Pleistocene Bathyal Molluscan Assemblages from Southern Italy

Home , Akritogyra, Calliotropis, Callumbonella, Cantrainea, Ennucula, Fissurisepta

Rivista Italiana di Paleontologia e Stratigrafia volume luj numero 3 tavole 1-10 pagine 389-426 Dicembre 1997

PLEISTOCENE BATHYAL MOLLUSCAN ASSEMBLAGES FROM SOUTHERN ITALY

ITALO DI GERONIMO & RAFAEL LA PERNA

Key-llbrds: Bathyal benthos, Molluscs, Pleistocene, Mediterra- greater depth, probably not exceeding 1,000 m, is suggested for the nean, Southern Italy, Systenatics, Paleoecology. other two. Taxonomic affinities with nonheast Atlantic and more gene- Riassunto. Sono state studiate quattro faune a molluschi batiali rally with World Ocean deep-sea molluscan faunas are remarkable. provenienti da depositi pleistocenici dell'italia meridionale (Calabria The Plio-Quaternary evolution of the deep Mediterranean benthos is e Messina). Vengono segnalate 136 specie, 24 sono trattate nella pane discussed. sistematica e 35 sono illustrate. Le seguenti nuove combinazioni vengono proposte: Solariella marginulata (Philippi, 1844), Ipbitus tenuisculptus (Seguenza, 1876), Introduction. Benthomangelia tenuicostdta (Seguenza, 1879), Cbrysallid,a rnicroscalaria (Seguenza, 1876), Ennucula corbuloides (Seguenza, 1877), Ennucula Although Plio-Pleistocene Mediterranean molluscs rotundata (Seguenza, 7877), Tbestyleda cuspiddtd (Philippi, 1844), Ka- have been investigated since the last century, the deep- tadesmia confusa (Seguenza, 1877), Austrotindaria pusio (Philipp| sea have 1844), Austrotindaria salicmsis (Seguenza, 1877). I generi Fissurisepta forms been disregarded for a long time. Diffi- Seguenza, 1862, Solariella'Wood, 1842, Ennucula kedale, 7931, The- culties in finding deep-sea deposits, located mainly in styleda lredale, 7929, Ledella Verrill 8c Bush, 1897, Yold.iella Verrill & areas affected by strong Plio-Quaternary tecronics, lo'w Bush, 1892, Batlryspinula Filatova, 7958, Katadesrnia Dall, 7908, Au- density of deep-sea macrofaunas, and taxonomic com- strotindaria Fleming, 1948 e Cadulus Philippi, 1844 sono tassonomi- camente commentatì. plexities, have all probably restrained paleontological in- I caratteri composizionali delle faune studiate, ed in particola- vestigation. Pioneer contributions were made in the late re la rtcchezza, in nuculoidi, corrispondono a quelli tipici delle asso- 19th century by Giuseppe Seguenza, who investigated ciazioni malacofaunistiche profonde. Una profondità di sedimentazio- extensively the Plio-Pleistocene deposits southern Ca- ne di 500-600 m è indicata per due delle faune ed una maggrore, of probabilmente entro i 1.000 m, è suggerita per le altre due. labria and Messina, describing many deep-sea species Notevoli sono le affinità tassonomiche con 1e malacofaune and correctly interpreting their paleoecological meaning profonde de1 nord-est atlantico ed oceaniche in generale. Ijevoluzione (Di Geronimo, t991, pp. 137-143). Many of the fossil de1 benthos profondo mediterraneo durante i1 Plio-Quaternario viene discussa. species described by Philippi (1836, 1844) from the same area have been shown to come from deep-sea deposits as Abstract. Four Pleistocene bathyal molluscan assemblages from well. Seguenza referred most species to the "Astian", a southern Italy (Calabria and Messina area) were studied. One hun- middlelate Pliocene stage now in disuse. On the basis dred and thirty-six species were recorded. Twenty-four were classified and described in detail and thiny-five were illustrated. of modern stratigraphic knowledge, the Astian deposits The following new combinations are proposed: Solariella mar- of Seguenza are known to be middle-late Pliocene to ginulata (Philippi, 7844), Ipbitus tenuisculptus (Seguenza, 1876), Bmr early-middle Pleistocene in age. honangelia tenuicostdtd (Seguenza, 1879), Chrysalli.cJa miuoscalaria More recently, several papers (some by amateur (Seguenza, 1876), Ennucula corbuloid.es (Seguenza, 7877), Ennucula rotundata (Segtenza, 1.877), Tltestyleda cuspidata (Philippi, 7844), Kata- malacologists) have been devoted to deep-sea Pliocene desntia confusa (Seguenza, 1877), Austrotindaria pusio (Philippi, tsaa), and Pleistocene molluscan faunas of ltaly, mainly dea- A ustrotindaria salicensis (Seguenza, 1877). Comments concerning the ling with their composition and paleoecology (Di Gero- taxonomy of Fissurisepta Seguenza, 1862, Solariella Vood, 1842, En- nimo, 1.979; Robba, 1981; Rindone Yazzana, 1,989; nucula lredale, 1931, Tbestyleda Iredale, 1929, Ledella Yerctll & Bush, & 1897, Yoldiella Verrill & Bush, 1897, Bailryspínula Filatova, 1958, Ka- Micali & Villari, 1989, 1991.; Tabanelli, 1.993, 1997; tadesmia Dal1, 1908, Aus*otindd.ria Fleming, 1948 and Cadulus Pht- D'Alessandro & De Marco, 1993; Palazzi & Villari, lippi, 1844 are included. 1994, 1.996; Yazzana, 1.996; Barrier et al., 1996, efc.). The assemblages are dominated by nuculoids and fit the general compositional pattern of the deep-sea molluscan communities. A Other works dealing with several benthic inverte- paleodepth of 500-600 m is inferred for two assemblages, whereas a brates, have helped to outline the Plio-Quaternary evo-

Istituto Policattedra di Oceanologia e Paieoecologia, Università di Catania, Corso Italia 55,I-95729 Catania. 390 I. Di Geronirno & R. La Perna

CALABRIA stctLY

Fig. 1 - Locationof thesampledsites(triangles): A.Lazzàro andValloneCatrica; B.Furnari,C.Bianco(I.G.M.maps,modified).

Iution of the deep Mediterranean benthos (Benson, Materials and methods. 1.972; Gaetani & Saccà, 1984; Zibrowíus, 7987, I99la,b; The faunas studied come from four Pleistocene pelitic outcrops Roux et al., 1988; Barrier aL, 7989; Perna, 1994; et La (Fig. t). One site is located ca. 2 km norrheasr of Furnari (Messina). '1995; Moissette & Spjeldnaes, Di Geronimo et al., 1996; Two other sites are located in southern Calabria (Messina Strait area), Di Geronimo 6c La Perna, 1996, 1997; Rosso & Di Ge- near Lazzàro and at Vallone Catrica (ca. 2 km southeast of Lazzà,ro). A fourth site is ca. 4 km north ofBianco (fonian coast). ronimo, in press). All of these papers support an "ocea- Faunas were collected zla sediment bulk samples and manual nic" model of the Piio-Pleistocene deep-sea benthos, picking at outcrops. About 10 dmr of sediment were smpled at Fur- markedly different from the Recent one. Climatic chan- nari, Vallone Catrica and Lazzàro, whereas ca. 40 dm3 were sampled at ges and geodynamic events at Gibraltar have been in- Bianco, due to the 1ow density of macrofauna. Bulk sampling is hìghly recommended in paleoecological stu- creasingly interpreted as the main factors contributing dies (Di Geronimo & Robba, 1976; Robba, 1978), as it allows quanti- to the faunistic and oceanographic evolution of the deep tative data related to the original paleocommunities (relative abundan- Mediterranean. ces, size-class distributions, etc.) to be obtained. Bulk sampling is also comparable with sampling Recent shelled assemblages using a grab or Taxonomic studies of Plio-Pleisrocene deep-sea boxcorer. On the other hand, large and rare species are more easily moiluscs have, however, only occasionally been attemp- collected by hand. ted. Many species have never been revised, and mosr Bulk samples were disaggregared in warer and washed through knowledge is based solely on living species. a 250 1tm mesh. Molluscs were sorted from the whole dried fraction coarser than 500 prm, whereas only small ?lmounts of the fine fractions The present deals work with four Pleistocene ba- were examined. thyal molluscan assemblages from a classical area, i.e. Fossils were generally well preserved and did not show signs of the southern Calabria and Messina regions. Systematic, postmortem wear. Some panicularly thin and fragile shells were lar- gely fragmented, such as Seguenzia monocingulata, ecological and biogeographical aspects of several species Delectopecten vitreus and Abra longicallus, as well as the long-shelled scaphopods Graptactne are reported and discussed in the context of an evolutio- agili and Entalina tetragond. Shel1 repairs, and evidence of drilling and nary model of the deep Mediterranean benthos. boring by endobionts, were also obserted, mainly in large specìes. Bathyal Molluscan frorn Soutbern ltaLy 391

Concerning the species addressed in our systematic study, the Pleistocene in age, as shown by foraminiferal (Violanti, material examined is generally abundant (up to several hundred speci- 1988) and magnetostratigraphic data (Aifa e'r. a1.,1987). mens), except for a few rare species. Material fron other localities in The Plio-Quaternary sequence along the southern Italy and from some Nonh Atlantic stations $tIuséum Na- Jonian tional d'Histoire Naturelle, Paris; MNHN) was also used for cornpari- coast of southern Calabria ("Jonian Basin post-orogenic sons. The material studied is housed in the Museo di Paleontologia Complex" of Ogniben, 1973),where the Bianco ourcrop dell'Università, Catania. is located, may be summarized as follows: early Pliocene whitish marls ("Trubi"), middlelate Pliocene bluish Geological setting. marls, Pleistocene greyish marls, yellowish silts and calcarenites. The Plio-Pleistocene marls and silts are deep- The Plio-Quaternary sedimentation in the Messi- sea deposits. The sample studied comes from the upper na Strait area was strongly affected by tectonics, which part of the greyish marls. split the substrate into several fault-controlled compartments (Barrier, 1987; Ott d'Estevou et al., 1987). Strati- Composition and paleoecology. graphic sequences show sharp lithological and bathyme- tric changes, as well as signs of synsedimentary tecronis, Overall, 136 species were recovered from the four such as gravity flows, olistoliths, fault escarpments and sites (Tab. 1), most of them from bulk samples. The ri- neptunian dikes (Vachard et al., 1,987; Barríer er a1., chest faunas are those from Furnari (96 species, 1,847 1.987; Barrter et al., t996). Extensional movements led ro specimens) and from Bianco (24 species, 1,109 speci- a rapid sinking of compartments and to deep-sea sedi- mens), whereas the faunas from Vallone Catrica (55 spe- mentation. Subsequent uplifting raised the bathyal depo- cies, 605 specimens) and Lazzà,ro (56 species, 305 speci- sits as much as 400-500 m above the present sea-level mens) are less abundant. Several small-sized species were (Barrier, 1987; Montenat & Barrier, 1987). detected from the fine fractions, such as some gastro- More distai Tyrrhenian and Jonian sectors of sout- pods informally listed as "skeneimorph" (sensu \flarén, hern Calabria seem to have been affected by less intense 1992). Abouf l3o/o oÍ the species occur in all of the bulk Plio-Quaternary tectonics, as demonstrared by weaker sampie assemblages, although with different degrees of synsedimentary deformations. Strong vertical displace- dominance. The trochid Solariella mdrginulatd, the nu- ments are recognizable in these areas as well, related to culoids Ennucula corbuloides, Austrotindaría pusio, Ba- an increasing uplift of the Peloritani Mounts during the tlryspinula excisa, Katadesmia confusa, lcdella Tnessanensls Pleistocene (Fabbri et al., 1980). and the scaphopods Graptacme agilis, Cadulus ouulum The geology of the Furnari area was studied by and Entalina tetragona are among the most frequent spe- Kezirian (1992) who recognised the "Furnari block", a cies. compartment a few km wide, controlled by two fault The dominance of nucuioids (40o/o to 47o/o) and systems (N30"E and N140'E). Shelf calcarenites and scaphopods (6o/o to 23o/"), the scarciry of heterodonts, silts overlay the Miocene substrate, whereas two N-S consisting almost exclusively of the micro morphtc Kel- faults affect the northern margin of the biock, defining liella abyssicola, and the occurrence of several taxa, such a graben ca. 200x500 m in size. The graben is filled with as Fissurisepta, Cantrainea, So lariella, S egwenz ia, Benth o- grey bathyal marls, while on the fault escarpments re- mangelia, Ledella, Yoldiella, Batlryspinula, Katadesmia, mains of deep-sea sessile faunas are preserved. Foramini- Batlryarca, Limops is, Delectopecten, Cyclopecten, Cadulus, feral assemblages indicate an early Pleistocene age. resulted in a compositional pa.ttern closely fitting the The geology of Vallone Catrica was described by "taxonomic structure" of the soft-bottom deep-sea mol- Barrier (1987). A fault escarpment, involving the Mioce- luscan communities (Clarke, 1962; Knudsen, 7979; ne substrate, is encrusted by deep-sea sessile faunas. The F{ickman, 1984; Rex, 1981). escarpment is draped by early Pleistocene grey marls Nuculoids account for one of the dominant tro- containing bathyal faunas. Faulting caused a progressive phic groups, i.e. the infaunal detritus-feeders. Epifaunal deepening of the basin, as suggested by the early coloni- detritus-feeders are also weil represented by trochids and zation of the escarpment by deep-shelf faunas ("yellow rissoids. Scaphopods are microcarnivorous, preying on corals") and the subsequent settlement of bathyal faunas benthic foraminifers (Bilyard, t974; Poon, 1987). Filter- ("white corais"). feeders are scarce and consist mainly of epifaunal (ar- Barrier (1987) also studied the sequence of Laz- cids, anomiids, pectinaceans) or seminfaunal (Limopsi) zàro, where some shallow water units, followed by ba- species, whereas Kelliella abyssicola is the only abundant thyal marls (ca. 13 m thick) and deltaic conglomerates heterodont. Carnivorous fa;r-:a are typically represented are distinguished from the base to the top. Boulders fell by neogastropods. Some parasitic eulimid and pyrami- from the faulted basement during the deep-sea sedimen- deliid species are also present. tation, becoming "rocky islands" on the muddy bottom A strong bathyal mud-dwelling smck, accounting (Barrier et aI., 1.996). Bathyal marls are early-middle for 77-90o/o of the bulk sample assemblages, can be reco- 392 L Di Geronimo G R. La Perna

BF L c Gastropoda cocculinomorph sp. 0.3 Notocrater so. 0.8 Fissurisepta papillosa Seguenza,'1 862 t.o 0.6 Fissurlsepfa rostrata Seguenza, 1 862 + 0.2 Puncturella noachina (Linne, 1 771 ) 0.5 Pu ncturella granulata (Seguenza, 1 862) 0.3

Solariella marginulata (Philippi, 1 844) 0.6 o. l 3.8 Danilia otaviana (Cantraine, 1 835) + P u tz ey s i a w i se ri (Calcara, 1 842) 0.3 0.2 4 Callumbonella suturale (Philippi, 1 836) 0.9 1 + + Cantrainea peloritana (Catraine, 1 835) + + + Homalopoma emulum (Seguenza, 1 876) 0.2 Paruiturbo so. 0.1 Akritogyra conspicua (Monterosato, 1 880) Llssofesfa minima (Seguenza, 1 876) Llssofesfa gittembergerl (Van Aartsen & Bogi, 1988) Llssofesfa tu rrita (Gaglini, 1 987) Llssofesfa sp. A rk Llssofesfa sp. B Trochaclis versiliensis Warén, Carrozza & Rocchini, '1992 Orbitestella sarsi (Bush, 1897) Rugulina monterosatoi (Van Aartsen & Bogi, 1986) *.

Ci rso n el I a romette n si s (Granata-G ril lo, 1 877) 0.7 U.J Moelleriopsis /?essanensis (Seguenza, 1 876) + U.J 0.6 skeneimorph sp. A v.z skeneimorph sp. B skeneimorph sp. C 0.1 Brookula (?) sp Seguenzia monocingulafa Seguenza, 1 876 6.7 0.3 Haloceras carinata (Jeffreys, 1 BB3) 0.1 Alvania cimicoides (Forbes, 1844) 8.4 0.7 Alvania subsoluta (Aradas, '1847) 4.3 4.1 2.0 Alvania zetlandica (Montagu, 1 81 5) + Pseudosetia tu rgida (Jeffreys, 1 870) u.ó Obfuse//a sp. 1.5 1.2 Aporrhais serresranus (Michaud, 1 828) 0.1 Torell ia del icata (Philippi, 1 844) + + Capulus ungaricus (Linne, 1758) 0.1 Polin ices fusca (Blainville, 1 825) 0.4 0.2 + Strobiligera brychia (Bouchet & Guillemot, 1978) + + Eumetula aliceae (Dau2enberg & Fischer, 1896) 0.4 0.1 Aclis attenuans Jeffreys, 1883 0.4 0.3 0.2 Aclis bicincta Seguenza, 1879 0.1 G raph is g racilis (Monterosato, I 87 4) 0.2 1.0 0.7 U.J Melanella spiridioni(Dautzenberg & Fischer, f 896) 0.2 Granulina sp. 0.4 Claviscala richardi(Dautzenberg & de Boury, 1897) + Opaliopsis atlantis (Clench & Turner, '1952) + lphitus tenuisculptus (Seguenza, 1 876) 0.1 Pagodula echinata (Kiener, '1840) '1.6 '1.9 + Trophonopsis muricata (Montagu, 1 803) 0.1 Coralliophila richardi (Fischer,'1 BB2) + Coralliophila squamosa (Bivona, 1 838) T Fusinus rostratus (Olivi. 1792) + 0.2 Batlryal Molluscan from Southern ltaly 393

B F L c Nassarlus /rma (Dillwin, 1817) 2.4 I Nassarrus edwardsi (Fischer, 1 882) 0,1 | Nassarrus cabrierensis (Fischer & Tournouer, 1873) 0.'1 + I Profundinassa sprnulosa (Philippi, 1 844) 0.1 | Amphissa acutecostata (Philippi, 1 844) 0.7 4.4 0.3 + | "Babylonella" profunda (Tabanelli, 1 985) 0.3 41 91 | Be nth o m a n g el i a te n u icosfafa (Seg u en za, 1 87 9) l-t z.c 0.3 I Tu rriclavus h arpul a (Brocchi,'1 81 4) 0.1 | D rill iol a e me n data (Monterosato, 1 87 2) 0.2 05 | Taranis moerchi (Malm, 1 863) 0.'1 Taranis cf . borealis Bouchet & Waren, 1980 1.3 I + I Gymnobela torquata (Philippi, 1 844) 0.2 l Spirotropis confusa (Seguenza, 1 879) 0.3 Mitrolumna smithi (Dautzenberg & Fischer, 1896) 0.3 + Teretia feres (Reeve, 1844) 0.2 Pleurotomella packardi Verrill, 1 872 0.'1 0.1 0,2 Pleurotomella eurybrocha (Dautzenberg & Fischer, 1896) 0.2 0.1 Heliacus alleryi (Seguenza, 1 876) 0.2 Heliacus zancleus (Seguenza, 1 876) + Ammonicera rofa (Forbes & Hanley, 1850) * * Ammonicera fischeriana (Monterosato, I 869) * Ch rysallida brattstroemi Warén, I 99'1 u.b NA 0.7 Ch rysal I id a sfefanlsi (Jeffreys, I 869) Chrysallida microscalaria (Seguenza, 1876) ot Chrysallida sp. 0.1 Eulimella scillae (Scacchi, 1835) no 0.2 Eulimella ventricosa (Forbes, I 844) 0.1 0.1 .1 Odostomia acuta Jeffrevs, 1848 0 1.1 Odosfomla sp. 0.4 Ondina sp. 0,2 * * Tj ae rnoe i a exq u isita (Jeffreys, 1 883) Japonacteon pusillus (McGillivray, 1 843) 0.1 Acteonidae sp. 0.2 44 Pyrunculus obesiusculus (Brugnone, 1877) 0.8 t. I ^a 0.2 Ringicula leptocheila Brugnone, 1 873 0.'1 0.1 Cylichna a/ba (Brown , 1827) 0.1

Bivalvia Nucula sulcata (Bronn, 1831) 0.1 2.4 3.0 3.2 Ennucula corbuloides (Seguenza,'1 877) 0.1 0.5 7.2 z.o Ennucula rotundata (Seguenza, 1 877) 0.1 3.0 1.2 Ennucula decipiens (Philippi, 1844) + Ennucula tenuis (Montagu, 1808) 0.4 1.3 'Yoldia' minima (Seguenza, 1 877) 0.1 1.6 0.6 Phaseolus ovatus Seguenza, 1877 0.4 0.2 Microgloma tumidula (Monterosato, 1880) * * * Microgloma pusilla (Jeffreys, 1 879) 0.2 0.3 Microgloma sp. 2.0 Au strotinda ri a pusio (Ph ilip pi, 1 844) 8.5 2.1 ó_ó o.o Thestyleda cuspidata (Philippi, 1 844) 0.6 0.8 1.6 Bathyspin ula excr'sa (Philippi, 1 844) 28.9 30.0 15.6 1.4 Katadesmia confusa (Seguenza, 1 877) 2.1 lo 1.6 1.2 Ledella messanensls (Jeffreys, 1 870) 1.9 1.7 5.2 tJ. / Yoldiella micrometrica (Seguenza, 1877) 0.2 Yoldiella philippiana (Nyst, 1 845) 0.7 + 394 I. Di Geronimo & R. La Perna

B F L c Yoldiella seguenzae Bonfitto & Sabelli, 1995 1.4 0.3 t.o 27 Asperarca nodulosa (Mùller, 1776) 0.6 z.ó 0.8 Bathyarca grenophia (Risso,'1 826) 0.'1 0.'1 ?n 6.9 Limopsis pygmaea (Philippi, 1836) 1"9 z.+ Limopsis minuta (Philippi, 1836) + u.z + Limopsis aurita (Brocchi, 1814) + Limopsis sp. + Cyclopecten hoskynsi (Forbes, 1 844) v.z 0.9 Delectopecten vitreus (Gmelin, I 79'1 ) 1.8 12.1 Karnekampia bruei (Payraudeau, 1 826) + n? 0.2 Spondylus gussonii Costa O.G., 1829 + 0.3 + Pododesmus aculeatus (Mueller O.F., 1776) 0.4 0.2 8.2 zz.o Acesta excavata (Fabricius, 1779) 0.2 Notolimea crassa (Forbes, 1844) 0.1 0.1 0.2 Leptaxinus ferruginosus (Forbes, 1 844) U.J Digitaria digitaria (Linne, 1758) 0.1 Plagiocardium papillosum (Poli, 1 795) 0.'1 Abra longical/us (Scacchi, 1834) 0.5 1.0 0.3 Kelliella abyssicola (Forbes, 1 844) 1.4 u.o 4.8 1.4 Timoclea ovafa (Pennant, 1777) u.z Poromya granulata (Nyst & Westendorp, 1839) Spinosipell a acuticostata (Philippi, 1 844) 0.2 + Cuspidaria rostrata (Spengler,'1 793) 1.2 0.3 u.z Cardiomya philippi (Seguenza, 1 876) + 0.2

Scaphopoda Graptacme aglls (Sars M. in Sars G.O., 1872) 5.2 1.2 J.J 5.2 Cadulus ovulum (Philippi, l B a) 2.3 0.6 t.o 1.7 Cadulus diploconus Seguenza, 1876 n? G ad il a jeffreysi (Monterosato, 1 875) 0.5 1.5 1.0 2.0 Entalina tetragona (Brocchi, 1 81 4) '15.0 2.4 1.6 2.0

Total of soecimens í lno 1,847 305 AAN

I XD. I Systematic list of the studied faunas, relative abundance of species and total amounr of specimens from bulk-samples (B : Bianco, F : Furnari, L: Lazzàro, C - Vallone Catrica). lJnscored species are from the fine fraction (<5OO pm) (stars) and from manual .;"t;"" 1".^."."ì gnised. Most nuculoids and scaphopods, together with from deep-sea fossil faunas are difficult. In the present many other bivalves and gastropods, can be referred to case, this is due mainly to the wide bathymerric range this stock. The remaining portion consists mainly of of most of the extant species. Furthemore, the abundant mud-dwelling species ranging from the outer-shelf to the occurrence of extinct species, such as SolarielLa margtnu- bathyal zone, such as Ahtania cimicoides, Nucula swlcata, lata, Batlryspinula excisa, Tl:estyleda cuspidata, Austrotin- Kelliella abyssicola and "Gadila" jeffreysi. A minor stock daria pusio, Katadesmia confusa and Cadulus wwlurn, of sessile bivalves (Asperarca nodulosa, Batlryarca greno- makes paleobathymetric comparisons with the Recent pbia, Spondylus gussoníi, Pododesmus aculeatus, Acesta ex- assemblages even more difficult. Nevertheless, a depth caaata) can be distinguished within the dominant freeli- not shallower than 500 m is clearly suggested by the ving epifaunal and infaunal species. Sessile bivalves are taxonomic composition and ecological structure. This more frequent at Furnari, Lazzàro and Vallone Catríca, value is also indicated by the occurrence in all samples due to the nearby hard-bottoms (fault-escarpments and of. Ennucula corbuloides and Yoldiella seguenzae, whose boulders) or to coarse skeletal remains on the bottom. A present depth range begins at about 500 m (see below). few shelf coarse-bottom species, i.e. Digitaria digitaria, Di Geronimo & La Perna (1996) regarded the increasing Plagiocardium papillosum and Timoclea wata, occtr as dominance o[ Batlryspinula excisa in Pleistocene beds as rare juvenile valves in two samples. related to depth. On this basis, the molluscan faunas As noted by Moissettte & Spjeldnaes (1995) and from Bianco and Furnari should be regarded as the dee- by Di Geronimo et al. (in press), paleodepth inferences pest. The occurrence of Seguenzia monocingulata in Bathyal Molluscan from Soutbern ltaly 395 both sites, and of Haloceras carinata at Furnari, also sug- by generally smaller species, lower than 3 mm), and gests a greater depth, probably within 1,000 m. On the high-conical forms, consisting of larger species. In the other hand, it is worth noting the scarce occurrence at first (most typical) group, shells are generally thicker

Lazzàro and Vallone Catríca of Yoldiella philippiana, a and more heavily sculptured, as in -E papillosa and E gra- species which "replaces" the allied Y seguenzae at depths nulosa, as well as other Atlantic species (tenuicola Dall, shallower than 500-600 m. A paleodepth greater îhan 1927, trifolium Dall, t881., agulbasae Clarke, I96t) and 1,000 m for the Furnari and Bianco assemblages cannot the Pacific F. undwlau Okutani, 1964. The Aúantíc acu- be excluded, although no data presently exist to support minata'Watson, 1883, the Pacific festioa Crozier, 1966 this conclusion. and the southern onychoides Herbert & Kilburn, 1986 all belong to the second group, togerher with .E rostrata Seguenza, 1862 (see below). These species share thin, glossy and finely granulous shells. Such a morphological Systematic descri ption grouping could be useful for a subgeneric arrangement of Fissurisepa. Class G ast ro p o d a Cuvier, t797 Distribution. The only "Recent" Mediterranean re- Subclass P ro s o b ran c h i a Milne Edwards, 1848 cord is that by Taviani (1974), who reported both ,F. papillosa and ,F. granulosa as subfossil shells from a 9OO Family F i s s u r e I I i d a e Flemrng, 1822 m deep Tyrrhenian station. Nordsieck (1968) also reported both species from the Northeast Atlantic, but his papillosa Fissurisepta Seguenza, 1862 drawing of F. papillosa is not very reliable. Pl. 1, Íis. 1,2,3 Seguenza (1862) originally reported E papillosa as a Miocene fossil from Rometta (Messina), whereas he L862 Fissurisepta papillosa Seguenza, p. 10, fig. 2,2a, 2t:. referred it to the deep-sea "Astian" faunas in a later 1974 Fissurisepta papillosa - 40, fig. 1a, 1b, 1c. Taviani, p. work (Seguenza, !879). 1.98A Fissuriseptapapillosa - \Varén, p. 14, pl. 2,fig. 19,20. 1983 Fissurisepta papillosa - Ghisotti & Giannini, fig. 5. E papillosa is common in Pleistocene bathyal deposits from southern Ita|y (La Perna, 1994), being fre- Description. Small, iow-conical, laterally compres- quent where a coarse biogenic fraction occurs on pelitic sed shell, with apical hole and internal septum. Ovate substrates (Barrier et al., t996). base. Early coiled stage lost. Apical hole roughly triangular to polygonal. Septum consisting of a thin lamina, slightly curving when inserting on shell sides. Sculpture Fissurisepta rostrata Seguenza, 1862 of small tubercles loosely arranged in radial rows to ran- Pl. 1, fig. 4, s domly scattered. Short ill-defined radial ridges extending downward from tubercles. Rugose comarginal micro- 1862 Fissurisepta rostrdtd. Seg.uenza, p. 10, fig. 3, 3a, 3b, 3c. sculpture. Height up to 1.9 mm, base 2,8x2.0 mm. 1896 Fissurisepta crossei Dauf.zenberg & Fischer, p. 492, pL 22, frg. 15 . Taxonomy. This is the type-species of Fisswrisepu Seguenza, 1862. Regarded previously as a subgenus of Description. Small, high-conical, laterally compres- Puncturella l-owe, !827, Fissurisepta was ranked definiti- sed shell with apical hole and internal septum. Ovate base. Early coiled stage lost. Posterior apex. Oyate api vely to full genus status by Co'wan (1969) (see also 'ù7a- rén, 7972 and Corselli & Bernocchi, 1,993). cal hole. Anterior slope convex, posterior slope concave. Septum consisting of a thin straight lamina, on Remarks. Jeffreys (1883a) described F. granulosa inserting from the Norwegian coast, correcting a previous identi- shell sides and extending well beiow midway of shell fication as papillosa Seguenza. In the same paper, he re- height. Sculpture of fine granules, roundish to ovate, ali- ported F. papillosa from the Portugal coast as well. Wa- gned in irregular radial rows, becoming crowded and rén (1980) noted that this last papillosa is different from randomly scattered towards the base. Smooth apical por- the one described by Seguenza, and is probably an un- tion. Faint sulcus, internally corresponding to septum described species. insertion, extending on both sides from apex. Height up F. papillosa and F. granulosa are closely related, the to 5.8 mm, base 4.0x2.8 mm. latter differing mainly in having smaller tubercles, regu- Remarks. F. rostrata is similar to the North Atlan- iarly aligned in radial rows (see Taviani, L974). A rugose tic E acuminata (]watson, 1883), which differs in being microsculpture also occurs ìn F. granulosa. shorter and iess curved. It is also very similar to F. ony- Comparing the sheil shape of several Fissurisepa choides Herbert & Kilburn, 1986 (South Africa, 250-430 species, Ghisotti & Giannini (1983) distinguished two m), which differs in being more curved and with a shor- morphological groups: low-conical forms (characterised ter septum. 396 I. Di Geronimo G R, La Perna

Two specimens from the Biscay Bay (THALASSA sion in \íarén, 1993) and S. marginulafa is even more Exp., St. 413, 48o03N/08o29F-,805 m, MNHN), contai- similar to the type-species, when compared with other ning the soft body , were examined. They closely resem- smooth-shelled species referred to Solariella. In Calliotro- ble both E uossei Dautzenberg & Fischer, 1869 (from pls, however, many spiral rows of sharp tubercles occur Azores in 1,022-!,385 m) and the fossil Mediterranean on the last whorl. In the type-species C. ottoi (Philippi, specimens. Therefore, F. crossei should be regarded as a 1844), the protoconch is slightly protruding and inrolled synonym of E rostrata. (Colman & Tyler, 1988; Warén, 1991), as seen in the Distribution. The Recent distribution of F. rostrata New Zealand species (Marshall, 1979). In S. marginwla- ranges from Biscay Bay south to the Azores, excluding ta, the protoconch is rather flat and regularly coiled, as the Mediterranean. Jeffreys (1883a) reported E rostrata in other Solariella species (Herbert, t987;Varén, 1,993). from 740-1,095 fms. off Portugal. He also reported the Calliotropis and Solariella were allocated in distinct "Travailleur" (Biscay Bay, I,093 fms.) and the "Challen- subfamilies by Hickman & Mclean (1990), mainly on ger" (Bermudas, 1,,375 m) findings. The latter proved to the basis of soft body features. be based on F. acuminaa by Pérez Farfante $9a7). Remarks. The only Atlantic species showing a cer- In deep-sea Pleistocene deposits F. rostrata frequen- tain resemblance in shape and sculpture to S, marginula- tly occurs together with F. papillosa. ta is S. amabilis (feffreys, 1865), recently treated by Wa- rén (1993). Another seemingly closely related species Family Trocbidae Rafinesque, 1815 was reported by Dautzenberg & Fischer (1896, p. 178, pL. 20, figs. 15-17) from the Azores as S. cinca (Philippi, Solariella marginulata (Philippi, 1844) comb. nov. 1836). k is, however, quite distinct from Philippi's species (see holotype \Warén, 1993) and Pl" 1, fig. 6,7 in it might be an undescribed species, if not a peculiar form of S. amabi- 1844 Tiochus marginul atus Philippi, p. 227 , pl. 28, tig. 4. /ls. This species, together with S. marginulata and S. L846 Solarium nitidum Aradas, p. 172, pl. 1, fig. 9a,b. amabilis, seem to represent a group of allied eastern 1986 Líscbkeia marginulata - Micali Ec Villari, fig. 2. Atlantic deep-sea species. As suggested by \Warén (pers. 1994 Calliotropis rnarginulata - Palazzi & Villari, Írg. 22, 31. 1996 Lischeia marginulata - Yazzana, pl. 1, fìg. 2a.b. comm.), the western deep-sea species Solariella (Suaoo- trocbu) lubrica (Da11, tSSt) (holotype in Quinn, 1929), Description. Medium-sized, trochoid, solid shell. also shows a certain resemblance to S. marginulata, but Protoconch ca. 1,.5 whorls, finely pitted, 350 pm large. is distinguishable by its markedly convex and smoother Flat teleoconch whorls, numbering up to 5. Suture whorls. deep. First 3 whorls with axial faintly prosocline riblets, S. cincta was also described from the Plio-Pleisto- intersected by 3-4 spiral cords. Following whorl with a cene of southern Italy. The east Atlantic Gibbula inop- subsutural series of sharp tubercles, a fainter suprasutu- tanda Incard, 1.897 was recently considered a synonym ral one, and axial riblets. Last whorl with a subsutural (Varén, 1993). series of sharp tubercles, two peripheral keels (the up- Tiocltus marginulatus was originally reported from permost sometimes granulated) and basal granulated "prope Messina". The original drawing, description and cords bounding a wide funnel-shaped umbilicus. Parietal origin of Solariwrn nitidum Aradas, 1846 from Gravitelli lip thickened by a strong granulated funiculus. Periphe- (lr4essina) leaves no doubt about its synonymy with 7. ral and basal part frequently displaying faint spiral stri rnarginulatus. ae. Roundish aperture. Height up to 7 mm. Height to Distribution. S. marginwlata is known only from breadth ratio ca. 1.0 in adult specimens. deep-sea Pleistocene deposits of southern Italy, where it Taxonomy. This species may appear somewhat dif- is one of the most frequent species. ferent in morphology from the low-spired and strongly As "Calliotropis marginulatltrs", if has been regar- keeled type-species of Solariella S.V. Wood, t842 (5. ma- ded (Di Geronimo et al., L996; Di Geronimo & La Per- culaa S.Y. \Wood, 1842), showing instead a certain re- na, 1997) as a cold stenothermic species, extinct in the semblance to Calliotropls L. Seguenza, 1,903. Actually, Mediterranean. The present revision supports this pa- Solariella is still used in a rather broad sense (see discus- leoecological interpretat ion.

PLATE 1

Fig. 1,2,3 - Fissurisepta papillosa Seguenza, 1.862. Lazzàro. 1. Lateral view, 1.5 mm. 2. Apical view, scale bar : 250 pLm. 3. Sculpture detail, scale Der : IUU Um. Fig.4,5 - Fissurisepta rostrdtaSegùenza, 1862.Lazzàro.4.Laterzl view,4.5 mm. 5. Sculpture detail, scale bar : 50 pm. Fig.6,7 -Solariellamarginulata (Philippi, 1844).Furnari.6.6.1 mm.T.Protoconch.^"1. 1""': ?óÒrr- Pl. 1 BatlryaL Molluscan from Southern Italy 397 398 I. Di Geronimo & R. La Penra

Famiiy S e gu e n z i i d a e Yerriil, 1884 As usual, Seguenza described some "varieties", namely elegans (with well-defined axial striae), and elata

Seguenzia monocing ulata Seg,a,enza, 187 6 (higher than the typical form). Another Seguenziidae, Ancistrobasis (Phi- Pl. 2, fig. s-8 reticulaa lippi, 1844), was described as a fossil from southern Italy.

1.876a Seguenzia monocingulata Seguenza, p. 188. It is presently known from the Atlantic (\X/arén, 1991). 1994 Seguenzia monocingulata - Palazzi 6c Villari, fig. 7. Distribution. "5. monocingulata" was reported from the Atlantic by some authors (\latson, 1886, Daurzen- Description. Small, conical-globose, thin, fragile berg, 1927, etc.), but the actual value of these records shell. Prominent protoconch, ca. one'whorl, finely gra- can be evaluated only by examining the material. Quinn nulated, 270-300 pm large. Teleoconch of 5 whorls in (1983) listed S. monocingulata withour comment on fully grown specimens. Spire whorls v/ith a sharp sli- synonymy or its present distribution. ghtly supramedian keel (shoulder keel), beginning soon S. monocingulata is rather common in deep-sea de- after protoconch and a faint subsutural and suprasutural posits from southern ltaly. Di Geronimo & La Perna cord. Last whorl with a shoulder and a peripheral keel, (1996) reported it as S. formosa. and 5 basal cords. Axial sculpture of faint, sigmoidal ri- biets, evenly spaced, crossing numbe- not spiral cords, Famiiy H a I o c e r a t i d a e Varén & Bouchef, l99l ring 12-13 per mm on last whorl. Whorl profile concave between spirals. Suture mostly hidden by sutural cords. Haloceras carinata (]effreys, 1883) Convex base, becoming flatter to slightly concave around the umbilicus. Umbilicus closed in adults, nar- Pl. 4, fig. 2,9 row in juveniles. Concave and tooth-bearing columeilar 1883b Cithna carinata p. 111, pl. 20, [ig.9. lip. Outer lip broken in all specimens examined. Height Jeffreys, 1991 Haloceras carinata - Warén & Bouchet, 137, Íig.33, 16, 58-61, up to ca. 4.0 mm; height to breadth ratio 1,.3 in fully 10s-108. grown specimens; last whorl to shell height ratio 0.6. !993 Haloceras carinata - Bouchet & Varén, p. 722, fig. úA8, 17A9, 1719, 1723. Remarks. An important unresolved puzzle invol- ves genus the Seguenzia, as Seguenza (1876a) and Jeffreys Remarks. One fully grown but partly broken and (1826) proposed it almost simultaneously. The priority one postJarval specimen were found in the sample from of Segwenzia Jeffreys on Seguenzia Seguenza was proven Furnari. by Marshall (1983) on the basis of ICZN rules, but ac- The deep-sea family Haloceratidae was recenrly cording to Warén (pers. comm), is pen- priority still described extensively by Warén & Bouchet (1991). ding. About twenty species are presently known from the Beginning with (1877, 1885), S. monocin- Jeffreys Atlantic, Indian and Pacific Oceans, five of them occur- gulata 1-876 S. Seguenza, and forntosa Jeffreys, 1876, have ring in the northeast Atlanric. been regarded as synonyms. Flowever, in spite of a gross Distribution. H. carinata was reported originally resemblance (see illustration of S. in formosa Jeffreys, from Portugal ín 994 fms. (|effreys, 1883b). It is an 1876), the two species are quite distinct. Inforrnosa, "ren amphiatlantic species, ranging from 27-42"N in 700-2,075 threadlike riblets on the base" occur, whereas monocin- m (\Warén & Bouchet, 1991; Bouchet &.\X/arén, 1993). gulata is characterized by five cords (as also noted by The specimens from Furnari are the first record Seguenza). In the former, the sigmoidal axial striae are of Haloceratidae from the Mediterranean area. crossed by "fine close-set spiral lines, producing a rericu- late appearance", whereas only sharp cords or keels oc- Family E p i t o n i id a e Berry, 79lQ cur in monocingulata. On the basis of present knowledge of shell morphology and sculpture in Seguenzia and Seguenziidae (Quinn, 1983; Marshall, 1983, I99I), lphitus tenuisculptus (Seguenza, 1876) comb. nov. these differences cannot be disregarded. Pl. 4, fig. 4

PLATE 2 Fig. 1 - Ahsania subsoluta (Aradas, 1847).Lazzà,ro,2.1 mm. Fìg.2 - Alpania zetland.ica (tr4ontagu, 1815). Furnari, 3.0 mm. Fig. 3 - Pyrunculus obesiusculus (Brugnone, 1877). Vallone Catrica, 3.5 mm. Fig. 4 - Graphis gracills (Monterosato,1874 ex Jeffreys ms.). Furnari, 2.1 mm, Fig. 5-8 - Seguenzia monocingulata Seguenza, 1876. Bianco. 5. Aboral view of a panly broken juvenile, 1.6 mm. 6. Detail of fig. 5. 7. Apical view, scale bar : 200 pm. 8. Protoconch detaìI, scale bar - 25 pm. rqq Pl. 2 Bathyal Molluscan from Southern ltaly 400 I. Di Geronimo & R. La Perna

1876b Stylotrochus tenuisculptus Seguenza, p. 186. distribution, whereas dregeri was a shelf species. Furr- 1925 Cithna marsballi Sykes, p. 190, pl. 9, fig. 4,4a. hermore, the stratigraphic distributions are different (see 1986 Ipbitus narsballi - Bouchet 6c Warén, p. 492, fig. 11,37, 1158, 1t59. below). 7994 Stylotrochus tenuisculptus - Benolaso &Palazzt, unnumb. fig. Actually, Babylonella Conrad, 1865 does not represent a good systematic position for either of these spe- Remarks. Bertolaso &. Palazzi (1994) lirst recogni- cies. As stressed by Verhecken (1986), the type-species of sed the synonymy between Stylonochus tenuisculptus Se- Babylonella has two well-defined columellar folds and gueîza, 1876 and Cithna marshalli Sykes, 1925. They re- apertural lirations, whereas both the columelia and the garded lphitus Jeffreys, 1883 as a synonym of Stylotro- inner lip are smooth in dregeri and in profunda. Consid- chus Seguenza, 1,876, which is preoccupied by Stylotro- erable shape differences also occur. On the other hand, chus Haeckel, 1,862. Admete Króyer in Móller, 1842 is markedly buccini- Two other lpbitus species are kno'wn from deep-sea phorm in shape (e.g. see the northeast Atlantic species Pleistocene deposits, i.e. L asperatus (Segtenza, 1,876) l: in Bouchet & Warén, 1985). A berter systematic posi- I. tuberatus (feffreys, 1883)l and L papillosocínctus (Se- tion for dregeri, profwnda and several related Miocene gueîza, 1876) (see Bertolaso & Palazzr, 1,994; Palazzi &. and Pliocene species (see Pavia, 1.976; Davoli, 1995) Villari, 1.996). The former is also known from Mediter- should be traced among the present-day Indo-Pacific ranean Late Glacial submerged beds (Taviani & Sabelli, deep-sea cancellariids (e.g. see Verhecken, 1997). 1982). The latter is closely related to L cancellatus Distribution. "8." profunda is found rarely in Plei- Dautzenberg & Fischer, 1896 from the Azores. stocene bathyal deposits. It ranges from (Middle)-Late Distribution. I. tenuiscwlptars is known presently Pliocene to Pleistocene, whereas "B." dregeri ranges from from Portugal and the Gibraltar Strait, its depth range Late Miocene to Early Pliocene. being probably not shallower than 500 m (Bouchet & Warén, 1986). Family T w r r i d a e Swainson, 1840 s./.

Famiiy Cancellariidae Gray, 1853 Mitrolumna smithi (Dautzenberg & Fischer, 1896) Pl. 3, fig. ltPl. 4, Íis.7 "Babylonella" profunda Tabanelli, 1985

Pl. 3, fig. 8 1896 Mitrornorpha sntitbi Dautzenberg & Fischer, p. 432, pL, 15, fig. 19 . 1980 Mitrolurnna smitbi - Bouchet & lX/arén, p.78, fig, 161.

1985 Balrylonella nassiformis profunda Tabanel\i, p. 21, frg. t-3. 1993 Admete dregeri profundus - Tabanelli, fig. 4. Remarks. Although in the typical form the last whori is axially smooth, a faint axial sculpture occurs Remarks. This species was originally described rarely among the specimens examined. The protoconch (Tabanelli, 1985) as a bathyal Late Pliocene subspecies of of M. sntitbi has 2.5 whorls, as in the common shallow- Babylonella nassiformis (Seguenza, 1879). Later, the same wafer M. oliaoidea (Cantraine, 1835), but it is larger in author (Tabanelli, 1993) reported it as lAdrnete dregeri the former (680 pm) than in the latter (SOO pm). profundus, following the remarks by Petit (1986) and Distribution. M. smithi is known presently from Verhecken (1986), about the preoccupied status of Can- the Azores, in 400-800 m (Bouchet & Varén, 1980). cellaria nassiforrnis Seguenza. The latter aurhor proposed This is the first record from the Mediterranean area. ldmete (s.1.) dregeri (Hoernes 6c Auinger, 1890) as the valid name for this species. Flowever, there are several Benthomangelia tenuicostata reasons to rank " Balrylonella" profunda to full species sta- (Seguenza, 1879) comb. nov. tus. It has a prevailing axial sculpture, in contrasr to the Pl. 3, fig. 2;PL.4, fìg.1. markedly cancellate sculpture in " Babylonella" dregeri. The protoconch shows many spiral threads ín profunda, 1879 Mangelia tenuicostd.td Seguenza, p. 258. and a few (3-a) profunda in dregeri. "B." had a bathyal 1994 Mangelia tenuicostd,td, - Palazzi 6c Villari, tíg. 17 , 18, 29 .

PLATE 3 Fig. 1 - Mitrolumna srnitbi (Dautzenberg & Fischer, 1896). Furnari, 6.1 mm. Fig.2 - Benthomangelid tenuicostdtd (Seguenza, 1879). Furnari, 4.7 mm. Fig. 3 - PleurotorneLla eurybrocha (Dautzenberg 8c Fischer, 1896). Furnari, 4.8 mm. Fig.4,5 - Ampbissad.cutecostd.ta (Philippi, 1844). Bianco.4.6.1mm.5.5.8 mm. Fig.6,7 - Thraniscf. borealis Bouchet 6c $ilarén, 1980. Bianco. 6.4.2mm.7.3.2mm. Fig. 8 - "Babylonella" profunda Tabanelli, 1985. Bianco, subadult, 3.3 mm. Batbyal Molluscan from Soutbern ltall 401 402 I. Di Geronimo ù R. La Perna

Description. Medium-sized, roughly biconical, cf. 1980 Thranis borealis Bouchet & S7arén, p.78, fíg. 162, 270. 1989 Tàranis borealis - Rindone k Yazzana, fig. 7a. rather solid shell. Protoconch of 3.5 convex whorls, 760- 960 pm large, with opisthocline riblets on ending por- Remarks. The scant Pleistocene material examined tion. Teleoconch of 3-4 whorls in fully grown speci- so far differs from the North Atlantic specimen repor- mens, with a shoulder one third from the adapical sutu- ted by Bouchet & Warén (1980), mainly in having the re. Axial sculpture of strong, fatnúy opisthocline ribs peripheral keel in a lower position. Some (Pl. 3, Fig. 7) (ca. 10 per whorl), flexuose and almost lameilar on are as slender as the Atlantic specimen, whereas others ramp. Spiral striae, arranged as alternating stronger and are larger and more squat (Pl. 3, Fig. 7; see also Rindo- fainter ribs, crenulating axial ribs and lacking on ramp. ne 8icYazzana, 1989, fig. 10). A clear-cut distinction be- Well-incised, sinuose suture. Narrow, elongate aperture, tween these forms (both occurring in the same assem- vzith a terminal rib. Deep subsutural sinus. Height up to biages) is not apparent, and these differences might be 6.5 mm. Height to breadth ratio 1 9. Last whorl to shell intraspecific (e.g. see the variability of Thranis moerchi height ratio 1.5 in adults. 'Warén noted by Bouchet & Warén, 1980). Vhether the Pleisto- Taxonomy. (pers. comm.) suggested Bentbo- cene material really belongs to T boreali.s is not fully mangelia Thiele, 1925 as the suitable systematic position understood. for the present species. Benthomangelia is rather similar Distribution. T borealis is known presently from to Mangelia Risso, 1826, the former differing from the the Swedish and Norwegian coast (150-1,980 m) and latter by having a thinner and markedly shouldered (bi- from the western Atlantic (Bouchet & Warén, 1980). conical) shell, fainter and more flexuose axial ribs, becoming almost lamellar on the ramp, a well-developed spi- Subclass H b n c h i 1840 ral sculpture and a deeper subsutural sinus. Furthermo- ete ro ra a GrayJ.E., re, Mangelia typically has a shallow-water distribution, Family Pyramidellidae Gray J.E., 1840 whereas Bentltomangelia is a bathyal to abyssal genus. B. rnacra ('Watson, 1881), a northeast Atlantic to Ghrysallida microscalaria (Seguenza, 1876) comb. nov. deep-sea (Bouchet Mediterranean species & \farén, Pì. 4, fig. 8 1980), is rather close to B. tenuicosuta.It differs in being more slender, high-spired, and less sculptured than the 1876a Thrbonilla (Pyrgulina) microscàlaria Segrenza, p. 92. present species. B. tenuicosaa also shows the typical protoconchal sculpture of the planktotrophically develo- Description. Very small shell, markedly thick for píng Benthomangelia species (see Bouchet & \Varén, the genus. Shape somewhat scalariform. Smooth proto- 1980, fig. 2I2,2I3). conch, strongly intorted and protruding, consisting of Remarks. A "Mangelia tenuicostata (Brugnone, slightly more than one visible whorl. Two convex teleoconch whorls with ill-defined spiral striae 1862)" is reported in the Mediterranean Mollusca chec- and strong ort- hocline ribs (14-15 last klist (Sabelli et al., 1990). This probably originates from in whorl), weakening toward the base and entering Nordsieck's (1977) report of "Mangelia tenuicosau Brr- the umbilicus. Deep, almost channelled suture. Ovate aperture. Absent gnone, 1868", clearly based on Pleurotoma attenudtd v^r. columellar fold. Height up to 1.25 mm. Last to tenuicosta Brugnone, 1862. whorl shell height ratio 0.8. Remarks. in the brief original description, Seguen- Distribution. B. tenuicostara is known only from za stressed a resemblance "Pyrgulina pygmaea", hi.s Pleistocene bathyal deposits of Messina and Calabria. fo species differing in having a shorter shell, more crowded ribs and a well-defined umbilicus. He marked it with an Taranis cf. borealis (Bouchet \flarén, 1980) & "s", to indicate a deep-sea species. The species regarded Pl. 3, fig. 6,7 as ngmaed is most lrkely Chrysallida stefanisl (feffreys,

PLATE 4 Fig. 1 - Benthomangeliatenuicostata (Seguenza, 1879). Furnari, protoconch, scale bar : 200 pm. Fig.2 - Haloceras carindta feffreys, 1883). Furnari, protoconch, scale bar : 200 pm. Fig. 3 - Gymnobela torquatd (Philippi, 1844) l: G. abyssorum (Locard, 1897)1. Furnari, protoconch, scale bar : 200 pm Fig. 4 - Iphitus tenuisculptus (Seguenza, 1876). Furnari, protoconch, scale bar : 250 pm. Fig. 5 - Pleurotomella eurybrocba (Dautzenberg & Fischer, 1896). Furnari, protoconch, scale bar : 200 pm. Fig.6 - Amphissa d.cr.ftecostdta (Philipphi, 1844). Bianco, protoconch, scale bar : 250 pm. Fig.7 - Mitrolumna srnitbi (Dautzenberg & Fischer, 1896). Furnari, protoconch, scale bar : 200 pm. Fig.8 - Cbrysallidamicroscalaria (Seguenza, 1876). Furnari, 1.1 mm. Fig. 9 - Haloceras carinata (]effreys, 1883). Furnari, 2.9 mm. Fig. 10 - Aclis attenuans Jeffreys, 1883. Lazzàro,4.9 mm. Pl. 4 Bathyal Molluscan from Soutbern ltaLy 403 404 I. Di Geronimo & R. La Perna

1869), for a long time misidentified as C. pygmaea (Gra- by Monterosato himself, in I-ocard's collection (Tiavail- teloup, 1838) (see Linden Er Eikenboom,1992), and also leur and Tàlisman exp., MNHN). It was said to be "un occurring in deep-sea Pleistocene deposits. C. pygmaea is peu plus ventrue" than P, caatus. The name was never a Miocene species, recently renamed C. interita by Lin- used by Monterosato in published works, bur was intro- den & Eikenboom (1992). duced by Brugnone (1877) îor the fossil species from Fi- C. rnicroscalaria can be easily distinguished from carazzi (Palermo), listed by Monterosato (1877) as "Cyli- C. stefanisi, being shorter, thicker, roughly ribbed, and cltna wata Jeffreys". in lacking spiral ribs. P ooatus (Warén, 1980, pl. 6, Íig. 14; Bouchet, Distribution. C. microscalarh is known only from 1975, pl. 2, fig. d) differs from P. obesiusculus, being'. Pleistocene deep-sea deposits of southern ltaly, where it more slender and less pyriform in shape. The aperture is is rather common. also narrower in its upper parr than in P watul The species reported as Cylichna waw Jeffreys by \flatson (1886, p. 664, 49, Chrysall ida brattstroemi \X/arén, 199 | pI. fig.9) from some Atlantic stations seemingly fits the description of P. obesiusculus. 7997 Cbrysallida brattstroem.i !/arén, p. 100, fig. 32a-c,33d. Bonfitto et al. (1994) illustrated a markedly ovate 7994 Cbrysallida brattstroemi - Palazzi 6c Villari, Íig. 11, 12. Pyrunculus from a Late Glacial Mediterranean assemblage, dubiously regarding it as P obesiwsculus. The speci- Distribution. This is the most frequent Chrysallida men actually falls within the variability range of P. obe- in bathyal Pleistocene deposits Qalazzí &. Yillari, 1994; siusculus. Di Geronimo et al., in press). Distribution. P obesiusculus is common in Pleistoce- It is known presently from western Norway ne bathyal assemblages. Its "Recent" Mediterranean fin- (found living in 200-270 m; \farén, 1991). Empty shells dings are probably based on Late Glacial shells. are known from deep Mediterranean waters fVarén, 1991; Bonfitto et a1., 1994), probably originating from Class Bivalvia Linnè, 1258 Late Glacial beds. Subclass P roto b ra n c h i a Pelseneer, 1889 Nucw. Subclass O p i st o b ra n c h i a Milne-Edwards, 1848 Family lidae Gray, 1824 Family Retusidae Thiele, 1931 Ennucula corbuloides (Seguenza, 1877) comb. nov.

Pl. s, fig. 1.-7, Is, 16 Pyrunculus obesiusculus (Brugnone, 1,877)

Pl. 2, Íis. 3 !877a Nucula corbuloid.es Seguenza, p. 92. 1877b Nucula corbuloides - Seguenza, p. 9, p1. 1, fig. 3, 3a, 3b. tó/ / LyucDnd ooestuscurd brugnone, p. J7. pl. I, tlB. / 1986 lcionucula corbuloides - Di Geronimo & Bellagarnba, pl. 3, fig. 1994 Pyrunculus sp. - Bonfitto et a1., p. 148, fig.25. 8.9. 1996 Cylicbna obesiuscula - Pùrzzi & Villari, p. 262, Íig. 95. 1992 Nuculoidea bushae (pmrtim ?) - Rhind & Al1en, p.73, fig.9-rc. 1996 Nuculoma corbuloides - Ptlazzi & Villari, fig. 140. Remarks. As recently noted by Palazzi & Villari (1996), the Pyrunculrzs species occurring in deep-sea Plei- Description. Small, thin, somewhat inflared, ovate- stocene beds is different from P. watus Seffreys, 18ZO), elongate, inequilateral shell. Umbo orthogyrate, large, ri- and should be referred to as ? obesiusculus (Brugnone, sing well above the dorsal margin. Iil-defined lunuie and 1877). This attribution was confirmed by Jeffreys escutcheon. Moderately truncate posterior margin. Sur- himself, to whom Brugnone sent his material (Brugno- face with faint concentric ridges. Chevron-shaped teeth, ne, 1.877). numbering 4-5 anteriorly and 5-6 posteriorly in 3 mm Bouchet (1975) reported the presence of a speci- long valves. Ovate, weakly impressed muscle scars. men of "Cylicbna obesiuscula Monterosato", labelled so Smooth inner margin. Vell-developed condrophore, ver-

PLATE 5 Fig. l-7,15,76-Ennucula corbuloid.es (Seguenza, 1877). l. Lazzàlo, 3.5 mm. 2. Lazzàro,3.4 mm. 3. Lazzàro, 3.4 mm. 4. Ibero-Maroccan Gulf (BALGIM Exp., St. D\161,35"31N 7'26V1, 1,222 m, MNHI$, 2.9 mm.5.Lazzà,ro,2.5 mm. 6,T.Tyrrhenian Sea (St. BS 78-14, NE Sardinia, 1,7A7-1,293 m),3.2 mm. 15. Tyrrhenian Sea (EOCUMM95, St. 16, Eolian Archipelago, 1,340 m),2.0 mm. 16. Lazzà,ro,2.1mm. Fig.9-14 - Ennucula rotunddtd. (Seguenza, 1877). 9. Vallone Catrica, 2.2 mm. 10. Lazzà.ro, 1.8 mm. Il, 12. La,zzà,ro, 1.9 mm. 13, 14. Lazz\ro, 1.9 mm. Fig. 8 - Ennucula aegeuris (Forbes, 1844). Tyrrhenian Sea (EOCUMM95, St. 32, Eolian Archipelago, 248 m), 3.2 mm. Pl. 5 Batlryal Molluscan from Southern ltaly 445 446 I. Di Geronimo & R. La Perna

tically projecting from the hinge-plate. Maximum ante- finding them closely resembling their Pleistocene and ro-posterior length ca. 4 mm. Roundish prodissoconch, Recent Mediterranean counterparts, 270-310 pm in length. Young specimens of E. aegeensz.s (Forbes, 1844) (P1. Taxonomy. Three genera, namely Nuculoma Cos- 5, fig. 8), a common species in outer-shelf and upper- smann, 1907, Leionucula, Quenstedt, 1930 and Ennucula siope MediterraÍeafl stations, may be misidentified as E Iredale, 193I, were used for the Recent ovate, smooth- corbuloides. In individuals of the same size. the valves of margined nuculids. Allen 6c Hannah (1986) regarded E. aegeensis are less inflated, thicker and more oblique Nuculoma as the only valid genus, Maxwell (1988) trea- than in E. corbuloides. Furthermore, the resilifer in E ted Ennucula as the suitable genus (see also Schenck, aegeensis is smaller and oblique, becoming almost verti- 1.934, 1939), whereas Bergmans (199L) regarded Ennucu- cal only in large valves (up to Z-8 mm). Young speci- la as a senior synonym of l-eionucula. Rhind & Allen mens of E. corbuloide.r are somewhat rounded in shape (1992) referred some deep-sea smooth-margined nuculids and might be mistaken {or Ennucula rotundda (Seguen- with orthogyrate umbo and a vertical resilifer to Nucu- za, 1877) (see below). loidea Ylllliam & Breger, t9I6, and used Nuculoma for Distribution. E. corbuloides is one of the most fre- opisthogyral species with an oblique resilifer. ìfe agree quent deep-sea nuculids in Pleistocene deposits (La Per- with Maxwell (1988) in considering the type-species of na, 1994; Di Geronimo et a1., in press). It is known Nuculoma (|urassic) too different from the modern from Mediterranean stations deeper than 500 m (Di Ge- smooth-margined nuculids. The type-species of Nuculoi- ronimo et al., 1995; La Perna, unpubl. data). dea (Late Paleozoic) is also quite different, whereas the The BALGIM stations in the Ibero-Moroccan type-species of Leionucula (Cretaceus) appear rather clo- Gulf, where E. corbuloides occurs, range from 540 to se to îhe modern species. Since protobranchs, and nucu- 2,000 m. lids in particular, are markedly conservative (Allen, 1978), strong evolutionary restraints in shell morpholo- Ennucula rotundata (Seguenza, 1877) comb. nov. gy to be expected. The lack of a radial structure in ^re Pl. s, fig. e-14 the outer shell layer, accounting for the smooth-margin (Van De Poel, 1955), probably evolved several times (e.g. 7877b Nucula corbuloides var. rotundata Seguenza, p. 9, pl. 1, fig. 3c, polyphyletically through paedomorphosis; see Gofas & Salas, 1996). On these grounds, Ennucula is much more suitable for the Cenozoic species, than a Mesozoic or Description. Small, thin, somewhat inflated, ovate- triangular, inequilateral shell. Opisthogyrate ri- ^1.1"" o".r," umbo, Remarks. Among the Atlantic deep-sea nuculids, sing moderately above the dorsal margin. Ill-defined lu- E. corbuloides'was recently reported by Rhind & Allen nule and escutcheon. Moderately tnrncate posterior (1992) as Nuculoidea bushae (Dollfuss, 1898) t: Nucula margin. Surface with faint concentric ridges. Roughly subwata Verill & Bush. 1898 : Nucula bushi nom. nov. chevron-shaped teeth, numbering 7-8 aîteriorly and 5 Doilfuss, 1898 : Nucula busbae emend. Schenck, 19391. posteriorly in 2 mm long valves. Roundish-ovare, im- E. bushae, however, is most probably different from E. pressed muscle scars. Inner margin smooth. Short con- corbuloides. The drawings of Nucula subwau by Verill drophore, faintly projecting from the hinge-plate. Maxi- & Bush (1898, p. 852, pL 81, fig. 8, pl. 83, fig. 5) show a mum antero-posterior length ca. 2.2 mm. Roundish pro- nuculid with a thicker and longer hinge, and a marke- dissoconch, ca. 160 pm in length. dly lower umbo, compared with E corbuloides. \lhether Remarks. E. rotundata was described by Seguenza all the material examined by Rhind & Allen (1992) as a "shorter and more rounded" variety oÍ Nucula cor- from many eastern and western North Atlantic stations buloides.It was also well illustrated. belongs to Verrill & Bush's or Seguenza's species is It is not understood whether E. granulosa (Verrill, unknown. Anyway, E. corbuloides does occurs in the 1884) (see Rhind & Allen, 1992 as Nuculoma granulosa) North Atlantic, and it is probably widespread is a synonym of E. rotundata, although some differences throughout the European basin, as reported by Jeffreys may point to two distinct species. Judging from the dra- (1879). \fie examined some Atlantic specimens (ibero- wings and description by Rhind & Allen (1992), the Moroccan Gulf, BALGIM exp., MNHN; Pl. 5, Fig. 4), North Atlantic E. granwlosa differs from E. rotundata ín

PLATE 6 Fig. 1-4,9-11-Tbestyled.a cuspidata (Philippi, 1844). 1, 2,Lazzàro,8.1 mm. 3.Lazzàro,5.i mm. 4.Lnzàro,3.2 mm. 9. Furnari, hinge detail from a juvenile (3.5 mm), scale bar:200 pm. 10. Prodissoconch, scale bar : 50 pm. 11. Prodissoconch detail, scale bar:20 Fm. Fig. 5, 6 - Ledella messanensis (Jeffreys, 1879). Vallone Catrica, 4.4 mm. Fig. 7, 8 - Ledella nicotrae (Seguenza, 7877). Gnmmtchele (SE Sicily, early Pleistocene), 3.7 mm. PI. 6 Batbyal Molluscan from Soutbern ltaly 447 408 I. Di Geronimo & R. La Perna

having a rounded to subquadrate outline, whereas E. ro- oblique resilifer. Unequal adductor scars, the posterior tundata is ovate-triangular, as well as a little more equila- much smaller and elongate, both close to the hinge teral than E. granulosa. Anterior teeth are more nume- ends. Indistinct pallial scar. Ovate, net-like sculptured rous (up m 8) and more close-set in E. rotundata than in prodissoconch, ca. 230 m in length,. maximum shell E. granulosa (up to 6). Iength 10 mm. Both E granulosa and E. rotundata resemble E Taxonomy. The type species of Tbestyleda lredale, corticata (lvloller, 1842) from Greenland (see Schiotte Ec 1929 is leda ramsayl Smith, 1885, a deep-sea species Varén, 1992). from New South 'ù7ales. Thestyleda was regarded as a E. rotundaa can be easily distinguished from juve- subgenus (e.g. Clarke, 1961; Okutanì, 1962) or as a nile stages of E. corbuloid.es, being markedly opisthogy- synonym (Allen Ec Flannah, 1986) of Nuculana Link, ral and sub-triangular in shape. Anterior teeth are smal- 1807, but it is worth keeping these long-shelled nucula- ler and more closely set in E rotundata. nids with a slender and bent rostrum, distinct from Nw- The fossil "Nucula tenuis var. subrotunda" of Bru- culana. The partly lamellar and overlapping teeth (Pl. 6, gnone (1877, pl. t, fig.3; as var. rotundata in the text, p. Fig. 9) are also distinctive o{ Tltestyleda, and are comple- 20) is most probably E. aegeensis, whereas "Lionucula tely chevron-shaped in Nuculana. Flowever, the shape corbwloides rotundaa (Seguenza)" of Nordsieck (1971, of Thestyleda juveniles closely resembles Nuculana (P1.6, fig. Z) is E. corbuloides. Fig. a) and the net-like prodissoconch sculpture is simi- Distribution. E. rotundaa is known only from lar in the two species (Pl. 6, Fig. 10, 11; see N. rninua La southern Italy deep-sea Pleistocene deposits. It often oc- in Ockelmann &'Warén, in press, and comments by Perna, press). curs together with E corbuloides, as reported also by Se- in gtreîza (1877b). Thestyleda shows a world-wide deep-sea distribution, whereas Nucwlana has a mainly boreal shelf distribution. Several deep-sea species may be referred to The- Family N u c u I a n i d a e H.& A. Adams, 1858 styleda, such as Leda cordyla Dall, 1908, L. longicaudata Thiele, 1912, Nuculana scalata Prashad, 1932, N. louiseae Thestyleda cuspidata (Philippi, 1844) comb. nov. Clarke, 1961, N. subscalata Okutani, t962. A marked fig. r-4' 9-1r Pl. 6, cold-water adaptation of. Tbestyleda is obvious in the Antartic T. longicaudarz (see Dell, 1990 as Propeleda lon- 1844 Nucula cuspilata Philippi, p. 47, pl. 15, fig. 8. gicaudata). 7994 Nuculana cuspidata -Pa.lazzi & Villari, fig. 48,49. 1996 Nuculana (Costelloleda) cuspiddtd' - Palazzi' 6c Villari, fig. 134, Propeleda lredale, 1924 was used also for these 135, 14r. long-shelled nuculanids. Flowever, both Propeleda and the related Poroleda FIutton, 1893 (see Maxwell, 1988) Description. Moderately large, convex, elongate, differ markedly from Thestyleda in having smoother, ovate shell, posteriorly extending in a long, anteriorly flatter and almost unkeeled shells. In addition, the ro- bent and blunt rostrum. Ventral margin regularly con- strum is more squat and iess bent than in Thestyleda. vex to slightly concave towards the rostrum tip. Concave The Mio-Pliocene " Icda" claoata (Calcara, 1841) and the postero-dorsal margin. Small, barely rising, opisthoryra' Plio-Pleistocene "l-eda" hoernesi Bellardi, 1875, seem in- te umbo. Sculpture made of widely spaced concentric stead to pertain to Propeleda. ridges, anteroventrally obsolete, and two keellike rostral Distribution. Originally reported from the Lamato ridges, long and flat postero-dorsal area. Concentric rid- River valley (Calabria), T cuspidaa was reported by Se- ges turn at right angles to the rostrum, becoming sli- guenza (1877b) from many southern Italy "Astian" Ioca- ghtly sinuous, stronger and doubled in number. Space lities. It is really one of the most common nuculoids in between rostral ridges barely concave. Internal mid-ro- Pleistocene bathyal deposits (La Perna, 1994; Palazzi &. strum ridge from umbonal area to posterior tip. Taxo- Villari, 1994, 1996; Di Geronimo et al., in press). dont, moderately thick hinge, with 13 anterior and 2l posterior teeth in 9 mm long valves. Chevron-shaped, close-set teeth, becoming somewhat lamellar and over- Ledella messanensis (feffreys, 1870) lapping in the proximal area. Deep, triangular, slightly Pl. 6, fig. s, 6

PLATE 7 Fig. l,2,7A-72-.Yoldia' minima (Seguenza, 1877). l.Lazzàro, 1.4 mm. 2.Lazzàro, 1.6 mm. l0.Lazzàro. Hinge detail, scale bar: 200 pm. 11. Lazzlro. Prodissoconch, scale bar:50 ttm. 12.Lazzàro. Prodissoconch detail, scale bar : 10 pm. Fig. 3, 4 - Microgloma turnidula $4onterosato, 1880)' 3. Bianco, 0.8 mm. 4. Bianco, 0.2 mm. Fig. 5, 6 - Yoldiella segumzae Bonfitto & Sabelli, 7995. Lazzàro, 3.9 mm. Ftg.7-9 - Bafiryspinula excisa (Philipp| 1844).7,8.La22àto,7.i mm.9. Furnari, hinge detail from a fully grown valve, scale bar : 500 pm' Pl.7 Bathyal MoLluscan from Southern ltaly 449 410 I. Di Geronimo & R. La Perna

1870 Leda acuminata Jeffreys, p. 69. precisely define the taxonomic status of this form, 1877b Leda (Junonia) acuminata - Seguenza, p. 15, pl. 3, fig. 15, 15a-c. which corresponds to L. nicotrae (Seguenza, 1877). L879 Leda tnessanensis Jeffreys, p. 576 (nom. nov.). 1978 Yoldiella acurninata - Warén, p. 215, fig. 10, ll. The origin of the European lrdella lineage dates 1978 Yoldiella messanmsis - Varén. p. 218, fig. 2a, 21. back to the late Miocene, with Z. perffinis (Seguenza, 1981 Ledella messanensis - Di Geronimo 6c Li Gioi, p1.2, {tg.5,6. 1877) and L, glabra (Laghi k Palazzi, t99t). L. serninu- 1986 Nuculana (Ledella) pusio - Laghi, pl. 3, fig. 1-7 (not fig. 5); pl. +, (Seguenza, rls. l-). lum 1,877) is a Pliocene bathyal species (re- ú8; Yotdiettd messanmsis - '$larén, p. 329, fig. 3e, lla,b. ported by Robba, 1981, pl. 11, fig. 2, 3 as Nuculana 1989 Izdella acurninata - Allen 6c Hannah, p. 153, fig. 57-60. messanensis). In Pleistocene bathyal deposits anorher spe- 1994 Nuculana rnessanensis - Palazzi 8c Villari, fig. 54, 56. cies, l. rectidorsata (Seguenza, t877) Pi Geronimo et al., in press) occurs, of which the northeast Atlantic Z. Taxonomy. l*della bushae'Warén, 1978 was propo- folini (Yarén, 1978) may be a synonym. sed by \flarén (1981) as type-species oÍ Ledella Verrill Ec Distribution. The Recent distribution of L. messa- Bush, 1892. Later, Allen & Flannan (1989) regarded L. nensis tanges from Norway south to the Azores and the bushae as a synonym of L. wltima (Smith, 1885). Mediterranean, at depths ranging between 200-2,000 m Among the species treated by Sl'arén (1978, L989) (\farén, 1989; Allen 6c FIannah, 1989). It is one of the under the genus Yoldiella, "Yoldiella" messanensis stands most common nuculoids in Mediterranean Pleistocene out as a notable exception, as it has a moderately thick to Recent bathyal faunas. shell, a sharp rostrum and a well-defined subrostral sinus. Although this species may appear somewhat different from the thick, short and bluntly rostrate kdella Yoldiella seguenzae Bonfitto & Sabelli, 1995 bushae or L. ultima, it is also quite different f.rom Yol- Pl. 7, Íig. 5, 6 diella lucida (Lovén, 1846), the type-species of Yoldiella 1877b Yoldia abyssicola - Seguenza, p. 20, pl. 5, fig.28. Verrill & Bush, 1897. The rostrum and subrostral sinus \994 Yold.iella producta -Palazzi E Villari, p. 99, fig. 58,59,66. are good taxonomic features to distinguish lcdella from 1995 Yoldiella seguenzde Bonfitto & Sabelli, p. 24, fig. 5, 6, 7 , 9, 10, 11. Yoldiella, as also noted by Verrill Er Bush (1898). In Ze- della, the rostrum may be more or less sharp, elongate Taxonomy. Yoldiella seguenzae, as well as the shal- or weakly keeled, and always enhanced or outlined by a lower Y philippiana (Nyst, 1845), the Atlantic Y lenticu- sinus. The rostrum tip is typically turned down in lr- la @Ióller, 1842), Y propinqua (Leche, 1878), the Arctic della and slightly upturned or straight in Yoldiella.Fu,rr.- Y tarnara (Gorbunov, 1946) (all treated by 'Warén, hermore, Yoldiella has thinner, flatter and smoother val- 1989), and the Tertiary Íossrl Y pygmaea (Minster,1837) ves, with a shorter and more delicate hinge. Allen & (see Janssen, 1979) belong to a homogeneous group Hannah (1989), Kilburn (1.994) and Allen et al. (1995) which differs from the Yoldiella type-species and allied recently followed the same taxonomic approach fior Le- species in having thicker and more convex valves with a della and Yoldiella. blunt and "low" rostrum. The hinge is also longer and Remarks. Jeffreys (1879) replaced Leda dcunltndta relatively thicker. Allen et aL (1995) treated this group in with Seguenza's m.s. messanensis (feffreys, 1870), belie- part as Portlandia Mòrch, 1857, bur the type-species P ving that Nucula acuminata Von Buch, 1833 and N. acu- arcticd (Gray, 7824) is quite distinct from these species. rninata Eichwald, 1.853 were to be placed in Leda. Remarks. Seguenza (1,877b) reported this species as

\flarén (1978) kept the Recent acumina.ta distinct Yoldia abyssicola Torell, listing " Leda producu Montero- from the fosstl messanensis,bvf later (\larén, 1989) refer- sato" and "Yoldia striolata Brugnone" as synonyms. Z. red both the fossil and Recent material Ío messanensis. producta, first published as a synonym (Monrerosato, Actually, L. messanensls is markedly variable in outline, 1825), was never used as an available name and the com- thickness and sculpture, as reported by Varén (1978, bination proposed by Palazzi & Villari (1994), t.e. "Yol- 1989). The Pleistocene specimens parallel Recent exam- diella producta (Seguenza, 1877)", is unjustified. ples in shell variability. Nevertheless, the Pleistocene up- Distribution. Y seguenzae is known from the Medi- per-slope specimens differ notably from those found terranean and northeast Atlantic in 500-2,000 m (Bonfit- deeper, being shorter, thinner, more convex and less ro- to & Sabelli, 1995; La Perna, unpubl. data). It is also strate (Pl. 6, Fig. 7, 8) than the typical L. messanensrs. one.of the most abundant bathyal nuculoids in Pleisto- Additional material is currently under study to more cene deposits (Di Geronimo et a1., in press).

PLAIE 8 Fig. 1-10 - Katadesmia confusa (Seg.uenzt, 1877). 1, 2. Archi, 5.4 mm, 3,4. 5.1 mm. 5. Furnari, 5.1 mm (postero-dorsal margin partly broken : Yoldia confusa var. major Seguenza, 1877). 6. Furnari, 4.6 mm. 7. Lazz'aro,5,2 mm. 8, 9. Archi, juvenile, 2.5 mm (:Neilo pbaseolinus Seguenza, 1877). rc.Lazzàro, hinge detail from a fully grown va1ve, scale bar : 500 pm. Pl. 8 Batlryal Molluscan from Southern ltaly 412 I. Di Geronimo & R. La Perna

Bathyspinula excisa (Philippi, 18aa) Desciiption. Small, rounded-ovate, rather convex,

Pl.7, fig.7, 8, 9 moderately thin shell. Neither carination, nor rostra- tion. \fleakly prominent, orthogyrate, anterior to midli- 7844 Nucala excisa Phil'ippi, p. 46, pl. 15, ftg. 4. ne umbo. Surface with faint concentric striae and lowly 7989 Spinula excisa - \larén, fig. 1.2d, e. stepped growth increments. Moderately thin, taxodont 1993 Spinula excisa - Tabanelli, fig. 5. 1994 Spinula excisa -Palazzi & Villari, fig.57,68. hinge, with short, weakly chevron-shaped teeth, numbe- 1996 Baúryspinula excisa - Di Geronimo. & La Perna, p. 108, pl. 1, fig. ring 8 anteriorly and ll posteriorly in 2 mm valves. 1-10; pl. 2, fig. 1-3. Triangular, deep ligament pit, located beneath the beak. Roundish adductor scars, equal in size. Indistinct pallial Taxonomy. Because of the preoccupied status of scar. Ovate, wrinkled prodissoconch, ca. 150 pm large. Spinula Dall, 1908, Filatova & Schileyko (1984) replaced Maximum shell length 2.5 mm. it with Batlryspinula Filatova, 1958, originally proposed Taxonomy. A conservative allocation in "Yoldia" rs as a subgenus of Spinula. They also proposed Acutispinu- maintained, as it is still difficult to find a suitable syste- la as a replacement name for the subgenertc rank of Ba- matic position for this species, both at the genus and tbyspinula calcar DaIl, 1908, i.e, the type-species of Spl- family level. ")1" rninima shows a weak shell-shape re- nula. The present species belongs to the subgenus Bdtlry- semblance to Sarepta A. Adams, 1860 and Psuedoglo- spinula, like all the Recent Atlantic species, although Ba- to mus DaIl, 1898, whose type-species were tbyspinula and Acutispinula probably deserve a full gene- recently illu- ric status. strated by Ockelamnn & Warén (in press), Sarepta re- In the systematic framework by Allen and co-wor- sembles Yoldiella morphologically fVarén, pers. 'whereas kers, Bathyspinwla is referred to the subfamily Spinulinae comm.), the hinge of Pseudoglomus shows affí- Alien Ec Sanders, 1982 (now Acutispinulinae), within nities to malletids, tindariids and neilonellids. the family Nuculanidae. Maxweli (1988) maintained this An unusual wrinkled prodissoconch sculpture oc- allocation. Filatova & Schileyko (1984) raised Ledellinae curs in this species. Allen & Sanders, 1982 to a f.amily rank, referring Batlry- Distribution. "Y" minirna is common in Pleistoce- spinwla to it. According to a recent classification scheme ne bathyal deposits (La Perna, 1994;Di Geronimo er a1., of nuculoids by Ockelmann & IX/arén (in press), Batlry- in press). It is also found as old shells from Mediterrra- spinwla is allocated within an undivided Nuculanidae fa- nean deep-sea stations (Di Geronimo & Li Gioi, 1981; mily. La Perna, unpubl. data). According to Jeffreys (1879), ir The ligament features of Batlryspinwla are similar also occurs in the Atlantic. to those of the malletiids. A mainly external ligament juveniles occurs in mature specimens, whereas show a Family Malletiidae H. &A.Adams, 1858 well-developed inner pit (Di Geronimo & La Perna, 1,996: pl. 1). Both in Batlryspinula (P1.6, Fig. 9) and in malletiids (P1. 7, Fig. 10) adult specimens show a central Katadesmia confusa (Seguenza, 1877) comb. nov. hinge area resembling a well-developed resilifer. Actual- Pl. 8, fig. 1-10 ly, it consists of an edentulous gap, in which the ligament holds only a small portion close to the beak. L877a Yoldia confusa Seguenza, p.96. 1877a Neilo pbaseoltnus Seguenza, p. 96. Remarks. Historical review, description and distri- 1.877b Yoldia pelLucid.a - Seguenza, p. 19, pl. 4, frg. 25, 25a, bution of B. excisa were reported and discussed by Di 1877b Yoldia confusa - Seguenza, p. 20, pl. 4, Írg. 24, 24a, 24b, 24c. Geronimo & La Perna ft996\. 1,877b Neilo phaseolinus - Seguenza, p. 23, pl. 5, fig. 31,31a,3lb.

Description. Elongate-oval, inequilateral, inflated, "Yoldia" minima (Seguenza, t877) moderateiy thin shell. Posterior margin extending in a PL.7, fig. 1., 2, 1.0-12 blunt rostrum. Orthogyrate umbo. Faint dorsal keel, running from umbo to rostrum tip. Glossy valve surfa- 1.877a Yoldia rninirna Seguenza, 96. ce, with lines and concentric close 1877b Yoldia minima - Seguenza, p. 18, pl. 5, fìg.27, 27 a,b. faint grovth ridges 7987 Yoldia minima - Di Geronimo & Li Gioi, pl.2, fig. 1,2. to the ventral margin. Narrow hinge-plate. Chevron-sha-

PLAIE 9 Fig. 1-6, ll- Austrotindariapusio (Philippi, 1877). L Bianco,5.1 mm,2. Furnari,4.3 mm.3,4. Bianco,3.9 mm.5. Furnari,4.1 mm.6. Bianco, .juvenile, 1.3 mm. 11. Bianco, hinge detail from a fully grown valve. Fig.7-9 - Austrotindaria salicensis (Seguenza, 1877).7. Sa1ìce (Messina, early Pleistocene), 4.1 mm. 8, 9. Rometta (\4essina, middle Pliocene), 4.5 mm, Fig. 10 - Austrotind.aria latior fiefÍreys, 1876). Tyrrhenian Sea (St. BS 78-14, NE Sardinia, 1,707-1,293 m), 4.8 mm. Bathyal Molluscan from Soutbern ltaly 413 I. Di Geronimo & R. La Perna

ped teeth, 9-10 anterior and l4-I5 posterior in 5 mm rently based on a large valve with the postero-dorsal long valves. Outer anterior and posterior ligament fur- margin partly broken (P1. 8, Fig. 5). Seguenza also de- rows, the former short and ill-defined, the latter well-described Neilo phaseolinus, a smail nuculoid which proved fined, as long as tooth series. Shallow and elongate liga- to be a juvenile stage of K. confusa (Pl. 8, Fig. 8,9). ment pit beneath the beak. Oval muscle scars, the ante- Laghi (1986, pl. 3, fig. 5) illustrated a USNM spe- rior one a little larger. Deep pallial sinus. Antero-dorsal cimen of K, confusa from "Pliocene, Messina" as Nucula- U-shaped scar. Posterior region marked internally by an na (kdella) pusio (Philippi, 1844) (:bdella messanensis, abrupt decrease in convexity. Ovate, weakly rostrate ju- see below), among other îrue rnessanensis. Palazzí & Vil- veniles. Maximum antero-posterior length 6.5 mm. larí (1994, fig. 53) also reported a valve of K. confusa as Bean-shaped prodissoconch, 240-260 pm in length. "Nuculana messanensis" from the Pleistocene of Messina. Taxonomy. Bouchet & \flarén (1979) and \Varén They also illustrated " Neilo phaseolinus" . (1989) distinguished three malletiid genera, i.e. Malletia Three Atlantic species (see Sanders & Allen, 1985) Desmoulins, 1832, Pseudornalletia Fischer, 1886 and Ka- are markedly close to K. confusa, i.e. K. pallida (Smith, tadesrnia Dall, 1908, whereas Sanders Ec Allen (1985) 1885), K. polita (Yerrill Ec Bush, 1898) and K. malita treated the latter two as synonyms of Malletia. In Kata- (Sanders Er Allen, 1985). The cosmopolitan K. cuneata desmia the shell is elongate and posteriorly pointed, (|effreys, 1826) is the only Kaudesmia presently occur- whereas it is ovate and more equilateral in the southern ring in the Mediterranean. It more closely resembles the and shallower Malletia, and subrectangular in Pseudomal- type-species K. oincula (Dall, 1908) than the coffisa- letia. Katadesmia typically comprises bathyal and abyssal grouP. species. Features useful in distinguishing K. confusa from The U-shaped anterior scar is a peculiar and disre- the frequently co-occurring lcdella messanensis, include garded malletiid feature, as noted by Knudsen (1970) rn the shell outline (no subrostral sinus in Katadesmia), the "NucttldnA" pallida (Smith) (:K. pallida). This shell fea- slightly upturned rostrum tip and the markedly inflated ture appears to be related to soft-body anatomy, as the sheli. hind-gut, making an anterior loop, does not penetrate Distribution. Seguenza (1.877a,b, t879) reported this the mantle (Sanders & Allen, 1985). species from several "Astian" deep-sea sites near Messina Marked growth-related shape changes, iike those and southern Calabria. It is one of the most frequent observed in K. confusa, were reported by Sanders & Al- nuculoids among the Pleistocene bathyal faunas. It was len (1985) for several malletiids. recently reported from the Archi section (southern Ca- Remarks. Briefly introduced by Seguenza (1877a), labria) (Di Geronimo et a1., in press), where some of it was soon after reported as Yoldia pellucida (Philippi) the specimens illustrated here were collected. and Yoldia confusa (Seguenza 1,877b). Philippi's drawing (18aa) of Nucula pellucida shows an ovare nuculoid, an- Family N e i I o n e I I i d a e Shileyko, 1989 teriorly and posteriorly rounded (it is probably synonym of "Yold.ia" longa Bellardi, 1.875). Surprisingly, Se- gveîza stressed a close resemblance between his speci- Austrotindaria pusio (Philippi, 1844) comb. nov. mens and Philippi's species. Seguenza illustrated both L Pl. 9, fig. 1-6, 11 pellucida and Y confusa as very similar to each other, the latter showing a small ligament pit. Subsequently, Se- 1844 Nucula pusro Philippi, p. 47, p|.15, fig 5. guerrza (1879) resolved the pvzzle, reporting his species !986 Psudoneilonella salicensis - Laghi, p. 191, p1. 5, fig. la, b, 3a,b, 4, 5a,b, 6a,b,c,7a,b,c. as "Y confusa Seguenza : L. pellucida Seguenza (non lù/aftn, 7989 Neilonella pusio - p. 252, frg. 76e, f . Nucwla Phil.), )a lucida Segtenza (non Lovén)'. The last 1993 Neilonella pusio pasio - Tabanelli, fig. 10. species, however, was only in part misidentified, as it 1994 Neilonella pusio -Palazzr & Villari, îig.60,61,63,67. was iilustrated correctly (Seguenza, 1,877b, pl. 5, fíg. 26). The form reported by Seguenza as Y confusa var. Description. Triangular-ovate, inequiiateral, rather major, showing a slightiy upturned rostrum, s/as appa- thick sh,ell. Rounded anterior side, posterior side obtu-

PLATE 10

Fig. 1-6, la- Cad.ulus wulurn (Phtlippi, 1844). 1. Bianco, 3,2mm,lateral viev 2,3. Furnari,3.0 mm, lateral (2) and dorsal (3) view. 4. Sa1ìce,3.O mm, lateral view (this form may correspond to C. salicensis Seguenza, 1879). 5. Lazzàro,3.3 mm, lateral view. 6. Vallone Catnca, 3.0 mm, lateral view. 10. Furnari, apical detail. Fig.7-9,11.- Cadulus d.ttmudtus Monterosato, 1875.7,8. Bovalino, 3.3 mm, lateral (7) and dorsal (8) view. 9. Grammichele,2.g mm,lateral vrew. 11. Bovalino, apical detail. Ftg. t2-1,4 - "Gadila" jrffr"yti (ì\4onterosato, 1875).12, 13.Lazzàro,4.1 mm lateral (12) and dorsal (13) view. 14. Furnari, 4.5 mm, lateral view. Fig. 15, 16 - Cadulus diploconus Seguenza, 1876. Bianco, 3.0 mm, lateral (15) and dorsal (16) view. Pl. 10 Batlryal Molluscan frùfti Southern ltaly 415 416 I. Di Geronimo & R. La Perna

sely angulate. Regularly convex ventral margin, poste- ria and "Neilonella" species reported by Knudsen (1979). rior margin slightly convex. Orthogyrate umbo, ante- In A. pusio, the ligament pit is well-developed in juveni- rior of midline. Sculpture of fine concentric ridges, of- les (P1. 8, Fig. 6), becoming more and more reduced in ten becoming obsolete postero-dorsally. Veakly defined adults (Pl. 9, Fig. 4, 5) to the point of being quite over- lunule, escutcheon absent. Roughly ovate muscle scars, grown by teeth (Pl. 8, Fig. 11). the anterior scar a little larger. Pallial sinus well-incised Remarks. Philippi (1844) described Nwcwla pusio aligned with posterior angulation. Moderately large from Bianco, virtually the "same locality" of the present hinge-piate. Chevron-shaped teeth, numbering 1O-11 an- work. teriorly, 12-13 posteriorly in 4 mm long valves. Liga- According to Laghi (1986), a wrong interpretation ment pit well-defined in juveniles, ill-defined to absent of N. pwsio Philippi originated from Seguenza (1877b), in adults. Short ligament furro% extending from beak who misidentífied l,eda (Saturnia) pu.slo when describing as far as fifth-sixth posterior tooth. Maximum shell "rLar salicensls". Laghi therefore regarded N. pusto as cor- iength ca. 5 mm. Bean-shaped prodissoconch, 170-210 responding to the species commonly known as l-edella i- ì---tL "- or Yoldíella rnessanesis (see above), and ranked "var. sali- Taxonomy. In gross shell morphology, this species censis" as Pseudoneilonella salicensi.s. F{owever, Philippi resembles Neilonella corpulenu (Dall, 1881), type-species described in detail its species, which do not show any of Neilonella Dall, 1881, from the Caribbean area. Both definite resemblance to Ledella messanensis (see also Wa- have ovate and concentrically ridged shells, with a main- rén,1989). ly external ligament, which is amphidetic in corpulenu Yar. salicensi.s has had a notable success since its and opisthodetic in pusio. Thís feature was well descri- description. Jeffreys (1879) referred to it as Leda pusio bed by Dall (1881, 1886) and is also obvious from the var. latior. Locard (1889) reported both Leda pusio l: holotype illustrated by Laghi (1986). Noting this diffe- Yoldiella semistriata (feffreys, 1579)l and lcda salicensis rence, Laghi (1986) proposed that Pseudoneilonella f: Austrotindaria latior (|effreys, t876) : A. striolaa should replace the preoccupied Saturnia Seguenza, L877. (Brugnone, 1876) sensu Warén, 1989] from the Atlantic. ju- F{owever, Pseudoneilonella Laghi, 1986 appears as a Laghi (1986) treated tt as Psewdoneilonella salicensis, whe- nior synonym of Austrotindaria Fleming, 1948. The reas \Warén (1989) put salicensis in synonymy with Leda same difference also led Maxwell (1988) to question the striolata Brugnone, 1876 and with Leda pusio var. latior synonymy between Austrotindaria and Neilonella repor- Jeffreys, 1876. Beîore attempting to give any value to ted in the literature (e.g. Allen 6c Hannah, 1986). var. salicensis. it should be stated that the Pleistocene The type-species of Austrotindaria is A. utrighti material of A. pusio examined closely resembles the Fleming, 1948 (Recent, New Zealand). Notable differen- typical form illustrated by Philippi, except for some ces, in addition to ligament characteristics, occur be- shallower (upper-slope) populations, which may be l-eda tween Neilonella and Austrotindarla, as stressed by Fle- striolaa Brugnone, 1876 (see below). Yar. salicensis was ming (19a8). Neilonella has a slightly concave, postero- said to be thicker, narrower, more convex and with dorsal side, resulting in a slightly upturned "rostrum", more gibbous umbos, than the typical form. 'We exami- which is slightly downturned in Austrotindaria, due to ned a few specimens closely resembling var. salicensis, the convex postero-dorsal side. The hinge is more equila- from Salìce (typeJocality, early Pleistocene) (Pl. 8, fig. teral in Neilonella than in Austrotindarla. "No defined 7) and Rometta (middle Pliocene) (Pl. 8, fig. 8, 9). It lunule or escutcheon" are present rn Austrotindaria, should be noted that typical valves of A. pusio also oc- whereas both are well-defined in N. corpulenta. cur at Salìce. The early Pliocene valves illustrated by Since neilonellids and tindariids show strong simi- Robba (1981) also seem to agree with salicensis. These larities in shell morphology, one may suspect that Au- observations support the hypothesis that two distinct s*otindaria is a tindariid. Tindariids differ mainly from species exist, one Pliocene, ranging up to the early Plei- neilonellids by lacking siphons (and a pallial sinus) (San- stocene (A. salicensi), and one Pleistocene (A. pusio). ders & Allen,1977; 'ù7arén, 1989). No mention of a pal- Flowever, more Pliocene material still needs to be exa- lial sinus was given when Fleming (19a8) desutbed Au- mined. suotindarid. Flowever, judging from the drawings and Pleistocene material seemingly referable to lcda description ("Hinge-plate strong, wide at either end, in- striolaa Brugnone, 1876 (described from Pleistocene de- terrupted below the beaks by a gap occupied by the nar- posits near Palermo) is currently under study. Shells are row base of the ligament"), the hinge of A. arigbti is narrower, iess triangular and more regularly ridged than very similar to that of A. pusio. A gap or a shallow pit A. pusio. They are markedly different from A. salicensis, in the hinge is actually characteristic of neilonellids; the and also different in outline from A. latior ffeffreys, tindariid hinge is uninterrupted beneath the umbo (San- 1876). For these reasons A. latior is considered distinct ders 6c Allen, 1977; see also \Warén, 1989 andthe Tinda- f.rom L. striolau Brugnone. Bathyal Molluscan from Southern ltaly 417

Three species similar a A. pusio are known from ly, this group resembles neither Gadila s.s. nor Cadulus the east Atlantic. One of them is A. latior (|effrey$. k is s.s., although it is probably closer to Gadila than Cadu- known from the North Atlantic in the 200-3,100 m lus. depth range, and is also found rarely in Late Glacial Me- According to the most recent classification of diterranean assemblages (Pl. 8, Fig. 10). As noted by'ùla- Scaphopoda (Steiner, 1992; Scarabíno, 1,995), Cadulus is rén (1989), Pseudoneilonella montanaroe Laghi, 1986 is a allocated to the order Gadilida Starobogatov, 1974, fami- synonym of A. " striolatA" , í.e. of A. latior, but the small ly Gadilidae Stoliczka, 1.868, subfamily Gadilinae Sto- fossil valves from Sicily (Laghi, 1986, pl. 9, fig. 5a, b, 8a, liczka, 1868. b) belong to A. pusio. Another species is known as Nei- Remarks. A certain variability has been observed lonella guineensis (Thiele, 1931) from the east Atlantic among the material examined from Pleistocene deep-sea (4' N) to Southeast Asia, in 2,278-4,0t8 m (see Knud- deposits (several hundred shell$. Most of them are sen, 19ZO). The third species was described by Métivier strongly swollen (ength/diameter:1.6-L7) and with the (1982) as Nucwlana foresti {rom the Azores in 3,360 m. apical markedly narrower than the oral opening (Pl. 10, The original shaliow-water records (58 and 22 fíg. 1,-a). They closeiy resemble the type specimen from fms.) of A. urigbti are striking when compared with the Crotone (south Calabria), as originally illustrated. Some general distribution of Austrotindaria and the neilonel- specimens are more slender (ength/diameter:1.7-1.9), iids. with the apical and oral diameters more simiiar, or with Distribution. A. pusio is abundant in Pleistocene the "neck" weakly defined (Pl. 10, fig. 5, 6). The occur- deep-sea deposits. rence of intergrading specimens in the same samples The late Miocene report of Neilonella pusio by strongly suggests a single species. Sacco (1898) "fide Doderleini" was based on Ledella gla- A distinct species, so far misidentified as C. wu- bra (Laghi k. Palazzi, 1991) (see Laghi, 1986; Laghi 8r lwm, occurs in shallower (upper-slope) Pleistocene depo- Palazzi,l99t). sits from southern kaly (P1. 10, fig. 7-9). It resembles the slender form of C. ooulurn, except for a markedly Class S c a p h o p o d a Bronn, L862 barrel shape Qength/diameter up to 1.9) and an apical and oral opening that are roughly equivalent. It is also Family Gadilidae Stoliczka, 1868 generally smaller than C. wulum, with more obiique apical and oral openings and a more central swelling. Cadulus ovulum (Philippi, 18aa) The apical crown is often well-preserved, s.hereas it is Pl. 10, fig. 1-6, 10 frequently worn out in C. wulunt, probably because the lobes are more deeply incised than in C. wulum (P1. 10, 1.844 Dmtalium wulurnPhilippi, p. 208, p|.27, fig.21. fig. 10, 11). The sheiis illustrated by Di Geronimo 1979 Cadulus wulum - Caprotti, p. 244, pl. 14, fig. 5-8. (1979, pL.7, fig. 2a-c) and Pavia (1991, 9, 10) 1997 Cadulus ooulum - Pavia, pl. 9, fig. 6,9, fi. pl. fig. 6, 1994 Cadulus oaulum -Palazzi &- Vìllari, fig. 41. as C. wulum belong to this species. Pavia also reported a scatter-plot (fig. e) suggesting two distinct species. The Description. Globose, swollen, smooth, shiny name Cadulus attenudtus Monterosato, t875 should be shell. Slightly anterior maximum diameter. Roundish used for this species. C. wwlum var, attenudtd was repor- section. Ventral side much more prominent than the ted as a nolnen nudum by Monterosato (1872) from the dorsal side in lateral view, rapidly sloping to the apex Pleistocene beds of Ficarazzi (Palermo). Subsequently, and gently curving to the oral opening. Slightly ovate, he reported it as a synonym of C. cyathus (De Cristofori coronate apical opening, with inner callus. Oblique, Er Jan, 1832) (see below) and briefly described it ["più shortly constricted oral opening, larger than the apical piccolo del precedente (C. ouulum), nè così gibboso nel opening. Length up to 3.5 mm; length to maximum dia- mezzo della sua parte dorsale", Monterosato, 1875]. The meter ratio 1.6-1..9. Ficarazzi beds contain deep-sheif to upper-slope faunas, Taxonomy. This is the type-species oÎ Cadulus Phí- among which truly bathyal species are scarce, compared lippi, 1844. Two Cadulu.r groups can be distinguished: a with the faunas studied in the present work (La Perna, typical group, with globose shells and a crownJike api- in press). Similar faunas occur in the deposits where the cal opening, and a spindle-shaped group, with a simpie specimens illustrated here were collected (La Perna, apex. A third group consists of slender, more ore less t996). It is also worth noting that Seguenza (1879) re- bent, centrally swelling and thin-walled species, lacking ported "C. cyatbus : C. wwlumvar. dttenuatd Montero- the apical callus (e.g. jrffr"yti Monterosato, subfusiformis sato" from several southern Italy localities, keeping it Sars, sauridens'W'atson, sirnillimus'Watson). This group, distinct from C. wwlwm. including both shelf and deep-sea species, is treated C. gibbus Jeffreys, 1883 from Biscay Bay ("allied to either as Gadila or Cadulus by different authors. Actual- C. wulum of Philippi, but much smaller and not so 418 I. Di Geronimo & R. La Perna

oval, and the ends are equal in size"), appears to resem- Taxonomy. This species belongs to the spindle-sha- ble C. attenuatus. The Miocene C. taurwolum saccoiPa- ped, non-coronate Cadulus group. vía, 1991., is markedly barrel-shaped and less swollen Remarks. Palazzi & Villari (7994) recently repor- than C. dttenuatus. ted this species, tentatively regarding C. tumi^dosus Jef- Although accurately described resta mi- ["Creseis freys, 1883 as a synonym. Judging from the original dra- nima, cyathiformiinflata (alt. 2 mm, lat. 1 mm"], C. wing, C. (?) tumidosus is quite distinct from the presenr cldtbus is an enigmatic still species. Pinna (197I) illustra- species. On the confrary, two Indo-Pacific deep-sea spe- ted the original material, consisting of a badly broken cies described by Scarabino (1995) closely resemble C. shell from an unknown locality. Ir is probably not a diploconus, i.e. C. sofi.ae and C. labeyriei. C. rossoi descri Pleistocene fossil, like species all the in De Cristofori & bed by Nicklès (1979) from Senegal in 150-250 m is also Jan's coliection. It might be the same as C. wulum, as roughly similar to C. diploconus, but it lacks the inner supported by Caprotti (1,979); however, since the Mioce- apical callus, and thus probably belongs to the third ne and Pliocene deep-sea Cadwlus are still very poorly " Cadulus" group previously mentioned. known, it is not advisable to regard the well-described Distribution. C. diploconus is known only from and now well-known C. wulum as a junior synonym of Pleistocene bathyal deposits of Calabria and the Messina C. cyathus, involving also the type-species of Cadwlus area, where it is found rarely. itself. Distribution. Although C. wulum is listed among Discussion. the Mediterranean Mollusca checklist (Sabelli et al., 1990), tt should be regarded as absent from the Mediter- Two noteworthy faunal groups occur among the ranean, if not extinct. The Atlantic "finding" by Fi- faunas studied. The first group consisrs of North Atlan- scher, quoted by Jeffreys (1S83a) and locard (1898) may tic species, i.e. Fissurisepta rostrata, Puncturella noachina, be based on a particularly inflated specimen of C. gib- P granulata, Callumbonella suturale, Haloceras carinata, bus. The Mediterranean report by Acton (off Naples, see Pseudosetia turgida,'Torellia delicaa, Eumetula aliceae, Jeffreys, 1883a) and those by Caprotti (1979) and Gagli- Melanella spiri.díoni, Claoiscala richardí, Opaliopsis atlan- ni (1986), are likely based on reworked shells (or tis, Ipbitus tenuisculptus, Tàranis cf. borealis, Mitrolumna dredged from old sediments). The specimens examined smithi, Plewrotomella eurybrocha, and Pleurotomella pac- by Gaglini (1986) were said to be fossils or subfossil-loo- kardi. Seguenzia monoc i ngu I a a, Pyrun c u I u s obesiusculus, king and some to come from "a few metres". Bonfitto et Chrysallida brattstroemi, Ennucula rotundata and "Yol- al. (I99a) recently listed C. wulum from a deep-sea dia" minima may also be added. Except for other spe- Tyrrhenian assemblage in which several Pleistocene species mentioned in the present work, other Atlantic species were also found. cies from Pleistocene bathyal deposits (La Perna, 1994) According to Pavia (1991), Sacco's report (1897) of deserve to be mentioned, such as Micropilina minuta C. oaulum from the norchern Italy late Miocene may be Warén, 1989, "Diodora" tenuiclathrata (Seguenza, 1862) based on C. taurwolum, although this material was lost. (: Glypbis edzuardsi Dautzenberg & Fischer, 1896), Cal- C. ar.,ulum was also reported from Pliocene depo- liostonta maurolici (Seguenza, 1876) (: Gibbula obesula sits (Bernasconi, t996), although no reliable Pliocene Locard, 7898), 'Tiochus" luciae (Seguenza, 1876) (: Gib- shell has been illustrated so far. bula bannonis Locard, 1898), and Calliotropis ottoi (Phi- lippi, 1844). This list is not exhaustive, as several fossil species still need to be revised and compared with their Gadulus diploconus Seguenza, I876 Atlantic counterparts. Pl. 10, fig. 1s, 16 \X/ith a few exceptions, such as Puncturella noachina, Iivíng at shelf depths in northern latitudes, mosr of 1876c Cadulus diploconus Seguenza, p. 266. these Atlantic species have a deep-sea distribution 1994 Cadulus diploconus - Palazzi & Villari, frg. 42, 43. throughout their latitudinal ranges. Several Atlantic species have been reported also from deep thanatocoenoses Description. Spindle-shaped, slightly bent, smoorh, and referred to the Late Glacial age (Di Geronimo & Li shiny shell. Maximum diameter at mid-height of shell, Gioi, 1981; Di Geronimo & Bellagamba, 1986; Bouchet where an obscure angulation occurs. Roundish section. & Taviani, L989, Corselli & Bernocchi, 1990; Bonfitto Ventral side regularly convex in lateral vieq gradually et al., 1994, etc.). sloping to apical and oral openings; dorsal side centrally \Whether Atlantic deep-sea species presenrly enter swelling. Slightly ovate apex, with inner callus. Roun- the Mediterranean is a matter of discussion (see below). dish, oblique oral opening, larger than the apical ope- Larval ecology is undoubtedly an important factor ning. Length up to 3.0 mm; length to maximum diame- (Bouchet & Taviani, 1989, 7992,1993). Apart from the ter ratio ca. 1.8. deep oceanographic conditions, some biotic factors may Bathyal Molluscan from Southern ltaly 419

also bias the Mediterranean distribution of "Atlantic" An Atlantic affínity of the deep Mediterranean species. This is the case of Acesta excAt)Ata, Iphitus aspera- Plio-Peistocene benthos has been reported for several tus and Coralliophila ricbardi (see Ghisotti, 1979; Taviani faunistic groups, such as ostracods, scleractinians, bra- & Sabelli, 1982; Taviani & Taviani, 1986), all associated chiopods, stalked crinoids and bryozoans (see references with deep-sea scleractinian communities, presently decli- in Introduction). Benson's report (1972) desewes special ning in the Mediterrane î. According to Corselli & Ber- mention, being the first modern work to stress the pa- nocchi (1993), the disappearance of Pwncturella granulau leooceanographic meaning of psychrospheric Plio-Plei- was due to the loss of its trophic supply, i.e. the deep- stocene taxa from the Mediterranean. sea porifer communities. On the other hand, some spe- On these grounds, a model of the Plio-Quaterna- cies regarded previously as Atlantic, such as Ennucula ry evolution of the deep Mediterranean benthos has corbuloides, Yoldiella micromenica, Kaadesmia cuneata, been outlined (La Perna, 1994; Di Geronimo et al., prove instead to be common components of eastern and 1,996; Di Geronimo & La Perna, 1996). According to western Mediterranean bathyal assemblages (fanssen, this model, many deep-sea cold species appeared and 1989; Di Geronimo et al., !995, La Perna, unpubl. successfully spread in the Mediterraneaî since the mid- data), although currently found only as empty shells. dlelate Pliocene. This phenomenon should be seen wi- The other faunistic group was recognised more re- thin a generalised development of the deep oceanic cently (La Perna, 1,994). It consists of extinct species benthos following the Northern Hemisphere cooling which are closely related to Atlantic species or to Ocea- trend (Berggren, 1972; Shackleton & Opdyke, 1977, nic taxa in general. As shown for Homalopoma emulum Thunnel & \flilliams, 1983). The "absence" of a sill at (Di Geronimo & La Perna, 1997) and for Batlryspinula Gibraltar during the Plio-Pleistocene, supported by geo- excisa (Dr Geronimo & La Perna, 1996), these species logical data (Bousquet 6c Philip, 1976; Drllon er. aI., have no Recent counterparts among the deep Mediterra- 1980), aliowed the Mediterranean basin to have full nean benthos. Other species should be referred to this hydrological and faunistic exchanges with rhe Ocean (or group, such as Fissurisepa papillosa, Solariella tnarginula- much wider than those present today). At that time, the ta, Profundindssa spinulo sa, Benth omangelid ten ui co stata, North Atlantic-Mediterranean benthos had lost most of Awstrotindaria pusio, Thestyleda cuspidata, Katadesmia the Mesogean deep-sea taxa (see Barrier et al., I99I; La confusa, Cadulus wulum and Cadulus diploconus. It is Perna, in press). also worth reporting other species, such as the trocha- Dramatic changes occurred in the Late Pleistoce- ceans Cal I io sto rna sdyanum (Seguenza, 187 6), Cal I io stoma ne, when the oceaniclike communities disappeared. Co- formosissimum (Seguenza, 1876) and Microgaza solarioi- lalongo & Pasini (1988) dated the end of psychrospheric des (Seguenza, 1876), all showing close relationships conditions in the Mediterranean to the upper part of with North Atlantic deep-sea species. The pseudococcu- early Pleistocene. In any case, rich oceanic bathyal com- linrds Notocrater sp. and Coccullinella sp., also deserve to munities are known from Pleistocene deposits in Cala- be noted. Close phyletic relationships between living bria ranging from 1.1 M.y. to 600-700 k.y. (MNN 19e- and fossil species are obvious, although any lineage re- MNN 19f Biozones of Rio et al., 1990) (Di Geronimo construction is difficult or merely theoretical due to the et al., in press). A compression phase at Gibraltar since lack of data concerning Atlantic fossil deep-sea faunas. 1 Ma (Bousquet & Philip, 1976; Dillon et a1., 1980) The presence of archaeogastropods, nuculoids and seems to have played a strong role in isolating the Medi- scaphopods among this oceanic stock is cleariy related terranean from the Atlantic Ocean, and in determining to the non-planktotrophic larval development shared by its present-day oceanographic features. A main role in these groups. the fate of bathyal communities has been ascribed (Di These species have been regarded as cold-stenot- Geronimo & La Perna, 1997) to warm homothermic hermic due to their oceanic affinities (Di Geronimo et conditions which developed during Late Quaternary in- al., 1996; Di Geronimo & La Perna, 1996, 1997). A ran- terglacial phases. A model of increasing homothermy, ge of roughly 1O-8 to 4oC can be inferred for them, ac- however, seems to fit better the increasing threshold-ba- cording to the thermal range accounted for the psycro- sin conditions, related to the sill uplift. As known from sphere (e.g. Benson, 1972, t975). the literature (e.g. Allouc, 1982, Delibrias & Taviani, Other deep-sea species, such as Chrysallida micro- 1985) the deep Mediterranean benthos flourished again

scalaria and " Babylonella" profunda seem to have evolved in parr during the Late Pleistocene glacial phase from shallower ancestors, as outlined for the Pagodula (lVùrmian), when the oceanic Plio-Pleistocene species vaginaa-echinau líneage by La Perna (1996). A similar had already been lost. trend probably involved Cadulus. C. ooulum appears to \X/hat kind of deep-sea benthos was selected by represent the cold deep-sea term of a lineage ranging such processes? A scarcity of endemisms, a lack of cold- from the Miocene taurwolum to the Pleistoceîe attenua- stenothermic species, iow diversitlz, low density and bio- tus. mass, and a high degree of eurybathy, have been stressed 420 I. Di Geronimo & R. La Perna

often as characteristic of the deep Mediterranean bent- Katadesmia cuneata (>800-1,000 m). In rhese cases, per- hos (Perès & Picard, !964; Menzíes, 1,973; Fredj & Lau- sistence since the Plio-Pleistocene seems more likely biea 1985; Bouchet & Taviani, 1992,1993; Harmelin Er than a present-day introduction. A larval inflow of d'Hondt, 1993), although a slightly different model was many planktotrophically developing species from the outlined by Bellan-Santini et al. (1992). adjacent Atlantic Ocean is a reasonable hypothesis, but Bouchet & Taviani (1992, 1993) proposed a model the presence of an autochtonous, although scarce, deep depicting the deep Mèditerranean benthos as mainly Mediterranean benthos cannot be re.iected. It consists of represented by "pseudopopulations" of Atlantic species. species which endured better than others oceanographic In other words, the deep Mediterranean benthos is sup- changes during the Late Quaternary. plied with a meroplanktonic inflow from the Atlantic aia the Gibraltar threshold. Planktotrophic larvae of deep-sea species can bypass the sill thanks to vertical on- Achnozaledgernmts. togenetic migrations (Bouchet, 1,976; Borchet & Fontes, lVe wish to express our thanks to Serge Gofas and Philippe 1981), but the deep hydrological conditions prevent full Bouchet (lt4uséum National d'Histoire Naturelle, Paris), who made benthic development or reproduction. This model was available the material from the BALGIM and THALASSA Expeditions kindly assisted applied originally to turrids, a molluscan group in and one of us at MNHN. \le also thank Anders Sfarén (Naturhistoriska Riksmuseet, Stockholm), for which planktotrophy is widespread. It is difficult, how- suggestions concerning taxoflomy and help with the bibliography. ever, to use this theory to explain the occurrence of Orazio Torrisi (C.N.R., Catania) assisted in SEM photography. non-planktotrophically developing species in the Medi- Earlier versions of the manuscript were reviewed by Anders Warén (Naturhistoriska Riksmuseet, Stockholm) and Elio Robba (Milan terranean, whose depth-range begins beyond the well University). sill-depth e.g, Ennucula corbuloides and Yoldiella seguen- Financial suppoft was provided by M.U.R.S.T. Grants to Prof. zae () 500 m), Yoldiella miuomenica (>600-700 m) and Italo Di Geronimo (40o/o,6A0/ù.

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