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A Largely Cryptic, Adaptive Radiation of Parasitic Snails: Sibling

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A Largely Cryptic, Adaptive Radiation of Parasitic Snails: Sibling 4 A largely cryptic, adaptive radiation of parasitic snails: sibling species in Leptoconchus (Gastropoda: Caenogastropoda: Coralliophilidae), associated with specific coral hosts (Scleractinia: Fungiidae) Adriaan Gittenberger and Edmund Gittenberger A largely cryptic, adaptive radiation of parasitic snails: sibling species in Leptoconchus (Gastropoda: Caenogastropoda: Coralliophilidae) associated with specific coral hosts (Scleractinia: Fungiidae) This text is not issued for purposes o f zoological nomenclature (see ICZN Art. 8.2.) Adriaan Gittenberger1 and Edmund Gittenbergeri2 1 National Museum ofNatural History, P.O. Box 9517, NL 2300 RA Leiden. 2 Institute o f Biology, University Leiden, P.O. Box 9516, NL 2300RA Leiden The Netherlands. E-mail: gittenberger affytatur alis. nnm. nl Key words: parasitic snails; coral reefs; coral/mollusc associations; Coralliophilidae; Leptoconchus; Sclerac­ tinia; Fungiidae; Indo-Pacific Abstract Molecular analyses ................................................................. 67 Taxonomical implications ..................................................... 68 A large, cryptic, adaptive radiation is revealed. Fourteen Lepto­ Systematic part........................................................................ 68 conchus species (Gastropoda: Coralliophilidae) that live associ­ Leptochonchus Riippell, 1834......................................... 68 ated with a variety of mushroom coral species (Scleractinia: The genus ............................................................................. 68 The species............................................................................... 70 Fungiidae) are provisionally described as new to science, i.e. L. inactiniform is................................................................. 70 Leptoconchus inactiniformis, L. inalbechi, L. incrassa, L. incy­ L. inalbechi ......................................................................... 70 closeris, L. infungites, L. ingrandifungi, L. ingranulosa, L. inli­ L. incrassa............................................................................ 71 max: L. inpileus, L. inpleuractis, L. inscruposa, L. inscutaria, L. L. incycloseris..................................................................... 73 intalpina, L. massini. [These names will be made available in the L. infungites......................................................................... 73 near future.] Although their identities as separate gene pools are L. ingrandifungi ................................................................. 74 convincingly demonstrated by molecular data, most of these L. ingranulosa..................................................................... 75 species cannot be identified unequivocally on the bases of only L. inlimax ............................................................................. 77 conchological characters. Shell shape and sculpture are only L. inpileus............................................................................. 78 partially diagnostic because of the interspecifically strongly L. inpleuractis..................................................................... 79 overlapping character states and the large phenotypic plasticity. L. inscruposa....................................................................... 81 Environmental conditions, sexual dimorphism and probably L. inscutaria........................................................................ 82 protandry may affect shell size, shape and sculpture in ways that L. intalpina........................................................................... 82 are still insufficiently known. Flowever, in accordance with the L. m a ssin i............................................................................. 83 molecular data, the ecological data, i.e. host species preferences, Discussion and conclusions ...................................................... 85 do reveal the identity of the various gastropod parasite species Acknowledgements ..................................................................... 86 that were found to be associated with only one or a restricted R eferences..................................................................................... 86 number of fungiid species and have large ranges, similar to those of their hosts. None of the host coral species was found to be associated with more than one Leptoconchus species. Introduction Contents Within the gastropod family Coralliophilidae Chenu, 1859, the genus Leptoconchus Rüppell, 1834, is Introduction ................................................................................... 61 extreme in various ways. The snails live in bore-holes Material and methods.................................................................. 63 in corals, locked up there for most of their lives. A Morphology................................................................................ 63 host specificity was mentioned already by Deshayes Fieldw ork .................................................................................. 63 (1863: 124), but several taxa were introduced in the DNA extraction and sequencing ......................................... 64 Sequence alignment and phylogenetic analyses .............. 65 past without any details on the associated coral host Systematic descriptions.......................................................... 67 species. That means that a potential clue to their R esu lts ............................................................................................ 67 identity is not always available. This is essential since 62 A. Gittenberger, E. Gittenberger - A largely cryptic, adaptive radiation o f parasitic snails 120 Localities 10. Fig. 1. The Indo-Pacific region, from the Red Sea to the Hawaiian Archipelago, illustrating the research localities 1-10. 1, MarsaNakari, c. 350 km S of Hurghada, Egypt (Red Sea); 2, Oman; 3, Vilamendhoo Island, Ari Atoll, Maldives; 4, Phiphi Islands, Krabi, Thailand; 5, Palau; 6, Siladen and Bunaken Islands, N Sulawesi, Indonesia; 7, Togian Islands, E Sulawesi, Indonesia; 8, Spennonde Archipelago, SW Sulawesi, Indonesia; 9, Wakatobi, SE Sulawesi, Indonesia; 10, Bali, Indonesia. in the course of evolution from free-living to boring distinguished by Massin and Dupont (2003). These snails, the shells have lost most potentially diagnostic authors, while summarizing the state of the art in characters of shape, sculpture or colour pattem. Apart Leptoconchus systematics and ecology, distinguished from that, there seems to be a great phenotypic nine so-called Operational Taxonomic Units (OTUs plasticity in relation to environmental factors like 1-9), without clarifying the taxonomic status of these size and other characters of the coral host. We entities. observed a conspicuous sexual dimorphism. The Since only shell morphology and anatomy do not variation in conchological characters might be even result in unequivocal results, an additional discriminating more confusing when Leptoconchus species are tool had to be introduced. Here we describe the results protandric hennaphrodites indeed, as was convincingly of a molecular analysis, on die basis of DNA sequencing suggested by Richter and Luque (2004). data for many snails that were identified as Leptoconchus The animals have to be collected with a hammer, spec. This research material was collected from since only by breaking the coral hosts to pieces they several fungiid hosts, in a vast range, from the Red become available for study. This implies that for both Sea in the west to Palau in the east (fig. 1). It turned technical and ethical reasons large series of specimens, out that the OTU's distinguished by Massin and which are a prerequisite to study the variation in shell Dupont (2003) are not always equivalent to separate shape, could not be acquired. As a consequence of gene pools, i.e. species. In several cases an OTU all this, the genus is still poorly known, despite the turned out to be composed of more than one species, fact that Massin (1982,1983) published some useful which are often not even sister taxa or monophyletic reviews of our current knowledge on Leptoconchus groups. It has to be concluded that Leptoconchus is and closely related genera. much more diverse than hitherto thought. On the This paper deals mainly with the Leptoconchus basis of our results we may additionally conclude species that are associated with mushroom corals that most probably a relatively high number of species (Fungiidae). On the basis of general shell shape, i.e. remains to be discovered and described. height/width ratio, the shell surface, which is either For the moment being, not all the Leptoconchus smooth or not, the presence versus absence of an species that emerge from the molecular analyses can operculum, and the location of the bore-hole, on be characterized morphologically. Here we illustrate either the upside or the basis of the mushroom coral and describe the shell of the future holotype, which disc, some species or species groups have been is always a relatively large, in all probability female Parasitic gastropods and their coral hosts - Chapter 4 63 snail. Whenever possible the associated, smaller, alcohol 96%. A superficial analysis did not result in male shell is also described and the largest shell the discovery of species specific anatomical details. measurements found are added. The intraspecific Because the snails do not possess a radula or jaws, variation remains unknown. that potential source
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