20
Facies (2006) 52: 361–380 DOI 10.1007/s10347-006-0048-2 ORIGINAL ARTICLE Margret Steinthorsdottir · Scott Lidgard · Eckart H˚ akansson Fossils, sediments, tectonics Reconstructing palaeoenvironments in a Pliocene–Pleistocene Mediterranean microbasin Received: 27 February 2005 / Accepted: 6 June 2005 / Published online: 29 April 2006 C Springer-Verlag 2006 Abstract Tectonic displacement and small-scale tsunamis apparently affected deposition of the Kolymbia limestone, Cape Vagia, Rhodes, Eastern Mediterranean. Coarse beds interrupt the sequential build-up of this Pliocene–Pleistocene bryomol limestone. Celleporid bryozoans, bivalves, and brachiopods dominate these beds. The palaeoecology of the thicket-forming Celleporaria palmata is re-evaluated and subsequently revised. The limestone comprises two parasequences in a transgressive systems tract, and deposition occurred at palaeodepths between 30 and 120 m. At intervals, tectonic movements lowered relative sea level and sent slumps of shallow-water fauna downslope. The depositional history was validated using independent sets of data: sediment structure and grain size, palaeobathymetry using bryozoan growth forms and occurrences of modern representatives of bryozoans and other taxa, basin configuration, and regional tectonics. Concordance of these lines of evidence provides a means of evaluating confidence in palaeoenvironmental inferences. Keywords Celleporaria . Bryozoans . Palaeoecology . Cool-water carbonates . Eastern Mediterranean . Pliocene–Pleistocene M. Steinthorsdottir · S. Lidgard () Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA e-mail: slidgard@fieldmuseum.org Tel.: +1-312-665-7625 Fax: +1-312-665-7641 E. H˚ akansson Geological Institute, University of Copenhagen, Øster Voldgade 10, 1350 Copenhagen K, Denmark Present address: M. Steinthorsdottir Department of Geology, Trinity College, Dublin 2, Ireland Introduction Formation of carbonate sediments outside the tropical realm has become a widely observed phenomenon (Lees and Buller 1972; Nelson et al. 1988; James 1997; Wright and Burchette 1998; Mutti and Hallock 2003). Bryozoan- rich facies are common in these sediments and beds formed mostly or entirely by bryozoan skeletons are not infrequent. Studies of cool-water carbonates in both modern and past environments are now revealing more subtle differences in tectonic settings, admixtures of skeleton-producing or- ganisms, palaeoecological and taphonomic patterns, and resulting facies architectures. Factors influencing carbon- ate production are more numerous than previously thought and typical cool-water associations (foramol, rhodalgal, and bryomol) do occur in tropical settings (Pomar et al. 2004). Still, the principal models for cool-water carbon- ate deposition and the overwhelming majority of examples come from just two well-studied regional contexts: large open ocean platforms and shelves such as those in South Australia and New Zealand, and ramps such as those in the Western Mediterranean (Hayton et al. 1995; Henrich et al. 1995; James 1997; Pomar 2001; Pedley and Grasso 2002; Lukasik and James 2003), though the latter are less open, with lower wave energy, less tidal flux, shallower storm- wave bases, and clearer water. Detailed geological studies of cool-water carbonate depositional histories in smaller, more closed basins and in palaeooceanographic settings with relatively sluggish circulation and normally lower en- ergy regimes such as those of the Eastern Mediterranean are few in number. Similarly, while recent analyses of tectonic controls (Butler et al. 1997), taphonomic factors (Brachert et al. 1998; Yesares-Garcia and Aguirre 2004), and actual- istic palaeobathymetry (Moissette 2000a, b; Scarponi and Kowalewski 2004) have progressed toward greater objec- tivity in interpreting cool-water carbonate deposition and facies architectures, few case studies have attempted to synthesize these different lines of evidence. The aim of the present study is to infer and validate the depositional history of a highly fossiliferous bryomol

Fossils, sediments, tectonics

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Page 1: Fossils, sediments, tectonics

Facies (2006) 52: 361–380DOI 10.1007/s10347-006-0048-2

ORIGINAL ARTICLE

Margret Steinthorsdottir · Scott Lidgard ·Eckart Hakansson

Fossils, sediments, tectonics

Reconstructing palaeoenvironments in a Pliocene–Pleistocene Mediterraneanmicrobasin

Received: 27 February 2005 / Accepted: 6 June 2005 / Published online: 29 April 2006C© Springer-Verlag 2006

Abstract Tectonic displacement and small-scaletsunamis apparently affected deposition of the Kolymbialimestone, Cape Vagia, Rhodes, Eastern Mediterranean.Coarse beds interrupt the sequential build-up of thisPliocene–Pleistocene bryomol limestone. Celleporidbryozoans, bivalves, and brachiopods dominate these beds.The palaeoecology of the thicket-forming Celleporariapalmata is re-evaluated and subsequently revised. Thelimestone comprises two parasequences in a transgressivesystems tract, and deposition occurred at palaeodepthsbetween 30 and 120 m. At intervals, tectonic movementslowered relative sea level and sent slumps of shallow-waterfauna downslope. The depositional history was validatedusing independent sets of data: sediment structure andgrain size, palaeobathymetry using bryozoan growth formsand occurrences of modern representatives of bryozoansand other taxa, basin configuration, and regional tectonics.Concordance of these lines of evidence provides a means ofevaluating confidence in palaeoenvironmental inferences.

Keywords Celleporaria . Bryozoans . Palaeoecology .Cool-water carbonates . Eastern Mediterranean .Pliocene–Pleistocene

M. Steinthorsdottir · S. Lidgard (�)Field Museum of Natural History,1400 South Lake Shore Drive,Chicago, IL 60605, USAe-mail: [email protected].: +1-312-665-7625Fax: +1-312-665-7641

E. HakanssonGeological Institute, University of Copenhagen,Øster Voldgade 10,1350 Copenhagen K, Denmark

Present address:M. SteinthorsdottirDepartment of Geology,Trinity College,Dublin 2, Ireland

Introduction

Formation of carbonate sediments outside the tropicalrealm has become a widely observed phenomenon (Leesand Buller 1972; Nelson et al. 1988; James 1997; Wrightand Burchette 1998; Mutti and Hallock 2003). Bryozoan-rich facies are common in these sediments and beds formedmostly or entirely by bryozoan skeletons are not infrequent.Studies of cool-water carbonates in both modern and pastenvironments are now revealing more subtle differencesin tectonic settings, admixtures of skeleton-producing or-ganisms, palaeoecological and taphonomic patterns, andresulting facies architectures. Factors influencing carbon-ate production are more numerous than previously thoughtand typical cool-water associations (foramol, rhodalgal,and bryomol) do occur in tropical settings (Pomar et al.2004). Still, the principal models for cool-water carbon-ate deposition and the overwhelming majority of examplescome from just two well-studied regional contexts: largeopen ocean platforms and shelves such as those in SouthAustralia and New Zealand, and ramps such as those in theWestern Mediterranean (Hayton et al. 1995; Henrich et al.1995; James 1997; Pomar 2001; Pedley and Grasso 2002;Lukasik and James 2003), though the latter are less open,with lower wave energy, less tidal flux, shallower storm-wave bases, and clearer water. Detailed geological studiesof cool-water carbonate depositional histories in smaller,more closed basins and in palaeooceanographic settingswith relatively sluggish circulation and normally lower en-ergy regimes such as those of the Eastern Mediterranean arefew in number. Similarly, while recent analyses of tectoniccontrols (Butler et al. 1997), taphonomic factors (Brachertet al. 1998; Yesares-Garcia and Aguirre 2004), and actual-istic palaeobathymetry (Moissette 2000a, b; Scarponi andKowalewski 2004) have progressed toward greater objec-tivity in interpreting cool-water carbonate deposition andfacies architectures, few case studies have attempted tosynthesize these different lines of evidence.

The aim of the present study is to infer and validatethe depositional history of a highly fossiliferous bryomol

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limestone from the Pliocene–Pleistocene in a characteristictectonic microbasin in the Eastern Mediterranean. A sec-ondary aim is to infer the life mode and habitat of bryozoanthickets built by Celleporaria palmata in this limestone.Our approach is synthetic. We combine literature-basedlocal and regional data to establish a tectonic context,then stratigraphical, palaeontological, and sedimentolog-ical data to reconstruct the palaeoenvironment and thedepositional history. We first place the island of Rhodesin a regional geologic framework, and then summarize thestratigraphy and environment of the study area, Cape Va-gia. We describe the stratigraphic column of the Kolymbialimestone, dividing it into facies and beds. The results offield and laboratory work are then listed with interpreta-tions of these data. We infer the mode of deposition of eachbed, based on sedimentological data and palaeoecologicalevidence derived from the fossils. The different fossilgroups and the various sets of palaeoecologic/bathymetricindications are outlined separately in an attempt toconvey their input objectively. The fossil assemblages areanalyzed within this perspective and then evaluated usingbathymetric data from extant members of the assemblages.The data are then brought together to reconstruct the de-positional history of the Kolymbia limestone. The creationof the basin topography is inferred and the depositionalframe, within which evidence from all organisms mustfit, is established. Our interpretations are supported bya suite of independent lines of evidence that are congru-ent with each other, providing a validating test to ourhypothesis.

Geological setting

The Greek island of Rhodes in the Aegean Sea is at theeastern end of the sedimentary part of the Hellenic arc(Fig. 1A). The arc is one of the southernmost nappes of theAlpine system on the southern rim of the Anatolian Plate inan active tectonic regime (McKenzie 1978; Pirazzoli et al.1996; Kontogianni et al. 2002). The tectonic frameworkis dominated by collision of the Arabian and Africanplates with Eurasia (Oral et al. 1995). The African plateis presently subducting below the Aegean plate along theHellenic arc, involving complex vertical movements andcrustal extension (Dewey et al. 1986). The Hellenic Trenchin front of the subduction zone is followed northwards bya sedimentary arc and then a volcanic arc. The AegeanSea and particularly the Hellenic arc are seismicallyactive and among the most tsunamigenic areas in Europe(Perissoratis and Papadopoulos 1999; Dominey-Howes2002). Compared to the Western Mediterranean, wherethere is scant evidence for seismically triggered slope insta-bilities, the Eastern Mediterranean region is characterizedby frequent failures and sediment movements on beddingplanes (Mienert et al. 2002). The Eastern Mediterranean iswarm-temperate, transitional to the true reef environmentsof the tropics, as indicated by the presence of banks ofthe zooxanthellate coral Cladocora caespitosa in someprotected situations in the Aegean (Henrich et al. 1995). Inthe Pliocene, the area may have been even warmer, as thesea-surface temperatures may have been slightly higherthan today (Haywood et al. 2000; Nebout et al. 2004).

Fig. 1 A The position of Rhodes in the Aegean Sea as well as thelocation of the study area, the bay south of Cape Vagia, on the eastcoast, between Rhodes’ two main cities (after Spjeldnaes and Mois-sette 1997). B The sketch map shows the general configuration of

the Cape Vagia microbasin, position of major faults, the distributionof the deposits and the most important topological features, as wellas the position of localities 1–4 and the distribution of the studiedsamples

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Rhodes consists mostly of Mesozoic and Caenozoicrocks, the local basement, which were folded and faultedlate in the Alpine orogeny. A Late Pliocene and Pleis-tocene postorogenic sediment cover of varying thicknessoverlies the basement (Mutti et al. 1970; Meulenkampet al. 1972; Hanken et al. 1996). Tilting occurred in theLate Pliocene, raising northeastern and lowering south-western Rhodes. The area apparently became divided intoblocks with differential vertical movements bounded bynormal faults (Mutti et al. 1970; Pirazzoli et al. 1983, 1989;Hanken et al. 1996; but see Kontogianni et al. 2002). Fur-ther rise and differential tectonic movements continue to thepresent day (Meulenkamp et al. 1972; Benda et al. 1977).This regime exposes Rhodes to rapid, localized, tectoni-cally driven sea-level changes in addition to regional Plio-Pleistocene glacial eustacy (Kovacs and Spjeldnaes 1999;Hansen 2001).

In 1996, Hanken et al. proposed a new formal lithos-tratigraphy using facies mapping. They recognized threemajor transgression–regression cycles, subdividing thestratigraphic succession into three formations: the LindosAcropolis, Rhodes, and Kritika formations (Fig. 2). Therehas been little Holocene sedimentation. A number oflittoral notches, commonly between five and eight, havebeen incised within the 3.75 m zone above the present sealevel, recording post-glacial sea-level fall as well as activecoastal uplift. The maximum Holocene transgression isabout 4 m (Pirazzoli et al. 1989; Hanken et al. 1996;Kontogianni et al. 2002). The Eastern Mediterraneanin general is an enclosed, microtidal and calm sea. Thesubsurface topography of Rhodes is generally very steep,which in other settings has been reported as setting thebase wave and swell-wave level at quite shallow depth,approximately 30–80 m (Hageman et al. 2003).

The study area, the bay south of Cape Vagia, is a mi-crobasin opening to the south (Figs. 1B and 3A). The ex-posed succession of Kolymbia limestone and Lindos Bayclay is draping or onlapping Jurassic meta-limestone base-ment (Hanken et al. 1996) that is hard, dark grey, andseverely eroded. Karstification, marine mechanical erosionexpressed by surf caves and notches, and marine bioerosionare apparent. The basement is locally broken into blocksthat rest at the foot of solid basement ridges. Cape Vagiasensu stricto is approximately 45 m high, bound by a majornormal fault to the ESE (see Fig. 1B). The basement head-land to the west is about 15 m high with large basementblocks at the foot of the headland, forming cryptic envi-ronments (caves) between them. To the north the basementrises gradually to about 30–40 m, obscured by later de-posits, vegetation, and building constructions. These base-ment highs were sites of a “carbonate factory” for the lime-stone deposited in the basin.

The basin was divided into four localities, based on dif-ferences in facies architecture and composition (Fig. 1B).In the western part of the basin, the Kolymbia limestone isexposed in a small section close to sea level, succeeded bythe Lindos Bay clay, which forms a steep cliff (locality 1).The largest portion of the succession is exposed laterallyin the northern part of the bay (locality 2). Towards the tip

Fig. 2 The stratigraphy of the Plio-Pleistocene succession ofRhodes. Three formations are recognized. The deposit studied here isthe Kolymbia limestone, part of the Rhodes formation (from Hankenet al. 1996)

of Cape Vagia, the Kolymbia limestone drapes the steepbasement (locality 4). In the southwestern part of the basin,it onlaps large basement blocks (locality 3). For this study,the majority of samples are derived from locality 2, dueto the superior exposures and stratigraphic range of thatlocality (Fig. 3B).

The Kolymbia limestone is represented by four facies(Kl 1–4) and the Lindos Bay clay by one (Lbc 1). Outof numerous samples collected in the field, 11 sampleswere analyzed in detail. The position of the samples inthe basin is marked in Fig. 1B and their distribution in thestratigraphical column is illustrated in Fig. 4.

Material and methods

A simple geological map was constructed since detailedtopographical maps are unavailable (due to strenuousrelations with nearby Turkey) (Fig. 1B). Three partial

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364

Fig. 3 Two photographs showing: A the configuration of the CapeVagia basin, with basement, deposits, and the main locality indicated,and B a close up of locality 2, showing the appearance of facies in

the field and the relationship between the Kolymbia limestone andthe Lindos Bay clay

stratigraphic section logs were combined into a compositelog, correlating the distinct beds across the basin (Fig. 4).Representative bulk samples of 0.5–1.5 kg were collectedthroughout the study area. The sediment has undergonediagenesis and some beds are reworked, making speciesidentification difficult and time-consuming for somefossils, primarily bryozoans. Effort was therefore focusedon clarifying the composition and origin of the fauna of thecelleporid beds (bed 4 and 10), as well as the coarse shellbeds, the oyster-pectenid bed (bed 6) and the Gryphus bed(bed 8). Two samples were included from the bryozoanlimestone, which separates the coarser beds as well asconstitutes the matrix in those. The most effective methodfor separating the fossils was treatment with a solution ofGlauber’s salt (Na2SO4·10H2O), poured over a sample,which was then frozen and thawed 10 times. Sampleswere then sieved through four different mesh sizes (4,2, 1, and 0.5 mm), dried, weighed, and studied using amicroscope. The 1–2 mm fraction was in most cases splitinto four subsamples using a designated sample-splittingmachine, minimizing the risk of losing information. Thethree largest fractions were sorted into different faunal andnon-faunal groups, which were weighed in order to estab-lish the relative abundance of each group. Bryozoans wereclassified to genus or species level when possible and 23nominal species of cheilostomes and cyclostomes were ex-amined using a scanning electron microscope (SEM). Thelarger fossils, the bryozoan C. palmata, the brachiopodsGryphus vitreus and Novocrania anomala, the bivalvesPecten jacobeus and Ostrea lamellosa were identified andinformation about their modern ecological preferences

gathered. Rhodoliths, echinoderms, and Lindos Bay claybryozoans were examined for ecological purposes only.

Stratigraphy

The Kolymbia limestone is exposed along the east coastof Rhodes, including its type section at Cape Vagia. Thelithology is dominated by highly fossiliferous limestone,mostly rudstone, sometimes grading into floatstone(Cuffey 1985) and reworked limestone facies associations.The limestone is generally fining upwards, with increasingcontent of terrigenous silt and clay. Ichnofossils are abun-dant, and together with body fossils and lithology, theyindicate an increasing water depth towards the top (Hankenet al. 1996). Deposition of the limestone during a majortransgression is also a conclusion of subsequent studies(Moissette and Spjeldnaes 1995; Spjeldnaes and Moissette1997; Hansen 2001). The transgression moved across asteep, irregular coastline. Facies architecture, composition,and palaeontology were strongly influenced by localpalaeo-relief. The sessile benthic communities providedlocal sources of carbonate grains for the deposition ofcarbonate sediments below wave erosion depths. As thetransgression continued, carbonate production rates did notkeep up with the rising sea level and drowning occurred,resulting in gradation of the Kolymbia facies group into theLindos Bay clay. However, this drowning is likely to haveoccurred at slightly different times in individual basins,reflecting the size and distribution of local carbonatefactories.

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365

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Page 6: Fossils, sediments, tectonics

366

Fig. 4 A composite log, constructed using three partial logs, cor-relating the more distinctive beds across the basin. The log showsthe full stratigraphic range of approximately 6 m of the Kolymbialimestone at Cape Vagia

The lower boundary of the Kolymbia limestone is anunconformable contact with the basement. It representsa sequence boundary. Where the boundary can be ob-served, the Kolymbia limestone onlaps and locally drapesthe basement. A sparsely exposed basal conglomerate,the Haraki limestone (Hanken et al. 1996; Bromley andHanken 2003) is thin and yellowish with pebble-size base-ment clasts. Whether or not this conglomerate representsa distinct stratigraphic unit is not considered in this paper,where we treat it as part of facies 1 (Table 1). Approximately6 m of the stratigraphic column of the Kolymbia limestoneis present in the basin (Fig. 4). The main constituent is a ho-mogeneous limestone of yellowish gray color, consistingof fine matrix supporting abundant macrofossils, mainlybranches of bryozoans (facies Kl 1). A muddy version ofthe limestone is observed distally in the basin (facies Kl2). At intervals, coarser, distinct beds, containing high con-centrations of fossils, interrupt the homogeneous limestone.These beds contain large celleporid bryozoans, oysters, bra-chiopods, bivalves, and echinoids (facies Kl 3 and Kl 4).There is a gradual upward transition from the Kolymbialimestone into the terrigenous silts and clays of facies Lbc1, the Lindos Bay clay. This transitional boundary repre-sents the local drowning of the Kolymbia limestone carbon-ate factory prior to the maximum flooding in the lower partof the Lindos Bay clay. There is no visible interfingeringbetween the Kolymbia limestone and the Lindos Bay clayin the Cape Vagia basin. The boundary was reported to bepositioned approximately 70 cm below a characteristic redash bed by Hanken et al. (1996) or where the carbonate con-tent falls below 50% (Bromley and Hanken 2003). In con-trast, we observed a clear and consistent lithological change90 cm beneath the red ash bed at Cape Vagia. Above thisboundary, the sediments are fine-grained, forming a steep,near vertical cliff. Below this boundary the sediments arecoarser and form flat beds with some inclination reachingbelow the present sea level (Fig. 3A and B). We also notethat assessment of percentage carbonate content in the fieldis unworkable. Given the sharp boundary we observed inthe study area, we used this lithologically based boundary.

Magnetostratigraphic studies and the occurrence of theforaminifer Hyalinea balthica at the type locality indicate aLate Pliocene age for the Kolymbia limestone (Løvlie et al.1989). The presence of the Pliocene indicator Echinolam-pas sp. and the bryozoan genus Metrarabdotos, which isreported to disappear from the Mediterranean at the begin-ning of the Pleistocene (Moissette et al. 2002), support thisconclusion.

Facies descriptions and interpretations

In the study area, the Rhodes Formation is represented bytwo facies groups: the Kolymbia limestone facies group andthe Lindos Bay clay facies group. The Kolymbia limestonefacies group comprises a wide spectrum of lithologies onRhodes, from pure siliciclastics to pure carbonates, with avariety of grain sizes. In the Cape Vagia region four facies

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Fig. 5 The retrogradational stacking pattern is characteristic of mostof the Cape Vagia stratigraphy. The individual beds can be inferredfrom the composite log in Fig. 4

are distinguished; Facies Kl 1: bryozoan limestone, FaciesKl 2: muddy limestone, Facies Kl 3: celleporid beds, andFacies Kl 4: shell beds. The Lindos Bay clay facies groupcomprises one facies: facies Lbc 1. All facies are describedand their deposition inferred in Table 1.

Facies architecture and sequence stratigraphy

Non-tropical shallow-water carbonates can form deposi-tional sequences similar to siliciclastic ones and therebydiffer substantially from their tropical counterparts, but di-rect comparison of the controlling factors for non-tropicalcarbonates and siliciclastics should be made with care (Bet-zler et al. 1997; Pedley and Grasso 2002). Draping is oneof two stacking patterns in the Kolymbia limestone. It isdeveloped exclusively on steeper slopes. Drapes are dom-inated by reworked limestone and transgressive limestonefacies associations. Lower parts often have basal breccia-conglomerates. Due to steep slopes, the carbonate factorywas limited in area, producing only minor amounts ofcarbonate. There is often a gradual change from drapingto aggradational or retrogradational architecture. In short,drapes reflect local topography and restricted carbonateproduction (Hansen 2001). At locality 4, the basementsteepens towards the tip of the cape and fossils in the drapedbeds become less abundant in the same direction. Retrogra-dation, the second stacking pattern, is well developed on in-termediate basement slopes. It shows a succession of faciesthat indicates gradual deepening upwards (localities 1, 2,and 3) before changing into the deeper Lindos Bay clay. Thelong-term trend implied is that carbonate production wasnot sufficient to keep up with rising sea level (Hansen 2001).

Figure 5 illustrates the general facies stacking pattern atCape Vagia. It indicates an overall increase in depth, pass-ing into deep-water silts and clays at the top. This patternis punctuated by event beds, which mix shallow and deeperwater faunal components in coarse beds. In terms of se-quence stratigraphy, the Kolymbia facies group is largelyreferred to the transgressive systems tract (van Wagoneret al. 1988) with an ill-defined maximum flooding surfacein the lower part of the Lindos Bay clay. The stratigraphiccolumn of the Kolymbia limestone at Cape Vagia, repre-sented by the composite log (Fig. 4), may be regarded astwo retrogradational parasequence sets within the trans-gressive systems tract, the first one includes beds 1–8 andthe second one includes beds 9–15.

Palaeoecology

The Kolymbia limestone is extremely rich in macrofos-sils. Bryozoans dominate the 0.5–4 mm grain size frac-tions and locally often the >4 mm fraction as well. Bra-chiopods and bivalves are also abundant. Rhodoliths, echi-noids, foraminifers, gastropods, and arthropod fragmentsoccur as subdominant components. This abundance in fos-sils, many of them having modern representatives, providesan opportunity to extract palaeoecological information andto check environmental inferences from each group againstthe others. Bryozoans are by far the most diverse and abun-dant of the fossils in the Kolymbia limestone and are themain focus of the palaeoecological analysis. The celleporidthickets in the Kolymbia limestone represent a very differ-ent scale of occurrence from the rest of the bryozoan fauna.We treat them separately as well as together with otherbryozoan occurrences. Other fossils from the limestone aremainly employed as additional validation tests of the in-ferred setting in which the limestone was deposited.

Bryozoans

The bryozoan fauna is consistent through facies in termsof species present and the approximate abundance of eachof those species. About 23 species are the dominant com-ponents and perhaps five more species occur in smaller butpersistent amounts. Spjeldnaes and Moissette (1997) re-ported 64 bryozoan species from the Kolymbia limestoneat Cape Vagia, including those from a single bulk sample.They reported 12 species found exclusively encrusting thecelleporids; these species were not included in this study.Disregarding these 12 species, about 30 species from theirlist are still unaccounted for. There are also some substan-tial discrepancies between the list determined in this studyand that of Spjeldnaes and Moissette (1997). One of themost common bryozoan species in all the samples studiedhere (Umbonula sp. 1) is not present in their list. Further-more, these authors only report one distinct bed with C.palmata in the Kolymbia limestone, whereas in fact thereare two.

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Page 9: Fossils, sediments, tectonics

369

Recognition to species level was impeded somewhat bythe preservational state of fauna. Most bryozoan coloniesare smothered in fine-grained sediment (mud-fraction) andare slightly re-crystallized. Bryozoans in a sample from afossil cave are however in better preservational state andconfirm the unexpectedly low number of species foundin the samples from the basin. Twenty-two abundant andeasily recognized bryozoan species were studied in de-tail and are listed in Table 2. An additional species, Me-trarabdotos moniliferum, is very abundant in a samplefrom the cryptic cave, but present only sporadically in thebasin.

We apply several different techniques to infer envi-ronmental provenance from the bryozoans, includingdifferent approaches to comparative growth-form analysesand inference of palaeobathymetry using depth rangeoverlaps from extant species. We consider data showingrelative dominance of different colony growth forms,using species counts and relative abundance as separatemetrics. We consider separately species occurrences indifferent limestone strata and facies, then apply depthranges of living conspecifics using an assumption ofuniformitarianism. Modern bathymetric range data arederived from published literature and studies of moderncelleporid thickets occurring elsewhere. The results for theKolymbia limestone are compared to results for bryozoansfrom the above lying Lindos Bay clay. We then summarizethe results derived from this comprehensive study ofthe bryozoan fauna. Bathymetric and other ecologicalattributes of non-bryozoan taxa are used principallyin post-hoc comparisons that either validate or disputepalaeoecological inferences drawn from the bryozoans.

Bryozoan growth forms as ecological indicators

The growth forms of bryozoan colonies correlate to vary-ing extents with the environments in which they live (Stach1936; Smith 1995). A number of different types of growth-form classification schemes have been put forward over theyears (e.g., Stach 1936; Lagaaij and Gautier 1965; Schopf1969; McKinney and Jackson 1991; Hageman et al. 1997,1998; Kaselowsky et al. 2005). One of the most commonlyused is the archetypal classification, based on comparisonto archetype growth forms, where each group is given thename of an archetypal bryozoan taxon. This rationale issimple and often yields interpretable results when appliedto fossil material. The disadvantages include the somewhatarcane terms and lack of consensus on which growth formsto include in wider groups and which to assign to their owngroups. Therefore, some archetypal classifications are min-imalist, with fewer than 10 morphogroups while others aremore complicated with more than 20 morphogroups. Themost severe problem of this classification is that it presumesthat bryozoan colony form is conservative or invariantwithin species, which usually it is not (McKinney and Jack-son 1991). Nor have the palaeodepth predictions of archety-pal classifications been confirmed in a wide range of mod-ern environments. An archetypal classification is used in the

Fig. 6 An archetypal classification of bryozoan growth forms. Sevenmorphological forms are distinguished, each named after archetypeforms (revised after Moissette 2000b)

present paper, due to its simplicity and prevalence, but alsoon grounds of precedence and comparison, as the classifica-tion has been applied to fossil bryozoan assemblages fromthe studied area before (Moissette and Spjeldnaes 1995;Spjeldnaes and Moissette 1997). An archetypal schememodified after Moissette (2000b and references therein) isgiven later showing the most commonly used names for theseven colony forms occurring in this study (Fig. 6).

If a predictive relationship between growth-formdominance in assemblages and (palaeo)environment canbe established, it can be an effective tool in predictingpalaeoenvironments in unknown settings, consideringthe great abundance of bryozoans today and in thefossil record. Relative species richness and abundance ofbryozoan colony forms do change with changes in the en-vironment, notably depth (McKinney and Jackson 1991).Bathymetric distributions of changing growth-form domi-nance patterns are thus used to interpret palaeobathymetry.This method was developed by Stach (1936) and has sincebeen modified and applied by numerous workers (e.g.,Schopf 1969; Hageman et al. 1998; Moissette 2000a,2000b). However, two types of metrics, relative taxonomicrichness and relative abundance, have sometimes beenconfused in the literature. Relative taxonomic richnessusually refers to the proportion of species with differentgrowth forms in a given environmental setting. Relativeabundance usually refers to the proportion of skeletal mass,volume, or living biomass in different growth forms in agiven environmental setting. Environmental settings forboth these metrics have then been abstracted as ecologicalparameters such as depth, current regime, grain size,or bottom type. In this study, we apply both metrics inindependent trials to infer palaeoenvironmental conditions.Moissette (2000b) used relative abundance of each colonyform as a measure of the depth interval at which a givenassemblage lived (rather than using taxonomic richnessand presence/absence). These inferred preferences are usedhere.

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Taxonomic patterns of colony form dominance

The distribution of colony growth forms within the studiedspecies is given in Table 2. Within the model of Moissette(2000b), the encrusting membraniporiform and cellepori-form colonies prefer solid and loose substrate, respectively,whereas both the growth forms are typically more abun-dant in environments with moderate to strong currents andslow sedimentation. The membraniporiforms predominatein shallow water, the celleporiforms live in slightly deeperwater. The adeoni-, vincularii-, and reteporiform coloniesare more abundant on solid substrates, with low to moderatecurrents and slow sedimentation, at moderate depths. Thevinculariiforms are typically more abundant in deep envi-ronments. The cellariiforms colonies prefer solid or loosesubstrate, low to moderate currents, slow to moderate sedi-mentation at shallow to moderate depths. The lunulitiformsare more abundant on loose substrates, endure moderatecurrents and slow to rapid sedimentation mostly at moder-ate depths. When our data are evaluated using this model,the inferred ecological conditions that result include a hardsubstrate, low to strong currents, slow sedimentation rate,and moderate depth.

Abundance patterns of colony form dominance

Abundances were determined for 11 samples in four abun-dance categories (Table 2). These categories, rather thancontinuous metrics, were used based on results from a pre-vious study (Steinthorsdottir 2001). One of the conclusionsof this earlier study is that whereas continuous metrics tendto show high variance and potential sample bias, categori-cal metrics reduce variance and show stronger consistencyamong workers. Abundances were determined from thethree most studied facies, facies Kl 1 (bryozoan limestone),Kl 3 (celleporid beds), and Kl 4 (shell beds). Adeoni- andvinculariiform bryozoans dominate in all three. There isno clear trend in the distribution of species between thefacies or samples, but there is a definite difference in thespecies abundance from sample to sample. When apply-ing the growth-form model of Moissette (2000b) to thebryozoan growth forms using abundance as a parameter,instead of mere presence or absence of species, the envi-ronmental factors for adeoniform and vinculariiform bry-ozoans are accentuated. These are a preference for solidsubstrates, low to moderate currents, and slow sedimenta-tion at moderate depth.

Bathymetry based on extant bryozoan species’ ranges

Known bathymetric ranges of extant species also presentin the fossil assemblage were compared as a means of re-constructing palaeobathymetry (Fig. 7). The bathymetricranges were divided into 10 m intervals to 260 m, belowwhich only five eurybathymetric species prevail. The num-ber of species in each interval was calculated and the largestoverlap of species established. All species are present at the

depth of 60 m and the largest overlap of bathymetric rangesoccurs between 30 and 80 m. Some species have also beenassigned a “preferred” bathymetric range, in which theyare most commonly or abundantly found. Registration ofnumber of species within each bathymetric interval wasrepeated, weighting the preferred bathymetric ranges dou-ble (i.e., counting the species twice within the preferredbathymetric range and once within the normal bathymetricrange). The resultant largest overlap was also 30–80 m.

Bryozoan reefs and thickets

A growing literature exists on reefs and thickets con-structed by bryozoans (e.g., Cuffey et al. 1977; Bradstockand Gordon 1983; Gordon et al. 1994; Henrich et al. 1995;Hageman et al. 1998; Batson 2000; Betzler et al. 2000;Pedley and Grasso 2002; Cranfield et al. 2003; Lukasikand James 2003; Cocito 2004; James et al. 2004). Thesestructures range from micro-scale patch reefs (Scholz andHillmer 1995) to large continuous build-ups, sometimeseven dominating carbonate production completely (Haytonet al. 1995; Cocito and Ferdeghini 2001). The range of bry-ozoan constituents in terms of size, form, and phylogenyis also wide. The settings in which these structures occurare likewise variable and the controlling factors are many.Celleporaria sp. is a common builder of thicket-forminglump- or tree-shaped colonies (Bradstock and Gordon1983; Gordon et al. 1994; Hageman et al. 1998; Batson2000; Pedley and Grasso 2002; Grange et al. 2003). Withmore new studies emerging, the controlling ecologicalfactors seem to become more complicated and thereforesimilar bryozoan structures appear in different geologicalsettings. In a study of the environmental controls on Celle-poraria, Hageman et al. (2003) demonstrate that celleporidthickets can be found at different depths and settings, butall are associated with mesotrophic conditions, moderatesedimentation rates, and relatively low energy (belowswell-wave base). These authors also considered a mud-siltsubstrate to be an important controlling factor, but sincethe colonies grow as epibionts, the encrusted organismsas much as the depositional environment influence Celle-poraria distribution patterns. Celleporid recruitment andgrowth strategy is seemingly an opportunistic utilizationof resources when present (Hageman et al. 2003). In thispaper, we use the celleporids in ecological interpretations,in line with the rest of the fossils.

C. palmata thickets

Large-scale bryozoan thickets in the Kolymbia limestoneare essentially monospecific, constructed by C. palmata.The colonies are celleporiform, constructed by chaotic,frontally budded zooids, forming massive, multi-layeredarborescent structures (Fig. 8). The branches retain moreor less the same thickness throughout their length, anddo not fuse to three-dimensional structures or have heavy,more nodular bases. The celleporid branches are 1–4 cm

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Fig. 7 Bathymetric ranges of extant species of bryozoans in theKolymbia limestone. Darker intervals represent preferred bathymet-ric range. Largest overlap (dotted vertical bar) is between 30 and80 m; all species are present at 60 m depth (data from Cheetham

1967; Hayward and Ryland 1979, 1985; Moissette 1988; Moissetteand Spjeldnaes 1995; Moissette et al. 2002; Pouyet and Moissette1992; Ryland and Hayward 1977; Spjeldnaes and Moissette 1997;Zabala and Maluquer 1988)

thick and each segment is up to 15 cm long. The branchdiameters are bimodal, 1–2 cm and 3–4 cm. Neitherjuvenile colonies, nor bases from which the bryozoanscould have grown, were observed. Relatively few small,nodular colonies were found. A somewhat flattened stemhole occurs in the middle of many of the branches, witha slightly greater abundance in thicker branches thanthinner ones. This cylindrical stem hole is uniformlyabout 2 mm thick and represents an organism, whichthe colonies encrusted and used as support for initialgrowth.

The ecology of C. palmata interpreted from the literatureand our observations can be summarized as follows: thecelleporids initiated their growth upon cylindrical organ-isms, probably gorgonians, which supported and anchoredthem on the hard substrate of the basement. The habit ofgrowing as epibionts on other organisms is a common strat-egy for bryozoans (e.g., Cocito et al. 2000; Hageman et al.2000). Indeed, some species of Celleporaria seem to recruitonly as epibionts (Hageman et al. 2003). Though the indi-vidual branches are now “broken”, this fragmentation maybe due to their original configuration as pseudo-cellariiform(falsely jointed) as well as to compaction and pressure fromoverlying deposits. Some of the branches have been brokenin situ, as indicated by the broken counterpart continuingon the other side of the neighboring branch. Despite care-ful observation and measurements, no preferred orientationfor the celleporid bryozoan branches could be detected.Colonies anchored on firm substrates may have required

Fig. 8 One of the most spectacular colonies of C. palmata in theKolymbia limestone (photo: Margret Steinthorsdottir)

strong disturbance to be detached and transported more orless intact.

Spjeldnaes and Moissette (1997) inferred a depth prefer-ence for the celleporids of 30–50 m. This inference was

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based on modern bathymetric ranges of membranipori-form bryozoans species encrusting them. The epibiont at-tachment of colonies on gorgonians is similar to the ex-tant celleporid species Turbicellepora avicularis, whichbuilds pseudo-cellariiform colonies (Pouyet 1973). Thegorgonian may have been similar to Lophogorgonia cerato-phyta, which has the appropriate height, construction, andsubstrate preference in the Mediterranean. Several trace-making species were present, among those Culicia woodiiand Gastrochaena (Rocellaria) dubia indicate fairly shal-low water (Bromley and Asgaard 1993; Tillbrook 1997).

Bryozoans from the Lindos Bay clay

Samples from the Lindos Bay clay were not analyzed in thepresent study, but data on the bryozoans from that depositare given in Moissette and Spjeldnaes (1995). The authorsinfer depths of about 400 m for the lower parts of the clay,decreasing upwards to about 250 m, using data obtainedfrom bryozoans, ichnofossils, and other fossils. However,when using their data on known bathymetric ranges of ex-tant bryozoan species found in the Lindos Bay clay for con-structing an overlap, in the same way as for the bryozoansfrom the Kolymbia limestone, the largest overlap was foundto be between 100 and 600 m. When constructing an over-lap using only bathymetric ranges of bryozoans found inthe Cape Vagia section, the same overlap of 100–600 mwas found. The difference in largest overlap in bathymet-ric ranges between the Kolymbia limestone and the LindosBay clay supports the deposition model of transgression atCape Vagia.

Palaeoenvironment inferred from bryozoans

Although there is no overriding trend among different fa-cies or samples, there are small variations in the abun-dances of species, some of which can help in the inter-pretation of facies and reconstruction of the depositionalhistory. The consecutive bathymetric ranges of the studiedbryozoan species range over 4300 m. Yet all ranges coin-cide at 60 m and the largest overlap of species is between30 and 80 m. This result is also consistent with inferencesfrom growth forms based on both taxonomic richness andabundance. The bulk of the bryozoan fauna present in theKolymbia limestone at Cape Vagia thus probably livedin a range around 30–80 m. The full range of speciesis conservatively estimated to 10–120 m. The fauna isquite mixed, with shallow- and deep-water forms depositedtogether.

Brachiopods

There are a number of brachiopod species in the Kolymbialimestone. The most distinctive of these are G. vitreus,deposited in a shell-rich bed (facies Kl 4, bed 8) anddistributed unevenly in small amounts in facies Kl 1

sediment, and N. anomala, encrusting on celleporids orbasement. G. vitreus at Cape Vagia are rather thick-shelledand differ in this respect from specimens normallydescribed in the literature. Typically this brachiopod liveson the shelf and in the bathyal zone on hard substrates. Itis associated with N. anomala at depths from 120 to morethan 1000 m in the Mediterranean today, perhaps witha bathymetric preference for approximately 150–300 m(Logan 1979; Logan et al. 2004). Emig (1989) and Emigand Garcıa-Carrascosa (1991) studied the distribution ofG. vitreus along the Mediterranean continental margin anddefined five zones of density versus depth. The G. vitreusbiocoenosis (bathyal detritic sand, dominated by thebrachiopod) is reported to form a belt at depths of approx-imately 100–250 m. The distribution of the biocoenosisis directly related to moderate bottom currents, movingperpendicular to the slope. The density per square meterof G. vitreus in bed 6 is approximately 100 specimens/m2,thus intermediate to zones 2 and 3 of Emig (1989), atapproximately 120–140 m depth. The bathymetric dis-tribution of the thick-shelled specimens in the Kolymbialimestone may be shifted upwards relative to the normalthin-shelled form (Spjeldnaes and Moissette 1997).

Bivalves

A detailed palaeoecological study from the Pliocene ofSouthern Spain showed that in general, bivalve assem-blages from shallow, coarse-grained deposits constructstrongly ribbed, inflated shells, like the shell of P. ja-cobeus (Aguirre et al. 1996). P. jacobeus can sometimespredominate in coarse-grained sediments, such as coarse-to-medium sand and calcirudites. When found in calciru-dites, the other main components are oysters, coralline redalgae, mytilids, arciids, serpulid worms, balanids, and bry-ozoans (Aguirre et al. 1996). P. jacobeus currently livesat depths of 25–183 m (Hallquist and Hansen 1997). Thespecimens in the Kolymbia limestone were probably livingdistally in the basin at depths of ∼ 50–120 m on facies Kl1 sediment bottom.

O. lamellosa typically constructs banks in shallow wa-ter. Tropical littoral oysters grow from the surface down to10–20 m (Laborel and Laborel-Deguen 1996). The recentspecies Ostrea edulis, which is quite similar to O. lamel-losa, lives in environments with normal salinity, in veryshallow water (Laurain 1984). The oysters at Cape Vagiawere living in shallow water at the slopes or on top of thebasement carbonate factory. They encrusted basement andthe shells of one another.

Lithophaga lithophaga bores into hard, vertical to over-hanging surfaces, creating the ichnofossil Gastrochaeno-lites torpedo. The Mediterranean species L. lithophaga isrestricted to extremely shallow marine environments andis considered an excellent indicator of upwards co-seismicmovements (Laborel and Laborel-Deguen 1996). Its opti-mal zone of occurrence is immediately below sea level,giving it a very narrow range in the microtidal Mediter-ranean Sea. It is abundant at depths from 0 to 1 m;

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population density decreases as depth increases to about10 m, below which it is rarely found. L. lithophaga is in-tolerant of sedimentation and the optimum orientation ofthe boring is therefore horizontal, whereas the preferredsubstrate is vertical (Bromley and Asgaard 1993). The ich-nofossil G. torpedo is abundant in the basement limestoneat Cape Vagia, indicating wide fluctuations of sea levelwithin the basin.

Gastrochaena (Rocellaria) dubia is represented bythe characteristic “figure-of-8” shaped ichnofossil Gas-trochaenolites isp. on C. palmata colonies in this studyand in the Pliocene Coralline Crag (Tillbrook 1997). G.(Rocellaria) dubia is also indicative of shallow conditions,ranging down to several tens of meters (Bromley and As-gaard 1993).

Rhodoliths

Rhodoliths are dependent for growth on photosynthetic al-gae and bacteria and thus form within the euphotic zone.Morphologically the rhodoliths in the Kolymbia limestoneare similar to Lithophyllum racemus, which lives on hardsubstrate and sandy bottom in shallow water in the Mediter-ranean today (Riedl 1983). Lithophyllum sp. in a studyfrom the Holocene of the Mediterranean occurred at depthsranging from very shallow to just below 20 m. Corallinealgal build-ups were associated with several large gorgoni-ans (Sartoretto et al. 1996). In the Kolymbia limestone,the rhodoliths are always found abundantly in associationwith the celleporid thickets in facies Kl 3 and in somewhatsmaller amounts in the other facies. They do not encrustthe bryozoans, which may indicate a distinct and probablyshallower provenance (cf. Ferdeghini et al. 2000; Corda andBrandano 2003; Pomoni-Papaioannou et al. 2003). Theyprobably lived on the top and upper slope of the basementcarbonate factory and were transported downslope togetherwith the celleporids.

Echinoderms

Echinoderms are abundant in the Kolymbia limestone, withregular echinoids especially abundant in the transportedfacies Kl 3 and Kl 4. Regular echinoids are particularlycommon today on hard substrate within the euphotic zone,where they graze on algae (Smith 1984). The presence ofregular echinoids is further indicated by the presence ofGnatichnus isp., the stellate grazing trace fossil made byregular echinoids, on transported basement blocks in fa-cies Kl 3 and Kl 4. Echinocyamus pusillus is present inlarge part of the Kolymbia limestone stratigraphic column.E. pusillus lives in the Mediterranean today at depths ofapproximately 20–50 m (Riedl 1983; Philippe 1984). Echi-nolampas sp. is also found in the Kolymbia limestone. Thegenus Echinolampas lives today only in subtropical seas.Generally they live between 10 and 500 m, with preferencefor 30–50 m (Philippe 1984).

Depositional history

Basin creation

Steep normal faults bounding the Cape Vagia basin dipto the east; the cape is the footwall block and the basinbottom is the hanging wall block of the eastern fault(Fig. 1B). The overall geometry is that of a half-graben,creating a microbasin between the raised footwall blocks.Pre-transgressive karstification also sculpted the topogra-phy in the basin, creating a rugged surface. Large basementboulders at locality 3 are interpreted to have been displacedwhen the main normal faults became active. They weresubsequently deposited at the foot of the next raised block.Downslope sliding or small-scale tsunamis triggeredby earthquakes are known transport mechanisms in theMediterranean (e.g., Papazachos et al. 1985). Mastronuzziand Sanso (2000) recorded boulders along the Ionian coast(southern Italy, in a setting somewhat similar to Rhodes),of up to 80 tonnes in weight, which had been transportedat least 40 m. Mastronuzzi and Sanso (2000) note thatcatastrophic events are a neglected but important agent indevelopment of coastal morphology.

Depositional framework

We reconstructed the depositional framework and inferredpalaeobathymetry of the Cape Vagia bay and the Kolymbialimestone by synthesizing topographical, sedimentologi-cal, and palaeoecological data. The highest basement ridgeextends approximately 40 m above the most proximal partof the Kolymbia limestone (Fig. 9). At the top of the base-ment ridge, the greatest water depth is restricted to about80 m. This depth is an approximate limit for oysters and

Fig. 9 The depositional framework of the Kolymbia limestone.Water depth at the most proximal locations was between 10 and80 m and at distal locations between 50 and 130 m

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of peak abundances for the light-dependent algae creatingrhodoliths. The Kolymbia limestone was deposited belowwave base, inferred by us to be approximately 50 m. Thisis therefore the minimum depth at the proximal part of thedeposit. The maximum water depth at the proximal part ofthe deposit is 120 m (80 m + 40 m), constrained by to-pographical evidence. Minimum water depth at the top ofbasement ridge is 10 m (50 m – 40 m). The maximum waterdepth in the most distal part of the basin is approximately130 m. Currents across the sea floor are indicated both bythe open geometry of the basin and banks of G. vitreus. Allfossils found fit conformably into this depositional frame.

Deposition of the Kolymbia limestone

The deposition of the Kolymbia limestone in the CapeVagia basin took place during a large-scale transgression,punctuated by smaller-scale tectonic movements and/or eu-static sea-level fluctuations. The overall trend of the strati-graphic section is retrogradational stacking with deepeningand fining upward of sediments. Coarse event beds inter-rupt this pattern before disappearing towards the top of thesection. The coarse beds contain shallow-water fauna. TheKolymbia limestone then grades into the deep-water claysand silts of the Lindos Bay clay.

The first bed deposited at the onset of transgression wasthe muddy limestone of facies Kl 2. At the same time,oysters and bryozoans grew on the basement slope. Small-scale tectonic disturbance(s) are the inferred mechanismfor downslope mass movement, transporting the oystersand bryozoans into deeper water and depositing bed 2. Wa-ter depth continued to increase and the bryozoan limestoneof facies Kl 1 accumulated as bed 3. At the same timeproximally in the basin, rhodoliths and oysters were liv-ing at or near the top of the cape ridge, while C. palmatainhabited the bottom of the slope. The water depth wasapproximately 50–60 m distally in the basin and 10–20 mproximally. Slumps and debris flows, again inferred to forc-ing by tectonic movement, carried fossils downslope intothe basin, mixing shallow with deeper water fauna and de-positing bed 4 of facies Kl 3. As transgression continued,bryozoan limestone (facies Kl 1) was again accumulatedin the basin (bed 5). Water depth was now approximately30–40 m proximally and 70–80 m distally. P. jacobeusformed large communities proximo-distally, with some G.vitreus distally. Echinoids were a common, but minor com-ponent. Oysters were living most proximally. Storms orsmall-scale tectonic event(s) again resulted in mixing fau-nal components in bed 6, representing facies Kl 4 shellbeds. Pectenids and brachiopods were living on the sand-silt bottom initially whereas the oysters were transported.Depth was now approximately 80 m proximally and 120 mdistally. Bryozoan limestone with some echinoids and otherminor components accumulated again in bed 7. With ris-ing water depth, the ecological conditions for G. vitreusbecame increasingly favorable and the brachiopod becameabundant. A few oysters still inhabited the topographicallyhighest sites in the basin.

Sea level was lowered up to 50 m between beds 8 and 10.It may be that the main mechanism involved was eustaticsea-level fluctuation. Tectonic movements probably alsoplayed a role in downslope movements. Shallower watersediments and fauna were mixed with the G. vitreus com-munity at the basin bottom, forming bed 8. Water depth wasnow similar to that before the deposition of bed 4. Bryozoanlimestone accumulated as the sea level rose, forming bed 9.Oysters and rhodoliths inhabited the most proximal basin,while C. palmata thickets again formed on the slope ofthe cape ridge. A small-scale tectonic movement may haveagain created a debris flow, transporting the faunal compo-nents to deeper water and mixing them with the bryozoanlimestone, depositing bed 10 of facies Kl 3.

After deposition of bed 10, transgression continued.Hereafter only bryozoan limestone (beds 11–15) was de-posited until facies Kl 1 graded into the Lindos Bay clay.No beds of shallow faunal composition were deposited, butcontinuation of small-scale tectonic movements or stormsare indicated by beds with a comparatively larger propor-tion of bryozoan branches (beds 11, 13, and 15). At the timeof deposition of these five last beds, the water depth musthave exceeded depth boundaries indicated by rhodolithsand oysters proximally in the basin, as well as celleporids,pectenids, and G. vitreus more distally in the basin. Beds11–15 have not been studied in detail. However, it is likelythat the bathymetric range of the bryozoan fauna in thesebeds will reflect their deeper water affinity and bridge thegap between the largest bathymetric range overlap of 30–80 m for the underlying part of the Kolymbia limestone andthe largest bathymetric range overlap of 100–600 m for theoverlying Lindos Bay clay.

Discussion

Deposition of the celleporid beds

The depositional model for the Kolymbia limestone in-ferred here is similar to interpretations given by Spjeld-naes and Moissette (1997) and Moissette and Spjeldnaes(1995). However, there are important differences betweenour models. Spjeldnaes and Moissette (1997) interpret thecelleporid beds as autochthonous, living directly on the softbottom. Their interpretation rests on two assertions; (1) theconfiguration of the thicket deposits rules out transport,and (2) the celleporid thickets are analogous ecologicallyto the “reefs” formed by the living azooxanthellate coralLophelia.

First, we do not find compelling evidence for the assertionthat the configuration of branches precludes downslopetransport. Slump deposits in poorly consolidated silty andclayey low-angle inclined seabeds are not uncommon phe-nomena in modern seas. Such deposits are perhaps mostoften related to seismic activity and may leave epibenthicskeletal remains largely intact in a different location.Synsedimentary block faulting, slumping, and mass sedi-ment movements in debris flows are also well documentedin tectonically active Mediterranean basins, particularly in

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the Aegean (Perissoratis and Papadopoulos 1999; Brachertet al. 2001; Mienert et al. 2002; Lykousis et al. 2003).Outsize basement blocks up to 20 cm often accompany therelatively intact celleporid branches (colonies in excess of25 cm, see Fig. 8), along with rhodoliths and oysters fromshallow water. These blocks, rhodoliths, and large skeletalfragments, as well as intermediate skeletal grains, floattogether in a silt–clay matrix. The lack of sorting, hugegrain size variance, and diverse provenance of basementand skeletal elements appear to be strongly indicative oftransport en masse without significant winnowing, ratherthan in situ growth and deposition of celleporids on amore or less homogeneous soft seabed. Fossils can betransported some distance without showing major signsof damage, especially if transported in a mud slump.Large, branching colonies of some fragile bryozoansare seasonally detached and transported relatively intactby wave and current action from nearshore to subtidaloffshore environments (Pederson and Peterson 2002).

Second, while we also recognize some gross structuralsimilarities between Celleporaria thickets and the rigidframework of the living deep- and cold-water coralLophelia (Freiwald 2002), we note that these groupsof organisms are phylogenetically, ecologically, andmechanically quite different. Living Lophelia reefs haverecently been found in the Ionian Sea (Taviani et al. 2005),but they are growing on firmground basement substratesand at greater depths (500–700 m) than envisioned in theearlier model’s analogy. A much closer analogy on allbases of comparison exists with thickets formed by livingand fossil congeneric bryozoan species (Cuffey et al. 1977;Bradstock and Gordon 1983; Gordon et al. 1994; Tillbrook1997; Pedley and Grasso 2002; Hageman et al. 2003;Lukasik and James 2003; Taylor et al. 2004; and others).While species of Celleporaria in these studies have nodularor mound-like colony forms in addition to branching ones,there are some ecological commonalities associated withtheir occurrences in various shallow to outer shelf settings.Most occur below storm-wave base and at the low end of,or below, the euphotic zone. Some actualistic studies haveindicated an increased occurrence of Celleporaria thicketsin an inferred “mesotrophic” gradient of nutrient availabil-ity between oligotrophic and eutrophic conditions, or atolerance for relatively low nutrient conditions. Hagemanet al. (2003) report that some thicket-constructing speciesof Celleporaria in South Australia are tolerant of silty ormuddy conditions, although occurrence as epibionts onfistulose sponges and on hard substrates are also reported.Celleporaria species from Tasman Bay, New Zealand, alsooccur in turbid conditions, though at a shallower depthrange (10–35 m) in lower energy environments (Bradstockand Gordon 1983). Lower wave energy conditions,microtidal fluctuations, and sluggish bottom circulationare conditions that broadly characterize the EasternMediterranean today, and presumably began to developfollowing the post-Messinian reflooding of the entirebasin.

Spjeldnaes and Moissette (1997) suggest a depth of about40 m for the celleporid thickets, indicated by known mod-

ern bathymetric ranges of epizoic bryozoan species. Pres-ence of boulders of Jurassic limestone, as well as oys-ters and rhodoliths from shallow water, were explainedby these authors as washed down into the thickets fromsteep basement slope by heavy storms or tsunamis afterearthquakes. However, the same authors (Moissette andSpjeldnaes 1995) had, in a study on bathymetric indica-tions provided by bryozoans, disregarded all samples fromthe Kolymbia limestone at Cape Vagia because of the mix-ing of deeper and shallow-water fauna. Their interpretationof the autochthonous deposition of the celleporid thicketsalso conflicts with interpreting the stratigraphically adja-cent Gryphus bed, as G. vitreus is a deep-water species.If the celleporid thickets were embedded in situ, a depthdecrease of approximately 100 m would have to have tran-spired in the time period represented by 30 cm of facies Kl 1bryozoan limestone. The limestone shows no signs of suchrapid shallowing. It would be expected that such a signifi-cant transition would provide a more apparent sedimento-logical signature. Rapid shallowing of sea level caused byeustatic sea-level fall and perhaps tectonism did occur be-tween deposition of the Gryphus bed and of the uppermostcelleporid bed. However, by replacing the habitat of thecelleporid thickets to the basement slope in our model, thedifference in depth (about 40 m) becomes more easily ex-plained. We also believe that the intermixed rhodolith, oys-ter and celleporid gravel-sized grades, sand-sized bryozoancolonies and silt/mud matrix are explained more parsimo-niously by a common shallow-water origin, with slump ordebris flow transport en masse. The perhaps slightly lesserdepths at Cape Vagia may be explained by different eco-logical conditions for the thicker shelled specimen of G.vitreus found at that locality, as also noted by Spjeldnaesand Moissette (1997).

Large, robust branching colonies would have neededrather expansive bases to support and stabilize themselveson soft poorly consolidated silt–mud substrate, but suchbases are absent. Where actualistic studies have shownlarge celleporids growing on muddy substrates, the colonyforms have been lattice-like or globular rather than tree-like. For example, Australian Celleporaria form thicketsby originating their growth upon sponges, which stabilizethem on the soft bottom (Hageman et al. 2003). These thick-ets form three-dimensional boxworks, by fusing branchesand thereby constructing a self-supported structure. This isnot the case here. We propose that the distribution of organ-isms upon which the Celleporaria colonies initiate theirgrowth may be as important a factor as the nature of thesedimentary substrate per se. Indeed the colonies require anassociated erect organism to be present before thickets canbe formed, the so-called “ghost frame-builders” of Cocitoet al. (2000). When no such organism is present, Cellepo-raria form lumps on the sea floor, which are more stable ona soft bottom, and do not form erect thickets. Spjeldnaesand Moissette (1997) interpret the cylindrical hole in thecenter of many celleporid branches to be the trace of anorganism upon which the colony formed at the beginningof its growth, possibly a gorgonian. The celleporids areinferred by us to have initiated colony growth epizoically

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on gorgonians or similar organisms. Epizoic growth isapparently a common, underappreciated facet of carbonateproduction and establishment of large individual coloniesby bryozoans (Hageman et al. 2000). In addition, certainCelleporaria species are apparently selective of erect,upright substrates, and hollow cylindrical openings areconspicuous in both fossil and modern samples (Gordonet al. 1994; Hageman et al. 2003). Gorgonians would haveprovided the celleporids with support by anchoring on hardsubstrate, which, in the case of growth on soft bottom, couldbe provided by secondary hard bottoms such as basementblocks or oyster shells. However, such small substrates arefairly patchy and unstable on a soft, current-swept bottom,which does not seem to be in accordance with the large,widely distributed thickets. This problem can be avoidedby accepting the transported nature of the celleporidthickets and inferring their provenance from the basementslope, where hard substrates would have been morecommon.

Tectonic movements as depositional agent

The coarse beds in the Kolymbia limestone, interpretedas tectonic event beds, occur at regular intervals sepa-rated by about 20–30 cm of limestone of normal faciesKl 1 composition. These intervening beds are all of moreor less identical texture and composition. Such regularlyspaced events, causing shallow and deeper water faunato mix, can be attributed to periodically reoccurring tec-tonic movements. Even in recorded history, earthquakesare sometimes reported to occur at regular intervals, thelength of which depends on the seismo-tectonic area con-sidered (Pirazzoli et al. 1994; Perrissoratis and Papadopou-los 1999). Kontogianni et al. (2002) suggest a modifiedearthquake cycle model for Rhodes, reflecting small upliftsalternating with major uplifts and subsidence as a conse-quence of reactivation of different faults of the same faultzone. Earthquake-generated tsunamis are a well-knownphenomenon in the Mediterranean region (Perrissoratisand Papadopoulos 1999; Pirazzoli et al. 1999; Massari andD’Alessandro 2000; Dominey-Howes 2002; Yalciner et al.2002; Lykousis et al. 2003) and are reported to transport anddeposit geological material, sometimes even massive boul-ders (Papazachos et al. 1985; Mastronuzzi and Sanso 2000).However, deposits derived from tsunamis in the Aegeanare often both thin and limited in lateral extension, prob-ably due to coastal morphology (Dominey-Howes 2002;Papadopoulos et al. 2004). In this seismically active areaRhodes is still experiencing uplift. The vertical displace-ment was superimposed on a transgression and is recordedby Holocene notches, excavated by wave action at sea level(Pirazzoli et al. 1989, 1996, 1999). Some of the coarse bedsmay also be interpreted as storm deposits, e.g., bed 6, butthe overall regularity of the thick, coarse beds seems toindicate a tectonic drive behind the events, rather than aclimatological one.

Bryozoans as ecological indicators

The evidence presented here on bryozoans, based both ontheir growth forms and on bathymetric information pro-vided by extant species, validates the depositional frameset independently by topographic restrictions and ecolog-ical information from other organisms. The concordanceof these independent suites of data contributes to a vali-dating test of our depositional model. The consistency oftwo different types of data (colony growth forms, actual-istic depth constraints combining a number of individualspecies) from the bryozoans is also significant. These datasubstantiate the value of bryozoans as palaeobathymetricindicators. The deeper water affinities of the Lindos Bayclay are also consistent with the largest bathymetric over-lap of bryozoans. The bryozoans alone, however, would notprovide as detailed information about each bed, and therebythe depositional history of the Kolymbia limestone, as doesthe combined information from bryozoans and other fos-sils at Cape Vagia. It is also relevant to note that diageneticoverprints, such as the loss of aragonitic forms noted above,can potentially reduce the precision of our palaeobathy-metric estimates. For example, if patterns of bathymetricrange overlaps from aragonitic bryozoans would have pro-vided more tightly constrained palaeodepth estimates, andthese forms were absent due to diagenetic dissolution, theremaining calcitic forms would represent only a subsam-ple of the original fauna. This type of taphonomic over-print highlights the importance of utilizing multiple linesof palaeoecological inference.

Considerable progress has been made by recent stud-ies emphasizing quantitative multivariate methods, includ-ing statistical resampling, to enhance the objectivity, pre-cision, and associated error margins of palaeoecologicaland palaeobathymetric interpretations of cool-water car-bonates (e.g., Jimenez and Braga 1993; Brachert et al.1998; Ferdeghini et al. 2000; Lukasik et al. 2000; Moissette2000b; Hageman et al. 2003; Scarponi and Kowalewski2004; Yesares-Garcia and Aguirre 2004). We concur witharguments supporting the value of these approaches, butalso acknowledge that there is considerable cost in timeand effort that cannot be afforded in many field studies.We suggest that all of these methodological advances stillrely on one form or another of inductive reasoning. The re-sulting interpretations are essentially based on patterns ofassociation or correlation, with some degree of uncertaintybeing unavoidable. That reasoning may be from the generalto the specific, as when depth ranges of living taxa accumu-lated over regional or global scales are applied to specificcase studies, in which case the variance associated withmodern observations is factored out. The reasoning mayalso be from the specific to the general, as when resultsfrom a limited number of actualistic studies are assumedto be uniformly true in all cases, or when the ecologicalparameters of one species are extrapolated to all membersof a genus or higher taxon. An implicit, but necessary,assumption of uniformitarianism may include both typesof induction. This assumption may be especially relevantdue to anthropogenic effects such as trawling and other

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forms of bottom fishing, that have drastically altered pat-terns of occurrence of benthic species and are especiallydestructive to biogenic constructions of long-lived colo-nial animals (Watling and Norse 1998; Collie et al. 2000;Grange et al. 2003). The present study is no exception inall these regards. We hope to have shown that bryozoans dohave value as palaeobathymetric and palaeoenvironmentalindicators. At the same time, we believe that the confidenceassigned to such palaeoecological interpretations or expla-nation is proportional to the degree of concordance of manyindependent lines of inference (Oreskes et al. 1994; Rykiel1998; Salmon 1998).

Conclusions

• The Cape Vagia basin is an extensional microbasin, de-veloped as a half-graben located between two majornormal faults. The upper slope area was the site of acarbonate factory, producing coarse skeletal material fordeposition toward the basin center, where finer materialalso accumulated.

• The Kolymbia limestone constitutes four lithofacies. Thestratigraphical column is fining upward, dominated bybryozoan limestone, interrupted at regular intervals bycoarse beds, containing mixed shallow and deeper wa-ter fauna. The facies architecture of the limestone isretrogradational on intermediate slopes and draping onsteep slopes. The Kolymbia limestone is bounded belowby a type 1 sequence boundary and above by a marineflooding surface, placing it in the transgressive systemstract, where it comprises two parasequences.

• The fauna of the coarse beds includes large brachiopods,bivalves, and the bryozoan C. palmata, forming thick-ets. The bryozoan colonies apparently initiated theirgrowth epizoically, probably on gorgonians, which an-chored them to the basement slope, where they grew atpalaeodepths of ∼ 30–50 m. Detachment and downslopetransport occurred during deposition.

• The growth-form distribution of the 22 classified bry-ozoan species indicates a moderately deep-water habi-tat, in low to moderate currents and slow sedimentationon solid substrates. The subset of these species that areextant have their greatest bathymetric overlap between30 and 80 m.

• A depositional frame was constructed using the topogra-phy of the basin as well as ecological data derived fromthe fossils at Cape Vagia. The water depth at the time ofdeposition of the Kolymbia limestone was 10–80 m atthe most proximal sites and 50–130 m close to the basincenter. All the studied fossils and living representativesfit conformably within the depositional frame.

• The deposition of the Kolymbia limestone took placeduring a major transgression, punctuated by smaller tec-tonic events, triggering downslope mass movements.

• Bryozoans proved to be significant ecological indicatorswhen interpreting the overall depositional history, butdid not yield information as detailed as that provided bysome other fossils.

• Ecological, bathymetric, sedimentologic, and tectoniclines of evidence provide independent validation of adepositional framework and history.

Acknowledgements Richard Bromley, Rikke Bruhn, and Ulla As-gaard (University of Copenhagen, Denmark) provided useful discus-sions, and Patrick Wyse Jackson (Trinity College, Dublin, Ireland),the editor A. Freiwald and anonymous reviewers commented criti-cally on the manuscript. We gratefully acknowledge their help. Wealso thank Hotelejer Anders Mansson og hustru’s mindelegat Foun-dation and the Danish-Icelandic Society for generous support offieldwork.

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