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Cretaceous Research 32 (2011) 472e479

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Cretaceous Research

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Taphonomy and palaeoecology of the gastropod fauna from a Late Cretaceousrocky shore, Sweden

Anne Mehlin Sørensen, Finn Surlyk*

Department of Geography and Geology, University of Copenhagen, Øster Voldgade 10, Dk-1350 Copenhagen K, Denmark

a r t i c l e i n f o

Article history:Received 22 January 2010Accepted in revised form 16 March 2011Available online 24 March 2011

Keywords:GastropodsTaphonomyPalaeoecologyRocky shoreLate CretaceousSweden

* Corresponding author.E-mail addresses: anne@geo.ku.dk (A.M. Sørensen

0195-6671/$ e see front matter � 2011 Elsevier Ltd.doi:10.1016/j.cretres.2011.03.003

a b s t r a c t

A gastropod fauna comprising 17 species, each represented by a limited number of specimens, isdescribed from a Late Cretaceous, late early Campanian rocky shore at Ivö Klack, southern Sweden. Thegastropod fauna is associated with the most diverse ancient rocky shore fauna ever found. However, thelow gastropod species diversity compared to the faunas of modern rocky shores is ascribed to tapho-nomic factors, notably dissolution of the aragonitic shells, but the predominance of epifaunal herbivoresis indicative of a guild structure similar to that found on modern rocky shores. The presence of drill holesassigned to the ichnospecies Oichnus simplex suggests the former presence of muricid gastropods whichhave not been found as body fossils. A single drill hole is referred to Oichnus paraboloides and wasprobably made by a naticid gastropod. The infaunal mode of life of naticids makes preservation of suchdrill holes difficult, since the majority of infaunal prey such as burrowing bivalves has aragonitic shellswhich are not preserved. The relatively high number of species in comparison to many other LateCretaceous rocky shore faunas, offers an opportunity to compare gastropod guild structure at Ivö Klackwith modern counterparts, even though taphonomic processes such as mechanical destruction andaragonite dissolution have played an important role in the fossil gastropod assemblage.

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1. Introduction

Modern rocky shores are inhabited by dense aggregations ofhighly diverse biota which encrust and cling to the substrate in theturbulent environment. Gastropods normally make up a significantpart of the fauna and herbivorous species are predominant in theperiodically exposed intertidal zone along the shore (Brehaut,1982;Levinton, 2001). Marine snails constitute the largest and mostdiverse molluscan class and have many different ways of feeding,including herbivory, carnivory, omnivory, scavenging, browsing,deposit and suspension feeding. This allows them to inhabit a widerange of subenvironments with different food sources (Clarkson,1993; Ruppert et al., 2004).

The knowledge of ancient rocky shore faunas is relativelylimited since such settings represent sites of erosion rather thandeposition, and the majority of faunal elements present will fallvictim to taphonomic processes. Many species with hard parts areunlikely to be preserved due to ongoing wave surge and resultingmechanical destruction. Organisms which encrust or bore may bepreserved in life position, whereas organisms which cling to the

), finns@geo.ku.dk (F. Surlyk).

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rocks or are attached with soft pedicles or a byssus such asgastropods, brachiopods, some bivalves, and echinoids experiencepost-mortem transportation, wear and fragmentation. Anothermajor taphonomic process is carbonate dissolution especially ofaragonite shells which in both modern and ancient settings takesplace in marine waters during very early shallow burial or even onthe sea floor. In modern environments, this dissolution results fromacidity caused by microbially mediated reactions associated withthe decay of organic material (Cherns and Wright, 2000; Wrightet al., 2003). In many limestones moulds are the only evidence ofthe former presence of aragonitic shells and the scarcity of certainaragonite-shelled molluscan groups may therefore be a largelytaphonomic and not an ecological effect (Cherns andWright, 2009).

A well-exposed Late Cretaceous rocky shore is preserved on theisland of Ivö in southern Sweden (Fig. 1) (Surlyk and Sørensen,2010). It has yielded a rich and diverse invertebrate fauna whichcombines elements from the intertidal and subtidal rocky zonesand from the associated shallow water carbonate sediments. Snailsconstitute an important faunal group since they include some ofthe few herbivores preserved, and are the sole group of predatorsthat left traces (drill holes) to document their feeding on inverte-brates. The producer of the holes can be deduced from drill holemorphology (Bromley, 1981). Holes made by naticid gastropodshave a larger external then internal diameter with parabolic walls

Fig. 1. Simplified geological map of north-eastern Skåne, southern Sweden, showing the Kristianstad Basin and the location of Ivö Klack. Modified from Norling and Bergström(1987) and Lindgren and Siverson (2002).

A.M. Sørensen, F. Surlyk / Cretaceous Research 32 (2011) 472e479 473

compared to drill holesmade bymuricids (Fig. 2). Naticids normallyprey on infaunal molluscs and feed infaunally although somespecies do hunt on the sea floor. Muricids on the other hand prey onepifaunal taxa such as limpets and oysters and their boreholes arecylindrical and perpendicular to the shell surface (Fig. 2A)(Bromley, 1981). Most gastropod shells are composed of aragonitewhich is not preserved at Ivö Klack, explaining the low number ofspecimens.

The aim of the present study is to present an overview of thisrare gastropod material, to interpret modes of life by comparisonwith selected extant taxa, and to assign the gastropod fauna toguilds in order to interpret their ecological role. It has not beenattempted to undertake a taxonomic revision due to the characterof the material but only a classification sufficient for ecologicalinterpretation of the gastropod fauna.

2. Geological setting

A late early Campanian rocky shore is exposed in a disusedkaolin quarry, Ivö Klack, on the northern part of the island Ivö,located in the lake Ivösjön in the northern part of the KristianstadBasin in north-eastern Skåne, southern Sweden (Fig. 1). During theLate Cretaceous, the global sea level was up to about 100 m higherthan today (Kominz et al., 2008) and the basin was transgressedrepeatedly, resulting in the formation of an archipelago with lowislands and peninsulas (Surlyk and Christensen, 1974; Christensen,1975; Surlyk, 1997; Surlyk and Sørensen, 2010). A rocky shore wasdeveloped during transgression as the deeply weathered basementwas swept clean by wave action during transgression. The topo-graphy of the steep shore comprised hummocks and boulders of

gneiss onlapped by oyster banks associated with a highly diversefauna. A biological zonation of the cementing epifauna on the largeboulders was recognised by mapping and reflects different pref-erences for shelter, exposure and light (Surlyk and Christensen,1974). The carbonates deposited on and around this rocky coastbelong to the uppermost lower Campanian Belemnellocamaxmammillatus belemnite biozone, c. 81e80 ma (Christensen, 1975).

3. Material

The present study is based primarily on material collected by A.Rosenkrantz in 1928, C.C. Hansen in 1947 during quarrying, and byW.K. Christensen in 1966e1967 just after quarrying stopped. Allspecimens are housed in the collections of the Natural HistoryMuseum of Denmark, Copenhagen; some specimens bear the prefixGMI (Geological Museum, Ivö). The gastropod fauna comprisesa total of 17 species, 14 of which are assigned to four genera, andthree remain unidentified. Each species is represented by only oneor a few specimens, illustrating the paucity of thematerial (Table 1).Most shells were originally composed of aragonite with theexception of four species, Patella sp., Otostoma sp. A, B, and C. Theoriginal pigmentation of the shells in Otostoma sp. B and C isretained. The aragonitic shells were dissolved during burial,explaining the preservation as moulds. Only one external mould ofan aragonitic species is found in a cemented part of the limestone.The absent outer morphology of the shells makes taxonomicassignment difficult or impossible since internal moulds of thick-shelled species commonly differ considerably in appearance fromthe original shell. The mode of life and habitats of the fauna areinterpreted on the basis of a comparison with modern relatives.

Fig. 2. Gastropod boreholes in the brachiopod ‘Rhynchonella’ triangularis (Wahlenberg, 1821) from Ivö Klack, A. ichnospecies Oichnus simplex Bromley, 1981, probably produced bya muricid gastropod; B. ichnospecies Oichnus paraboloides Bromley, 1981, probably produced by a naticid gastropod.

A.M. Sørensen, F. Surlyk / Cretaceous Research 32 (2011) 472e479474

4. The gastropod fauna

Dimensions given are either shell height (apex to base) orlargest diameter. Species assignments are based on Knight et al.(1960), Abdel-Gawad (1986, 1990), Jung (1987), Popenoe et al.(1987), Griffin and Hünicken (1994), Elder (1996), Perrilliatet al. (2000), Squires and Saul (2001), Kiel (2001, 2006), Kieland Bandel (2001, 2003, 2004), Pacaud (2004), Del Río andMartínez (2006), and Saul and Squires (2008).

Campanilid sp. APlate 1, Fig. 8

Two specimens with large-sized, turreted shell, up to 42 mm indiameter; both specimens incomplete; whorls rounded probablywith sutural shelf, and almost three times wider than high;periphery rounded; aperture oval; reconstructed apical angle 35�;internal mould preservation, so no ornament visible.

Possible juvenile of Campanilid sp. APlate 1, Fig. 6

One specimen with medium-sized, turreted shell, up to 24 mmin diameter; whorls widely separated, indicating a thick shell, threetimes wider than high, andwith rounded to slightly angular profile;periphery rounded; umbilicus present; preserved as internalmould, so no ornament visible.

Table 1Guild structure of the gastropod fauna from Ivö Klack. A ¼ aragonite, C ¼ calcite.

Subclasses Species

Caenogastropoda Campanile sp.Caenogastropoda Campanilid sp. ACaenogastropoda Campanilid sp. BNeritimorpha Otostoma sp. ANeritimorpha Otostoma sp. BNeritimorpha Otostoma sp. CPatellogastropoda Patella sp.Archaeogastropoda Trochid sp. aff. ChilodontaArchaeogastropoda Trochid sp.

Caenogastropoda Volutodermidae sp. ACaenogastropoda Volutodermidae sp. B

Caenogastropoda Gyrodes sp.Caenogastropoda Neomesogastropoda aff. Naticidae sp.Caenogastropoda Naticidae sp.

Campanilid sp. BPlate 1, Fig. 4

Three specimens with large-sized, turreted shell, up to 50mm indiameter; all specimens incomplete; whorls flat to slightly convexwith deep incised abapically pointing reconstructed suture, whorlsare relatively widely separated, indicating a rather thick shell, andalmost twice as wide as high; periphery rounded; reconstructedapical angle 23�; internal mould preservation, so no ornamentvisible; aperture not visible.

Campanile sp.Plate 1, Figs. 5 and 7

Two specimens with large-sized, turreted shell, up to 45 mm indiameter; both specimens incomplete; whorls slightly convexadapically with deeply incised, abapically pointing reconstructedsuture, indicating a rather thick-shelled species; whorls almostthree times wider than high; periphery rounded; reconstructedapical angle 40� and 46�; aperture shaped almost like a parallelo-gram; internal mould preservation, so no ornament visible.

Gyrodes sp.Plate 1, Fig. 14

One specimen with medium-sized shell, 25 mm in height anddiameter; globular shape, flat spire; periphery and sides rounded;

Abundance Composition Guild

2 specimens A Epifaunal herbivore3 specimens A3 specimens A1 specimen C-A1 specimen C-A1 specimen C-A2 specimens C-A1 specimen A1 specimen A

2 specimens A Epifaunal carnivore1 specimen A

1 specimen A Infaunal carnivore8 specimens A1 specimen A

Plate 1. All specimens in natural size except Figs. 11, 12, 13, and 16. Fig. 1 aeb: Neomesogastropoda aff. Naticidae sp. (GMI 3715). Mould, a) apical view, b) side view. Fig. 2 aeb:Naticid sp. (GMI 3716). Mould, a) apical view, b) side view. Fig. 3 aec: Trochid sp. aff Chilodonta sp (GMI 3719). Mould, a) apical view, b) side view showing the imprints after a jaggedouter lip c) side view showing the maximumwidth of the shell. Fig. 4: Campanilid sp. B (GMI 3702). Side view of mould. Fig. 5: Campanile sp. (GMI 3704). Side view of mould. Fig. 6:Juvenile Campanilid sp. A (GMI 3705). Side view of mould. Fig. 7: Campanile sp. (GMI 3703). Side view of mould. Fig. 8: Campanilid sp. A (GMI 3701). Side view of mould. Fig. 9 aeb:Juvenile Neomesogastropoda aff. Naticidae sp. (GMI 3708). Mould, a) apical view, b) side view. Fig. 10: Unidentified (GMI 3720). Side view of mould. Fig. 11: Unidentified (GMI 3718).Top view of mould. Fig. 12 aeb: Otostoma sp. A (GMI 3706). a) Apical view, b) apertural view. Fig. 13 aeb: Otostoma sp. B (GMI 3707). a) Apical view, b) apertural view. Fig. 14 aeb:Gyrodes sp. (GMI 3710). Mould, a) apical view, b) side view. Fig. 15: Volutodermidae sp. B (GMI 3712). Side view of mould. Fig. 16: Trochid sp. (GMI 3713). Apical view. SEMphotography. Fig. 17 aeb: Volutodermidae sp. A (GMI 3711). Mould, a) side view, b) apical view. Fig. Fig. 18 aeb: Otostoma sp. C (GMI 3717). a) Side view to show the width of theshell, b) side view to show the shoulder of the shell. Fig. 19: Patella sp. (MO 132640). Apical view. Fig. 20: Unidentified (GMI 3721). Side view of mould.

A.M. Sørensen, F. Surlyk / Cretaceous Research 32 (2011) 472e479476

ornament consisting of collabral ribs; incomplete and preserved asinternal mould.

Naticidae sp.Plate 1, Fig. 2

One specimen with medium-sized shell, 34 mm in height and44 mm in diameter; globular shape, flat spire and two whorls;periphery and sides rounded; umbilicus present; aperture halfcircle formed; preserved as internal mould, so no ornament visible.

Neomesogastropoda aff. Naticidae sp.Plate 1, Fig. 1

Five specimens with medium-sized shell, up to 37 mm in heightand diameter; globular shape, flat spire and two and a half whorls;periphery rounded, sides slightly angular with deeply incised abap-ically pointing suture; aperture large and angular; all specimensincomplete, preserved as internal mould, so no ornament visible.

Possible juvenile of Neomesogastropoda aff. Naticidae sp.Plate 1, Fig. 9

Three specimens with medium-sized shell, up to 11 mm inheight and 20 mm in diameter; circular outline, flat spire and morethan two whorls; sides rounded to angular; all specimens incom-plete, preserved as internal mould, so no ornament visible.

Otostoma sp. APlate 1, Fig. 12a, b

One specimen with small, hemispherical shell, 9 mm in heightand 8mm in diameter, comprising twowhorls; spire depressed, butrapidly expanding into globose body whorl; aperture large, oval inoutline; periphery rounded, but with irregular outline on accountof ornament, consisting of protruding radial ribs in upper half ofwhorl and of rounded tubercles on lower half.

Otostoma sp. BPlate 1, Fig. 13a, b

One specimen with small, hemispherical shell, 8 mm in heightand 6mm in diameter, comprising twowhorls; spire depressed buthigher than in Otostoma sp. A, expanding rapidly into globose bodywhorl; aperture large, oval in outline; periphery and sides rounded;ornamentation consisting of fine radial ribs; original pigmentationpreserved as dark colouration of the adapical part of whorls.

Otostoma sp. CPlate 1, Fig. 18

Whorl fragment of one specimen, sharp shoulder; collabral ribsand original pigmentation preserved as black band under shoulder;15 mm high and 18 mm wide.

Patella sp.Plate 1, Fig. 19

Two well preserved specimens with small shell, calcitic, up to5 mm in height and 15mm in length; oval outline, apex subcentral;outer surface with strong radial ribs with rounded spines andconcentric growth lines.

Trochid aff. ChilodontaPlate 1, Fig. 3

One specimen with medium-sized shell, 18 mm in height and17 mm in diameter, trochiform, with two whorls; periphery andsides rounded, whorls almost three times wider than high; bodywhorl with imprints of teeth on outer lip; incomplete andpreserved as internal mould, no ornament visible.

Trochid sp.Plate 1, Fig. 16

One specimen with small shell, 3 mm in height and diameter;turbiniform with three whorls; periphery and sides rounded;ornament of collabral ribs; rubber cast of an externalmould, lackingapertural features.

Volutodermidae sp. APlate 1, Fig. 17

Two specimens with medium-sized, fusiform shell, up to 15mmin diameter, more than two whorls; periphery subrounded toangular, probably on account of nodes or spine ornament onshoulder; sides near straight; aperture slender, elongate andsharply narrowed posteriorly; both specimens incomplete andpreserved as internal mould, so no details of ornament visible.

Volutodermidae sp. BPlate 1, Fig. 15

One specimen with medium-sized, fusiform shell, 18 mm indiameter; periphery rounded, sides near straight; larger than vol-utid sp. A, and lacking ornament around shoulder; incomplete andpreserved as internal mould.

GMI 3718Plate 1, Fig. 11

One specimen with small shell, 6 mm in diameter; just a single,rounded whorl preserved. The shell was probably elongate andturreted; incomplete and preserved as internal mould, so noornament visible.

GMI 3720Plate 1, Fig. 10

One specimen with medium-sized shell, 10 mm in diameter,fusiform; periphery and sides rounded; bodywhorl with row of pitsafter teeth halfway up after growth stop; imprints of teeth on outerlip; incomplete and preserved as internal mould, so no ornamentvisible.

GMI 3721Plate 1, Fig. 20

One specimen with medium-sized shell, 15 mm in height; tur-biniform; periphery and sides rounded; body whorl with imprintsof radial ribs c. 0.1 mmwide; incomplete and preserved as internalmould.

5. Results

The identified gastropod species belongs to four subclasses, theCaenogastropoda, the Neritimorpha, the Archaeogastropoda, andthe Patellogastropoda. They are subdivided into three guilds on thebasis of interpreted feeding strategy and position relative to thesubstrate. All identified gastropod taxa are assigned to a guild,following comparison with modern relatives (Table 1) (The

Fig. 3. Number of gastropod species represented on different modern and LateCretaceous rocky shore settings.

A.M. Sørensen, F. Surlyk / Cretaceous Research 32 (2011) 472e479 477

Paleobiology Database, 2000; Thack, 2005, 2007). The epifaunalherbivorous guild comprises the highest number of species and isrepresented by nine species. The infaunal carnivore guild includesthree species, and the epifaunal carnivores are represented by twospecies (Table 1).

Some carnivorous gastropods drill holes in shells of otherinvertebrates, including conspecific and other gastropods. A total of31 boreholes were found in brachiopod and bivalve shells; nearlyall are assigned to the ichnospecies Oichnus simplex Bromley, 1981

Fig. 4. Small tube dwelling polychaetes encrusting the muscle

which most likely was produced by muricid gastropods (Fig. 2A).However, no body fossils of this group have been found. A singleborehole found in a valve of the brachiopod ‘Rhynchonella’ trian-gularis (Wahlenberg, 1821) is referred to Oichnus paraboloidesBromley, 1981. Judging from its morphology, it probably wasproduced by a naticid gastropod (Fig. 2B). Omnivores, scavengers,suspension and deposit feeding gastropods have not been recog-nised in the present material.

6. Discussion

The gastropod fauna comprising a total of 17 species is ofa relatively low diversity compared to modern rocky shore faunasin Italy, Oregon, California and southern Australia, from where 94,54, 41 and 65 species have been recorded, respectively (Fig. 3;Garrity, 1984; Goddard, 1997; Kelaher, 2003; Terlizzi et al., 2003).Thus, modern rocky shore gastropod faunas show very high speciesdiversities. This may also have been the case during the LateCretaceous where the most diverse rocky shore gastropod faunaknown includes 46 species from the Cenomanian of Germany (Kieland Bandel, 2004). A high diversification among gastropods hasalso been noted throughout the Mesozoic, suggesting that gastro-pods already had reached a high degree of specialisation during theLate Cretaceous and were able to inhabit both intertidal and sub-tidal settings (Knight et al., 1960; Sepkoski, 1990).

The abundance of moulds after former aragonite-shelledmolluscan species at Ivö Klack indicates a slow dissolution ofaragonite within the carbonate sediment (cf. Cherns and Wright,2000). Epifaunal encrustation on oyster muscle-scars, after disso-lution of the aragonitic layer indicates that dissolution alsooccurred on the sea floor or possibly under very shallow burial andsubsequent reworking of the sediment during storms, exposing theoyster shells at the sea floor (Fig. 4). Since the majority of shellygastropods are epifaunal the low number of specimens of eachgastropod species probably reflects severe taphonomic loss due toboth mechanical destruction of shells and aragonite dissolution.Gastropods become dislodged from the substrate right after deathand the shells were damaged and abraded during wave action andstorms, resulting in more rapid and easy dissolution. The slightlyhigher number of specimens of infaunal species may illustrate

scar of an oyster after dissolution of the aragonitic layer.

Fig. 5. Pie diagram of the identified gastropod species from Ivö Klack showing thedistribution of species among subclasses.

A.M. Sørensen, F. Surlyk / Cretaceous Research 32 (2011) 472e479478

better preservation when aragonitic species are already buried inthe sediment after death. However, the diversity is high comparedto many other Cretaceous rocky shores, e.g. from Saxony in easternGermany (four species; Pietzsch, 1962); New Zealand (threespecies; Crampton, 1988), and Mexico (one species; Lescinsky et al.,1991), but low compared to Mülheim in western Germany (46species; Kiel and Bandel, 2004); all of these faunas have also beensubject to taphonomic loss (Fig. 3). The higher species diversity atIvö Klack and Mülheim compared to the other Cretaceous rockyshores may be due to long-term quarrying of the carbonates atthese localities giving access to new exposures of the carbonateswhich yielded the (semi) vagile, non-cemented fauna. The muchhigher diversity at Mülheim may be due to better preservation ofaragonitic species in the hard limestone.

The guild structure at Ivö Klack appears to be relatively similar tothat found in modern settings, despite the drawback caused bytaphonomy and the paucity of material (Table 1). The majority ofspecies is assigned to theepifaunalherbivorousguildwhich is also thedominant category on modern rocky shores. Only two limpet shellshave been collected at Ivö Klack. Their shells are composed of bothcalcite and aragonite, making them suitable for fossilisation. Theywere probably more abundant on the shore, being well adapted toclinging to rocks even during strongwave or current action similar totheir modern relatives. Their preferred habitat was high in the surfzone and this probably led to early destruction of their shellsfollowing death. The other two guilds comprise infaunal andepifaunal carnivores, represented by three and two species, respec-tively. Carnivorous gastropods are also documented through theirdrill holes in invertebrate shells. Most drill holes recognised can beascribed toOichnus simplexBromley,1981,probablymadebymuricidswhich today are abundant in shallow waters where they prey onepifaunal forms. No body fossils representing this group have beenfound at Ivö Klack but they must almost certainly have been present.One drill hole probably produced by an infaunal naticid gastropodwas found in a brachiopod shell, indicating that thesemainly infaunalspecies may also have hunted epifaunal prey. They normally feedinfaunally butmost shells of infaunalmolluscanpreyare composedofaragonite and are preserved as internalmoulds, thus obliterating anydrill holes originally present in the associated shell. Diverse infaunalbivalve and echinoid faunas are recorded, indicating that the sedi-ment was well oxygenated. Consequently, it may be expected thatgastropods were also able to live within the sediment. The twoNaticidae species and the naticid species, Gyrodes found at Ivö Klack,are thus interpreted to have lived infaunally similar to most modernnaticids. Omnivores or scavengers have not been recognised in thisstudy. Thegastropodguildstructure is thus similar tomodernsettingsalthough species diversity is interpreted to have been considerablyhigher in the original fauna.

The studied fauna belongs to four subclasses (Fig. 5), theArchaeogastropoda which are usually rocky shore inhabitants

(Bandel and Wedler, 1987; Hickman and McLean, 1990 in Kiel andBandel, 2004), the Neritimorpha which includes many commonrocky intertidal species (Ruppert et al., 2004), and the Patellogas-tropoda which are also common in the rocky intertidal areas(Ruppert et al., 2004). The majority of the gastropod species belongto the Caenogastropoda, a subclass including gastropods able to livein environments from rocky shores into silty, especially soft-bottom, habitats (Ruppert et al., 2004). Species from this subclassfound at Ivö Klack belongs to families which are found in otherintertidal to shallow subtidal Late Cretaceous environments (Kiel,2001; Kiel and Bandel, 2004). The invertebrate fauna found at IvöKlack from themore distal parts of the rocky shore succession awayfrom the gneiss surface comprise fossils at all stages of fragmen-tation and disintegration from complete and well preserved shellsto sand- and pebble-sized angular fragments. The fossil fragmentsare mainly angular, indicating limited or no transport (Surlyk andSørensen, 2010). The Caenogastropoda found at Ivö Klack is thusbelieved to have lived in the intertidal to shallow subtidal envi-ronment on the associated carbonate sediments and boulders alongthe rocky shore.

7. Summary and conclusions

A gastropod fauna from the late early Campanian rocky shore atIvö Klack, southern Sweden comprises 17 species. The diversity islow compared to modern rocky shore faunas, undoubtedly due totaphonomic factors, such as fragmentation by waves and dissolu-tion of aragonitic shells. The diversity is, however, significantlyhigher than most other Cretaceous rocky shore faunas, probablyreflecting that quarrying of the carbonates has given continuousaccess to new carbonates, yielding the (semi)mobile, non-cemen-ted fauna at Ivö Klack.

Three guilds are identified; nine species are assigned to theepifaunal herbivorous guild, three to the infaunal carnivores, andtwo to the epifaunal carnivores. This distributionmatches well withthe guild structure of modern rocky shore gastropods. Omnivores,scavengers, suspension and deposit-feeders are not recognised inthis study.

Two types of gastropod drill holes are recognised: Oichnussimplex probably produced by muricids, and O. paraboloides,probably produced by naticids, based on the morphology of thedrill holes. Muricids are not represented by body fossils but weremost likely present since the drill holes are common. Only a singleexample of O. paraboloides and one naticid species is recognised.The rarity of O. paraboloidesmay be due to the infaunal mode of lifeof naticids and themouldic preservation of their infaunal aragoniticmolluscan prey.

Acknowledgements

The present study was supported by a grant from the DanishNatural Science Research Council (FNU). We thank John Jagt forcommenting on an early version of the manuscript, Steffen Kiel forhis very helpful advice on gastropod taxonomy and for his and ananonymous reviewer’s constructive reviews which improved themanuscript, Thomas Hansen for discussion of the taxonomy,and Rikke Harlou for preparing scanning electron micrographs ofTrochid sp.

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