TEMPERATURE REQUIREMENTS FOR MATURATION OF GONADS OF NORTHERN OYSTERS
Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red...
9
Facies (2007) 53:319–327 DOI 10.1007/s10347-007-0110-8 123 ORIGINAL ARTICLE Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea Martin Zuschin · Christian Baal Received: 18 December 2006 / Accepted: 18 April 2007 / Published online: 22 May 2007 © Springer-Verlag 2007 Abstract The shell of a living specimen of the Indo- PaciWc gryphaeid giant oyster Hyotissa hyotis was colo- nized by numerous encrusting, boring, nestling and baZing taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamaria- cean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such diVerences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seaXoor. The cryptic habitats support a species assemblage of crustose algae and forami- nifera that, on exposed surfaces, would occur in much deeper water. Keywords Coral reef · Bioerosion · Encrustation · Cryptic habitats · Palaeoecology Introduction Biogenic hard substrates include a broad variety of mineral- ized organic tissues (Taylor and Wilson 2003) which can be summarized as shellgrounds sensu Dodd and Stanton (1990). Such shell grounds are frequently colonized in modern and fossil marine environments (e.g., Stachowitsch 1980; Kidwell and Gyllenhal 1998; Zuschin et al. 1999). They can be infested during life or after death of the host, depending on whether the shell is external or internal and whether the life habit of the host is epifaunal or infaunal (Taylor and Wilson 2003). Hard substrata are frequently studied by palaeoecologists because encrusting and boring organisms retain their original positions on the substrate after fossilization and the problems of transport and assem- blage mixing are minimized (Brett 1988). Distribution pat- terns of colonizing organisms, however, typically diVer between exposed and cryptic surfaces of inorganic and bio- genic marine hard substrata (e.g., Palmer and Fürsich 1974; Surlyk and Christensen 1974; Nebelsick et al. 1997; Rich- ter et al. 2001). The presence of such diVerences can be used to infer life orientation of extinct fossil organisms if they were infested during their lifetime (e.g., Bordeaux and Brett 1990; Lescinsky 1995; Leighton 1998). Lack of such polarization, in contrast, is expected to occur on rolling hard substrata (Lee et al. 1997) or on frequently overturned substrata such as rhodoliths (Foster 2001). Oysters are typical accessory components in fossil and Recent coral reefs (Crame 1986; Zuschin et al. 2001). The vast majority of such reef-associated bivalves are small (mostly in the size range of a few centimetres), but fre- quently colonized by sclerobionts (e.g., bryozoans, serpu- lids, crustose red algae; terminology after Taylor and Wilson 2002). The gryphaeid oyster Hyotissa hyotis (Linnaeus, 1758) occurs throughout the tropical and sub- tropical waters of the Indo-PaciWc (Slack-Smith 1998; Zus- chin and Oliver 2003a, b) and was recently introduced to the western Atlantic (Bieler et al. 2004). This species is exceptional in that it can reach a large size (up to several decimetres; Stenzel 1971). Although it is typically mas- sively encrusted, its role as a habitat-providing structure for sclerobionts is unexplored. The colonizing organisms are diVerentiated into functional groups, including binding M. Zuschin (&) · C. Baal Department of Palaeontology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria e-mail: [email protected]
Transcript of Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red...
Facies (2007) 53:319–327
DOI 10.1007/s10347-007-0110-8ORIGINAL ARTICLE
Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea
Martin Zuschin · Christian Baal
Received: 18 December 2006 / Accepted: 18 April 2007 / Published online: 22 May 2007© Springer-Verlag 2007
Abstract The shell of a living specimen of the Indo-PaciWc gryphaeid giant oyster Hyotissa hyotis was colo-nized by numerous encrusting, boring, nestling and baZingtaxa which show characteristic distribution patterns. On theupper valve, sponge-induced bioerosion predominates. Onthe lower valve intergrowth of chamid bivalves and thickencrusting associations—consisting mostly of squamaria-cean and corallinacean red algae, acervulinid foraminifera,and scleractinian corals—provides numerous microhabitatsfor nestling arcid and mytilid bivalves as well as forencrusting bryozoans and serpulids. Such diVerencesbetween exposed and cryptic surfaces are typical for manymarine hard substrata and result from the long-term stableposition of the oyster on the seaXoor. The cryptic habitatssupport a species assemblage of crustose algae and forami-nifera that, on exposed surfaces, would occur in muchdeeper water.
Keywords Coral reef · Bioerosion · Encrustation · Cryptic habitats · Palaeoecology
Introduction
Biogenic hard substrates include a broad variety of mineral-ized organic tissues (Taylor and Wilson 2003) which can besummarized as shellgrounds sensu Dodd and Stanton(1990). Such shell grounds are frequently colonized inmodern and fossil marine environments (e.g., Stachowitsch1980; Kidwell and Gyllenhal 1998; Zuschin et al. 1999).
They can be infested during life or after death of the host,depending on whether the shell is external or internal andwhether the life habit of the host is epifaunal or infaunal(Taylor and Wilson 2003). Hard substrata are frequentlystudied by palaeoecologists because encrusting and boringorganisms retain their original positions on the substrateafter fossilization and the problems of transport and assem-blage mixing are minimized (Brett 1988). Distribution pat-terns of colonizing organisms, however, typically diVerbetween exposed and cryptic surfaces of inorganic and bio-genic marine hard substrata (e.g., Palmer and Fürsich 1974;Surlyk and Christensen 1974; Nebelsick et al. 1997; Rich-ter et al. 2001). The presence of such diVerences can beused to infer life orientation of extinct fossil organisms ifthey were infested during their lifetime (e.g., Bordeaux andBrett 1990; Lescinsky 1995; Leighton 1998). Lack of suchpolarization, in contrast, is expected to occur on rollinghard substrata (Lee et al. 1997) or on frequently overturnedsubstrata such as rhodoliths (Foster 2001).
Oysters are typical accessory components in fossil andRecent coral reefs (Crame 1986; Zuschin et al. 2001). Thevast majority of such reef-associated bivalves are small(mostly in the size range of a few centimetres), but fre-quently colonized by sclerobionts (e.g., bryozoans, serpu-lids, crustose red algae; terminology after Taylor andWilson 2002). The gryphaeid oyster Hyotissa hyotis(Linnaeus, 1758) occurs throughout the tropical and sub-tropical waters of the Indo-PaciWc (Slack-Smith 1998; Zus-chin and Oliver 2003a, b) and was recently introduced tothe western Atlantic (Bieler et al. 2004). This species isexceptional in that it can reach a large size (up to severaldecimetres; Stenzel 1971). Although it is typically mas-sively encrusted, its role as a habitat-providing structure forsclerobionts is unexplored. The colonizing organisms arediVerentiated into functional groups, including binding
M. Zuschin (&) · C. BaalDepartment of Palaeontology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austriae-mail: [email protected]
123
320 Facies (2007) 53:319–327
encrusters, destroyers, nestlers and baZers (Fagerstrom1987). The present study investigates the role of this large-sized bivalve as an ecosystem engineer (sensu Jones et al.1994), which provides a substratum for the attachment ofepibionts and refuges from predation, physical or physio-logical stress to other organisms (Gutiérrez et al. 2003).
Material and methods
One living specimen of H. hyotis was taken in May 1996from a shallow-water site (6 m water depth) in the NorthernBay of Safaga (Fig. 1). The study site was dominated by thesoft coral Sarcophyton sp., and the oyster was attached to adead and degraded massive coral colony (Fig. 2). The totaldry weight of the encrusted shell is 7,219 g; three quartersof the weight (5,450 g) are made up of the massivelyencrusted lower valve (Fig. 3). The upper (right) valve hasan anterior-posterior length of 22 cm and a dorsal-ventralheight of 27 cm. The corresponding values for the lower(left) valve are 22£29 cm. The size of the adductor musclescar is 7.5£6 cm (Fig. 4a, b). The maximum thickness ofthe upper valve is ca. 4.5 cm, that of the lower valve (in thearea of the ventral end of the ligament) exceeds 10 cm. Theannual growth increments in the ligament area can be usedto infer the ontogenetic age of oysters (Stenzel 1971) andsuggest an age of more than 40 years for this specimen(Fig. 5). The sclerobiont infestation of this shell has beenevaluated semi-quantitatively from visual inspection of thein situ shell at the seaXoor, from investigations and photo
documentation of the wet surface of the still living oysterimmediately after collecting, and from the dry shell of thedead oyster. Encrusting associations have been studied inthin-sections. Of special interest are diVerences between thelower and upper valve regarding taxonomic compositionand density of the sclerobionts.
Results
The main encrusting binders on the oyster are peyssonne-liacean and corallinacean red algae, bivalves, serpulidworm tubes, and vermetid gastropod shells. The maindestroyers are boring sponges and bivalves (Table 1). Theupper and the lower valve of the oyster, however, are colo-nized by diVerent taxa in diVerent intensities. As a generalfeature, the upper valve shows only scattered and thin cal-careous crusts. Bioerosion is a dominant feature of theupper valve and consists mainly of ubiquitous clionid bore-holes and a single lithophagine borehole (Fig. 6a, b). Incontrast, calcareous organisms densely encrust the lowervalve (Fig. 7b). The thickness of these crusts is highly vari-able but exceeds 6 cm at some places (Figs. 5 and 7). As isobvious in the cross-section (Fig. 5), the ubiquitous clionidboreholes on the lower valve are mainly covered byencrusting epizoobionts, except for marginal parts of theshell (Fig. 7b). Lithophagine boreholes are more abundantthan on the upper valve and preferentially occur in theencrusting associations (Fig. 8).
Vermetid gastropods and sponges are the main coloniz-ers of the upper valve (Fig. 6). Calcareous crusts are verythin and rare, but Wlamentous green algae are abundant.Semisessile soritid and planorbulinellid foraminifera andcalcareous tubes of polychaetes are found in low numbers,along with a single degraded valve of an encrusting chamidbivalve. A xeniid soft coral colony inhabited this valve(Figs. 2 and 3), and a small colony of the scleractinian coralMontipora sp. occurred on the dorsal part in associationwith a nestling mytilid bivalve (Septifer forskali). A juve-nile T. maxima was byssally attached to the ventral part ofthe upper valve (Fig. 6a, b).
The lower valve, in contrast, is densely covered withchamid bivalves (Chama cf. brassica) and a thick, exten-sive layer of additional encrusters (Fig. 7a, b). The latterconsist of squamariacean and corallinacean red algae,acervulinid foraminifera and scleractinian corals, with sub-ordinate contributions by the cheilostome bryozoan Smit-tina sp., serpulids and small oysters (Fig. 8). Theintergrowth of chamid bivalves and these other encrustersproduced a highly irregular, rugged surface, which isdensely covered with serpulid tubes and oVers many micro-habitats for crevice-dwelling limid bivalves (Lima pauci-costata, Ctenoides annulata). Empty serpulid tubes are
Fig. 1 Location map, general bathymetry of the study area (after Pillerand Pervesler 1989), and collection site of H. hyotis in the southwestchannel of the Northern Bay of Safaga
123
Facies (2007) 53:319–327 321
Fig. 2 H. hyotis attached to a dead and degraded massive coral colony in an environment characterized by the soft coral Sarcophyton sp. CbChama brassica; S unidentiWed sponge; Sa Sarcophyton sp.; Xe Xenia sp
Fig. 3 Ventral view of freshly collected wet shell, showing densely encrusted lower valve. Cb Chama brassica; Lp Lima paucicostata; S unidentiWed sponge; Tm juvenile Tridacna maxima. Anterior-posterior length of shell is 22 cm
123
322 Facies (2007) 53:319–327
Fig. 4 Internal views of dry shell. a Upper (right) valve. b Lower (left) valve. Ams adductor muscle scar; C crustose algae; Cb Chama brassica;Cl clionid boreholes; M Montipora cf. informis; P Porites sp.; Se unidentiWed serpulid. Scale in centimetres
Fig. 5 Cross-section of lower valve showing growth increments anddense incrustation. Acc accretion of encrusting associations; Cb Cha-ma brassica; Cl clionid boreholes; L ligament area; Li lithophagine
borehole; M myostracum of the adductor muscle scar. Note annualgrowth increments
123
Facies (2007) 53:319–327 323
occasionally settled by the nestling mytilid bivalve Septiferforskali, and some of the lithophagine boreholes arecolonized by the nestling arcid bivalve Barbatia cf. parva.
Other epizoozoans on the lower valve include a few vermetidgastropods and several small, unidentiWed sponge colonies.
Discussion
Oysters, due to their epifaunal life habit and high preserva-tion potential, are among the most frequently colonizedmolluscan substrata for sclerobionts in modern and ancientmarine ecosystems (Taylor and Wilson 2003). Most oystersin fully marine environments, however, do not exceed afew centimetres in size. In the Northern Bay of Safaga, thecomparatively few large oysters mainly occur in areas asso-ciated with localized high suspension load, for examplewhere slowing longshore currents accumulate particulateorganic matter from upstream seagrass beds and coral car-pets (Riegl and Piller 1997). Such large Hyotissa shells aretypically densely colonized by numerous zoobionts (thisstudy; pl. 12, Fig. 2 in Slack-Smith 1998; Fig. 3 in Bieleret al. 2004, personal observations at the Seychelles, Febru-ary 2002). It has been suggested that nutrient levels deter-mine the degree of fouling in living molluscs (Voight andWalker 1995). However, shell size also strongly aVectsinfestation in that larger shells typically support a higherdiversity and density of fouling species than smaller shells(see Gutiérrez et al. 2003 for a review). Moreover,cemented oysters also provide a stable hard substratum fora long period of time, which increases the likelihood of col-onization (for review see Taylor and Wilson 2003).
The quantitative diVerence among bioeroders and encr-usters between the upper, exposed and lower, cryptic valveagrees with results from other studies on marine hard sub-strata (e.g., Nebelsick et al. 1997; Richter et al. 2001). Thiscan be related to the stable position of the oyster on the sea-Xoor, which yields diVerent colonization patterns accordingto ecological preferences of the respective taxa. Illumina-tion and water turbulence are probably the main factorscontrolling their distribution, followed by competition forspace and food, and predation (Jackson and Winston 1982;Martindale 1992).
The main destroyers on the studied oyster shell are bor-ing sponges and bivalves, which are generally among themost important bioeroders in Cenozoic coral reef environ-ments (Hutchings 1986; Kleemann 1996; Perry 1998).Clionid boreholes strongly dominate on the upper andlower valve of Hyotissa, whereas lithophagine boreholesare more important on the lower valve. Apart from vermet-ids, binding encrusters were mostly found on the lowervalve. Crusts of coralline red algae are dominant compo-nents of modern shallow-water coral reef environments(Hay 1981), where they typically form thick crusts onexposed surfaces (e.g., Martindale 1992). Such crusts areubiquitous in Safaga (Rasser and Piller 1997), where they
Table 1 Colonizing organisms assigned to functional groups on theupper and lower valve of the shell
¡ absent, + rare, ++ common, +++ abundant
Upper valve Lower valve
Encrusting binders
Red algae
Peyssonnelia sp. ¡ +++
Sporolithon ptychoides ¡ +++
Foraminifera
Acervulina inhaerens ¡ +++
Haddonia sp. ¡ ++
Planorbulinoides cf. retinaculata ¡ ++
Sponges
UnidentiWed sponge colonies + +
Corals
Porites sp. ¡ +
Montipora cf. informis + ++
Blastomussa loyae ¡ ++
Bryozoans
Smittina sp. ¡ +
Serpulids
UnidentiWed species + ++
Mollusks
Juvenile Hyotissa ¡ ++
Chama brassica + +++
UnidentiWed vermetid gastropods ++ ¡Destroyers (borers)
Cyanophyceae
Microborings +
Sponges
Clionid traces (Entobia sp.) +++ +
Bivalves
Lithophaga nigra + ++
Nestlers
Foraminifera
Planorbulinella cf. larvata ¡ +
Sorites orbiculus ¡ +
Bivalves
Barbatia cf. parva ¡ +
Septifer forsskali + +
Lima paucicostata ¡ ++
Ctenoides annulata ¡ ++
Tridacna cf. maxima + ¡BaZers
Corals
Xenia sp. + ¡Sarcophyton sp. + ¡
123
324 Facies (2007) 53:319–327
can be found on shells of living gastropods and hermit crab-inhabited shells and where they frequently overgrow deadmolluscs (Zuschin and Piller 1997; Zuschin et al. 2000).
The low abundance of calcareous crusts on the upper valveof the studied Hyotissa therefore requires explanation andcould be due to massive infestation by bioeroding clionid
Fig. 6 External view of upper valve. a Freshly collected wet shell.b Dry shell. Note ubiquitous clionid boreholes on the dry shell. Cb Cha-ma brassica, Li Lithophaga boring, M M. cf. informis, S unidentiWed
sponges, Tm juvenile T. maxima, Ve unidentiWed vermetid gastropods.Scale in centimetres
Fig. 7 External view of lower valve: a freshly collected wet shell; b dry shell. Cb Chama brassica; Lp Lima paucicostata; S unidentiWed sponges,Se serpulid worm tubes. Note dense calcareous crusts on dry shell. Scale in centimetres
123
Facies (2007) 53:319–327 325
sponges. A more likely explanation, however, is that thecommon Wlamentous green algae outcompeted crustosealgae, which are more sensitive to sedimentation onexposed surfaces in this near-shore, turbid area (Fabriciusand De’ath 2001).
Encrusting associations typically show a zonation of spe-cies in response to water turbulence and light, and strongtaxonomic diVerences between exposed and cryptic habitatsare therefore the norm (Riedl 1966; Martindale 1992). Filterfeeders in the Northern Red Sea, for example, have a muchhigher biomass in cavities than on the reef surface (Richteret al. 2001). With respect to light penetration, cryptic habi-tats therefore support communities that would occur onexposed surfaces in much deeper water (Fagerstrom 1987).This is particularly true for the extensive encrusting associa-tions on the lower valve of Hyotissa and three of their domi-nant components: Peyssonnelia sp., Sporolithon ptychoides,and A. inhaerens. On exposed surfaces, these taxa occur atdepths below 20–40 m in Safaga (Rasser and Piller 1997);A. inhaerens was also reported from shallow-water cryptichabitats in the Northern Red Sea (Reiss and Hottinger1984). Among binding scleractinian corals, Blastomussaloyae seems to prefer turbid water, shaded bottoms andcryptic conditions (Head 1978; Sheppard and Sheppard1991; Veron 2000a). On the other hand, M. cf. informis, is avery common coral on reef slopes between 5 and 20 mdepth, with no reported preference for cryptic habitats(Sheppard and Sheppard 1991; Veron 2000b). Chamid biv-alves are among the most prominent encrusters of exposed
dead hard substrata in the northern Red Sea (Zuschin et al.2001; Zuschin and Stachowitsch 2007), but are even moreabundant in cryptic, more sheltered habitats (personal obser-vations). Other binders on the lower valve include cheilos-tome bryozoans and serpulids; although they were notidentiWed down to species level, in coral reef environments,these taxa generally seem to prefer cryptic and semicryptic,downward-facing, sediment-free habitats (Martindale1992). In contrast, solitary-loosely aggregated vermetid gas-tropods were mainly found on the upper valve. Such verme-tid gastropods can be quantitatively important componentsof Recent intertidal to shallow subtidal coral reefs and aretypically found on exposed surfaces (e.g., HadWeld et al.1972; Zuschin et al. 2001) because they prefer more agitatedwater conditions for their suspension-feeding life habit(Hughes and Lewis 1974, Kappner et al. 2000).
Nestlers are represented almost exclusively by byssatepteriomorph bivalves and mostly preferred the microhabi-tats on the lower valve of H. hyotis: limid bivalves werefound in crevices of the rugged surface, and small arcidsand mytilids occurred in empty serpulid tubes and lithopha-gine boreholes. Such cryptic life habits are typical for thesetaxa (HadWeld 1976; Morton 1983; Zuschin and Oliver2003a). In contrast, the zooxanthellate bivalve T. maxima islight-dependent (Goreau et al. 1973) and was thereforefound on the upper valve of Hyotissa. BaZers, Wnally, areonly of very subordinate importance and include soft-bodied octocorals (Sarcophyton sp. and Xenia sp.) on theoyster’s upper surface.
Fig. 8 Thin-section of encrusting associations of lower valve. Ai A. inhaerens, Li Lithophaga nigra bore hole, M M. cf. informis, P Peyssonneliasp., Se serpulid worm tubes, Sm Smittina sp., Sp Sporolithon ptychoides
123
326 Facies (2007) 53:319–327
Acknowledgements Abbas Mansour, Werner Piller and MichaelRasser helped with Weldwork. Thanks are due to Karl Kleemann, Gra-ham Oliver, Michael Rasser and Michael Stachowitsch for helpful dis-cussions. Karl Kleemann and Norbert Vavra assisted with speciesidentiWcations. The review of Franz T. Fürsich improved the mansu-script. Financial support was provided by FWF project P10715Geo toF.F. Steininger and by project H-140/2000 of the Hochschuljubilä-umsstiftung der Stadt Wien to M. Zuschin.
References
Bieler R, Mikkelsen PM, Lee T, Ó Foighil D (2004) Discovery of theIndo-PaciWc oyster Hyotissa hyotis (Linnaeus, 1758) in the Flor-ida Keys (Bivalvia: Gryphaeidae). Moll Res 24:149–159
Bordeaux YL, Brett CE (1990) Substrate speciWc associations of epi-bionts on Middle Devonian brachiopods: implications for paleo-ecology. Hist Biol 4:221–224
Brett CE (1988) Paleoecology and evolution of marine hard substratecommunities: an overview. Palaios 3:374–378
Crame JA (1986) Late Pleistocene molluscan assemblages from thecoral reefs of the Kenya coast. Coral Reefs 4:183–196
Dodd RJ, Stanton RJ Jr (1990) Paleoecology: concepts and applica-tions. Wiley, New York, pp 1–502
Fabricius K, De’ath G (2001) Environmental factors associated withthe spatial distribution of crustose coralline algae on the GreatBarrier Reef. Coral Reefs 19:303–309
Fagerstrom JA (1987) The evolution of reef communities. Wiley, NewYork, 600 pp
Foster MS (2001) Rhodoliths: between rocks and soft places. J Phycol37:659–667
Goreau TF, Goreau NI, Yonge CM (1973) On the utilization of photo-synthetic products from zooxanthellae and of a dissolved aminoacid in Tridacna maxima f. elongata (Mollusca: Bivalvia). J Zool169:417–454
Gutiérrez JL, Jones CG, Strayer DL, Iribarne OO (2003) Molluscs asecosystem engineers: the role of shell production in aquatic habi-tats. Oikos 101:79–90
HadWeld MG (1976) Molluscs associated with living tropical corals.Micronesica 12:133–148
HadWeld MG, Kay EA, Gillette MU, Lloyd MC (1972) The Vermeti-dae (Mollusca: Gastropoda) of the Hawaiian Islands. Mar Biol12:81–98
Hay ME (1981) Herbivory, algal distribution, and the maintenance ofbetween-habitat diversity on a tropical fringing reef. Am Nat118:520–540
Head SM (1978) A cerioid species of Blastomussa (Cnidaria, Sclerac-tinia) from the central Red Sea, with a revision of the genus. J NatHist 12:633–639
Hughes RN, Lewis AH (1974) On the spatial distribution, feeding andreproduction of the vermetid gastropod Dendropoma maximum.J Zool 172:531–547
Hutchings PA (1986) Biological destruction of coral reefs. Coral Reefs4:239–252
Jackson JBC, Winston JE (1982) Ecology of cryptic coral reef commu-nities. I. Distribution and abundance of major groups of encrust-ing organisms. J Exp Mar Biol Ecol 57:135–147
Jones CG, Lawton JH, Shachak SM (1994) Organisms as ecosystemengineers. Oikos 69:373–386
Kappner I, Al-Moghrabi SM, Richter C (2000) Mucus-net feeding bythe vermetid gastropod Dendropoma maxima in coral reefs. MarEcol Prog Ser 204:309–313
Kidwell SM, Gyllenhal ED (1998) Symbiosis, competition, and phys-ical disturbance in the growth histories of Pliocene cheilostomebryoliths. Lethaia 31:221–239
Kleemann K (1996) Biocorrosion by bivalves. PSZN Mar Ecol17:145–158
Lee DE, Scholz J, Gordon DP (1997) Paleoecology of a late Eocenemobile rockground biota from Otago, New Zealand. Palaios12:568–581
Leighton LR (1998) Constraining functional hypotheses: controls onthe morphology of the concavo-convex brachiopod RaWnesquina.Lethaia 31:293–307
Lescinsky HL (1995) The life orientation of concavo-convex brachio-pods: overturning the paradigm. Paleobiol 21:520–551
Martindale W (1992) CalciWed epibionts as palaeoecological tools:examples from the Recent and Pleistocene reefs of Barbados.Coral Reefs 11:167–177
Morton B (1983) Coral-associated bivalves of the Indo-PaciWc. In:Russel-Hunter WD (ed) The Mollusca, vol 6, Ecology. Academic,New York, pp 139–224
Nebelsick JH, Schmid B, Stachowitsch M (1997) The encrustation offossil and recent sea-urchin tests: ecological and taphonomic sig-niWcance. Lethaia 30:271–284
Palmer TJ, Fürsich FT (1974) The ecology of a Middle Jurassic hard-ground and crevice fauna. Palaeontol 17:507–524
Perry CT (1998) Macroborers within coral framework at DiscoveryBay, north Jamaica: species distribution and abundance, andeVects on coral preservation. Coral Reefs 17:277–287
Piller WE, Pervesler P (1989) The Northern Bay of Safaga (Red Sea,Egypt): an actuopalaeontological approach. I. Topography andbottom facies. Beitr Paläont Österr 15:103–147
Rasser M, Piller WE (1997) Depth distribution of calcareous encrust-ing associations in the northern Red Sea (Safaga, Egypt) and theirgeological implications. Proc 8th Int Coral Reef Symp 1:743–748
Reiss Z, Hottinger L (1984) The Gulf of Aqaba: ecological micropale-ontology. Springer, Berlin, vol 8, 354 pp
Richter C, Wunsch M, Rasheed M, Kötter I, Badran MI (2001) Endo-scopic exploration of Red Sea coral reefs reveals dense popula-tions of cavity-dwelling sponges. Nature 413:726–730
Riedl R (1966) Biologie der Meereshöhlen: Topographie, Faunistikund Ökologie eines unterseeischen Lebensraumes. Parey, Ham-burg, Germany, 636 pp
Riegl B, Piller WE (1997) Distribution and environmental controlof coral assemblages in Northern Safaga Bay (Red Sea, Egypt).Facies 36:141–162
Sheppard CRC, Sheppard ALS (1991) Corals and coral communitiesof Arabia. Fauna Saudi Arabia 12:170
Slack-Smith SM (1998) Order Ostreoida. In: Beesley PL, Ross GJB,Wells A (eds) Mollusca: the southern synthesis. Fauna of Austra-lia. vol 5. Part A. CSIRO, Melbourne, pp 268–282
Stachowitsch M (1980) The epibiotic and endolithic species associatedwith the gastropod shells inhabited by the hermit crabs Paguristesoculatus and Pagurus cuanensis. PSZN Mar Ecol 1:73–101
Stenzel HB (1971) Oysters. In: Moore RC (ed) Treatise on invertebratepaleontology, Part N, Mollusca 6, Bivalvia, vol 3. Kansas UnivPress, Lawrence, KS, USA, pp 953–1124
Surlyk F, Christensen WK (1974) Epifaunal zonation on an Upper Cre-taceous rocky coast. Geology 2:529–534
Taylor PD, Wilson MA (2002) A new terminology for marine organ-isms inhabiting hard substrates. Palaios 17:522–525
Taylor PD, Wilson MA (2003) Palaeoecology and evolution of marinehard substrate communities. Earth Sci Rev 62:1–103
Veron J (2000a) Corals of the world, vol 3. Australian Institute ofMarine Science, Townsville, 489 pp
Veron J (2000b) Corals of the world, Vol. 1. Australian Institute ofMarine Science, Townsville, 463 pp
Voight JR, Walker SE (1995) Geographic variation of shell bionts inthe deep-sea snail Gaza. Deep-Sea Res Part I 42:1261–1271
Zuschin M, Oliver PG (2003a) Bivalves and bivalve habitats in thenorthern Red Sea. The Northern Bay of Safaga (Red Sea, Egypt):
123
Facies (2007) 53:319–327 327
an actuopalaeontological approach. VI. Bivalvia. Naturhistoris-ches Museum, Vienna, 304 pp
Zuschin M, Oliver PG (2003b) Fidelity of molluscan life and deathassemblages on sublittoral hard substrata around granitic islandsof the Seychelles. Lethaia 36:133–149
Zuschin M, Piller WE (1997) Gastropod shells recycled: an examplefrom a rocky tidal Xat in the northern Red Sea. Lethaia 30:127–134
Zuschin M, Stachowitsch M (2007) The distribution of molluscanassemblages and their postmortem fate on coral reefs in the Gulfof Aqaba (northern Red Sea). Mar Biol (in press). Availableonline. http://www.springerlink.com/content/r7t5l732u2585664/.Cited 26 April 2007
Zuschin M, Hohenegger J, Steininger FF (2000) A comparison of liv-ing and dead molluscs on coral reef associated hard substrata inthe northern Red Sea: implications for the fossil record. Palaeoge-ogr Palaeoclimatol Palaeoecol 159:167–190
Zuschin M, Hohenegger J, Steininger FF (2001) Molluscan assem-blages on coral reefs and associated hard substrata in the northernRed Sea. Coral Reefs 20:107–116
Zuschin M, Stachowitsch M, Pervesler P, Kollmann H (1999) Struc-tural features and taphonomic pathways of a high-biomass epi-fauna in the northern Gulf of Trieste, Adriatic Sea. Lethaia32:299–317
123
