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J. Paleont., 81(4), 2007, pp. 746–753Copyright � 2007, The Paleontological Society0022-3360/07/0081-746$03.00

KYLLINIA PARENTALIS (NEOGASTROPODA: TURRIDAE), NEW GENUS ANDSPECIES FROM THE PLIO–PLEISTOCENE OF THE CENTRAL

MEDITERRANEAN AREAVITTORIO GARILLI AND LUCA GALLETTI

Dipartimento di Geologia e Geodesia, Universita di Palermo, Corso Tukory 131, 90134 Palermo, Italy, �nenti@tiscali.it� and Via Mulinello, 20,90046 Monreale, Palermo, Italy, �gallucpa@tiscali.it�

ABSTRACT—A new genus and species of turrid neogastropod, Kyllinia parentalis, is described based on three shells from the Pliocene (NorthItaly) and the Pleistocene (northwestern Peloponnesus) of the central Mediterranean area. The new taxon is compared to the buccinid (sensulato) genus Metula H. and A. Adams, 1853 with which it shows a few superficial similarities mainly limited to the latticed sculptural patternof the teleoconch. Kyllinia parentalis is placed in the turrid subfamily Mangeliinae on the basis of teleoconch characters (double sinuosity ofthe outer lip and the short, forward curved anterior canal) and protoconch (a characteristic pattern formed by narrow axial riblets giving acoronated suture to the larval shell). These taxonomic characters, together with the almost clathrate teleoconch sculpture, form an originalcombination within the Mangeliinae, thus justifying the description of a new genus. The Recent species Philbertia marchadi Knudsen, 1956(for which we consider the previous attributions to Philbertia Monterosato, 1884 and Diaugasma, Melvill, 1917 incorrect) from the Republicof Guinea and Angola closely resembles K. parentalis, but differs in having axial folds on early teleoconch whorls, a more slender shell, anda proportionally taller last whorl. Philbertia marchadi fits well with the characteristics of the herein described genus, so that we propose thenew combination Kyllinia marchadi (Knudsen, 1956). Kyllinia parentalis, a species originated in the Mediterranean Pliocene, is to be consideredas the ancestor of K. marchadi. Kyllinia parentalis should be regarded as a meaningful marker in recognizing warm Mediterranean Pleistoceneevents.

INTRODUCTION

THIS STUDY is a further contribution in a series focusing on themolluscan assemblages from the Plio–Pleistocene of the cen-

tral Mediterranean region (Garilli, 2004; Garilli et al., 2005). Ma-terial studied here comes from the Pliocene of Castell’Arquato(Arda Valley, Piacenza, northern Apennine, Italy) and from theupper part of the Plio–Pleistocene sequence outcropping near Kyl-lini (Ileia, northwestern Peloponnesus, Greece) (Fig. 1). The ma-terial consists of three shells identified as belonging to the familyTurridae Swainson, 1840 mainly on the basis of general shellshape, the presence of a well-marked anal sinus, and a character-istic protoconch sculpture shown by several species of Mangeli-inae Fischer, 1887. These specimens show an original combina-tion of morphological features that is not present in other Recentand fossil turrids, and thus Kyllinia parentalis n. gen. and sp. isdescribed.

The family Turridae (sensu Powell, 1966) is one of the mostdiverse gastropod families. Its geographical and bathymetrical dis-tribution is extremely wide (from the tropical to the polar regionsand from shallow to very deep waters) and geologic range extendsto the Cretaceous (Powell, 1942). In the Mediterranean area, themost diverse turrid associations are known from well-establishedgenera such as Acamptogenotia Rovereto, 1899, Bathytoma Har-ris and Burrows, 1891, Clavatula Lamarck, 1801, Genota H. andA. Adams, 1853, Turricula Schumacher, 1817, and many othersdid not survive climatic deterioration recorded in this region atthe beginning of the Quaternary. In 1993, Taylor et al. made sometaxonomic changes in the families Turridae and Conidae Rafin-esque, 1815, moving, among other subfamilies, the Daphnellinaeand Mangeliinae from Turridae to Conidae. Along with manyother malacologists, we do not accept this placement. We agreewith Rosenberg (1998), who showed that the results of phyloge-netic analyses provided by Taylor et al. (1993) were not repro-ducible, and concluded that the traditional classification shouldnot be abandoned.

STRATIGRAPHICAL INFORMATION

The specimen here designated as paratype 1 (labeled as ExColl. Migliorini, Castell’Arquato, Pliocene, F371B) was providedby Maurizio Forli and came from the well-known Lower Pleis-tocene deposits of the Arda Valley. Most of these outcrops wereconsidered to be the Piacenzian stratotype (Barbieri, 1967) untilCastradori et al. (1998) redefined the base of this Middle Pliocene

Stage at Punta Piccola (Agrigento, southern Sicily). Although noprecise stratigraphic position is available, the location of paratype1 (the village of Castell’Arquato) and the fact that blue muddysediment is present inside the shell aperture suggest that this spec-imen was collected from the lower member (Pliocene in age) ofthe Castell’Arquato Formation, rather than from the upper mem-ber (Pleistocene) of this formation (mainly consisting of biocal-carenitic outcrops), or from the Argille di Lugagnano Formationcropping out near the village of Lugagnano Val D’Arda (see Raffiet al., 1989 for locations and overall litho-stratigraphical infor-mation on the cited Formations).

Whereas the Piacenzian stratotype area (Barbieri, 1967) iswidely known to paleontologists (see Caprotti, 1976 for the as-sociated molluscan assemblages), the deposits near Kyllini, wherethe remaining material was collected, have only been superficiallydiscussed even though they provide exceptionally rich and wellpreserved molluscan associations deposited in shallow marine andbrackish waters. The sequence of Kyllini (Vounargon Formationaccording to Hageman, 1976) is part of an anticlinal structure,which originated by a diapiric intrusion of Triassic halite andgypsum (Christodoulou, 1969, 1971; Hageman, 1976; Underhill,1988). The holotype was collected at the base of a sandy-muddygrayish bed (the H6 layer of Garilli et al., 2005, UTM 34S511793E 4199216N) with marine phanerogam remains, laterallychanging to a sandy bed with Lucinoma boreale (Linnaeus, 1767)in life position, and cropping out about 20 m above a turritellid-rich blue-grayish muddy level containing the nannofossil taxa Ge-phyrocapsa sp. 3 and Reticulofenestra asanoi Sato and Takayama,1992 (Fig. 2), whose Mediterranean distributions are respectivelyrecorded between 0.990 Ma (uppermost part of Lower Pleisto-cene) and 0.584 Ma (Middle Pleistocene) (Castradori, 1993; Spro-vieri, 1993; Di Stefano, 1998) and between 1.122 Ma (early partof the Lower Pleistocene Sicilian Substage, Ruggieri et al., 1984)and 0.781 Ma (lower part of the Middle Pleistocene) (De Kaenelet al., 1999). The specimen designated as paratype 2 was collectedfrom a yellowish sandy bed (here named F14), rich in Aporrhaispespelecani (Linnaeus, 1758) and glycymerids, cropping outabout 70 m above a grayish muddy bed containing R. asanoi (Fig.2). Consequently, an upper Sicilian Substage or, more likely, aMiddle–Upper Pleistocene age and a late Lower to Middle Pleis-tocene age could be assigned to the holotype (H6 layer) and para-type 2 (F14 layer), respectively.

747GARILLI AND GALLETTI—NEW MEDITERRANEAN PLIO–PLEISTOCENE GASTROPOD

FIGURE 1—Holotype (GNHM-FID 18) and paratype 2 (MGUP-KIGR 001)type locations; CA � Castell’Arquato, KY � Kyllini. Modified from Garilliet al. (2005).

FIGURE 2—Partial columnar section of the upper part of Vounargon For-mation (Kyllini, northwestern Peloponnesus, Greece) illustrating the strati-graphic position of holotype (GNHM-FID 18) and paratype 2 (MGUP-KIGR001). Modified from Garilli et al. (2005).

SYSTEMATIC PALEONTOLOGY

Class GASTROPODA Cuvier, 1797Order NEOGASTROPODA Thiele, 1929

Family TURRIDAE Swainson, 1840Subfamily MANGELIINAE Fischer, 1887

Genus KYLLINIA new genus

Type species.⎯Kyllinia parentalis new species.Diagnosis.⎯Small, mitriform-metuliform shell. Protoconch

conical, lacking typical Daphnellinae diagonal sculpture, withcrenulated suture. Teleoconch with weakly convex whorls withoutclearly defined sutural ramp. Sculpture delicate, mainly spiral,weakly latticed. Last whorl large, occupying two-thirds of shellheight. Aperture tall and narrow, ovate to pyriform. Outer liparcuate, internally smooth and thickened near edge, with twobroad and shallow subequal sinuses, anal sinus broader and deep-er. Inner lip slightly arcuate, with no plications. Anterior canalshort, curved forward.

Description.⎯Small, cylindrical-elongate shell reaching about 11 mm inheight. Protoconch conical, with more than two weakly sculptured whorls andwith partially immersed nucleus; suture crenulate. Teleoconch whorls slightlyconvex, not shouldered and separated by narrowly and shallowly incised su-ture. Sculpture predominantly spiral, weakly latticed. Last whorl large, oc-cupying two-thirds of shell height. Aperture pyriform to ovate. Outer lip withtwo sinuses, upper sinus broader and deeper, lower sinus in position of strom-boid notch. Inner lip slightly arcuate and smooth. Anterior canal short andslightly curved forward.

Etymology.⎯Kyllinia is after the small village of Kyllini (northwestern Pel-oponnesus), type locality.

Occurrence.⎯As fossil, Kyllinia is only known from the studied material,which ranges in age from Pliocene to Pleistocene (more likely Middle–UpperPleistocene) of the central Mediterranean area.

KYLLINIA PARENTALIS new speciesFigure 3.1–3.12

Diagnosis.⎯Small, cylindrical-elongate, mitriform-metuliformshell. Protoconch polygyrate consisting of little more than twowhorls, separated by crenulated suture, crenulations starting from

about first whorl. Sculpture consists of microscopic pustules,rounded tubercles, and subsutural short riblets. Teleoconch withweakly convex whorls without clearly defined shoulder or suturalramp, separated by shallowly incised suture. Sculpture weaklylatticed, consisting of fine spiral cords crossed by very thin axialthreads. Last whorl occupying two-thirds of shell height. Aperture

748 JOURNAL OF PALEONTOLOGY, V. 81, NO. 4, 2007

749GARILLI AND GALLETTI—NEW MEDITERRANEAN PLIO–PLEISTOCENE GASTROPOD

←

FIGURE 3—Kyllinia parentalis n. gen. and sp., Plio–Pleistocene of central Mediterranean area. 1–4, 8–11, Holotype, GNHM-FID 18, Pleistocene of Kyllini(northwestern Peloponnesus, Greece), Vounargon Formation (upper part), 1, apertural view; 2, dorsal view; 3, lateral view; 4, profile of the outer lip illustratingthe two sinuses; 8, sculpture of the penultimate whorl; 9, suture and sculpture of the early protoconch whorls; 10, 11, protoconch, 10, illustrating protoconch/teleoconch (P/T) boundary (black arrows); 11, illustrating the sinuate scars in the early whorl (white arrows); 5, 6, 12, paratype 1, MZB-30001, MiddlePliocene of Arda Valley (Castell’Arquato, Piacenza, northern Italy), Castell’Arquato Formation (lower Member), 5, apertural view; 6, lateral view; 12,protoconch, illustrating P/T boundary (black arrows); 7, paratype 2, MGUP-KIGR 001, Pleistocene of Kyllini, Vounargon Formation (upper part), aperturalview. Scale bars: 2 mm in 1–3, 5, 6; 1 mm in 4, 7; 500 �m in 8; 200 �m in 10, 12; 100 �m in 11; 50 �m in 9.

tall and narrow. Outer lip smooth and arcuate interiorly, with twobroad sinuses, anal sinus deeper.

Description.⎯Sturdy, cylindrical-elongate shell up to 11.2 mm in height(paratype 1). Larval shell conical, polygyrate, consisting of 2.25–2.4 convexwhorls (counted according to Verduin’s 1977 method). Nucleus partially im-mersed. Early whorls sculptured with microscopic pustules forming two–threediscontinous spiral lines on abapical region (Fig. 3.9); very small roundedtubercles cover remaining surface towards protoconch/teleoconch boundary(Fig. 3.10, 3.11). Well-marked, moderately sinuate scars (protoconch 1/pro-toconch 2 boundary?) (Fig. 3.11) at about end of first whorl. From about firstprotoconch whorl, very short riblets start near suture, which consequentlyappears crenulate (Fig. 3.9). This gives a coronated appearance to most oflarval shell. Protoconch height ranges from 645 �m to 660 �m, maximumdiameter from 550 �m to 570 �m, nucleus diameter about 120–150 �m.Protoconch/teleoconch boundary clearly defined by sinuate scar (Fig. 3.10,3.11). Teleoconch consisting of about 2.7 (paratype 2), 4 (holotype), 5 (para-type 1) weakly convex whorls sculptured with fine spiral cords, numbering14–15 (holotype) to 30–31 (paratype 1) on penultimate whorl. Spiral cordscrossed irregularly by delicate, flexuous axial threads extending from sutureto suture, rendering portions of shell weakly clathrate (Fig. 3.8). Episodicgrowth pauses present, mainly on paratype 1. Whorls separated by narrow,shallowly incised sutures. Last whorl large, making up about three-fourths(paratype 2), two-thirds (holotype) and three-fifths (paratype 1) of shell height.Aperture tall, narrow, slender, almost ovate-pyriform in holotype and paratype2, more markedly ovate in paratype 1 (largest specimen), making up abouttwo-thirds of body whorl height. Inner lip moderately arcuate, covered bythin, very narrow callus. Outer lip internally smooth, thickened close to edge,curved and prosocirt in middle, with two broad, distinct sinuses in upper andlower parts (Fig. 3.4). Anal sinus deep and wide, lower sinus wide and low,in position of stromboid notch. Anterior canal rather short, slightly but clearlycurved forward. Before coating for scanning electronic microscopy, holotypeand paratype 1 preserved spiral pattern of reddish (probably originally red-brown) spots, particularly marked near suture.

Etymology.⎯The species name, the Latin adjective parentalis, is dedicatedto our parents and grandparents.

Types.⎯Holotype, GNHM-FID 18, Goulandris Natural History Museum,Kifissia, Athens, Greece, and paratype 2, MGUP-KIGR001, Museo GeologicoG. G. Gemmellaro, Palermo, Italy, from the Pleistocene of Kyllini (Ileia,northwestern Peloponnesus, Greece), upper part of Vounargon Formation.Paratype 1, 30001, Museo Zoologico Bologna (MZB), Italy from Cas-tell’Arquato and is probably from the Middle Pliocene (Piacenzian Stage),Catell’Arquato Formation (lower member), Arda Valley (Castell’Arquato, Pia-cenza, northern Italy).

Measurements.⎯Holotype, GNHM-FID 18, height (H) 6.7mm, diameter of last whorl (D) 2.2 mm, protoconch height (Hp)660 �m, protoconch diameter (Dp) 550 �m; paratype 1, MZB-30001, H 11.2 mm, D 3.4 mm, Hp 645 �m, Dp 570 �m; paratype2, MGUP-KIGR001, H 3.7 mm, D 1.4 mm, Hp 660 �m, Dp 560�m.

Occurrence.⎯Kyllinia parentalis is known only from the studied material,which ranges in age from Pliocene to Pleistocene (more likely Middle–UpperPleistocene) of the central Mediterranean area.

Discussion.⎯As we noted in the description (where we use theterm mitriform-metuliform), at first sight, the described taxon re-sembles the Buccinidae Rafinesque, 1815 genus Metula H. andA. Adams, 1853 (also attributed to Colubrariidae Dall, 1904, e.g.,Vermeij, 2001; Schmelz and Portell, 2003). The similarities be-tween Metula species and Kyllinia are mainly limited to the lat-ticed, almost clathrate teleoconch sculpture, to the coloration pat-tern, which in some Metula spp. consists of largebrownish-reddish spiral bands, and, as noted by J. A. Todd (per-sonal commun., 2004), to the presence of growth pauses. Thesefeatures also are present in some Turridae Daphnellinae Hedley,1922 species, e.g., Daphnella bartschi Dall, 1919 (see Abbott,1974, fig. 3411), D. antillana Espinosa and Fernandez-Garces,

1990, D. botanica (Hedley, 1918), and D. interrupta (Pease,1860) (for these species the reader is referred to images onhttp://www.femorale.com and http://www.gastropods.com), whichalso show an overall shell shape comparable to that observed inKyllinia. Mitrid species, however, lack distinct sinuses on the out-er lip, and always have prominent colummellar plicae. Severaldifferences allow the Kyllinia to be separated from the Metulagroup, including the buccinid (sensu lato) taxa. Species of thisgenus (see also the holotype of Buccinum clathratum A. Adamsand Reeve, 1850, type species of Metula, well figured in Emerson,1986, figs. 1, 2, and in Beu and Maxwell, 1987, pl. 30, fig. g–i)show a longer anterior canal that is slightly to markedly curvedbackward, a clearly pyriform aperture, and a particularly thick-ened, almost varicose and often internally denticulate outer lip.In some Metula species a very slight sinuosity is shown only inthe upper portion of the outer lip. In contrast, K. parentalis n. sp.has a shorter anterior canal, slightly curved forward, so that theprofile of the dorsal margin appears very similar to that of Daph-nelliinae and (particularly) to Mangeliinae (see Kilburn, 1992 and1994) species. The shell aperture, as a consequence of ontogeny,is markedly pyriform (as in Metula species) only in the holotype(probably a subadult specimen), but is ovate in paratype 1 (thelargest, adult specimen), which has a thinner outer lip and a moremarkedly cylindrical shell shape. Another ontogenetic variation,clearly visible between the holotype and paratype 1, is the numberof spiral threads on the penultimate whorl (14–15 vs. 30–31).

The double sinuosity in the outer lip, which is lacking in Me-tula and not very frequent in turrid taxa (see Powell, 1966), ismarked in all specimens of Kyllinia with a pattern similar to thatshown by some species of the turrid subfamily Daphnellinae (e.g.,Daphnella bartschi, D. interrupta) and, especially Mangeliinaespecies, e.g., Gingicithara lyrica (Reeve, 1846) (see Kilburn,1992, fig. 57), G. ponderosa (Reeve, 1846) (see Kilburn, 1992,fig. 173), Citharomangelia africana (Sowerby, 1903) (see Kil-burn, 1992, fig. 82), Leiocithara costellarioides Kilburn, 1992(fig. 103), L. musae (Thiele, 1925) (see Kilburn, 1992, fig. 114),L. perlucidula Kilburn, 1992 (fig. 121), L. porcellanea Kilburn,1992 (fig. 127), Pseudorhaphitoma sienna Kilburn, 1994 (figs. 5,8), P. drivasi Kilburn, 1994 (fig. 11), P. obturata Kilburn, 1994(fig. 43), P. stipendiarii Kilburn, 1994 (fig. 50), P. fuscescens(Thiele, 1925) (see Kilburn, 1994, fig. 60), and P. tetragona(Gould, 1860) (see Kilburn, 1994, fig. 73). All these species havea more or less marked and deep anal sinus and a very slightsinuosity (similar to the stromboid notch) in the lower part of theouter lip, exactly the pattern occurring in Kyllinia.

The above-mentioned differences from the Metula group areclearly shown when comparing Kyllinia to the following species(for which J. A. Todd, personal commun., 2004, noticed a link toKyllinia), today considered as belonging in Metula (Emerson,1986; Beu and Maxwell, 1987): the Mediterranean Mio–PlioceneMurex mitraeformis Brocchi, 1814 (Fig. 4.1–4.3, from the Plio-cene of Altavilla, northwestern Sicily, Ruggieri et al., 1967; Rug-gieri and D’Arpa, 1993) and the Recent Antemetula elongata(Dall, 1907) (Fig. 4.4, 4.5) from the Philippines. Also other Mio–Pliocene species from outside the Mediterranean area such as Me-tula miocenica Schmelz and Portell, 2003 and M. roberti Olsson,1967 (in Schmelz and Portell, 2003, figs. 1, 2, respectively), showthe same features.

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FIGURE 4—Fossil and Recent species of Metula H. and A. Adams, 1853for comparison with Kyllinia n. gen. 1–3, Metula mitraeformis (Brocchi,1814), from the Pliocene of Altavilla (northwestern Sicily), housed in theDipartimento di Geologia e Geodesia, Universita di Palermo (DGUP), DGUPAT PA 023/484, in lateral (1), apertural (2), and dorsal (3) views. 4, 5, Metulaelongata (Dall, 1907), from Aliguay, Diplog, Philippines�200 m, DGUP ALPH 001/485 in lateral (4) and apertural (5) views. Note the typical Metulaanterior canal curved backwards (1, 4). Scale bars: 1 cm.

Kyllinia shows affinities in sculptural pattern with Daphnobela,a genus erected by Cossmann (1896, p. 93, pl. 5, figs. 30, 31)who cited ‘‘Bucc. junceum, Sow. Eoc,’’ from the Eocene (Bar-tonian) of England, as the type species. In reality, this species was

described by Solander in Brander (1766) as Murex junceus (seealso Sherbon, 1902), and thereafter only discussed by J. Sowerby(1822). Following Wenz (1938–1944), who placed Daphnobelaas a subgenus of Metula, Dolin et al. (1980, see also Le Renard,1995), describing D. juncea curryi (from the Paris Basin), statedthat this genus, originally placed in the Turridae, should be con-sidered as belonging in the Buccinidae Rafinesque, 1815. Weagree with Schmelz and Portell (2003), who considered Daph-nobela as a synonym of Metula. ‘‘Metula juncea,’’ as reported bySchmelz and Portell (2003, fig. 3) from the type locality, appearsto be a typical Metula: its aperture is pyriform, its anterior canalis curved backwards, and the outer lip is very slightly sinuous inthe upper part only.

Also, the normal shell size of Metula species is much larger.The smallest species (height 15 mm), originally described as Me-tula (Minitula) minor by Olsson and Bayer (1972), was placed inthe Columbellidae Swainson, 1840 (close to genus NassarinaDall, 1889) by Beu and Maxwell (1987); so that few Metula spp.are 20–25 mm in height and rarely as small as 17 mm. Notably,M. (M.) minor, the only species of the subgenus Minitula Olssonand Bayer, 1972, in our opinion correctly moved from the Metulagroup, has a short, straight anterior canal (see also Schmelz andPortell, 2003, p. 13) comparable with that of Kyllinia. Differencesare emphasized when comparing Kyllinia to other buccinid (s.l.)taxa (e.g., Bartschia Rehder, 1943; Colubraria Schumacher, 1817;Iredalula Finlay, 1926; Kanamarua Kuroda, 1951) more or lessrelated to Metula (Ponder, 1968; Beu and Maxwell, 1987), andsome of them are not similar to Kyllinia. Differences betweenKyllinia and Metula species are even observed in protoconch siz-es: Metula mitraeformis (Brocchi, 1814) and M. elongata Dall,1907 show a protoconch almost twice as large as the protoconchof Kyllinia.

The crenulated, coronate shape of the Kyllinia protoconch su-ture, formed by narrow subsutural riblets present in the last whorland protruding over the suture, is to be considered as a veryimportant taxonomic character. A very similar protoconch pat-tern—‘‘the last whorl (at least) axially ribbed’’—was describedby Kilburn (1994) as most common within the Mangeliinae. Ineffect, this Kyllinia protoconch character is certainly the same asis present in the Mediterranean Cythara rugulosa (Philippi, 1844)(see Richter and Thorson, 1975, pl. 12, figs. 81, 82). Other in-disputable turrid species (s. Powell) have a very similar patternwith narrow axial ribs protruding above the suture (Mangelia nu-perrima (Tiberi, 1855) (see Bouchet and Waren, 1980, fig. 215),Eucithara ubuhele Kilburn, 1992 (fig. 21), Eucithara striatella(Smith, 1884) (see Kilburn, 1992, fig. 32), Gingicithara lyrica(Reeve, 1846) (see Kilburn, 1992, fig. 58), G. notabilis (Smith,1888) (see Kilburn, 1992, fig. 72), Citharomangelia africana (seeKilburn, 1992, fig. 83), Leiocithara porcellanacea Kilburn, 1992(fig. 128), Pseudorhaphitoma sienna Kilburn, 1994 (fig. 2), P.drivasi Kilburn, 1994 (fig. 13), P. epistomifer Kilburn, 1994 (fig.20), and P. ethekwini Kilburn, 1994 (fig. 40). This protoconchcharacter, which to our knowledge is not found outside the Tur-ridae, determines the correct systematic placement of Kyllinia. Ina similar case, Le Renard (1995) moved some confusing taxafrom Buccinidae to Conidae Rafinesque, 1815 (considered in thesense of Taylor, as including most of the classic turrids) solely onthe basis of protoconch characters. Also, the conical shape of thelarval shell and the partially immersed nucleus shown by Kylliniaare common in Mangeliinae (Kilburn, 1992, 1994).

Kyllinia parentalis n. sp. closely resembles the East Atlantic(from the Republic of Guinea and Angola, Rolan et al., 1998)species Philbertia marchadi Knudsen, 1956 (p. 526–527, fig. 3;holotype well-figured by Rolan et al., 1998, fig. 1), mainly insculptural pattern, profile of the outer lip and coloration, consist-ing of spiral darker rows in both species. Also the protoconch,having the characteristic Mangeliinae crenulated suture (see Rolanet al., 1998, fig. 4), is very similar to that of K. parentalis. The

751GARILLI AND GALLETTI—NEW MEDITERRANEAN PLIO–PLEISTOCENE GASTROPOD

Recent species, however, has a more slender shell, a proportion-ally taller last whorl, and distinct axial folds on the last whorl ofthe protoconch. We consider the attribution to Philbertia Monter-osato, 1884 (synonym of Raphitoma Bellardi, 1847) incorrect.Consistent morphological characters support this opinion. Boththe genus Raphitoma (type Pleurotoma hystrix De Cristofori andJan, 1832, see the well-figured lectotype in Pinna and Spezia,1978, fig. 1–1a) and Philbertia (type Pleurotoma philberti Mi-chaud, 1829) are characterized by markedly turrited shells usuallywith denticles on the inner part of the outer lip, which often bears(particularly in Philbertia) a deep reversed L-shaped anal sinus.In contrast, P. marchadi Knudsen, 1956 is cylindrical, with veryweakly convex whorls, and an interiorly smooth outer lip. Alsothe attribution of Rolan et al. (1998) to Diaugasma, Melvill, 1917,is not correct. In fact, this taxon, proposed as a subgenus of Daph-nella Hinds, 1844, has a diagonally cancellated protoconch sculp-ture. Furthermore, from a biogeographic point of view, the place-ment of P. marchadi, a western African species, in Diaugasma,an Indo-Pacific taxon, is not convincing. Thus, Philbertia mar-chadi fits well with the characters of the genus Kyllinia, and isto be included in this genus.

Kyllinia parentalis and K. marchadi (Knudsen, 1956) n. comb.definitely have characters similar to those of many Mangeliinaespecies and they belong in this turrid subfamily. Fewer similarcharacters are seen in the subfamily Daphnellinae, whose speciesare characterized by having a multispiral larval shell with a di-agonally cancellated (see Powell, 1966) protoconch 2. Further-more, the original combination of characters shown by the de-scribed species (the double sinuosity of the outer lip and thecoronated protoconch shape, together with the almost clathrateteleoconch sculpture) is not recorded in other taxa of Mangeliinae,which are characterized by a sculpture with predominant axialribs, and so justifies the description of the new genus.

CONCLUDING REMARKS

The genus Kyllinia n. gen. includes two species, K. parentalisn. sp., from the Pliocene of northern Italy and the Pleistocene ofnorthwestern Peloponnesus, and K. marchadi (Knudsen, 1956) n.comb., from the coasts of West Africa. Occurrences of Kylliniaindicate that this genus originated in the Pliocene of the Mediter-ranean area and migrated in to the eastern Atlantic where theliving species K. marchadi originated. The direct phylogeneticlink between K. parentalis and K. marchadi is clear when con-sidering the very close similarity of the two species.

The above mentioned biogeographic configuration, with an At-lanticward migration flow, perfectly agrees with the view of thewell-established affinity between the subtropical to tropical east-ern Atlantic molluscan assemblage and that of the MediterraneanBasin. As documented by several (Mediterranean) Pliocene to(east Atlantic) Recent taxa (limited to the area between Senegaland the Gulf of Guinea) (see Le Loeuff and Von Cosel, 1998 andMonegatti and Raffi, 2001), the marked onset of this affinity oc-curred after the Middle Miocene interruption of the connection ofthe ‘‘old’’ Mediterranean with the Indo-Pacific paleoprovince, atabout 14 m.y. (Raffi and Serpagli, 1993), and the Upper MioceneMessinian salinity crisis (Ruggieri, 1967; Hsu et al., 1972). It isnot easy to precisely establish the time when certain Mediterra-nean fossil taxa migrated to the eastern Atlantic, especially whenconsidering that re-introduction in the Mediterranean area couldbe possible. The stratigraphic data on Kyllinia (including K. mar-chadi, for which no fossil record is known), however, could in-dicate that migration of this genus occurred in a Pliocene to Mid-dle–Upper Pleistocene range. Further Mediterranean and(especially) Atlantic fossil records of Kyllinia should be helpfulin solving this question.

At the present time, the living species of the genus Kyllinia isfound along the African western typical tropical and southern re-gions (Le Loeuff and Von Cosel, 1998), where the sea surface

water temperatures are never cooler than about 26� C and 20� C(Le Loeuff and Von Cosel, 1998), respectively. In this view, whenconsidering the herein presented Pleistocene records, K. paren-talis should be regarded as a meaningful thermophilic climaticmarker, in the same way as other molluscs (such as Strombuslatus Gmelin, 1791 � Strombus bubonius Lamarck, 1822, Can-tharus viverratus Kiener, 1835, Conus ermineus Born, 1778 � C.testudinarius Hwass in Bruguiere, 1792; synonymies are accord-ing to the Western Atlantic Gastropod Species Database at TheAcademy of Natural Science of Philadelphia, available on http://erato.acnatsci.org), living around Senegal, which are known to becharacteristic of the Mediterranean late Middle to Upper Pleis-tocene warm marine isotopic stages 7 and (particularly) 5 (seeHearty et al., 1986). Thus, K. parentalis allows recognition ofwarm water climatic conditions in the Pleistocene F14 and H6Kyllini layers.

ACKNOWLEDGMENTS

Our special thanks to S. Palazzi (Modena, Italy), who in various waysstrongly and generously encouraged us in describing Kyllinia n. gen., as wellas providing bibliographic help and very helpful comments on the originalmanuscript. We are indebted to M. Crisci (Palermo, Italy) for picking theholotype, to M. Forli (Prato, Italy) for kindly giving paratype 1 from hiscollection, to E. Di Stefano and A. Incarbona (Dipartimento di Geologia eGeodesia, University of Palermo, Italy) for detecting the occurrence of thestratigraphical markers Gephyrocapsa sp. 3 and Reticulofenestra asanoi, andto F. Pollina (Palermo) for providing photos in Figures 5 and 6. We are alsograteful to J. K. Tucker (Illinois Natural History Survey, Brighton, USA) forkindly making available his unpublished catalog of worldwide fossil and re-cent turrids. Also W. Backhuys (Leiden, The Netherlands), A. G. Beu (Insti-tute of Geological and Nuclear Sciences, New Zealand), the BioSophia as-sociation (www.biosophia.it), T. Boone (Rosenstiel School of Marine andAtmospheric Science, University of Miami, USA), R. Giannuzzi-Savelli (Pa-lermo), F. Pusateri (Palermo), G. W. Schmelz (Naples, Florida, USA) and G.Spada (Paris, France) provided us with some relevant literature. We are in-debted to J. A. Todd (British Museum (Natural History), London) whoserevision of the earlier manuscript provided useful comments and allowed usto go deep into the question of the systematic placement of Kyllinia. Thanksare due to B. Sabelli (Dipartimento di Biologia Evoluzionistica Sperimentale,Bologna, Italy) and E. Vardala-Theodorou (Goulandris Natural History Mu-seum, Kifissia, Greece) for quickly providing catalog numbers. We are es-pecially grateful for constructive reviews by A. G. Beu, and P. Lozouet (Mu-seum National d’Histoire Naturelle, Paris) and for the kind editorial adviceand helpful comments provided by L. C. Anderson (Department of Geologyand Geophysics, Louisiana State University, USA) and L. Vermaas (Depart-ment of Geoscience, The University of Iowa, USA). This research was par-tially supported by M.U.R.S.T. funds, 60% to A. Greco (Dipartimento di Geo-logia e Geodesia, Palermo).

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