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

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,as particularllt 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'

particolarmente abbondante. Dal punto di vista stratigrafico, la maggior parte delle spe-genere 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.

Metrarab do to s (Bry ozou Cheilostomatida) frrof southern Italy,with description ofnew

Antonietta Rosso

A. Rosso, Dipart imento di Scienze Geologiche, Sezione di Oceanologia e Paleoecologia, Cat

Catan ia ( I ta ly ) ; rosso@unic t . i t

Plio-Pleistocenespecies

Univers i ty , Corso l ta l ia 55 , l -95129

and Pliocene (see Buge & Galopim de Carvalho, 1963)before becoming extinct in the area, seemingly during(or at the end of?) the Early Pleistocene. This trendsomewhat paral lels the evolut ionary history in theAmerican distribution pole where the genus radiatedmainly from 8 to 4 My &go, spl i t t ing into some 12spec ies , most o f wh ich were ex t ingu ished by the

Quaternary (Benton & Pearso t r , 2001; Cheetham,2001). Besides M. monil iferus, Milne-Edwards, 1836,the type species of the genus, numerous species havebeen described and/or recorded, mainly from Europeand, subord ina te ly , f rom a few loca l i t ies o f theMediterranean Africa (Duvergier, 1924; Pansera, 1932;Roger & Buge , 1946; Lagaarj , 1952; Buge , 1957;Reguant, 196l; Buge & Galopim de Carvalho, 1963;Galopim de Carvalho , 197 l) part ial ly reviewed byCheetham ( 1968).

Notwithstanding the about two hundred years ofin te res t in th is genus , we l l charac ter ised by theremarkable ovicells among other features, new species,such as M. pouyeti Marcopoulou-Diacantoni & Wuest,have been recently described (Marcopoulou-Diacantoni& Wuest , 1999) o r g iven in open nomenc la tu re(Moisset te, 1988).

INTRODUCTION

Metrarabdotos is a long l iv ing bryozoan genuswhose f i rst representat ives date back to the LateEocene. Fossil species are known from the European-North African (Old World) and the American (NewWor ld ) a reas , where bo th Recent and foss i lrepresentatives of the genus are present in both EasternPaci f ic and southern North Amer ica through theCaribbean (Cheetham et al., 1999). Presently, speciesnumber is increasing considerably as several new taxahave been recently identified, although yet not describedf rom Amer i can Neogene sed imen ts and l i v i ngcommuni t ies (Cheetham et a l . , 2001, ht tp: l lnmi ta.geology.uiowa.edu).

In the Old World more than a dozen species areknown. The o ldes t recorded spec ies is l i ke ly M.vigneauxi Cheetham (Cheetham, 1968), described fromFrench Lower O l igocene sed iments . M. o r isens isReguant (Reguant, 1990), from the Middle Eocene ofSpain, whose inferred ovicells actually corespond tobroken zooids, in fact, has been doubtfully placed withinthe genus. After a period of likely low diversification,Metrarabdotos well differentiated during Late Miocene

/.s.s.\' 0375-7633

t2 Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

Among these known Metrarabdotos species, onlyM. mon i l i fe rus has been recorded, somet imesdoubtfully, from Italy (Seguenza, 1879-80; De Stefani,I 882; Namias , I 891 ; Nev ian i , 1891, 1894, 1900a, b ;De Angel is d'Ossat & Neviani, 1896; Annoscia, 1963).C h e e t h a m ( 1 9 6 8 ) d i d n ' t e v a l u a t e s u c h r e c o r d ssuggesting that a heterogeneous assemblage of speciescould had been quoted under the name M. moniliferus,the unique recorded also in more recent papers (Poluzzi& Padovani, 1984: Barrier et al., 1987; Di Geronimo eta l . , 1 9 8 7 ) .

A careful revision of the already available materialand newly collected specimens from several localities,all from Plio-Pleistocene of southern ltaly, allowed fivedifferent species to be identif ied. Three of them arenew, whereas two have been doubtfully attributed toalready known species. The aim of this paper is todescribe all of them, outl ining their geographical ands t r a t i g r a p h i c a l d i s t r i b u t i o n a n d d i s c u s s i n g t h e i rpalaeoecological meaning. The actual presence of M.moniliferus in the area remains to be proved.

MATERIALS AND METHODS

Studied samples come from sediments cropping outi n S i c i l y a n d C a l a b r i a ( s o u t h e r n I t a l y , c e n t r a lMedi terranean) and deposi ted dur ing Pl iocene andPleistocene in shelf-to-slope palaeoenvironments (Fig.I ). Detailed information can be found in Barrier et al.( 1987) for the Pavigliana section, in Di Geronimo et al.(1987, 1992) for the Ringigl io sect ion and in Rosso(2002a, b ) fo r the Capo Mi lazzo "ye l low mar ls " .Palaeobionomical reconstructions follow the schemeby Peres ( 1982). Recognised palaeocommunit ies areindicated as follows: DC - coastal detrit ic assemblage;DL - detr i t ic offshore assemblage; SGCF : coarsesands and fine gravels swept by bottom currents; VP: ba thya l mud assemb lage ; CB - - wh i t e co ra l s

0 50krnIg

Fig. I - Geographical location of the studied samples. Star - M.

e l egan t i s s imus : Open c i r c l e : M . i t a l i cus ; Squa re _ M .

pauciarmatus: Closed circle - M. cf. teixeirai: Triangle - M. cf .

elegans.

assemblage; PE : heterogeneous communities sensuDi Geronimo & Robba ( 1989).

Samples were rout inely treated: sediments werewashed and sieved and specimens sorted from residuescoarser than 500 pm. Measures and some photos wereperformed by means of a stereomicroscope equippedwi th a JVC TKl38l Video Camera and a Zeiss KSl003.00 program. Selected specimens were washed usingH,O, and diluted acetic acid to remove pelit ic particlesp f io i coa t ing fo r scann ing e lec t ron mic roscope.Measures have been taken following the scheme byCheetham ( 1968).

Described material and types of the new speciesare deposited at the Palaeontological Museum of theCatania University (PMC).

SYSTEMATICS

Family MErnaRABDorostDAE Vigneaux, 1949Genus Metrarabdotos Cani, l9l4

Type species: Eschara monil ifera Milne-Edwards,1 8 3 6

Metrarabdotos elegantissimus sp. nov.(Figs . 2a-d, Figs. 3a-f)

Etltmology Superlative; from the Latin adjectiveelegans, alluding to the distinct beauty of the slenderbranches.

Material - Cala S. Antonino West: sample I ( 1998).Holotype: a fert i le branch (PMC B. l3 a 21 .4.2003).Para types : about one hundred fe r t i le and s te r i lef ragments (PMC B. l3 b 21 .4.2003).

Addi t ional mater ia l : Cala S. Antonino Centre I( 1999): 32 fragments; 2 (1999): 14 fragments; 4 (1998):60 worn fragments. Cala S. Antonino East: sample 14(2000): 2 worn f ragments. Punta Mazza: sample 7(2000): 55 not wel l preserved f ragments; sample 8(2000): a single fragment.

All samples Late Pliocene in age; VP assemblageswi th a var iab le amount o f con taminants f rom CBassemblages and the local addition of shelf species.

Desc r i p t i on - Co lony e rec t , d i cho tomous l yb ranch ing a t l ong i n te rva l s , o r i g i na t i ng f r om anextremely small encrusting base sealed by the frontalthickening of zooids.

Branches roughly cylindrical in cross section fromabout 600 to I 000 pm reaching 1400 pm near the base.Zooids in four alternating longitudinal rows, those ofthe central ones opening on a single side and the lateralones along the margins, thus forming a frontal and adorsal surface. Bifurcating rows absent.

Zooids elongated hexagonal, those of the lateral rowsenlarged at mid length, their l imits forming a slightlyzigzagging line on the dorsal surface . Zooidal boundarieswell marked by raised rims and a median groove earlyin ontogeny, obscured by increasing calcification.

Frontal wall thin and slightly convex in new zooidsprogressively th ickened and f la t in the o lder ones.

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l l l ' c 1 - r l t t ' t t l l ' l t [ ' l l t l - l ' l c i s t o c c l ] c s L l c c c s s i t l t l - k l l ( ) \ \ l l L l \" \ c l l o u c u l c a l ' c ( ) u s n u l r ' 1 s " . u h r c h r l c l ' r o s i t c r l u l i c r t h c

l l ( ) s t \ 1 c s s r r r i u r t t t ' u t t s g r c s s i o t t . l ' i l l i n g l l t ' c r l t t t t s

r l c p l ' c s s i o r r s ( F o i s . | 9 8 9 . I 9 9 0 ) . S c r l i n t c n t s i . l t ' c n r . t i t t l r

e ( ) n s t i t r r t c r l b r p l u r t k t t l r t i c l i l t ' u n t i r t i l c r u r r ( ) ( ) , / c s l t l c u l l r

e ( )n tp r i s i r r g l i r s s i l r l c r ' ( ) r . l s [ - r cds u i t h u [ ' r t t n r l un t skc l c to t t so l ' l ' r c l r t h i c t ) t ' g r . r r r i s n r \ . s r r c h i r s r s i d i r l s . c c h i n o d c l ' n t s .

Scl lc l ' r l t r ' r )00 u n t| ( l ( ) ( ) 3 )o l ' r t i ( ) u \ o t t l l t c l i ' o t t t l t l

b r r o / o a n s . b c l r t h i c l i r t ' l u n i n i l c t ' u r t s l t n r l [ r t ' u c h i o l - l o t l s .\ l t ' l t ' ( t t ' ( t l t t f r t l t t . s t ' l t ' , g t u t I i s . r i n t t t . . \ P . l l ( ) \ . C ( ) l l l c s l l ' t l t l l

l u r c r s t ' c co ru lu tg t hc \ { I ' 15 u t t d \ l I ' � l 6 , / o l t cs ( \ ' i o l an t i .

l 9 l { 8 u n r l l l c r ' s . c o t n t n . ) . l a t ' s c l r ( ) \ c r ' l u l t p r n g n i t h t h c( i c l u s i r . r n S t a g c o l ' l { i o c t u l . 1 | t ) t ) - l ; .

l ' t t I ( ( ) ( ' t ' o l ( ) , g . 1 ' - S ; r c e i t t t c t t \ o l ' . l l t ' I t ' ( t t ' ( t l t t f r t I r t . s

t ' l t ' ,g tu t l i . s ' . s ' i r t t t t . s ' s l l . r . l ( ) \ . . c ( )n r l t l c tc l r l t [ rscn t l l ' o l t t l l u t ' cl ' t l r l r n t i r r i l ' c r l r n n r r r d s . \ \ c r c l ' r l r r t t r l i r r u I n t t l s t u | |( ) r ' g r . l n ( ) g c n i c s u r t d r l l t r c t ' s . t h c i t ' l t t - r t t t t d a n c c L r s L l l r l l r

l l o s i t i r c l r r c l u t c d t o t h c g r ' : . 1 \ c l l r l i ' l c t i o n a n t ( ) t t n t . . l - h c

n c \ \ s l l c c i c s . i n l ' u c t . l i l n n s c r c c t c o l t l t t i c s . u h i c h .l r l t l r t l r r g l r p ( ) s s c \ \ i r t s a s n r l t l I c n c t ' t t s t i r r g [ r l t s c r t t l t

c \cccdrng 2 nrn in r l i i tn rc tc t ' " r t cc r lc r l s r t i t i t t r l r l l l t ' gc h i t r t ls t r l rs t ra ta . , , \ r ' c lu t i r c l r snra l l n r rn rbc t ' o l ' l r i t s r . t l l rug t ' t ' t c t ' t t s .t ' c l l l ' c sc r r t r r r g l c ss t l u rn 0 .5 " , ( ) l ' t hc t t l t u l nu tn t - r c l ' . h i t r c

t

A. Rosso - Metrarabdotos.from Ptio-Pteistocene of southern ltalv, u'ith description o.f nev

been found. Also ovicells are rare and fertile branchesaccount for less than l%. Finally, regeneration afterfragmentation, a reproductive strategy common withinsome species belonging to the same genus (Cheethamet al., 2001, Cheeth&ffi, 2002), appears irrelevant ason l y two f ragmen ts show some ev idence o fregeneration. All this data could indicate that the specieswas present in the Capo M rlazzo Upper Pliocene deep-water sediments, although always represented by a fewcolonies. The deposi t ional palaeoenvi ronment waslocated in the epibathyal zone, as pointed out by studiedassemblages allowing a palaeodepth of 500-600 m tobe inferred (see Gaetani & Sacce, 1984; Gaetani, 1986;Violanti, 1988; Sciuto , 2003).

Metrarabdotos italicus sp. nov.(Figs. 4a-l)

|e87 y,"::'?:(:' ",;#;:'J{'; .' {,r$:'

Ine- Edwards ) - Bn nnI rn

Etymologv - Relating to the geographical area, whichthe species comes from.

Material - Pavigliana section sample 9,DL-VP 150-180 m, Late Pliocene. Holotype: a ferti le branch (PMCB. 14a,26.4.2003). Paratypes: the remaining ferti le andsteri le I l9 branches (PMC B. 14b,26.4.2003).

Pavigliana section: sample 4: 60 specimens, DC, 40m; Early Pl iocene; sample 8: 40 specimens, DC andSGCF, 100- 120 m, Late Pliocene.

C a l a S . A n t o n i n o c e n t r e : s a m p l e 2 ( 1 9 9 9 ) : l 0specimens VP and CB, 500-600 m, Late Pliocene (MPl5Zone).

Description - Colony erect, originating from anencrust ing base which gives r ise to a subcyl indricals tem, which in turns forms d ichotomous branches.Branches roughly cylindrical or nalrow, ribbon-like I -

2 mm in w id th , ra re ly more , compressed, be forebifurcating.

Zooids in 6- l0 alternating longitudinal rows, rarelyup to l2-14. Inser t ion of new rows by b i furcat ion,observed only on a few of the available small fragmentsand apparently not related with other features. The firstzooid of a new row proximally pointed, except for thosedistal to an ovicell.

Zooids rectangular elongated; strongly curved whenplaced laterally to an ovicell . Zooidal boundaries wellmarked by raised rims and a median groove early inontogeny, suddenly obscured by calcif ication.

Frontal shield umbonuloid with finely granular andsl ight ly convex surface, l imited by about 20 evenlydistributed marginal pores becoming less obvious afterthe thickening of the frontal surface late in ontogeny.Interareolar costulae not evident. Primary orifice hiddenwithin peristome with an arched, not well developed,pouch-shaped distal shelf. Peristome tubular, about 300pm in diameter, and not very raised except on zooidsof the lateral rows where sharply protrudes, up to 350pm; proximal interior with a longitudinal groove markedby two lateral ridges tending to converge at mid length

thus simulat ing two pair of pointed lateral denticlessubsequently placed inside the peristome. Secondaryorifice semielliptical, with a marked narrow and deepproximal sinus.

Ordinary avicularia paired, persisting on ovicellatezooecia; originating from the distalmost lateral pair ofareolae, distolaterally placed to the secondary orifice,the avicularian chamber lining the peristome, the rostrumperpendicular to the branch surface, proximally andinward directed. Rostrum pointed, elongated; cross barcomplete. On ovicelled zooids, avicularia are smallerand curved along the edge of the peristoffi€, lateral tothe ovicell, at about l13 of its high.

Special avicularia sensu stricto are absent althoughsome avicularia placed along branch margins and, morerarely, on zooids distal to ovicel ls could be sl ight lycurved and a bit larger than normal ones.

Fertile zooids longer than normal ones, sensibly andsuddenly enlarging, their maximum width at level ofthe modified peristomes. Orif ice crescent-l ike, with astraight distal lip edged by a smooth reversed flangeand deeply hidden by the smooth-to-lightly tuberculate,extremely in f la ted, prot ruding proximal l ip , whichlargely projects over the ovicell, when fully developed.Ovicell roughly rounded, arched-to-truncated distally;the ooecial cover moderately convex, sculptured bynumerous radial ribs separated by porous grooves andmargined by a row of areolae, similar to the frontalones .

Ances t ru la sea led by ca lc i f i ca t ion ; in a s ing leinstance an ancestrular triad, each part accounting fora sector of about 120o. is visible in basal view.

Measures ( pm)number mean

Zooidal length 33 l l04Zooidal width 26 393Fert i le zooidal length 8 1498Fert i le zooidal width 8 776Avicularian length 12 228Avicular ian width 12 122

range standarddeviat ion

990- t220 63.54330-520 42.87

1320-1630 101.38700- 820 37 .39200-320 37 .90I l0 - 150 12 .67

Variabilitv Variability involves branch flatteningand width, zoordal shape and dimensions. Secondaryca lc i f i ca t ion inc reas ing in on togeny sea ls zoo ida lboundaries and marginal pores and occludes orif icestransforming basal branches in strongly calcified stemswith a somewhat uniform surface only marked by light,irregular longitudinal furrows.

Remarks Metrarabdotos italicus sp. nov. roughlyresembles M. nyst i Lagaai j , 1952, for the sal ientperistomes, the somewhat costulate ovicell shield andthe wel l protruded proximal l ip of ovicel led zooids.Nevertheless, this latter species, seemingly with largerbranches, has prox ima l ly loca ted , s l igh t ly d is ta l l ydirected peristomial avicularia and a heavily costulateproximal l ip in ovicellate zooids. M. italicus sp. nov.recalls also M. eleganlissimus sp. nov. for zooidalcha rac te rs , f r om wh ich i t i s a l t hough eas i l yrecognisable, at first sight, for the branch shape andthe peculiar arrangement of zooids along them. Besides,M. italicus sp. nov. has sensibly smaller zooids (more

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l l ' o i n l c r t r l c z o o t t l \ . \ \ l r r c l r l l r c \ c n s i l r l r l o n g c t ' l r n r l l t [ - r o r c

r r l l u i r l c r ' . t l t l t n l l r c : t c r i l c ( ) n c \ l l t c o t ' l t l t l t : l l t l s l t c l l r s

r r l r f c r r . t r t t l t l r c \ c e ( r l t r l . t t ' r o r i l ' i e c u t \ l i l , r l i r t r t \ l l . n ( ) \

i l l ) l l c l r r s l l l l ' g c r l n r l n u u ' k c t l [ r r l l e ( ) n r l l l n t l r t c l r n a l ' r ' ( ) \ \ c t '

l ) r ' ( ) \ n ' n r . l l s r n L l : . I ' c t ' t s t t l n r n t l l t r t e r r l l r ' r i . r l l r c l t l t o g c t l t c t '

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t i t ( ) l ' c t l c r c l o p c t l l t t t t l : t ) n t c r i l u t t r l r l l c l ' c n t u t l c r l . l t l l r e c r l

u l r . l : l r g l r t l r l l r ' ( ) \ n n l r l p o : t l t o n . ( ) r r c c l l s : l t o t t l l n t ( ) r ' c

r l c r c l o p c r l . l r l l l l l r c n l l r n o l : e r r l p t r r t ' c r l l l r ' ( ) \ i r n l t l l r p . n l ( ) r ' c

1 t r o . j c c t i r t g ( ) t t t l t c ( ) ( ) c e n t I e ( ) \ c l ' . l ' t r t ' t l t c r ' t t t ( ) t ' c . l l t c

n l t ' r ' ( ) \ \ . r ' r t r t r o r t - l r l . , c l ' r n r n e l r c r t o g c t h c t ' r r t t l r t l t e : l t l t 1 ' r c

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i i \ r r ' r t l r t t ' r l r \ r u l P l e ( )

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. r l e l ' l t t r t ' e n t ' .

l l t . ' , r t ' t t t t ' a P l t t ' t l r o t ' t ' l t t t l e t l \ r t t t t l t l t ' ( ) . I . t l t ' l ' 1 t , , , - ' , . ' 1 1 . '

l . r r ' \ l l i l , / ( ) ( ) l t l l t l t ' l t l t ' t l t r ' i t t t ( ) l t ( ) l t l " t t r r t l i t t t t l t o l l ; t i 2 , , , r t , . 1 . .

I r ' t t l t l l \ r t t l l I l e ( )

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A. Rosso - Metrarabdotos.from Ptio-Pleistocene of southern ltalv, v'ith description of nev

d e v e l o p m e n t o f m a r g i n a l p o r e s a s w e l l a s t h emorphology of ovicells characterise this species amongother European cogeneric ones. Similarit ies could bestressed wi th an undescr ibed species f rom Rhodesf i gu red by Chee tham (2002 ) a l t hough spec imenexamination is needed to state conspecificity.

Distribution Metrarabdotos italicus sp. nov. hasbeen found in two nearby localities from southern ltaly:Pavigliana in southern Calabria and Capo Mrlazzo inthe north-eastern corner of Sicily. Stratigraphically, thespecies seems to be widely distributed in the Pliocene,from the Globorotalia puncticulata to the G. inflataZone in the Pavigl iana sect ion (Barr ier et al . , 1987).Within this time interval, probably in the MPl5 Zone,deposited also the Cala S. Antonino layers containingM. italicus sp. nov.

Paleoecologv - Most specimens of Metrarabdotositalicus sp. nov. come from sandy (mainly medium-to-coarse sands) sediments containing abundant skeletalremains of benth ic organisms (essent ia l ly mol luscs,bryozoans and subordinate brachiopods) point ing tomiddle- to-outer shel f envi ronments (Barr ier et a l . ,1987). Fossil associations allow to infer that sands werecolonised by coastal detr i t ic assemblages (DC) nearthe Inf ra l i t tora l -Ci rcal i t tora l bound l ty , and detr i t icoffshore assemblages (DL) at the shelf edge, testifyingdepths up to 150- 180 m. Pa laeoenv i ronments werecharacterised by strong bottom currents (Barrier et al.,1987). In a single case, sample 2 ( 1999) from Cala S.Antonino, a few specimens were found in deep-watersediments, though probably containing inputs fromnearby, shallower edge shelf environments. Like thepreceding species, also M. italicus sp. nov. seeminglydeveloped small sized colonies, arising from encrustingbases not exceeding 2 millimetres in the present material,thus exploiting coarse sandy or the rare fine gravellye lemen ts , as subs t ra tes . Sexua l p ropaga t i on wasprobably the unique reproduct ive st rategy for th isspecies as only a single doubtfully regenerated branchhas been observed; ovicelled fragments are commonwith an incidence of about 18% whereas bases are rare,not exceeding 2% of total fragments.

Metrarabdotos pauciarmatus sp. nov.(F igs . 5a-m)

1987 Metrarabdotos cf . moni l i . ferum (Mi lne-Edwards) - Dl

GEnoNtMo Er nr- . , tab. l .1992 Metrarabdotos sp. Dr GenoNlMo Er AL., tab. 3 Qtars).

EtvmologV - Relating to the scantiness of aviculariaand to their complete absence on most zooids.

Material - Ringiglio section: sample 2, PE l, about30 m. Holotype: a s ter i le branch wi th av icular ia onlateral zooids (PMC B. 15a,26.4.2003). Paratypes: ther e m a i n i n g 2 3 7 s t e r i l e f r a g m e n t s ( P M C B . l 5 b ,26.4.2003); Early Pleistocene.

R ing ig l io sec t ion : sample l : 80 spec imens, PEl ,about 20 m; sample 3: I specimen, PEI ,20-30 m; EarlyPle is tocene.

Description Colony erect formed by ribbon-likebranches l -4 mm in w id th , compressed, en la rgedbefore bifurcating. Trifurcation and irregular branchinggiving rise to slightly twisted branches common.

Zooids in 10-20, sometimes up to 30, al ternat inglongitudinal rows. Common insertion of new rows bybifurcation in zones of rapid widening; f irst zooids ofnew rows proximally cuneiform.

Zooids relat ively small , rectangular elongated orc lub-shaped, somet imes cons t r i c ted in the midd le .Zooidal boundaries markedly raised to form swollenth ick r ibs, more and more obvious wi th increasingcalcif ication, up to as wide as the proximal half of azooid, late in ontogeny. Zooids placed along the marginswider than the central ones. Frontal shield convex andcoarse ly g ranu lar tend ing to deve lop a few la rgetubercles aligned on the midline; l imited by l8-20 evenlydistributed large rounded marginal pores, except at thep rox ima l end where they become la rge r andquadrangular or often develop as a unique arched cleft,becoming more and more obvious late in ontogeny.Interareolar costu lae usual ly weak and per ipheral ,somet imes marked by granu le a rched a l ignments .Primary orifice marked by a small proximally projectingpouch-shaped distal shelf . Peristome low except onzooids of the lateral rows where it extends slightly onthe distal margin of the zooid and protrudes giving aserrated appearance to the branch margin; denticles, aproximal lateral pointed pair, shallowly set. Secondaryorif ice distally developing in a thin, proximally sinuate,round tubule.

Ordinary avicularia absent on most zooids, singleand sma l l , l a te ra l l y p laced to t he pe r i s tome andproximal ly directed, not easi ly dist inguishable frommarginal areolae. They are present in a very fewinstances on zooids of the central part of the branches,more common on lateral zooids and, above all, on themarginal broader ones, often paired with larger sizedspecial avicularra, characterised by a chamber extendingdistally beyond the orifice; rostrum pointed and slightlycurved around the margin of the or i f ice, d i rectedproximally.

Fertile zooids not present on the studied material.Ancestrulae and bases not observed.

Measures ( pm)number mean range standard

deviat ion7 60-940 44.98290-400 32.088 r 0 - 1 0 2 0 5 5 . 1 0420-580 46 .38t40-220 26.1|70- 130 16.76

Zooidal length 26Zooidal width l8Lateral zooidal length 23Lateral zooidal width l7Avicularian length I IAvicularian width 12

843329909484t6794

Var iab i l i t v B ranch w id th and number o flongitudinal zooidal rows varying to some extent aswell as zooidal size and shape. Nevertheless, variabil ityappears mainly related to ontogenetic stage, involvingincreasing second ary and extrazooidal calcif ication.Zooidal margins, raised but comparably thin in youngstages, become thick and swol len; surfaces granularat the beginning, can develop coarser granules up totrue tubercles along the mid l ine; zooidal interareolar

r 8 Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

costulae sometimes evolve into vert ical wal ls abovefrontal surfaces. The abnormal development of thela t te r two fea tures , en la rg ing , s t rong ly r i s ing andcoalescing to some extent, form patches of irregular" spongy " su r faces . Some f ragmen ts , seeming l ycorresponding to s tems or near-base branches, aredraped by u thin layer of extrazooidal calcification, thesur face marked by i r regu la r ly anas tomos inglongi tudinal r idges and furrows, local ly by coarsetuberc les.

Remarks - Metrarabdotos pauciarmatus sp. nov. iseasily recognisable amongst all the species known fromthe European-African area for the absence of aviculariain nearly al l zooids except the marginal ones. Whenpresent, ordinary avicularia are single. The presenceof a single avicularium is a feature shared only with M.pouyeti Marcopoulou-Diacantoni & Wuest from the LatePliocene of Greece, but no marginal special aviculariahave been repo r ted fo r t h i s spec ies . F ina l l y , M .terraconensis Reguant, 1960, from the Burdigalian ofSpain, showing somewhat similar zooidal characters(low peristomes and large peripheral areolae) completelylacks avicularia. Thus, it is l ikely that these three speciescould be closely all ied. Taking into account the trendabsence/presence of avicularia through single normalavicular ia versus normal p lus specia l av icu lar ia , anincreasing complexi ty can be envisaged, seeminglypointing to a phyletic relationship M. terraconensis-Mpouyet i -M. pauc ia rmatus . Never the less , fu r therknowledge is needed about distribution in time and spaceof Metrarabdotos species in the area, to put forwardmore valuable hypotheses.

Distribution - Metrarabdotos pauciarmatus sp. nov.has been found only in sediments cropping out alongthe Ringiglio section, near Mazzarino (central Sicily)which have been referred to the Early Pleistocene (DiGeronimo et al., 1987), namely to the uppermost partof the Emilian stage.

Paleoecology Metrarabdotos pauciarmatus sp.nov. comes from fine sands comprising an about 30%mud fraction; specimens reducing dramatically whenmud rises to about 50%. Bryozoan assemblages areoligospecific and mainly represented by the lunulitiformsReussirella reussiana (Manzoni) and Cupuladria gr.canar iens is (Busk) to wh ich B i f lus t ra savar t i i(Audouin) and Chaperiopsis annulus (Manzoni) add.Macrofaunas are largely constituted (ca. 90%) by thefree living serpulid Ditrupa arietina (Mueller) and thebivalve Corbula g ibba (Ol iv i ) , two f i l ter - feeding,opportunist ic species, whose dominance is indicat iveof the so called heterogeneous communities l iving inbiotopes largely influenced by water turbidity linked tohigh muddy inputs, mainly in basin instabil ity conditions(Di Geronimo et al . , 1987,1992; Di Geronimo & Robba,r ese).

M. pauciarmatus sp. nov. is exclusively representedby f ragmen ts , t he l a rges t o f wh i ch , t h ree t imesbranched, is about 2 cm high and 4 mm wide. I t islikely that colonies were larger and comparable in size

to those of other Neogene-to-Recent Mediterraneanadeonelliform species, which grow up to 10-20 cm inhigh, or more. It is worth noting that neither bases norovicells were found in the studied material. In addition,some of the largest fragments show some evidence oftw is t ing and i r regu la r b ranch ing , o f ten l inked toregenerat ion, usual ly by d is ta l budding. In a s ingleinstance a short branch arises from two adjacent zooidsby f ronta l budding. F inal ly , some f ragments showevidence of second dry, ext razooidal ca lc i f icat ionenveloping previously broken branch surfaces. All thispo in ts to hypothes ise as foss i l popu la t ions o f M.pauciarmatus sp. nov. seemingly exploited an asexuallyreproductive strategy, at least in the inferred, particularlys t ressed pa laeohab i t a t , whe re co lon i sa t i on l i ke l yhappened through the input of broken branches fromneighbour ing hard and/or coarse-gra ined bot toms,which survived and continued to grow by regenerationin the palaeobiotope. Similar adaptations have beensometimes observed for adeonel l i form species, suchas Smittina cervicornis (Pallas) and Adeonella calveti(Canu & Bassler).

Metrarabdotos cf. elegans Buge & Galopim deCarvalho. 1963

(Fie. 6t)

1963 Metrarabdotos elegans BucE & Gnloplvr DE CenvnlHo,p.162, p l . 1, , f igs. 3-4, text- f igs. l3-14.

1966 Metrarabdotos elegans Buge & Galopim de Carvalho -

Buce, p . 42 , p l . C , f ig . 5 .197 | Metrarabdotos elegans Buge & Galopim de Carvalho -

Gnloprv oE CnnvnLHo, p. l4 l , text- f ig. 20.1976 Metrarabdotos elegans Buge & Galopim de Carvalho -

Pouyer , p . 74 , p l . 12 , f igs . l -2 , 7 .1987 Metrarabdotos moniliferum (Milne-Edwards) - BnnRtER Er

nl., tab. I Qtars), non pl. 3 , f i9.4.1992 Metrarabdotos elegans Buge & Galopim de Carvalho - Er-

Hn: ln l r , p . 230, p l . l4 , f ig . 6 .1997 Metrarabdotos elegans Buge & Galopim de Carvalho -

HnvonNE & Molssrrre, f ig. 15 (2) .

Material - Pavigli ana section: sample l6b, VP, 500m, Early Pleistocene (Emilian).

Description - Colony seemingly formed by branches,at least up to 3 mm in width.

Zooids in 18, i rregular ly al ternat ing longitudinalrows, in the unique observed fragment comprising onlya m a r g i n . C o m m o n i n s e r t i o n o f n e w r o w s b ybifurcation, the first zooid proximally cuneiforrn.

Zooids relatively small, subrectangular or variouslyconstricted. Zooidal boundaries marked by a thin raisedrim. Frontal shield convex and granular limited by about20 marginal pores, separated by moderate interareolarcostulae. Peristome thin and raised, more developedand longitudinal ly channel led proximal ly. Secondaryorifice orbicular, with a proximal naffow and deep sinus.

Ordinary avicularia paired, proximo-laterally placedto the or i f i ce , the av icu la r ian chamber l in ing thepe r i s tome ; c ross ba r comp le te ; r os t rum po in ted ,upward and inward directed.

Fertile zooids, ancestrulae and bases not observed.

I t l ' ) ,

g - d

t&

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t s .

t t

3, n a

3t

rc

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'tj ' ,;r

. t 't .

(

i;

{t

20

Measures (pm)

Zooidal lengthZooidal widthAvicularian length

Avicularian width

number mean

9 9069 3748 1 9 58 9 6

Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

range standarddeviat ion

83 0- l 060 84.87310-450 s0 .03t7 0-220 t9 .278 0 - l l 0 l 1 . 8 8

Variabi l i ty Variabi l i ty relates to zooidal andavicular ian shape and s ize on the unique avai lablefragment.

Remarks The observed f ragment has beententatively assigned to M. elegans as it is the uniquespecies in the area wi th pai red la tera l av icular iap r o x i m a l l y l o c a t e d i n r e s p e c t t o t h e o r i f i c e .Nevertheless, the present fragment actually differs fromfigures and descriptions in l i terature for the zooidalshape , no t cons tan t l y sandg lass -shaped , and fo ravicularian shape and position, more elongated and notso prox ima l as in M. e legans . Ava i lab le da ta i sinadequate to state if it surely falls or not within M.elegans variability.

Distribution - Until now Metrarabdotos elegans wasknown f rom the westernmost Medi terranean areacoming back from the Messinian (Morocco: El Hajjaji,1992) and the Pliocene (Morocco and Portugal: Buge& Galopim de Carvalho, 1963; Galopim de Carvalho,197 l ; Spa in : Pouye t , 197 6 ; A lge r i a : Hamdane &Moissette, 1997).

Palaeoecology The species has been recordedfrom shelf environments and the not well-preservedfragment found in Pleistocene bathyal sediments couldhave been displaced or l ikely reworked from older(Pliocene) shallower sediments.

Metrarabdotos cf . teixeirai Buge & Galopim deCarvalho, 1963

(Figs. 6a-e)

1963 Metrarabdotos teixeirai BucE & GnloprM DE CnnvnlHo, p.157, p l . l . , f igs. l -2, text- f igs. l0- l I ;

1966 Metrarabdotos teixeirai Buge & Galopim de Carvalho -

BucE, p . 41 , p l . C , f ig . I ;197 | Metrarabdotos teixeirai Buge & Galopim de Carvalho -

Gnlopru oE CanvnlHo, p. 143, pl. 22, f igs. 2, 4, text-f ig.2 t .

1987 Metrarabdotos moniliferum (Milne-Edwards) - BenRIER Ernl., tab. I Qtars), non pl. 3 , frg.4.

Mate r i a l Pav ig l i ana sec t i on : samp le 4 , 78specimens, DC , 40 m; sample 5: 28 specimens, DCand SGCF, 50 m, Early Pliocene; sample 8: 9 specimens,DC and SGCF, 100- 120 m, Late Pliocene.

Description Colony erect. Branches ribbon-likeslightly enlarged before bifurcating, the largest about 3mm wide and 8 mm high.

Zooids in I 4-16, often up to 24, regularly alternatinglongitudinal rows; the disposition of orif ices simulatinga p innate pa t te rn a long branches. F i rs t zoo ids inbifurcating rows proximally cuneiform.

Zooids small, rectangular to club-shaped, sometimesundulate; laterally arched and proximally tapered, whenlateral to ovicells; somewhat raised distally. Zooidalboundaries raised and swollen, with a median furrow.Margin al zooids wider and slightly longer than the centralones. Frontal shield with nearly 20 small marginal poresevenly distributed, separated by interareolar costulaeon the per iphery; centra l area convex , granular .Pe r i s tome l ow . Second a ry o r i f i ce rounded andproximally sinuated.

Ordinary avicularia, paired or single, constant andpersisting on ovicell ate zooids; placed laterally and levelto the peristome, rarely sl ight ly distal ly; proximal lydirected; their chambers tubular, commonly broken.Rostrum pointed, cross bar complete.

Special aviculana larger, located on marginal andlor peri-ovicel late zooids, sometimes sporadical ly onzooids of the central rows, usually paired with a normalone. Their chamber extends distally beyond the orifice;the rostrum sharply pointed, directed proximally.

Fer t i le zooids commoh, p laced in centra l rows,often clustered (up to 5-6) on some branches, sensiblylarger and longer than normal zooids, suddenly andsharply enlarging at peristome level. Orifice invariablybroken; distal lip straight with a narrow reflected flange;proximal lip protruding and projecting, for at least l14,over the ov ice l l . Ooec ia l cover rounded- to -mi t re -shaped, sometimes truncated distally. Sculpture givenby tuberculate ribs alternating with perforate furrowsforming a costulate pattern.

An ancestrular tetrad is hardly visible on the uniquebase suffounded by extrazooidal calcification occludingorifices of basal zooids and making the surface windingsulcate.

Measures ( pm)

zooidarength ":*;

;:r, o1x,;:tl

Zooidal width l8 3 13 280-360 24.01Ferti le zooidal length 12 I 183 960- 1300 82.28Fert i le zooidal width 12 7l9 640-850 58.07Lateral zooidal length 9 816 720-970 80.48Lateral zooidal width 9 477 430-500 26.93Avicular ianlength 6 125 l10-140 13.78Avicularian width 6 80 70-90 6.32Special avicularian length l0 254 200-340 52.54Special avicularian width I 0 I 30 I I 0- 140 13.36

Variability - Zooidal and avicularian measures showa certain degree of variability, as well as zooidal shape.Ontogenesis, with increasing frontal and extrazooidalcalcif ication, accounts for branch thickening.

Remarks Present material has a bad preservationstate owing to heavy recrysta l l izat ion processes,preventing some characters to be observed. Specimenshave been referred to Metrarabdotos teixeirai, owingto zooidal shape, position of paired normal aviculariaand ovicell sculpture, although doubtfully, as sizes ofboth steri le and ferti le zooids are smaller that thosereported by Buge (1966) and Galopim de Carvalho(1971). The presence of special avicularia and marginalwider zooids observed in the studied specimens, but

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22

Paleoecologt - Metrarabdotos cf . teixeirai has beenfound in sandy sediments containing faunas belongingto the coarse sands and fine gravels under the influenceof bottom cuffents (SGCF) and the detritic bottom (DC)assemblages, thus test i fy ing deposit ion in soft , midshelf , current swept bottoms. Bryozoans are mainlyconstituted by Turbicellepora tubigera (Busk), Srn ittinacervicornis (Pallas), Melicerita charleswortht Milne-Edwards, Entalophoroecia deflexa (Couch) and Tbrviairregularis (Meneghini) and by minor percentages ofOnychocella marioni Jull ien, Tremopora radicifera(Hincks) and Hippopleur i . fera sedgwick i (Mi lne-Edwards) .

A single base has been found in the present materialbut, unlike most of the previously cogeneric describedspecies, M. cf. teixeirai shows a high incidence ofovicellate fragments, with a mean value of 320 , rangingfrom l4% (sample 5) to 77% (sample 8). This, joinedwith the absence of regenerated fragments, points to asexual reproductive strategy.

CONCLUSIONS

The present study allowed the identification of fivedifferent species belonging to the genus MetrarabdotosCanil, 19l4 from southern ltaly, namely central-easternSicily and southern Calabria.

Tab. I - Stratigraphic distribution of the "Old Word"- Canarian; Cz - Czech Republic; F - France; I :

Rumania; S - Spain; UK : United Kingdom, Y -

reference to Ages; (e) - Early; (l) - Late.

Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

Three of them (Metrarabdotos elegantissimtrs sp.nov. , M. italicus sp. nov. and M. pauciarmalas sp. nov.)are new, whereas the remaining two have been referred,although doubtfully, to M. elegans Buge & Galopim deCarvalho and M. teixeirai Buge & Galopim de Carvalho.The latter two species were previously known fromthe Iberian Peninsula and Morocco, with M. elegansreco rded f rom the A lge r i a t oo . A l l bu t no t M .p a u c i a r m a t u s l i v e d d u r i n g P l i o c e n e ; w i t h M .elegantissimu.r seemingly restricted to Late Pliocene andM. elegans, recorded since the Messinian, in Morocco,probably the long living species. The younger speciesseems to be M. pauciarmatus, which has been foundonly in Emilian (Lower Pleistocene) sediments. To thepresent knowledge, th is taxon could be one of theyounger or even the youngest one in the Old Worldarea.

A further species, M. moniliferus, the unique onepreviously recorded from ltaly is perhaps quite absentfrom the area. This taxon was quoted, mainly by 1800researchers, from Pliocene and Quaternary sediments(Seguenza, 1879-80; De Stefani , 1882; Namias, l89 l ;Neviani , I 891 , 1894, 1900a, b; De Angel is d 'Ossat &N e v i a n i , 1 8 9 6 ) a n d m o s t r e c e n t l y f r o m L o w e rPleistocene clays in southern ltaly (Calabrian, Venosa:Annoscia, 1963) and Crotonian sands from westernSici ly (Poluzzi & Padovani, 1984). Nevertheless, al lor, at least, part of the old material (only partly and

species of Metrarabdotos. A: Algeria; Au - Austria: B : Belgium; C - Crete; CaItaly; L : Libya, M - Morocco; P - Portugal; Po : Poland; R : Rhodes; Ru :

Yugoslavia. Asterisks indicate that in l i terature only the Epoch is given without

@(oo)

E(E.C,o(l)

()xf,aoq

A. Rosso - Metrarabdotos.from Ptio-Pteistocene of southern ltaly, v'ith description of nev

hard l y ava i l ab le ) c l ea r l y doesn ' t be long to M .moni l i ferus, as often a costulate ovicel l was beendescribed (see Seguenza, 1880: p. 208, for example).Moreover, both the most recent ( 1990s) records appeardoubtful, either for the bad preservation of specimens(abraded and showing broken ovicells as obvious alsofrom figures) either for the sedimentary context allowingreworking processes to be hypothesised. Otherwise,most records of M. moniliferus need to be re-evaluated(see Cheeth am,, 1968) and the distribution of this speciesseems presently restr icted to Pl iocene sediments ofsome European localities (U.K., the Netherlands, France,Iberian Peninsula: Lagaaij,, 1952; Buge , 1957; Buge &Galopim de Carvalho, 1963; Galopim de Carvalho,197 l ) and maybe North Afr ica (Morocco, Tunis ia:Galopim de Carvalho, l97 l) . Pleistocene specimensfrom the U.K. and the Netherlands, on the contrary,have been considered probably reworked (Cheetham,l e68) .

Spec ies o f Me t ra rabdo tos , wh i ch a re eas i l yd is t inguishable for thei r morphological characters,appear also well separated ecologically. Apart from M.elegans, for which a few data is avat lable, the threeP l i ocene spec ies p laced themse lves i n d i f f e ren tbathymetric horizons also exploiting slightly to sharplydiverse substrata. M. teixeirai seemingly lived on softs a n d s , i n m i d - t o - o u t e r s h e l f e n v i r o n m e n t s . T h i sdistribution was partly overlapped by M. italicus, yetextending towards major depths, to the shelf break.M. elegantissimus thrived in epibathyal bottoms withpalaeodepths of about 500-600 meters. I t could beworth noting as these three species line up subsequentlyin a two axes diagram considering zooidal width versrtszooidal length, both for sterile and fertile zooids. Theline up, from species with small-s rzed towards large-sized zooids, fits well with the inferred distribution fromshal low shel f waters to deep s lope waters. Wi th inspecies and within genotype zooidal size variability, bothin t ime and space , has been obse rved i n seve ra lcheilostomebryozoans and related to water temperature(see O'Dea & Okamura, 2000 for a rev iew). I f weassume that similar changes extend also to interspecific,within genus variabil ity, the progressively increasingzooidal size registered in the above-recorded species,could be l inked to physiological changes related todecreasing water temperature, at great depths, in anarea seemingly subject to upwellings.

Finally, M. pauciarmatus, seemingly the shallowestspecies, was capable of exploiting a particularly stressedpa laeoenv i ronmen t cha rac te r i sed by abno rma lsed imentary inputs . Broken f ragments seeming lysurvived and continued to grow adopting twisting andc o m b i n i n g s e x u a l a n d a s e x u a l r e p r o d u c t i o n , b yfragmentation, &S a winning adaptation strategy.

ACKNOWLEDGMENTS

Thanks are ex tended to A lan Cheetham ( Smi thson ianI n s t i t u t i o n , W a s h i n g t o n ) f o r d i s c u s s i o n s i - p r o v i n g t h emanuscript and to Pierre Moissette (Universite Claude Bernarde,Lyon). Dott. Annamaria Mannino (Palermo University) and MrCanzanella (Naples University) are kindly acknowledged forSEM photos. Paper financially supported by Catania University

Grants to Rosso, Palaeontological Research Group: contributionn.329.

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Manuscript received 05 April 2004Revised manuscript accepted l3 October 2004