Tribunale di Milano n. 479 del 15 Ottobre 1983 Spedizione in A.P. Art. 2 comma 20/C Legge 662/96 - filiale di Milano Maggio 2000 - spedizione n. 2/3 • 1999 ISSN 0394-7149 SOCIETÀ ITALIANA DI MALACOLOGIA SEDE SOCIALE: c/o Acquano Civico, Viale Gadio, 2 - 20121 Milano CONSIGLIO DIRETTIVO 1999-2000 PRESIDENTE: Riccardo Giannuzzi -Savelli VICEPRESIDENTE: Bruno Dell'Angelo SEGRETARIO: Paolo Crovato TESORIERE: Sergio Duraccio CONSIGLIERI: Mauro Brunetti, Renato Chemello, Stefano Chiarelli, Paolo Crovato, Bruno Dell’Angelo, Sergio Duraccio, Maurizio Forli, Riccardo Giannuzzi-Savelli, Mauro Mariani, Pasquale Micali, Marco Oliverio, Francesco Pusateri, Giovanni Repetto, Carlo Smriglio, Gianni Spada REVISORI DEI CONTI: Giuseppe Fasulo, Aurelio Meani REDAZIONE SCIENTIFICA - EDITORIAL BOARD DIRETTORE - EDITOR: Daniele BEDULLI Dipartimento di Biologia Evolutiva e Funzionale. V.le delle Scienze. 1-43100 Parma, Italia. Tel. + + 39 (521) 905656; Fax ++39 (521) 905657 E-mail : [email protected] CO-DIRETTORI - CO-EDITORS: Renato CHEMELLO (Ecologia - Ecology) Dipartimento di Biologia Animale. Via Archirafi 18. 1-90123 Palermo, Italia. Tel. + + 39 (91) 6177159; Fax + + 39 (9D 6172009 E-mail : [email protected] Marco OLIVERIO (Sistematica - Systematics) Dipartimento di Biologia Animale e dell’Uomo. Viale dell’Università 32. 1-00185 Roma, Italia. E-mail : [email protected] .it Italo NOFRONI (Sistematica - Systematict) Via Benedetto Croce, 97. 1-00142 Roma, Italia. Tel + + 39(06) 5943407 E-mail : [email protected] Pasquale MICALI (Relazioni con i soci - Tutor) Via Papina, 17. 1-61032 Fano (PS), Italia. Tel ++39 (0721) 824182 - Van Aartsen, Daniele Bedulli, Gianni Bello, Philippe Bouchet, Erminio Caprotti, Riccardo Catta- MEMBRI ADVISORS : Jacobus J. neo-Vietti, Renato Chemello, Fernando Ghisotti, Riccardo Giannuzzi-Savelli, Alberto Girod, Edmund Gittenberger, Folco Giusti, Gerhard Haszprunar, Mauro Mariani, Giulio Melone, Marco Oliverio, Giulio Pavia, Giuseppe Pelosio, Enrico Pezzoli, Winston F. Ponder, Elio Robba, Giuliano Ruggieri, Giovanni F. Russo, Lutfried von Salvini-Plawen, Gianni Spada, Marco Taviani, Anders Warén QUOTE SOCIAL1 1999-2000 Categorie di soci CEE EURO ESTERO EURO Enti, Istituti Sezioni 100.000 51,65 120.000 61,97 Sostenitori (minimo) 90.000 46,48 120.000 61,97 Ordinari 65.000 33,57 85.000 43,90 Giovani (fino 25 anni) 35.000 18,08 40.000 20,66 La quota sociale può essere corrisposta tramite ccp n. 28231207 intestato a: Società Italiana di Malacologia, viale Gadio 2, 20121 Milano. Indicare sempre la causale del versamento. Oppure versando la quota sul cc bancario n. 27/412 dell’Associazione, presso il Banco di Napoli, Ag. 86, Napoli, cod. ABI 1010, CAB 3486 (cod. per versamenti da filiali 0138), intestato alla Società Italiana di Malacologia. È possibile il pagamento anche trami- te carta di credito: CARTASÌ, VISA, MASTERCARD, EUROCARD, senza aggravio di spese, inviando i dati della Vostra carta per fax al n. 081.5514063 o per lettera al Segretario Paolo Crovato. Qualsiasi comunicazione o richiesta da inviare alla SIM deve essere indirizzata: Casella Postale 436-1 80100 Napoli. Bollettino Malacologico Roma, 2000 • Società Italiana di Malacologia , 35 (5-12) [1999] Distribution and Biogeography of the Recent Haliotidae (Gastropoda: Vetigastropoda) World-wide Daniel L. Geiger KEY words: distribution, biogeography, Haltotis, area cladogram. abstract: The distribution of all 56 abalone species is documented based on specimen records. The three models for the origin of the family (Pacific Rim, Indo-Pacific, Tethys) are evaluated. The area cladogram, which treats the distribution of the individual species as characters and the areas as taxa, is in general agreement with an unrooted phylogenetic tree of the taxa for which the geographic occurrence is superimposed. The basal node of the area cladogram is a fairly large polytomy uniting rather distant provinces such as the north Pacific, Australia, and Africa. A similar pattern emerges from the taxon cladogram. Accordingly, a very general Indo-Pacific origin emerges, best in agreement with the Indo-Pacific model for the origin of the family. The apparent discrepancy with the fossil record is discussed. Although the earliest fossils from the Upper Creatceous have been found in the Caribbean and in California, a major hiatus of on the order of 200 million years can easily account for the discrepancy between the earliest foss- sils and the biogeographical reconstruction, as well as for the large basal polytomy in the biogeographical analysis. All species are illustrated, including some distinct juvenile forms, and 30 photographs of live animals. riassunto: Distribuzione e Biogeografìa delle Haliotidae (Gastropoda: Vetigastropoda) recenti. Le Haliotidae comprendono 56 specie recenti descritte, dis- tribuite su scala mondiale, nelle aree temperate e tropicali di entrambi gli emisferi. La distribuzione di tutte le 56 specie è qui illustrata da mappe digitalizzate, sulla base dell'esame oggettivo di esemplari e dell'analisi critica dei dati di letteratura. Sono quindi esaminati i tre modelli proposti per l'origine della famiglia: "Pacific Rim", Indo-Pacifico e Tetisiano. Il cladogramma di area, che tratta le distribuzioni delle singole specie come caratteri e le aree come OTUs, è in accordo generale con un albero filogenetico delle specie (derivato dall'analisi cladistica di dati molecolari ed allozimici) su cui vengano sovrimposte le distribuzioni geografiche. Il nodo basale del cladogramma di area è una politomia piuttosto ampia che riunisce provincie relativamente distanti come il Nord-Pacifico, l'Australia e l'Africa. Un quadro simile emerge dal cladogramma dei taxa. Sulla base dell'analisi di tali cladogrammi emerge un'origine generale Indo-Pacifica del gruppo, che sembrerebbe supportare il modello Indo-Pacifico di origine delle Haliotidae. Vi è un'apparente discrepanza tra questi dati e la loro susseguente interpretazione da una parte, e il record fossile dall'altra. I fossili più antichi noti provengono infatti dal Cretaceo Superiore dei Caraibi e della California. Ciononostante viene discusso come uno iato del- l'ordine di 200 milioni di anni possa facilmente spiegare la discrepanza tra i fossili più antichi e la ricostruzione biogeografica più plausibile; si darebbe così anche ragione dell'ampia politomia basale nelle analisi cladistiche. Tutte le specie sono illustrate, con alcune distinte forme giovanili caratterizzate e 30 fotografie di animali viventi. Daniel L. Geiger, Molecular Systematic! Lab, Naturai History Museum of Los Angeles University, Los Angeles, CA 90007, USA; e-mail [email protected] INTRODUCTION to northeastern Australia has been identified by Talmadge (1963) Abalone are a family of marine gastropods consisting of 56 as a likely cradle of the family. From this nuclear distribution currently described species of world-wide distribution in abalone then dispersed in a star shaped pattern to the tropical and temperate waters of both hemispheres. During northwestern and then northeastern Pacific, to Australia and the Indian Ocean. basis of this model never explicit. revision work on this family I have recently evaluated all Recent The was made taxa, 200 at the species level and 17 at the genus level (Geiger, Model 2 (Fig. IB): Indo-Pacific. The highest present day 1998a). Some other taxonomic matters have been dealt with diversity of the family is found in the Indo-Malayan area. elsewhere (Geiger 1998b, 1999; Geiger & Stewart, 1998, Although it is well appreciated that the correlation of high present Stewart & Geiger, 1999). Knowledge of the distribution of day diversity with origin of the group in question is highly most species is sketchy, particularly due to the absence of problematic, it provides one possible center of radiation for the specimen based accounts. Most indications in the literature may family, and was discussed by LlNDBERG (1992) and BRIGGS be termed intuitive (e.g., Ubaldi, 1993, 1995) with limited (1999). exceptions (Geiger, 1996, 1999; Simone, 1998; Stewart & Model 3 (Fig. 1C): Tethys. From published chromosomal Geiger, 1999). In this second, larger treatment of the family count data Geiger & Groves (1999) considered an origin in the Haliotidae, I concentrate on the distributional pattern of all Tethys Sea. Since other Vetigastropoda and basal gastropods Recent species, including a biogeographical analysis. To date, have a relatively low chromosome number (2n = 18-20: studies on the biogeography ol the family are marked by their Patterson, 1967; Haszprunar, 1988) as compared to the anecdotal nature. Three hypotheses (Fig. 1) for the origin have values found in the Haliotidae (2n = 28-36: see Geiger & been proposed and have been discussed by Geiger & Groves Groves, 1999, for sources) a progression from low to high (1999) in a review of fossil abalone. values was considered. BlELER (1992) also concluded that the Model 1 (Fig. 1A): Pacific Rim. An arc spanning from Japan low chromosome number of patellogastropods is the > 57 < Daniel L. Geiger plesiomorphic condition within the archaeogastropod grade. It would suggest a radiation starting in the Tethys, today represented by the Mediterranean species Haliotis tuberculata Linnaeus, 1758, with 2n = 28, moving eastwards to the Indo- Pacific (2n = 32), and finally reaching the North Pacific (2n = 36). Here the three proposed models are evaluated using specimen based
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