Bollettino della Societd Paleontologica ltaliena, 44 (l), 2005, II-24. Modet

Metrarabdo to s (Bryozou Cheilostomatida)frr Plio-Pleistocene of southernItaly,with descriptionofnew species

AntoniettaRosso

A. Rosso,Dipartimento di ScienzeGeologiche, Sezione di Oceanologiae Paleoecologia,Cat University,Corso ltalia 55, l-95129 Catania( Italy); [email protected]

KEY WORDS - Metrarabdotos (Bryozoa), Svstematics, Mediteruanean area, Plio-Ple

ABSTRACT - The presence of Metrarabdotos irz Plio-Pleistocene deposits of Sicily ant dffirent species, three of which new, have been found: Metrarabdotos elegantissimus sp. / sp. nov. Other palaeo-populations hove been doubtfully referred to M. elegans Buge & G Galopim de Carvalho, already knownfrom the European-Mediterranean area. The recordec .from continental shelf (Circalittoral Zone) to the upper slope (Epibathyal), in bottoms cons are restricted to Pliocene, when the genus Metrarabdotos r,l,asparticularllt speciose in the t to be restricted to the Pleistocene. The presence ofM. moniliferus, the unique species prevt

RIASSLII{TO - lMetrarabdotos (Bryozoa,, Cheilostomatida) del Plio-Pleistocene de nuove specie] - Viene esaminata la presenza del genere Metrarabdotos in depositi plic meridionale. Questo genere ha due poti di distribuzione, localizzati nel l{uovo Mondo ( Mondo (Europa e Africa settentrionale) dove d documentato rispettivamente dall'Eocene at Italia meridionale sono state individuate cinque diverse specie, tre delle quali sono nuov italicus sp. nov. e M. pauciarmatus sp. nov. In particolare la prima i unica in tutto i subcilindrici, e per gli zooidi che individuano un lato frontale e uno dorsale. La secoi caratterizzata da dettagti morfologici a livello zooidale e zoarile e dai suoi rapporti mor. .facilmente individuabile per l'assenza di aviculari su quasi tutti gli zooidi non marginali dt riferiti, sebbene con dubbio a causa dell'esiguo numero di esemplari e/o del cattivo stato ' datt'area europea e africana: M. elegans Buge & Galopim de Carvalho e M teixeirai Bu sono ben separate anche per le loro esigenze ecologiche e si distribuiscono in orizzonti bati limitatomente al piano Circalitorale, e nell'Epibatiale, in.fondali da esclusivamente sabb' -genereparticolarmente abbondante. Dal punto di vista stratigrafico, la maggior parte delle spe Metrarabdotos erafortemente dffirenziato nell'areo. M. pauciarmatus, al contrari il tro record consente di ampliare al Quaternario la distribuzione di Metrarabdotos nell'r moniliferus, l'unica specie finora citata dall'ltalia, resta, invece, da provare.

INTRODUCTION and Pliocene(see Buge & Galopim de Carvalho, 1963) before becoming extinct in the area,seemingly during Metrarabdotos is a long living bryozoan genus (or at the end of?) the Early Pleistocene.This trend whose first representativesdate back to the Late somewhat parallels the evolutionary history in the Eocene.Fossil speciesare known from the European- American distribution pole where the genus radiated North African (Old World) and the American (New mainly from 8 to 4 My &go, splitting into some 12 World) areas, where both Recent and fossil species,most of which were extinguishedby the representativesof the genusare presentin both Eastern Quaternary(Benton & Pearsotr,2001; Cheetham, Pacific and southern North America through the 2001). BesidesM. moniliferus,Milne-Edwards, 1836, Caribbean (Cheetham et al., 1999). Presently,species the type speciesof the genus,numerous species have number is increasingconsiderably as several new taxa been describedand/or recorded,mainly from Europe havebeen recently identified, although yet not described and, subordinately,from a few localities of the from American Neogene sediments and living MediterraneanAfrica (Duvergier, 1924; Pansera,1932; communities (Cheethamet al., 2001, http:llnmita. Roger & Buge, 1946; Lagaarj, 1952; Buge, 1957; geology.uiowa.edu). Reguant, 196l; Buge & Galopim de Carvalho, 1963; In the Old World more than a dozen speciesare Galopim de Carvalho, 197l) partially reviewed by known. The oldest recorded speciesis likely M. Cheetham( 1968). vigneauxi Cheetham(Cheetham, 1968), described from Notwithstanding the about two hundred years of French Lower Oligocene sediments.M. orisensis interest in this genus, well characterisedby the Reguant(Reguant, 1990), from the Middle Eoceneof remarkableovicells among other features,new species, Spain, whose inferred ovicells actually corespond to such as M. pouyeti Marcopoulou-Diacantoni& Wuest, brokenzooids, in fact,has been doubtfully placedwithin have beenrecently described (Marcopoulou-Diacantoni the genus.After a period of likely low diversification, & Wuest, 1999) or given in open nomenclature Metrarabdotoswell differentiatedduring Late Miocene (Moissette,1988).

/.s.s.\'0375-7633 t2 Bollettino della SocietaPaleontologica ltaliana, 44 ( I ), 2005

Among these known Metrarabdotos species,only assemblage;PE : heterogeneouscommunities sensu M. moniliferus has been recorded, sometimes Di Geronimo& Robba( 1989). doubtfully, from Italy (Seguenza,1879-80; De Stefani, Sampleswere routinely treated:sediments were I 882; Namias,I 891; Neviani, 1891,1894, 1900a, b; washedand sievedand specimenssorted from residues De Angelisd'Ossat & Neviani, 1896;Annoscia, 1963). coarserthan 500 pm. Measuresand some photoswere Cheetham ( 1968) didn't evaluate such records performed by meansof a stereomicroscopeequipped suggestingthat a heterogeneousassemblage of species with a JVC TKl38l VideoCamera and a ZeissKSl00 could had been quoted under the name M. moniliferus, 3.00 program.Selected specimens were washedusing the uniquerecorded also in more recentpapers (Poluzzi H,O, and dilutedacetic acid to removepelitic particles & Padovani,1984: Barrier et al., 1987;Di Geronimo et pfioi coating for scanning electron microscope. al., 1987). Measureshave been taken following the schemeby A careful revision of the already availablematerial Cheetham( 1968). and newly collectedspecimens from severallocalities, Describedmaterial and types of the new species all from Plio-Pleistoceneof southernltaly, allowed five are depositedat the PalaeontologicalMuseum of the different speciesto be identified. Three of them are CataniaUniversity (PMC). new, whereas two have been doubtfully attributed to already known species.The aim of this paper is to describeall of them, outlining their geographicaland SYSTEMATICS stratigraphicaldistribution and discussingtheir palaeoecologicalmeaning. The actual presenceof M. Family MErnaRABDorostDAEVigneaux, 1949 moniliferus in the arearemains to be proved. Genus Metrarabdotos Cani, l9l4

Type species:Eschara monilifera Milne-Edwards, MATERIALS AND METHODS 1836

Studiedsamples come from sedimentscropping out Metrarabdotos elegantissimussp. nov. in Sicily and Calabria (southern Italy, central (Figs. 2a-d, Figs. 3a-f) Mediterranean)and depositedduring Pliocene and Pleistocenein shelf-to-slopepalaeoenvironments (Fig. Etltmology Superlative;from the Latin adjective I ). Detailed information can be found in Barrier et al. elegans,alluding to the distinct beauty of the slender ( 1987)for the Paviglianasection, in Di Geronimoet al. branches. (1987, 1992) for the Ringiglio sectionand in Rosso "yellow (2002a, b) for the Capo Milazzo marls". Material - Cala S. Antonino West: sample I ( 1998). Palaeobionomicalreconstructions follow the scheme Holotype:a fertile branch(PMC B. l3 a 21.4.2003). by Peres( 1982).Recognised palaeocommunities are Paratypes:about one hundred fertile and sterile indicatedas follows: DC - coastaldetritic assemblage; fragments(PMC B. l3 b 21.4.2003). DL - detritic offshore assemblage;SGCF : coarse Additional material: Cala S. Antonino Centre I sandsand fine gravels swept by bottom currents; VP ( 1999):32 fragments;2 (1999): 14 fragments;4 (1998): -- : bathyal mud assemblage;CB white corals 60 worn fragments.Cala S. Antonino East: sample 14 (2000): 2 worn fragments.Punta Mazza: sample 7 (2000): 55 not well preservedfragments; sample 8 (2000): a single fragment. All samplesLate Pliocenein age; VP assemblages with a variable amount of contaminantsfrom CB assemblagesand the local addition of shelf species.

Description - Colony erect, dichotomously branching at long intervals,originating from an extremely small encrustingbase sealedby the frontal thickeningof zooids. Branchesroughly cylindrical in cross section from about600 to I 000 pm reaching1400 pm nearthe base. Zooids in four alternatinglongitudinal rows, those of -leizz the centralones opening on a singleside and the lateral ones along the margins, thus forming a frontal and a \r::nrino I dorsal surface.Bifurcating rows absent. Zooidselongated hexagonal, those of the lateralrows enlargedat mid length,their limits forming a slightly 0 50krn Ig zigzaggingline on the dorsalsurface . Zooidal boundaries well marked by raisedrims and a median groove early - - Fig. I Geographical location of the studied samples. Star M. in ontogeny,obscured by increasingcalcification. Open circle : M. italicus; Square _ M. elegantissimus: Frontalwall thin and slightly convex in new zooids pauciarmatus: Closed circle - M. cf. teixeirai: Triangle - M. cf . elegans. progressivelythickened and flat in the older ones. ' tt itlt r/t'rct'tltliott of nt'v \1)1'1'11'1 li

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Scrlintcntsi.lt'c nr.tittlr lrltlrtlrrglr p()ssc\\irtsa snrltlI cnct'ttstirrg[rltsc rttlt e()nstitrrtcrlbr plurtkttlrticlilt'untirtilcrurr ()(),/cs ltlcullr c\cccdrng 2 nrn in rliitnrctct'"rtccrlcrl srtitittrlr lllt'gc hitrtl e()ntprisirrglirssilr lcr'()r.ls [-rcds u ith u['rttnrluntskclctotts strlrstrata.,,\ r'clutir clr snrall nrrnrbct'ol' lritsr.tllrugt't'tct'tts. ol' l'rclrthict)t'gr.rrrisnr\. srrch irs rsidirls.cchinodcl'nts. t'clll'cscrrtrrrglcss tlurn 0.5", ()l'thc ttltul nutnt-rcl'.hitr c A. Rosso- Metrarabdotos.fromPtio-Pteistocene of southern ltalv, u'ith description o.fnev been found. Also ovicells are rare and fertile branches thus simulatingtwo pair of pointed lateral denticles account for less than l%. Finally, regenerationafter subsequentlyplaced inside the peristome.Secondary fragmentation,a reproductivestrategy common within orifice semielliptical,with a marked narrow and deep some speciesbelonging to the samegenus (Cheetham proximal sinus. et al., 2001, Cheeth&ffi,2002), appears irrelevant as Ordinary aviculariapaired, persisting on ovicellate only two fragments show some evidence of zooecia;originating from the distalmostlateral pair of regeneration.All this datacould indicatethat the species areolae,distolaterally placed to the secondaryorifice, was presentin the Capo M rlazzoUpper Pliocenedeep- the avicularianchamber lining the peristome,the rostrum water sediments,although always representedby a few perpendicularto the branch surface,proximally and colonies. The depositionalpalaeoenvironment was inward directed.Rostrum pointed, elongated; cross bar locatedin the epibathyalzone, as pointedout by studied complete.On ovicelled zooids, aviculariaare smaller assemblagesallowing a palaeodepthof 500-600 m to and curved along the edge of the peristoffi€, lateral to be inferred(see Gaetani & Sacce,1984; Gaetani, 1986; the ovicell, at about l13 of its high. Violanti, 1988; Sciuto, 2003). Specialavicularia sensu stricto are absentalthough someavicularia placed along branchmargins and, more rarely, on zooids distal to ovicells could be slightly Metrarabdotos italicus sp. nov. curved and a bit larger than normal ones. (Figs. 4a-l) Fertilezooids longerthan normal ones,sensibly and suddenlyenlarging, their maximum width at level of ",;#;:'J{'; Ine-Edwards - BnnnI rn modified peristomes.Orifice crescent-like,with a |e87 y,"::'?:(:' ) the .'{,r$:' straight distal lip edged by a smooth reversedflange and deeplyhidden by the smooth-to-lightlytuberculate, Etymologv- Relatingto the geographicalarea, which extremely inflated, protruding proximal lip, which the speciescomes from. largelyprojects over the ovicell, when fully developed. Ovicell roughly rounded, arched-to-truncateddistally; Material - Paviglianasection sample 9,DL-VP 150- the ooecial cover moderatelyconvex, sculpturedby 180m, Late Pliocene.Holotype: a fertile branch(PMC numerousradial ribs separatedby porous groovesand B. 14a,26.4.2003).Paratypes: the remainingfertile and margined by a row of areolae,similar to the frontal sterile I l9 branches(PMC B. 14b,26.4.2003). ones. Paviglianasection: sample 4: 60 specimens,DC, 40 Ancestrula sealedby calcification; in a single m; Early Pliocene;sample 8: 40 specimens,DC and instancean ancestrulartriad, each part accounting for SGCF, 100-120 m, Late Pliocene. a sectorof about 120o.is visible in basalview. Cala S. Antonino centre:sample 2 (1999): l0 specimensVP andCB, 500-600m, LatePliocene (MPl5 Measures (pm) Zone). number mean range standard deviation Description - Colony erect, originating from an Zooidallength 33 ll04 990-t220 63.54 Zooidal width 26 393 330-520 42.87 basewhich gives rise to a subcylindrical encrusting Fertilezooidal length 8 1498 1320-1630 101.38 stem, which in turns forms dichotomousbranches. Fertilezooidal width 8 776 700-820 37.39 Branchesroughly cylindrical or nalrow, ribbon-like I - Avicularianlength 12 228 200-320 37.90 2 mm in width, rarely more, compressed,before Avicularianwidth 12 122 I l0-150 12.67 bifurcating. Zooids in 6- l0 alternatinglongitudinal rows, rarely Variabilitv Variability involves branch flattening up to l2-14. Insertionof new rows by bifurcation, and width, zoordal shape and dimensions.Secondary observedonly on a few of the availablesmall fragments calcification increasingin ontogeny seals zooidal and apparentlynot relatedwith other features.The first boundariesand marginal pores and occludesorifices zooid of a new row proximally pointed,except for those transformingbasal branches in strongly calcified stems distalto an ovicell. with a somewhatuniform surfaceonly markedby light, Zooids rectangularelongated; strongly curved when irregular longitudinal furrows. placed laterally to an ovicell. Zooidal boundarieswell marked by raised rims and a median groove early in Remarks Metrarabdotos italicus sp. nov. roughly ontogeny,suddenly obscured by calcification. resembles M. nysti Lagaaij, 1952, for the salient Frontal shield umbonuloid with finely granularand peristomes,the somewhatcostulate ovicell shield and slightly convex surface,limited by about 20 evenly the well protrudedproximal lip of ovicelled zooids. distributedmarginal poresbecoming lessobvious after Nevertheless,this latter species,seemingly with larger the thickening of the frontal surface late in ontogeny. branches,has proximally located,slightly distally Interareolarcostulae not evident.Primary orifice hidden directed peristomial avicularia and a heavily costulate within peristomewith an arched,not well developed, proximal lip in ovicellatezooids. M. italicus sp. nov. pouch-shapeddistal shelf. Peristometubular, about 300 recalls also M. eleganlissimus sp. nov. for zooidal pm in diameter,and not very raised except on zooids characters, from which it is although easily of the lateral rows where sharply protrudes,up to 350 recognisable,at first sight, for the branch shape and pm; proximal interiorwith a longitudinalgroove marked the peculiararrangement of zooidsalong them. 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Ilo :ct'lol': \lrrlpl.' t; A. Rosso- Metrarabdotos.fromPtio-Pleistocene of southernltalv, v'ith description of nev developmentof marginal pores as well as the Description Colony erect formed by ribbon-like morphologyof ovicellscharacterise this speciesamong branches l-4 mm in width, compressed,enlarged other Europeancogeneric ones. Similaritiescould be beforebifurcating. Trifurcation and irregularbranching stressedwith an undescribedspecies from Rhodes giving rise to slightly twisted branchescommon. figured by Cheetham(2002) although specimen Zooids in 10-20,sometimes up to 30, alternating examinationis neededto state conspecificity. longitudinal rows. Common insertion of new rows by bifurcationin zonesof rapid widening; first zooids of Distribution Metrarabdotos italicus sp. nov. has new rows proximally cuneiform. beenfound in two nearbylocalities from southernltaly: Zooids relatively small, rectangularelongated or Pavigliana in southernCalabria and Capo Mrlazzo in club-shaped,sometimes constricted in the middle. the north-easterncorner of Sicily. Stratigraphically,the Zooidal boundariesmarkedly raised to form swollen speciesseems to be widely distributedin the Pliocene, thick ribs, more and more obvious with increasing from the Globorotalia puncticulata to the G. inflata calcification,up to as wide as the proximal half of a Zone in the Paviglianasection (Barrier et al., 1987). zooid, late in ontogeny.Zooids placedalong the margins Within this time interval,probably in the MPl5 Zone, wider than the central ones. Frontal shield convex and depositedalso the Cala S. Antonino layerscontaining coarsely granular tending to develop a few large M. italicus sp. nov. tuberclesaligned on themidline; limited by l8-20 evenly distributedlarge roundedmarginal pores,except at the Paleoecologv - Most specimensof Metrarabdotos proximal end where they become larger and italicus sp. nov. come from sandy (mainly medium- quadrangularor often developas a unique archedcleft, to-coarsesands) sediments containing abundant skeletal becoming more and more obvious late in ontogeny. remainsof benthic organisms(essentially molluscs, Interareolarcostulae usually weak and peripheral, bryozoansand subordinatebrachiopods) pointing to sometimesmarked by granule arched alignments. middle-to-outershelf environments(Barrier et al., Primaryorifice markedby a small proximally projecting 1987).Fossil associations allow to infer that sandswere pouch-shapeddistal shelf. Peristomelow except on colonisedby coastaldetritic assemblages(DC) near zooids of the lateral rows where it extendsslightly on the Infralittoral-Circalittoralbound lty, and detritic the distal margin of the zooid and protrudesgiving a offshore assemblages(DL) at the shelf edge,testifying serratedappearance to the branch margin; denticles,a depthsup to 150-180 m. Palaeoenvironmentswere proximal lateral pointed pair, shallowly set. Secondary characterisedby strong bottom currents(Barrier et al., orifice distallydeveloping in a thin, proximally sinuate, 1987).In a singlecase, sample 2 ( 1999)from Cala S. round tubule. Antonino, a few specimenswere found in deep-water Ordinary avicularia absenton most zooids, single sediments,though probably containing inputs from and small, laterally placed to the peristomeand nearby, shallower edge shelf environments.Like the proximally directed,not easily distinguishablefrom precedingspecies, also M. italicus sp. nov. seemingly marginal areolae.They are present in a very few developedsmall sizedcolonies, arising from encrusting instanceson zooids of the centralpart of the branches, basesnot exceeding2 millimetresin the presentmaterial, more common on lateral zooids and, above all, on the thus exploiting coarsesandy or the rare fine gravelly marginal broader ones, often paired with larger sized elements,as substrates.Sexual propagationwas specialavicularra, characterised by a chamberextending probably the unique reproductivestrategy for this distally beyondthe orifice; rostrumpointed and slightly speciesas only a single doubtfully regeneratedbranch curved around the margin of the orifice, directed has been observed;ovicelled fragments are common proximally. with an incidenceof about 18% whereasbases are rare, Fertile zooids not presenton the studied material. not exceeding2% of total fragments. Ancestrulaeand basesnot observed.

Measures (pm) Metrarabdotos pauciarmatus sp. nov. number mean range standard (Figs.5a-m) deviation Zooidal length 26 843 7 60-940 44.98 1987 Metrarabdotos cf. monili.ferum (Milne-Edwards) - Dl Zooidal width l8 329 290-400 32.08 GEnoNtMoEr nr-.,tab. l. Lateralzooidal length 23 909 8r0-1020 55.10 1992 Metrarabdotos sp. Dr GenoNlMo Er AL., tab. 3 Qtars). Lateralzooidal width l7 484 420-580 46.38 Avicularianlength I I t67 t40-220 26.1| EtvmologV- Relatingto the scantinessof avicularia Avicularianwidth 12 94 70-130 16.76 and to their completeabsence on most zooids. Variabilitv Branch width and number of Material - Ringiglio section:sample 2, PEl, about longitudinal zooidal rows varying to some extent as 30 m. Holotype: a sterilebranch with aviculariaon well aszooidal size and shape.Nevertheless, variability lateralzooids (PMC B. 15a,26.4.2003).Paratypes: the appearsmainly relatedto ontogeneticstage, involving remaining237 sterile fragments(PMC B. l5b, increasing secondary and extrazooidalcalcification. 26.4.2003);Early Pleistocene. Zooidal margins, raised but comparablythin in young Ringiglio section:sample l: 80 specimens,PEl, stages,become thick and swollen; surfacesgranular about20 m; sample3: I specimen,PEI ,20-30 m; Early at the beginning,can develop coarsergranules up to Pleistocene. true tuberclesalong the mid line; zooidal interareolar r8 Bollettino della SocietaPaleontologica ltaliana, 44 ( I ), 2005

costulaesometimes evolve into vertical walls above to those of other Neogene-to-RecentMediterranean frontal surfaces.The abnormal developmentof the adeonelliformspecies, which grow up to 10-20 cm in latter two features,enlarging, strongly rising and high, or more. It is worth noting that neither basesnor coalescingto some extent, form patchesof irregular ovicellswere found in the studiedmaterial. In addition, "spongy" surfaces.Some fragments, seemingly some of the largestfragments show some evidenceof correspondingto stems or near-basebranches, are twisting and irregular branching, often linked to drapedby u thin layer of extrazooidalcalcification, the regeneration,usually by distal budding. In a single surface marked by irregularly anastomosing instancea short brancharises from two adjacentzooids longitudinal ridges and furrows, locally by coarse by frontal budding. Finally, some fragmentsshow tubercles. evidence of seconddry, extrazooidal calcification envelopingpreviously broken branch surfaces.All this Remarks - Metrarabdotos pauciarmatus sp. nov. is points to hypothesiseas fossil populationsof M. easilyrecognisable amongst all the speciesknown from pauciarmatussp. nov. seeminglyexploited an asexually the European-Africanarea for the absenceof avicularia reproductivestrategy, at leastin the inferred,particularly in nearly all zooids exceptthe marginal ones. When stressedpalaeohabitat, where colonisationlikely present,ordinary avicularia are single. The presence happenedthrough the input of broken branchesfrom of a single avicularium is a featureshared only with M. neighbouring hard and/or coarse-grainedbottoms, pouyeti Marcopoulou-Diacantoni& Wuestfrom the Late which survived and continuedto grow by regeneration Plioceneof Greece,but no marginal specialavicularia in the palaeobiotope.Similar adaptationshave been have been reported for this species.Finally, M. sometimesobserved for adeonelliformspecies, such terraconensisReguant, 1960, from the Burdigalian of as Smittina cervicornis (Pallas) and Adeonella calveti Spain, showing somewhat similar zooidal characters (Canu & Bassler). (low peristomesand largeperipheral areolae) completely lacksavicularia. Thus, it is likely thatthese three species could be closely allied. Taking into accountthe trend Metrarabdotos cf. elegans Buge & Galopim de absence/presenceof aviculariathrough single normal Carvalho. 1963 aviculariaversus normal plus specialavicularia, an (Fie. 6t) increasingcomplexity can be envisaged,seemingly pointing to a phyletic relationshipM. terraconensis-M 1963 Metrarabdotos elegans BucE & Gnloplvr DE CenvnlHo, pouyeti-M. pauciarmatus. Nevertheless, further p.162, pl. 1,,figs. 3-4, text-figs. l3-14. - knowledgeis neededabout distribution in time and space 1966 Metrarabdotos elegans Buge & Galopim de Carvalho Buce, p. 42, pl. fig. of Metrarabdotos species in the area, to put forward C, 5. 197 | Metrarabdotos elegans Buge & Galopim de Carvalho - more valuablehypotheses. Gnloprv oE CnnvnLHo,p. l4l, text-fig. 20. 1976 Metrarabdotos elegans Buge & Galopim de Carvalho - Distribution - Metrarabdotos pauciarmatus sp. nov. Pouyer, p. 74, pl. 12, figs. l-2, 7. has been found only in sedimentscropping out along 1987 Metrarabdotos moniliferum (Milne-Edwards) - BnnRtEREr the Ringiglio section,near Mazzarino (central Sicily) nl., tab. I Qtars), non pl. 3 , fi9.4. - which have been referred to the Early Pleistocene(Di 1992 Metrarabdotos elegans Buge & Galopim de Carvalho Er- Hn:lnlr, p. 230, pl.l4, fig. 6. Geronimo et al., 1987),namely to part the uppermost 1997 Metrarabdotos elegans Buge & Galopim de Carvalho - of the Emilian stage. HnvonNE & Molssrrre, fig. 15 (2).

Paleoecology Metrarabdotos pauciarmatus sp. Material - Pavigliana section: sample l6b, VP, 500 nov. comes from fine sandscomprising an about 30% m, Early Pleistocene(Emilian). mud fraction; specimensreducing dramaticallywhen mud rises to about 50%. Bryozoan assemblagesare Description - Colony seeminglyformed by branches, oligospecificand mainly representedby the lunulitiforms at leastup to 3 mm in width. Reussirella reussiana (Manzoni) and Cupuladria gr. Zooids in 18, irregularly alternatinglongitudinal canariensis (Busk) to which Biflustra savartii rows, in the unique observedfragment comprising only (Audouin) and Chaperiopsisannulus (Manzoni) add. a margin. Common insertion of new rows by Macrofaunas are largely constituted(ca. 90%) by the bifurcation, the first zooid proximally cuneiforrn. free living serpulidDitrupa arietina (Mueller) and the Zooids relativelysmall, subrectangularor variously bivalve Corbula gibba (Olivi), two filter-feeding, constricted.Zooidal boundariesmarked by a thin raised opportunisticspecies, whose dominanceis indicative rim. Frontalshield convex and granularlimited by about of the so called heterogeneouscommunities living in 20 marginal pores, separatedby moderateinterareolar biotopeslargely influencedby water turbidity linked to costulae.Peristome thin and raised, more developed high muddyinputs, mainly in basininstability conditions and longitudinallychannelled proximally. Secondary (Di Geronimoet al., 1987,1992;Di Geronimo& Robba, orifice orbicular,with a proximal naffow anddeep sinus. rese). Ordinary aviculariapaired, proximo-laterally placed M. pauciarmatus sp.nov. is exclusivelyrepresented to the orifice, the avicularian chamber lining the by fragments,the largest of which, three times peristome;cross bar complete; rostrum pointed, branched,is about 2 cm high and 4 mm wide. It is upward and inward directed. likely that colonieswere larger and comparablein size Fertile zooids, ancestrulaeand basesnot observed. Itl'),

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Measures (pm) Zooids small,rectangular to club-shaped,sometimes number mean range standard undulate;laterally arched and proximally tapered,when deviation lateral to ovicells; somewhatraised distally. Zooidal Zooidal length 9 906 830- l 060 84.87 boundariesraised and swollen, with a median furrow. Zooidal width 9 374 310-450 s0.03 Marginal zooidswider and slightly longerthan the central Avicularian length 8 195 t70-220 t9.27 Frontalshield with nearly20 small marginalpores Avicularian width 896 80-ll0 l1.88 ones. evenly distributed,separated by interareolarcostulae Variability Variability relates to zooidal and on the periphery; central area convex, granular. avicularianshape and size on the unique available Peristome low. Secondary orifice rounded and fragment. proximally sinuated. Ordinary avicularia, paired or single, constantand Remarks The observed fragment has been persistingon ovicellate zooids; placed laterally and level tentatively assignedto M. elegans as it is the unique to the peristome,rarely slightly distally; proximally species in the area with paired lateral avicularia directed; their chamberstubular, commonly broken. proximally located in respect to the orifice. Rostrum pointed, cross bar complete. Nevertheless,the presentfragment actually differs from Special aviculana larger, located on marginal andl figures and descriptionsin literature for the zooidal or peri-ovicellatezooids, sometimessporadically on shape,not constantly sandglass-shaped,and for zooids of the centralrows, usuallypaired with a normal avicularianshape and position,more elongatedand not one. Their chamberextends distally beyondthe orifice; so proximal as in M. elegans. Available data is the rostrum sharply pointed, directed proximally. inadequateto state if it surely falls or not within M. Fertile zooids commoh, placed in central rows, elegansvariability. often clustered(up to 5-6) on some branches,sensibly larger and longer than normal zooids, suddenly and Distribution - Until now Metrarabdotos eleganswas sharplyenlarging at peristomelevel. Orifice invariably known from the westernmost Mediterranean area broken;distal lip straightwith a narrow reflectedflange; coming back from the Messinian(Morocco: El Hajjaji, proximal lip protruding and projecting,for at least l14, 1992) and the Pliocene (Morocco and Portugal: Buge over the ovicell. Ooecial cover rounded-to-mitre- & Galopim de Carvalho, 1963; Galopim de Carvalho, shaped,sometimes truncated distally. Sculpture given 197l; Spain: Pouyet, 1976; Algeria: Hamdane& by tuberculateribs alternatingwith perforate furrows Moissette, 1997). forming a costulate pattern. An ancestrulartetrad is hardly visible on the unique Palaeoecology The species has been recorded basesuffounded by extrazooidalcalcification occluding from shelf environmentsand the not well-preserved orifices of basalzooids and making the surfacewinding fragmentfound in Pleistocenebathyal sedimentscould sulcate. have been displacedor likely reworked from older (Pliocene) Measures (pm) shallower sediments. ":*; o1x,;:tl Metrarabdotos cf . teixeirai Buge & Galopim de zooidarength ;:r, Carvalho, 1963 Zooidalwidth l8 313 280-360 24.01 (Figs. 6a-e) Fertilezooidal length 12 I 183 960-1300 82.28 Fertilezooidal width 12 7l9 640-850 58.07 Lateralzooidallength 9 816 720-970 80.48 1963 Metrarabdotos teixeirai BucE & GnloprM DECnnvnlHo, p. Lateralzooidal width 9 477 430-500 26.93 157, pl. l., figs. l-2, text-figs. l0- l I ; - Avicularianlength 6 125 l10-140 13.78 1966 Metrarabdotos teixeirai Buge & Galopim de Carvalho Avicularianwidth 6 80 70-90 6.32 BucE, p. 41, pl. C, fig. I ; Specialavicularian length l0 254 200-340 52.54 197 | Metrarabdotos teixeirai Buge & Galopim de Carvalho - Specialavicularian width I 0 I 30 I I 0-140 13.36 Gnlopru oE CanvnlHo, p. 143, pl. 22, figs. 2, 4, text-fig. 2t. 1987 Metrarabdotos moniliferum (Milne-Edwards) - BenRIEREr Variability - Zooidal and avicularian measuresshow nl., tab. I Qtars), non pl. 3 , frg.4. a certaindegree of variability, as well as zooidal shape. Ontogenesis,with increasingfrontal and extrazooidal Material Pavigliana section: sample 4, 78 calcification,accounts for branch thickening. specimens,DC , 40 m; sample5: 28 specimens,DC andSGCF, 50 m, Early Pliocene;sample 8: 9 specimens, Remarks Presentmaterial has a bad preservation DC and SGCF, 100-120 m, Late Pliocene. state owing to heavy recrystallization processes, preventingsome charactersto be observed.Specimens Description Colony erect. Branchesribbon-like have been referred to Metrarabdotos teixeirai, owing slightly enlargedbefore bifurcating,the largestabout 3 to zooidal shape,position of paired normal avicularia mm wide and 8 mm high. and ovicell sculpture,although doubtfully, as sizes of Zooids in I 4-16, often up to 24, regularlyalternating both sterile and fertile zooids are smaller that those longitudinalrows; the dispositionof orificessimulating reportedby Buge (1966) and Galopim de Carvalho a pinnate pattern along branches.First zooids in (1971).The presenceof specialavicularia and marginal bifurcating rows proximally cuneiform. wider zooids observedin the studied specimens,but l ',t. ^et . a rl Tft tr : l. vI e ; l" -b ,'

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nol rce(lrtlctl nr lrtcnrltrrc. \\otrlrl nrlrl.,ctlcrrt'ltlrlc thc l-rrrtnol lcltrt. :r.zc: rtl'lrolli :tct'tlc ltntl lcrtllc /()()trl: lu'e e \lunnlttton ol l) l)c t't'tlttct'ulllirr lr lrcltcr r'oltl'trlcnt'cllt :cfl:l[rlr llrrgcr nt \l ilttlit'rrx.u ltrelt .l]ttn r i.llort ct tlrc ru.ggc:tcrllrllrr[rrrtion. lt r\ \\ ortlr notnrs llrlrtlltlrorrglr nteltlcnec (:cc ltborc) ol'nc\cr elttstct'ctl lcrlrlc ztttttrlt :t)lnc ol llrc :flcr'rc\ ct'cctctl [.] Ilrrgc & ( rltl()l)nnrlc f ftc :1tceic: l: ltl:o :nltllu'lo \l lr'r',unlt't'tIltrgc & ( ( ( lrrrlrlltt) ( l()(r11\\ crc t'ctlcset'ilrcrl. ltl'tct' tltc llllpct' ['rr rlrl()llnr tlc ilr'\lrllro. tlrc lltllct'e ltlrle lct't:ctl [.) lltrgcr (rlrl()l)nn ( lrcctltluntlt)(,X) lrlrtll'rccrr prrt-rlrrlrctl. [.) tle zoorrl. rlrouirtg nri.u'gnutll)()r'c\. loeltllr nr tlott[rlc r'()\\\ ()- ( lrrrlrllro ( I I ). tltrs llrttcr lr.rtlrottlrtln't .tr1t1tlri.ut\ lrntllrr llrrgcr':rzcrlllcr'r:lonrilrlir\ re rrllu'ri.l nr lr tlr:tltlnto:t rnlirl'nLltltlnltl'ltlr.tt \l)ce tltl lt\ reLrlltt'ut luttl otltct'tltltrno:trt' l)()\tltrtn elrlrrle tcr':. \cr crtlrclcsr. :f)cenll ll\ rerrllrrl.r lutr c l-rccn rcecntlr rlcsct'tl-rctllot'thc elorclr rcllrtctl.'f)cerc: \l l )txtt'iltttliott- llr \()utltct'ltlllrlr \lt'lrtu'ttlttlrt/rtrt'I l,',,ttttlt't't lltrgc & ( rll()l)nl tlc ( ltt'rltllto. l'rr I i lllrllrlr /t'l\t'u'tu ltlts l'lccn liltrtttl onlr nt :crlrt't''tcltl:el'()plrlltg r I tt()l ) orrI llong tltc I'ltrtglnutlt :cettttn (:t)Lrtltcrrt('ltlltlrt'nt). \l t'I 1t'1\t'u'rt1rcelll: \l ttttltt'trxlrttt tlrl'l ct'cltec: ll'onrI lrr'l)-to-l.ltc l'lroecne ul irse(lJlrrr'rcr ct lrl lt)S- ) ,lt'eolr,. r()Ll:. rtre lt i.lstlrc tlt:tltlnto:l ll()\lltott ol lrrt'gcr' I't'crt()r.t\ r'ceot'rls t'clL't' to I'ltoecnc ll'ont l'ot'lttgltlltntl l)crr:tonrrlrllr \ rerrllr rnr luttl llrc lrlr:cnec tll tt'rrlr \lort)ee () (llugc. l()(r(r.lJrrgc .t ( ilrl()llnntlc ( ltt'rltllro. tlrilcrcnllrlctlrl)ce nrl tt\ rerrlltt'ul ()fr nlllrgntal tottrti. [.lt:t l ')(,I ( rlloninr rlc ( arr allro. l t)- l t 22 Bollettino della SocietaPaleontologica ltaliana, 44 ( I ), 2005

Paleoecologt - Metrarabdotos cf . teixeirai has been Three of them (Metrarabdotos elegantissimtrssp. found in sandy sedimentscontaining faunasbelonging nov., M. italicus sp. nov. and M. pauciarmalas sp. nov.) to the coarsesands and fine gravelsunder the influence are new, whereasthe remainingtwo have beenreferred, of bottom cuffents(SGCF) and the detritic bottom (DC) althoughdoubtfully, to M. elegansBuge & Galopim de assemblages,thus testifying deposition in soft, mid Carvalhoand M. teixeirai Buge& Galopim de Carvalho. shelf, current swept bottoms. Bryozoansare mainly The latter two specieswere previously known from constitutedby Turbicelleporatubigera (Busk), Srnittina the Iberian Peninsulaand Morocco, with M. elegans cervicornis (Pallas), Melicerita charleswortht Milne- recorded from the Algeria too. All but not M. Edwards, Entalophoroecia deflexa (Couch) and Tbrvia pauciarmatus lived during Pliocene; with M. irregularis (Meneghini) and by minor percentagesof elegantissimu.rseemingly restricted to Late Plioceneand Onychocella marioni Jullien, Tremopora radicifera M. elegans,recorded since the Messinian,in Morocco, (Hincks) and Hippopleuri.fera sedgwicki (Milne- probablythe long living species.The younger species Edwards). seemsto be M. pauciarmatus, which has been found A single basehas beenfound in the presentmaterial only in Emilian (Lower Pleistocene)sediments. To the but, unlike most of the previouslycogeneric described presentknowledge, this taxon could be one of the species,M. cf. teixeirai shows a high incidence of younger or even the youngest one in the Old World ovicellatefragments, with a meanvalue of 320 , ranging area. from l4% (sample5) to 77% (sample8). This,joined A further species,M. moniliferus, the unique one with the absenceof regeneratedfragments, points to a previously recordedfrom ltaly is perhapsquite absent sexual reproductive strategy. from the area.This taxon was quoted,mainly by 1800 researchers,from Pliocene and Quaternary sediments (Seguenza,1879-80; De Stefani,1882; Namias, l89l ; CONCLUSIONS Neviani, I 891, 1894,1900a, b; De Angelisd'Ossat & Neviani, 1896) and most recently from Lower The presentstudy allowed the identificationof five Pleistoceneclays in southernltaly (Calabrian,Venosa: different speciesbelonging to the genusMetrarabdotos Annoscia, 1963) and Crotonian sandsfrom western Canil, 19l4 from southernltaly, namely central-eastern Sicily (Poluzzi & Padovani,1984). Nevertheless, all Sicily and southernCalabria. or, at least, part of the old material (only partly and

o) (f) o) (o (f) (t) O) o) (o (o - (f) r O) o) (o r ;U) o) o (l) r o o (u o (o E o od (o (u- (E (E- z o) @ (f) O (E E @ @ @ o (E- s (o (t) o () o o) o) o) - c (o O E o o (! o) $ ui o E o @ o N o o E (o o L 'o- @ E o) E E .(\' o) (o N o) (\' E 'a o j ; (f) f (I' o o o- o) C 'o-E -c. a 3 o o I o t (u o) o f a o (t' C o .o o 'o (E (I' o E = o oo N trJ o C (E o) o I o o f ci E (U (I' t- P ro E o0 .C, o L o o a o- o o o o, C od od o- a o E, o (l) o) O o q a ci (t (U (l) q od o = a o o o f, (t, o .D c :i o o C E. o q f o) o, L 5 o o (U G' q q dl = (It .DE () (L q .o = (D c o, U' o o o cn CD ci o (t) f = o) .\ Q a q tr x q (! = o (E o o (t, E E dl o (u .n (! f, o q s J b g c s a () E e a c e o f 'o .4 's (! o (l) o o o (D = (u c q o <') o o, q (t q I q o e = f, la (! q e o CJ \ o o (u <}, o 's o j q o e o .S o o, q E o a- s o a- E o E E E E E E = E E E E E E E = E Pleistocene | (e) Gelasian R* BFUK-M* PSM- PMl" c I I

Pliocene Piacerzian R* BFUK-M* PSM- PMl* c (r?) I | (l)

Zanclean R* BFUK-M* PSM- PMI | (l)

Messinian B Ca?* A

Todonian SL LF F B Ca?*

Serravallian 2 2 Miocene Langhian ? 2

Burdigalian AuCzPoRuY S S ?

Aquitanian ? Chattian Oligocene Rupelian F "Old Tab. I - Stratigraphicdistribution of the Word"speciesof Metrarabdotos.A: Algeria;Au - Austria:B : Belgium;C - Crete;Ca - Canarian;Cz - CzechRepublic; F - France;I : Italy; L : Libya, M - Morocco;P - Portugal;Po : Poland;R : Rhodes;Ru : Rumania;S - Spain;UK : United Kingdom,Y - Yugoslavia.Asterisks indicate that in literatureonly the Epochis given without referenceto Ages;(e) - Early;(l) - Late. A. Rosso- Metrarabdotos.fromPtio-Pteistocene of southernltaly, v'ith description of nev hardly available) clearly doesn't belong to M. Grantsto Rosso,Palaeontological Research Group: contribution moniliferus, as often a costulate ovicell was been n.329. described(see Seguenza,1880: p. 208, for example). Moreover,both the most recent( 1990s)records appear REFERENCES doubtful, either for the bad preservationof specimens (abradedand showing broken ovicells as obvious also AnnosciaE. (1963).Antozoi e briozoidelle argille Calabriane di from figures)either for the sedimentarycontext allowing Venosa(Potenza). Geologica Romencl, 2: 215-278. reworking processesto be hypothesised.Otherwise, BarrierP., Casale V., CostaB., Di GeronimoI., Oliveri O. & RossoA. ( 1987).La sezioneplio-pleistocenica di Pavigliana most recordsof M. moniliferus needto be re-evaluated (Reggio Calabria).Bollettino della SocietoPaleontologica (seeCheeth am,, 1968) and the distribution of this species Italiana.25 (2): 107-144. seemspresently restricted to Pliocene sedimentsof BentonM.J. & PearsonP.N. (2001). Speciation in the fossil someEuropean localities (U.K., theNetherlands, France, record. Trendsin Ecolog, & Evolution, 16 (7): 405-411. Iberian Peninsula:Lagaaij,, 1952; Buge, 1957; Buge & Buge E. (1957).Les Bryozoairesdu Neogenede I'ouestde la Galopim de Carvalho, 1963; Galopim de Carvalho, Franceet leursignification stratigraphique et paldobiologique. national d'Histoire naturelle, n. s., 197l) and maybe North Africa (Morocco, Tunisia: Memoiresdu Museum serie C, Sciencesde Ia Terre,6 ( l): 435 pp. l97 l). Pleistocenespecimens Galopim de Carvalho, Buge E. ( I 966).Sur quelquesbryozoaires du Ndogenedu Maroc. from the U.K. and the Netherlands,on the contrary, Annales de Paleontologie,(lnvertebres), 52 ( I ): 19-48. have been consideredprobably reworked (Cheetham, BugeE. & Galopimde Carvalho A.M. ( 1963).R6vision du genre l e68). MetrarabdotosCanu 19l4 (Bryozoa-Cheilostomata).Rev. Species of Metrarabdotos, which are easily Fac. Ci2nciasLisboa, 2 ser.,C, I I (2): 137-196. distinguishablefor their morphologicalcharacters, CheethamA.H. ( 1968).Morphology and systematicsof the bryozoan genus Metrarabdotos.Smithsonian Misc. Coll., appearalso well separatedecologically. Apart from M. 133( I ): tzt pp. elegans, for which a few data is avatlable, the three CheethamA.H. (2001).Evolutionary stasis vs. change. In Briggs Pliocene speciesplaced themselvesin different D.E.G.& CrowtherP.R. (eds), Palaeobiology II, The bathymetrichorizons also exploiting slightly to sharply evolutionaryProcess and the fossilrecord, Species evolution: diverse substrata.M. teixeirai seemingly lived on soft 137-t42. sands,in mid-to-outershelf environments.This CheethamA.H. (2002).Asexual propagation in the cheilostomes Metrarabdotos and Coscinopleura: effects on genetic distribution was partly overlappedby M. italicus, yet variationand larval productivity.In Wyse JacksonP.N., extending towards major depths, to the shelf break. Buttler C.J. & SpencerJones M.E. (eds),Bryozoan Studies M. elegantissimusthrived in epibathyal bottoms with 2001. Proceedingsof the Twelfth InternationalBryozoology palaeodepthsof about 500-600 meters.It could be AssociationConference: 73-80. worth noting as thesethree species line up subsequently CheethamA.H., JacksonJ.B. & SannerJ. (2001).Evolutionary in a two axesdiagram considering zooidal width versrts significanceof sexualand asexualmodes of propagationin Metrarabdotosin central zooidal length, both for sterile and fertile zooids. The Neogenespecies of the bryozoan America.Journal o.fPaleontologv, 75 (3): 564-577. with small-srzed towards large- line up, from species CheethamA.H., JacksonJ.8., SannerJ. & VentocillaY. ( 1999). sizedzooids, fits well with the inferreddistribution from NeogeneCheilostome Bryozoa of TropicalAmerica: shallow shelf waters to deep slope waters. Within comparisonand contrastbetween the Central American speciesand within genotypezooidal size variability, both isthmus(Panama, Costa Rica) and the North-Central in time and space,has been observedin several Caribbean(Dominican Republic). In Collins L.S. & Coates cheilostomebryozoansand relatedto water temperature A.G. (eds.),A Paleobioticsurvey of Caribbeanfaunas from the Neogeneof the Isthmusof Panama.Bulletins of American (see & Okamura,2000 for a review). If we O'Dea Paleontologv,357: 159-192. assumethat similar changesextend also to interspecific, De Angelisd'Ossat G. & NevianiA., ( 1896).Corallari e Briozoi within genus variability, the progressivelyincreasing Neogenici di Sardegna.Bollettino della SocietaGeologica zooidal size registeredin the above-recordedspecies, Italiana, l5 : 571-598. could be linked to physiological changesrelated to De StefaniC. ( 1884).Escursione scientifica nella Calabria (1877 - - decreasingwater temperature,at great depths, in an 78) Jejo, Montalto e Capo Vaticano.Memorie della Regia Accademiadei Lincei (classedi ScienzeFisiche, Matematiche areaseemingly subjectto upwellings. e l,,laturali),serie 3, l8: l-292. Finally, M. pauciarmatus,seemingly the shallowest Di GeronimoI. & RobbaE. (1989).The structureof benthic species,was capableof exploitinga particularlystressed communitiesin relation to basin stability. Memorie palaeoenvironment characterised by abnormal dell'Accademiadei Lincei. 80: 341-352. inputs. Broken fragments seemingly Di GeronimoI., Raffi S. & RossoA. (1987).Dominanza di sedimentary "Popolamenti survived and continuedto grow adopting twisting and specieopportuniste nei Eterogenei"del Mazzarino(Sicilia centrale). Bollettino combining sexual and asexualreproduction, by Pleistoceneinferiore dr dell'Accademia Gioenia di ScienzeNaturali, Catania, 20 strategy. fragmentation,&S a winning adaptation (331):t29-166. Di GeronimoI., RossoA. & SanfilippoR. ( 1992).Bryozoans as ACKNOWLEDGMENTS sedimentaryinstability indicators.Rivista Italiana di Paleontologia e Stratigrafia.,98 (2): 229-242. Thanksare extendedto Alan Cheetham( Smithsonian DuvergierJ. (1924). Deuxidmenote sur les Bryozoairesdu Institution,Washington) for discussionsi-proving the Neogenede I'Aquitaine.Actes de la SocieteLinneenne, manuscriptand to PierreMoissette (Universite Claude Bernarde, Bordeaux.75: 5-50. Lyon). Dott. AnnamariaMannino (Palermo University) and Mr El Hajjaji K. (1992). Les bryozoairesdu Miocdne superieurdu Canzanella(Naples University) are kindly acknowledgedfor Maroc nord-oriental.Documents des Laboratoires de SEM photos.Paper financially supported by CataniaUniversity GeologieLvon, 123: 355 pp. 24 Bollettino della SocietdPaleontologica ltaliana, 44 (l ), 2005

FoisE. ( 1989).La successioneneogenica di CapoMrlazzo (Sicilia investigating palaeoseasonality. Palaeogeographv, NE). Rivista ltaliana di Paleontologiae Stratigrafia, 95 G): Palaeoc limatology, Palaeoecology, 162: 3 19-332. 397-440. PanseraO. ( 1932). Fossili miocenici di Casr Garabulli. Bollettino Fois E. ( 1990).Stratigraphy and palaeogeographyof the Capo della Societa Geologica ltaliano, 5l: 287-296. Milazzo area(NE Sicily, Italy): cluesto the evolutionof the PdrdsJ.M. (1982). Major benthic assemblages.1r Kinne O. (ed.),, southernmargin of the Tyrrhenianbasin during the Neogene. A comprehensive integrated treatise on life in oceans and Palaeogeographv,Palaeoclimato logy, Palaeoecology,7 8: 87- coastal waters. Marine Ecologv, 5. Ocean Management, part. I 08. l: 373-522, Wiley and Sons. GaetaniM. ( 1986).Brachiopod palaecommunities from the Plio/ Poluzzi A. & Padovani M.A. (1984). Osservazioni Pleistoceneof Calabriaand Sicilia (Italy). /r RacheboefP.R. paleoecologiche sui Briozoi delle sabbie crotoniane di S. & Emig C.C. (eds),Biostratigraphie du Pal6ozoique4: 281- Margherita Belice (Sicilia). Giornale di Geologia, serie 3,46 288. (2):gs-t t2. GaetaniM. & SaccirR. ( 1984).Brachiopodi batiali nel Pliocene PouyetS. (1976).Bryozoaires cheilostomes du Pliocene e Pleistocenedi Sicilia e Calabria.Rivista Italiana di d'Aguilas (Espagnemdridionale). Nouveau Archives du Paleontologiae Stratigrafia, 90 (3): 407-458. Museumd'Histoire l,,laturellede L1ton,14: 53-82. Galopim de CarvalhoA.M. (1971). Briozo6riosdo Tercirlrio ReguantS. ( 196l). Los Briozoosdel Ndogenoespafrol. Memorias Portugu0s(Cheilostomata do Neogdnicoda Orla Ocidental). del Instituto Gdobgico v Minero de Espafia, 62: 215-244. 176 pp. Centrode Estudiosde Geologiade Facultadede ReguantS. (1990). Metrarabdotosorisense n. sp. (Bryozoa Cienciasde Lisboa.Livraria Cruz,Braga. CheilostomataAscophora) del Eoceno Medio de Vic HamdaneA. & MoissetteP. (1997). Faunesde bryozoairesdu (Barcelona):una forma ramosasingular. Revista Espafiola de Pliocdnedu Saheloccidental algdrois (Algdrie). It{eues Paleontologia,5: 7 l-7 6. Jahrbuch .fiir Geologie und PalciontologieAbhandlungen, Rio D., SprovieriR. & Di StefanoE.(1994). The Gelasian Stage: 205: 69-96. a proposalof a new chronostratigraphicunit of the Pliocene Lagaaij R. ( 1952).The PlioceneBryozoa of the Low Countries series. Rivistaltaliana di Paleontologia e Stratigrafia, 100 and their bearing on the marine stratigraphyof the North ( I ): 103-t24. Sea. Mededelingenvan de GeologischeStichting, C, 5 (5): RogerJ. & BugeE. ( I 946).Les bryozoaires du Redonien.Bulletin 233 pp. de la SocieteGdologique de France, sdrie 5, 16: 217-230. Marcopoulou-DiacantoniA. & WuestJ. (1999).Les bryozoaires RossoA. (2002a).Brvobaculum carinatum sp. n., gen. r., & new du Pliocdnesupdrieur de Cr6te (Formationde Tsoutsouros, MediterraneanPliocene deep-sea bryozoan.lr WyseJackson Provincede Monofatsiou,SE d'Heraklion).Revue de P.N.,Buttler C.J. & SpencerJones M.E. (eds),Bryozoan Paldobiologie, I 8 (2) : 547-57 6. Studies2001 . Proceedingsof the Twelfth International Milne-EdwardsH. ( 1836).Observations sur les Polypiers fossiles BryozoologyAssociation Conference: 175-182, Dublin. du genreEschara. Annales de SciencesNaturelles, Zoologie, RossoA. (2002b). krataulopocella borealis gen et sp. nov., a Paris, 6: 321-345. deep-waterPliocene Lekythoporid (Bryozoa) from the MoissetteP. ( 1988).Faunes de Bryozoairesdu Messinien Mediterraneanarea. Memorie di ScienzeGeologiche, 54: 65- d'Algdrie occidentale.Documents des Laboratoires de 72. GeologieLyon, 102: 351 pp. SciutoF. (2003).Dati preliminarisulla ostracofaunapliocenica NamiasI. ( l89l ). Contributoai BriozoiPliocenici delle provincie di Capo Milazzo (Sicilia NE). Bollettino della Societd di Modena e Piacenza.Bollettino della Societd Geologica Paleontologicaltaliana, 42 (l): 179-184. Italiana- 9: 47 I -5 I 3. SeguenzaG. ( 1879-80).Le formazioniterziarie della Provincia NevianiA. ( I 891).Contribuzione alla conoscenzadei Briozoi di Reggio(Calabria). Regia Accademia dei Lincei, Memorie fossili italiani. Bollettino della Societd Geologica ltalianq, della Classedi ScienzeFisiche, Matematichee lt{aturali,serie l0: 99-148. 3, 6: 416 pp. Neviani A. ( 1894).Di alcuni Briozoi pliocenicidel Rio Landa Violanti D. (1988).I Foraminiferiplio-pleistocenici di Capo illustratida FerdinandoBassi nel 1757.Bollettino della Societd Mllazzo. Bollettino del Museo Regionaledi Scienzelt{aturali Geologicaltaliana, 12: 659-668. di Torino,6 (2): 359-392. Neviani A. ( 1900a).Briozoi neogenicidelle Calabrie. Palaeontographiqltalica, 6: ll5-266. NevianiA. (1900b).Briozoi terziane post-terziandella Toscana. Bollettino della SocietdGeologica ltaliana, 19 (2): 349-375. O'Dea & OkamuraB. (2000).Intracolony variation in zooid Manuscriptreceived 05 April 2004 size in cheilostomebryozoans as a new techniquefor Revisedmanuscript accepted l3 October 2004