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Habitat Preferences of Coral-Associated Wentletrap Snails (Gastropoda: Epitoniidae)

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Habitat Preferences of Coral-Associated Wentletrap Snails (Gastropoda: Epitoniidae) Contributions to Zoology, 82 (1) 1-25 (2013) Habitat preferences of coral-associated wentletrap snails (Gastropoda: Epitoniidae) Adriaan Gittenberger1, 2, Bert W. Hoeksema1, 3 1 Department of Marine Zoology, Naturalis Biodiversity Center, PO Box 9517, 2300 RA Leiden, The Netherlands 2 Institute for Environmental Sciences and Institute for Biology, Leiden University, PO Box 9516, 2300 RA Leiden, The Netherlands 3 E-mail: [email protected] Key words: coral-mollusc associations, Fungiidae, Indo-Pacific, parasitic snails, Scleractinia Abstract Coral densities and depth ranges .......................................... 18 Substrates ................................................................................... 19 Examination of about 60,000 scleractinian corals of the fami- Coral sizes and risk of host coral turnover ........................ 19 lies Dendrophylliidae, Euphylliidae and Fungiidae for the pres- Conclusions and recommendations ............................................ 20 ence of associated wentletrap snails (Gastropoda: Epitoniidae) Acknowledgements ........................................................................ 21 revealed various ectoparasitic life history strategies. Twenty References ........................................................................................ 21 Indo-Pacific wentletrap species were found, which were either host-specific or generalist. Most species were associated with mushroom corals, especially free-living species belonging to Introduction the Fungiidae. Snails showed different preferences with regard to their position relative to mushroom corals, the host’s size and its substrate. No preferences for depth were found. Infestation In light of the degradation of coral reefs and the poten- rates of mushroom corals in multi-species assemblages were tial for coral species extinctions (Carpenter et al., negatively correlated with coral densities, which indicates that 2008), understanding the biodiversity and ecological epitoniid veliger larvae may actively look for preferential hosts. complexity of reef communities and their species in- Indirect proof was found that burrowing shrimps remove any terdependences is essential (Castro, 1988; Stella et al., epitoniid that is on or underneath the mushroom coral under which they have their burrow. Fishes like wrasses and damsel- 2010, 2011; Hoeksema et al., 2012a). In the present fishes were seen to eat the snails the moment their host corals study corals of the scleractinian families Dendrophyl- were overturned, which suggests that the host corals may pro- liidae, Euphyl liidae, and Fungiidae with their associ- vide the snails with protection against predators. ated wentletraps (Gastropoda: Epitoniidae) are used to study such interrelationships. Epitoniids are generally known to live in association Contents with anthozoans (Cnidaria), such as species of Actinia- ria (Robertson, 1963, 1966, 1983; Perron, 1978; Schi- Introduction ........................................................................................ 1 mek, 1986; den Hartog, 1987; Kokshoorn et al., 2007; Material and methods ....................................................................... 3 Fieldwork localities and general methods ............................ 3 Hoeksema and Crowther, 2011), Zoantharia (Zahn, Sampling ........................................................................................ 3 1980; Robertson, 1981) or Scleractinia (Gittenberger Transect studies ........................................................................... 3 and Gittenberger, 2005). Morphological and molecular Measurement of coral sizes ...................................................... 8 studies have shown that a large, partly cryptic, adap- Overturned host corals ........................................................... 10 tive radiation occurred among these epitoniids (Gitten- Results ............................................................................................... 10 Abundance of infestations ...................................................... 10 berger et al., 2000; Gittenberger, 2003; Gittenberger Associations ............................................................................... 11 and Gittenberger, 2005; Kokshoorn et al., 2007). Their Coral densities and infestation rates ................................... 11 geographical distribution ranges track those of their Depth ranges .............................................................................. 11 hosts, with the highest numbers in the Indo-West Pa- Substrates and attachment ..................................................... 11 cific, especially in the Coral Triangle (Gittenberger Coral sizes .................................................................................. 13 Overturned host corals ............................................................ 14 and Gittenberger, 2005). Discussion ......................................................................................... 17 Although the genetic integrity of epitoniids can be The trophic role of coral-associated epitoniids ................ 17 demonstrated by molecular data (Gittenberger et al., 2 Gittenberger & Hoeksema – Habitat preferences of Epitoniidae 2006; Churchill et al., 2011), shell characters are only or ectoparasites (Gittenberger, 2003; Gittenberger and partially diagnostic in identifications at species level. Gittenberger, 2005). In the present study, they are con- In general, life habit and characters of the operculum, sidered parasites because they live a long period of the jaw, the radula and the spawn, characterize these their life history on a single host without killing it (see conchologically cryptic species much better (Gitten- Castro, 1988). berger and Gittenberger, 2005). These recent findings Most coral-associated wentletraps are found with contradict the hypothesis of Bouchet and Warén (1986: mushroom corals (Fungiidae), which are the main fo- 469) stating that little morphological variation among cus of this study. The Fungiidae consist of over 50 epitoniids may reflect a low degree of specialization. species exclusively in the Indo-Pacific (Hoeksema, A fundamental condition for the identification of coral- 1989, 1993a, 1993b, 2009, 2012b; Hoeksema and Dai, associated molluscs consists therefore of a good knowl- 1991; Gittenberger et al., 2011; Benzoni et al., 2012). edge of the available host coral species (Gittenberger These corals have a high survival rate during bleach- et al., 2000; Kleemann and Hoeksema, 2002; Gitten- ing events (Hoeksema, 1991a; Hoeksema and Mat- berger and Gittenberger, 2005, 2011; Hoeksema et al., thews, 2011; Hoeksema et al., 2012b), which is also 2012a). important for the survival of their associated fauna The snails may use their anthozoan hosts as food (Hoeksema et al., 2013). Most mushroom corals are source, as protection against predators, or shelter from free-living in adult phase and can easily be picked up turbulence. Due to their hidden position, observations and checked for parasites. Depending on shape, size of actual feeding epitoniids are scarce (Perron, 1978; and weight (Hoeksema, 1988, 1991b), such corals may den Hartog, 1987; Gofas et al., 2011), but nematocysts tumble downward along slopes, be pushed by bottom- in epitoniid gut contents indicate a predatory or para- dwelling animals, currents and waves, and they may sitic behaviour (Perron, 1978). Apparently, the snails even move themselves (Hubbard, 1972; Jokiel and are unaffected by the venom in these nematocysts. Cowdin, 1976; Yamashiro and Nishihira, 1995; Hence, epitoniids are either considered highly special- Plusquellec et al., 1999; Hoeksema and de Voogd, ized micropredators (Robertson, 1970; Hadfield, 1976) 2012). This mobility may help mushroom corals to Fig. 1. Research localities in the Indo- West Pacific: 1. Marsha Shagra and Marsa Nakari, c. 350 km S of Hurghada, Egypt; 2. Vilamendhoo Isl., Ari Atoll, Maldives; 3. PhiPhi Isl., Krabi, W Thailand; 4. Ok- inawa, Rykyu Isl., S Japan; 5. Cebu, Cen- tral Philippines; 6. Berau Isl., E Kaliman- tan, Indonesia; 7. Siladen and Bunaken Isl., N Sulawesi, Indonesia; 8. Togian Isl., Central Sulawesi, Indonesia; 9. Spermon- de Archipelago, SW Sulawesi, Indonesia; 10. Bali, Indonesia; 11. Flores, Indonesia; 12. Ambon, Indonesia; 13. Palau; 14. Great Barrier Reef, E Australia. Fig. 2. Research sites in the southern part of the Spermonde Archipelago, SW Sulawesi, Indonesia, consisting of patch reefs and a barrier reef: 1. Langkai; 2. Kapodasang (shoal); 3. Bone Tambung; 4. Kudingareng Keke; 5. Bone Batang; 6. Samalona; 7. Bone Baku (shoal); 8. Lae- Lae. Contributions to Zoology, 82 (1) – 2013 3 disperse and prevent lethal burial in sediment (Abe, species live in environments that are usually avoided 1939; Jokiel and Cowdin, 1976; Hoeksema, 1988; by divers, e.g. at relatively deep (> 25 meter) reef bases Chadwick-Furman and Loya, 1992; Gilmour, 2002; or shallow (< 2 meter) slopes and flats, or in areas with Bongaerts et al., 2012). It may also be functional in strong currents, murky water and sandy bottoms avoiding competition for space (Chadwick, 1988; (Hoeksema, 2012a). Hoeksema, 2004; Yamashiro and Nishihira, 1995; Hoeksema and de Voogd, 2012). However, movements Sampling by the host may also have implications for the sur- vival of the ectoparasitic snails, which will become The snails were collected and conserved in 70% or exposed to predators when the coral accidentally gets 95% ethanol for morphological or molecular
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