ALDROVANDIA 4 2008: 3 - 20 NOTES ON THE BELA BRACHYSTOMA COMPLEX, WITH DESCRIPTION OF A NEW SPECIES (MOLLUSCA, GASTROPODA: CONIDAE) Paolo Mariottini, Andrea Di Giulio, Carlo Smriglio, Marco Oliverio ABSTRACT Based on shell characters, the complex of extant and fossil species related to the Recent Mediterranean taxon Bela brachystoma (Philippi, 1844) is critically analysed. In spite of the fact that this species shows great shell variability, Recent shells are morphologically identical to the Pliocene shells. A lectotype of Pleurotomum granuliferum Brugnone, 1862, synonym of B. brachystoma, is herein designated. The relationship occurring between B. brachystoma and the Recent Atlantic species Bela exilis Ardovini, 2004, Bela africana Ardovini, 2004 and Bela ardovinii Mariottini & Oliverio n. sp. is also discussed. KEY WORDS Gastropoda, Conidae, Bela, new species, taxonomy, lectotype, Mediterranean Sea, Recent, Pliocene INTRODUCTION The epithet “turrids” has been largely used in a broad sense as relating to the largest radiation of marine gastropods, traditionally included in the familygroup Turridae (cf. Tucker, 2004), which is currently undergoing significant revision (Taylor et al., 1993; Kohn, 1998; Bouchet & Rocroi, 2005; Puillandre et al., 2008). Over 100 species of this group of neogastropods, sorted among 25-30 genera, are presently known from the Mediterranean Sea. Supraspecific systematics is still very unstable on a global scale and this is reflected also at the regional level (Powell, 1966; Nordsieck, 1977; Sabelli et al., 1992; Gofas & Le Renard, 2007). According to the most recent molecular phylogenetic analysis (Puillandre et al., 2008), a large part of the familylevel classification of “turrids” proposed by Taylor et al. (1993) on a morphological basis is also supported by DNA data. Although monophyly of the Mangeliinae is not supported in the analysis by Puillandre et al. (2008), this group belongs to a clade with closer relationship to Conus Linnaeus, 1758 than to other members of the “Turridae” s.l., a circumstance which therefore suggests their provisional inclusion into the family Conidae. The a-taxonomy of the Recent Mediterranean mangeliine species, largely based on shell characters, also suffers from the lack of comprehensive and modern revisions. Among the others, the mangeliine genus Bela Gray, 1847 was based on Murex nebula Montagu, 1803, the identity of which is uncertain and should be carefully reconsidered. The genus, as currently conceived, is based on the redescription by Powell (1966: 97), and consists of small Recent and Tertiary turrids with an eastern Atlantic distribution, including the Mediterranean Sea. As emphasised by Powell (1966: 98), this genus has been frequently considered as a “basket” for the allocation of a wide variety of unrelated turrids. As a consequence, the number of Recent species considered as valid varies a lot according to different authors (e.g. Nordsieck, 1977; Sabelli et al., 1992; Gofas & Le Renard, 2007). Among them, a group of morphologically similar species is herein revised. The best known one is Bela brachystoma (Philippi, 1844), originally described from the Mediterranean Sea, with a multispiral protoconch and a peculiar teleoconch sculpture. Brugnone (1862) described Pleurotomum granuliferum, based on material from the PlioPleistocene of Ficarazzi (Sicily) that is very similar to the Recent Mediterranean shells, and which we consider conspecific with B. brachystoma. Two species have been recently described from the eastern Atlantic: Bela exilis Ardovini, 2004 and Bela africana Ardovini, 2004. Regrettably, the latter two species were published in a journal which evidently did not provide a peer review, resulting in very poor descriptions, and subsequent taxonomic mistakes. Under these circumstances, comparative redescriptions of all the involved species became necessary. Ardovini (2004) identified also a possibly undescribed species from West Africa, which is herein described as new: Bela ardovinii n. sp. MATERIALS AND METHODS We based our morphological comparisons on hundreds of shells of B. brachystoma from different localities. We could examine material of P. granuliferum in the Monterosato collection (ex coll. Brugnone) at the Museo Civico di Zoologia di Roma (MCZR) and several shells of B. exilis, B. africana and B. ardovinii n. sp. In order to quantitatively evaluate the differences among the shells, morphometric variables such as diameter, number of whorls and 4 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO sculpture for the protoconch, and axial folds, length and width for the teleoconch, were measured. Moreover, the length/width ratio was calculated as a shape parameter. Unless otherwise stated, citation of unpublished names (as manuscript names, labels etc.) is not intended for taxonomic purposes. Abbreviations are as follows: CS-PM = collection Carlo Smriglio-Paolo Mariottini (Rome, Italy) MCZR = Museo Civico di Zoologia di Roma (Rome, Italy) MMP = Museo Malacologico Piceno di Cupra Marittima (Macerata, Italy) MNHN = Muséum National d’Histoire Naturelle (Paris, France) MO = collection Marco Oliverio (Rome, Italy) RA = collection Roberto Ardovini (Rome, Italy) SR = collection Stefano Rufini (Rome, Italy) D = diameter H = heigth lv = live collected specimen(s) sh = empty shell(s) SYSTEMATIC PART Conidae Mangeliinae Bela Gray, 1847 (The Annals and Magazine of natural History (1) 20: 270) Type species: Murex nebula Montagu, 1803, by subsequent designation (Gray, 1847). = Fehria van Aartsen, 1988 (type species: Ginnania taprurensis Pallary, 1904, by original designation) Remarks The identity of Murex nebula Montagu, 1803, is still uncertain and a neotype designation would be highly desirable to stabilise not only the species but also usage of the generic name. We maintain some doubts on the attribution to this genus of Pleurotoma brachystomum Philippi, 1844, which shows a very peculiar sculpture of both the protoconch and teleoconch as compared to the other species traditionally ascribed to Bela (e.g. Bela laevigata (Philippi, 1836) or B. menkhorsti van Aartsen, 1988) (Figs 143-144, 163). However, we prefer to provisionally maintain B. brachystoma within Bela s.l., pending on a future revision of the nominal genus. In contrast with most caenogastropods where the protoconch-teleoconch boundary is clearly marked by a discontinuity in sculpture, especially in species with planktotrophic larvae, the transition in Bela may be fairly gradual. In fact, the last part of the protoconch is often sculptured by collabral riblets which mask the real boundary. Bela brachystoma (Philippi, 1844) (Figs 1-77, 127-140, 151-161) Pleurotoma brachystomum Philippi, 1844 (Enumeratio molluscorum Siciliae: 169, 176, pl. 26, fig. 10) = Pleurotomum granuliferum Brugnone, 1862 = Raphitoma brachystoma var. comitasensis Fontannes, 1880 = Pleurotoma brachystoma var. curta Almera & Bofill, 1898 = Pleurotoma submarginata var. minor Almera & Bofill, 1898 = Pleurotoma brachystoma var. emporitensis Almera, 1907 Bela brachystoma apicalis Nordsieck, 1977 [taxon inquirendum] Original description. “Ph. testa turrita, fulvo - fusca; anfractibus convexis, superne angulatis, striis elevatis transversis; costis longitudinalis elevatis novem, ad suturam evanescentibus; apertura oblonga, dimidiam spiram vix superante; cauda brevissima; labro acuto. Rarissimum; specimina tria modo Neapoli legi. Testa 31/2/// alta, 11/2/// lata, turrita. Anfractus 8-9, propter costas versus ipsam suturam evanescentes subangulati, striis elevatis (circa 6-8 in anfractu penultimo) scabri; costae plicaformies, elevatae, interstitiis aequalis. Color rufus ad fulvum transiens. – Species valde distincta.” Types depositories. Type material of B. brachystoma is probably housed in the National Museum of Natural History (Santiago del Chile). Lectotype of P. granuliferum, here designated (in MCZR: H = 5.7 mm, D = 2.2 mm; figs 53a-b). Type localities. Italy, Central Tyrrhenian Sea, Naples (P. brachystomum); Sicily, Palermo, Ficarazzi (Upper Pliocene-Lower Pleistocene) (P. granuliferum). Material examined Recent. [Atlantic] 25 exx (sh), northeastern Atlantic, France, Gulf of Gascogne, Capbreton 88 sta. DE-01, 43°39.99’N 1°48.11’W, - 134 m, MNHN; 3 exx (sh), idem, sta. DR-29, 43°46.51’N 2°00.58’W, - 165 m, MNHN; 16 exx (sh), idem, DR-11, 43°22.77’N 1°59.18’W, - 94 m, MNHN; 8 exx (sh), idem, sta. DE-05, 43°57.42’N 2°05.16’W, - 164 m, MNHN. 16 exx (sh), Maroc, Agadir, R/V Vanneau 1923-1929 sta. 10, 29°54’N 9°58’W, - 110 m, MNHN; 3 exx (sh), idem, sta. 32, 34°01’N 7°32’W, - 145 m, MNHN. 1 ex. (sh), Mauritanie, R/V N’Diago sta. 239, 17°48’N 16°21’W, - 79 m, MNHN. BELA BRACHYSTOMA COMPLEX [Mediterranean] 40 exx (sh), Western Mediterranean Sea, France, Gulf of Lion, IFREMER/DEPRO 96 (R/V Europe) sta. chalut-10, 42°24.6’N 3°16.2’E, - 100/151 m, MNHN; 5 exx (sh), idem, sta. chalut-11, 42°09.4’N 3°22.5’E, - 350 m, MNHN; 5 exx (sh), Cap Béar, ECOMARGE 1984 sta. A61, 42°29’30”N 3°10’30”E, - 42 m, MNHN; 2 exx (sh), off Rhône delta, - 50/100 m, MNHN; 3 exx (sh), Arcachon, MNHN. 2 exx (sh), Spain, Estepona, 36°25’N 5°09’W, - 150 m, SR; 78 exx (sh), Italy, Tyrrhenian Sea, off San Vincenzo (LI), 43°05’N 10°24’E, - 34 m, CS-PM; 1 ex. (sh), off S. Marinella (RM), - 150/200 m, sediment in an old Roman dolium, 41°54.005’N, 011°47.666’E, MO; 38 exx (sh), off Fiumicino (RM), 41°43’N 12°06’W, - 80 m, CS-PM; 27 exx (sh), off Fiumicino (RM), 41°38’N 12°11’W, - 140 m, CSPM; 11 exx (5 juveniles) (sh), idem (in the gut content of Astropecten irregularis), MO; 2 exx (lv), 5 nm South of Fiumicino, - 25 m (in mud), MO; 3 exx (2lv, 1sh), Tor Paterno shoal (RM), - 150 m, MO; 24 exx (sh), Ponza Is. (LT), 40°51’N 12°55’W, - 40 m, CS-PM; 11 exx (sh), Adriatic Sea, off Civitanova Marche (MC), 43°18’N 13°46’E, - 45 m, CS-PM; 24 exx (sh), off Pescara, 42°31’N 14°12’E, - 50 m, CS-PM. 5 exx (sh), Croatia, Brac Is., 43°24’N 16°30’E, - 50 m, CS-PM. 19 exx (sh), Southern Mediterranean Sea, Libya, 110/150 m, CS-PM. Fossils. 2 exx, Italy, Latium, Guidonia (RM), 42°00’N 12°43’E (Pliocene), CS-PM; 3 exx, Calabria, Gallina (RC), 38°05’N 15°41’E (Pliocene), CS-PM; 125 exx (sh), Sicily, Ficarazzi (PA), 38°04’N 13°29’E (Upper Pliocene-Lower Pleistocene), labelled “P. granuliferum. var. parva”, coll. Monterosato (ex coll. Brugnone), MCZR; 48 exx, idem, Monte Pellegrino (PA), 38°04’N 13°29’E (Upper Pliocene-Lower Pleistocene), labelled “var. striiselevatioribus”, coll. Monterosato (ex coll. Brugnone), MCZR. Description Shell small for the genus, height 4.5-7 mm, width 1.7-2.5 mm, biconical, turriculate elongate, solid. Protoconch multispiral, dome shaped, of 2.3-2.4 convex whorls. First 1.6-1.9 apical whorls smooth, the remaining with reticulated sculpture of 4-5 granulose spirals (3 major spirals in the middle of whorl, 1 smaller subsutural and 1 smaller above the teleoconch suture) crossed by oblique axial riblets. Maximum diameter of protoconch 510-650 µm. Protoconchteleoconch transition not well marked. Teleoconch of 5-6 whorls, rounded, sutural ramp straight or very slightly convex, whorl sides gently convex. Last whorl about 2/5 of shell length. Axial sculpture of 8-9 prominent, slightly opisthocline, flexuous and narrowly rounded axial ribs, regularly spaced, with broader interspaces. Spiral sculpture of 915 major cordlets, with irregularly alternating smaller cordlets and interspaces of variable size. Each cordlet consists of a rows of densely packed rounded granules. Aperture narrow, ovate, about 1/3 of the shell height. Siphonal canal short, narrow and open, deviating on the left. Inner lip with a moderately developed parietal callus. Outer lip varicose. Anal sinus marked, arcuate on shoulder slope. 5 Animal with short head and two short tentacles. Black eyes on the external, thickened basal part of the tentacles, located on the distal third of their total height. Foot broad and long, slightly lobated anteriorly, tapering posteriorly. Background colour of the headfoot pinkish, semitransparent, with light yellow spots, and light yellow speckles on the proximal part of the tentacles. Siphon pinkish, semitransparent, with light yellow spots bordered by orange (Figs 33-34). Distribution Bela brachystoma is known from the entire Mediterranean Sea and the continental European and African coasts of the northeastern Atlantic. Fossil shells (“B. granulifera”) are known from several PlioPleistocene European outcrops (England, France, Spain), including Italian ones (Piedmont, Tuscany, Latium and Sicily) (Chirli & Richard, 2008). Remarks Bela brachystoma is a continental shelf species, easily distinguishable from all other members of the genus Bela s.l. by its very distinct shell sculpture. It has a multispiral protoconch with characteristic densely granulated spiral ribs. Fossil records of this species date back to Miocene (Chirli, 1997: 51), although records older than the Pliocene ones would probably deserve re-examination. Pleurotomum granuliferum was considered as a synonym of B. brachystoma by Monterosato (1872: 44; 1890: 186, “var. granulifera”) and Sabelli et al. (1992: 407). The examination of a large number of shells of P. granuliferum in the Monterosato collection (ex coll. Brugnone, MCZR) revealed that these shells are indistinguishable from B. brachystoma indeed. A lectotype for P. granuliferum is here designated from the type locality (Upper Pliocene-Lower Pleistocene of Ficarazzi, Palermo, Sicily, Italy) and is stored in MCZR (ex Brugnone, in Monterosato collection, no. 16980, three vials with three handwritten labels). Brugnone’s autographic labels refer to two forms (“var. parva” and “var. striiselevatioribus”) which have never been described, as far as we know, their names being therefore nomenclaturally unavailable according to ICZN (1999: art. 45.6.3-4). Nordsieck (1977: 44, pl. 11, fig. 86) described B. brachystoma apicalis, differing from the nominal subspecies in its paucispiral protoconch. Unfortunately, among the B. brachystoma s.l. material analysed we did not find any specimens fitting with Nordsieck’s description, nor could we examine the relevant type material. If it was confirmed that only the paucispiral protoconch allows distinction from B. brachystoma, we would rather consider it as a further 6 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO distinct species, sister to B. brachystoma, as in the cases of B. menkhorsti Van Artsen, 1988-B. zenetouae (Van Artsen, 1988) (cf. Mariottini et al., in press). Bela sp. (prope brachystoma) (Figs 78-90, 141-142, 162) Material examined 6 exx, Italy, Latium, Guidonia (RM), 42°00’N 12°43’E (Pliocene), CS-PM. Description Shell small for the genus, height 4.6-5.5 mm, width 1.8-2.3 mm, biconical, turriculate elongate, solid. Protoconch multispiral, dome shaped, of 2.4-2.5 convex whorls. First 1.60-1.70 apical whorls smooth, the remaining with reticulated sculpture of 4 major granulose spirals and 1 smaller granulose subsutural spiral crossed by oblique axial riblets. Maximum diameter of protoconch 610-620 µm. Protoconchteleoconch transition not well marked. Teleoconch of 5-6 whorls, weakly angulate, sutural ramp slightly concave, whorl sides gently convex. Last whorl about 2/3 of shell length. Axial sculpture of 7-8 prominent, slightly opisthocline, flexuous and narrowly rounded axial ribs, tending to fade out at the base, regularly spaced, with broader interspaces. Spiral sculpture of 9-15 major cordlets, with irregularly alternating smaller cordlets and interspaces of variable size. In each interspace one to three rows of rounded tiny granules, axially connected by a very fine threads. Aperture narrow, ovate, about half of the shell height. Siphonal canal short, narrow and open, deviating on the left. Inner lip with a moderately developed parietal callus. Outer lip varicose. Anal sinus marked, arcuate on shoulder slope. Colour of most shells whitish, one shell pale brownish. Distribution So far known only from the Pliocene outcrops of Guidonia (Latium, Central Italy). Remarks These shells are very similar to Recent Bela brachystoma. They differ in the more turriculated teleoconch outline, with a slightly larger shell length/width ratio mean (2.74 vs 2.61). The size of the protoconch is within the range of B. brachystoma, but with a higher number of the apical whorls (1.60-1.70 vs 1.30-1.40, cf. tab. 1). Given the great variation in shell features scored in B. brachystoma (Tab. 1), we refrain from describing this form as a new species until more material becomes available. It is worth noting that in the Pliocene outcrop of Guidonia we could not find any shells identifiable with the typical B. brachystoma, although we collected shells of a related species, Bela nitida Pavia, 1975 (Figs 81a-b), which frequently occurs sympatrically with B. brachystoma (Pavia, 1975; Cavallo & Repetto, 1992; Chirli, 1997, Chirli & Richard, 2008). Bela nitida shows several morphological characters of the shell, i.e. small size, teleoconch outline, protoconch sculpture, which are in close agreement with those of B. brachystoma, but it differs in the teleoconch sculpture, less scabrous, with fewer spiral cords and with smooth sutural ramp. Bela africana Ardovini, 2004 (Figs 91-93, 100, 103-110, 145-146, 164) Bela brachystoma africana Ardovini, 2004 (Malacologia 43: 8, fig. [unnumbered]) Original description. “La specie in esame [Bela exilis] appartiene ad un complesso di specie molto simili tra loro avente come riferimento Bela brachystoma (Philippi, 1844). La sottospecie africana di B. brachystoma presenta una protoconca più grande e più ampia con granulosità sul terzo giro sempre disposte su quattro file parallele più marcate e più definite. Il resto della conchiglia sempre percorsa da cordoncini spirali granulosi, fitti ma regolari, alcuni dei quali in rilievo (1 su 3). Le coste sono sinuose, circa sette per ogni giro. La colorazione è brunorossiccia uniforme, con banda mediana bruna. Peristoma e columella di colore scuro. Dovendo esaminare in ordine crescente secondo l’intensità di ornamentazione partendo da Bela brachystoma (Philippi, 1844), passando per Bela cycladensis (Reeve, 1845) e Bela brachystoma africana subsp. n., Bela exilis sp. n. si colloca all’estremo opposto, evidenziando caratteristiche morfologiche di specie ben distinta nell’ambito del gruppo.” Types depositories. Holotype stored in MMP; one paratype in RA. Type locality. Eastern Atlantic Ocean, Senegal, South of Dakar. Material examined Holotype, paratype and 12 exx (topotypes), Senegal, 30 miles South of Dakar, - 45 m (found mostly with the soft parts in the stomach of Astropecten sp.), CS-PM (topotypes). Description Shell very small for the genus, height 3.4-3.8 mm, width 1.5-1.6 mm, biconical, turriculate elongate, solid. Protoconch multispiral, dome shaped, of 2.6-1.7 convex whorls. Protoconch-I (embryonic shell) of BELA BRACHYSTOMA COMPLEX 0.4 whorls, separated by a demarcation from protoconch-II (larval shell). First 1.7-1.8 apical whorls apparently smooth, the nucleus with very fine striae, the remaining with reticulated sculpture of 5-6 granulose spirals (3 major, 1-2 smaller subsutural, 1 smaller suprasutural), crossed by oblique axial riblets. Maximum diameter of protoconch 760-780 µm. Protoconch-teleoconch transition not well marked. Teleoconch of 2.5-3 whorls, rounded, sutural ramp convex, whorl sides gently convex. Last whorl about 3/5 of shell length. Axial sculpture of 8-9 prominent, narrowly rounded axial ribs fading out at the base, regularly spaced, with equally sized interspaces. Spiral sculpture of 17-18 granulose cords, regularly spaces, with larger interspaces. Smaller granulose cordlets in most interspaces. Entire surface covered my microgranules. Aperture narrow, ovate, about 2/5 of the shell height. Siphonal canal short, broad and open, very slightly deviating on the left. Inner lip with a weak parietal callus. Outer lip not varicose. Anal sinus marked, arcuate on shoulder slope. Colour yellowish with white axial ribs, darker brown band in the middle of teleoconh whorl, base milk white, parietal callus brown. Distribution Only known from the type locality. Remarks This taxon was originally described as a subspecies of B. brachystoma by Ardovini (2004). Subsequently, Ardovini (2008), realising that it was worth of deserving species rank, redescribed it as a new species (as “Bela africana n. sp.”). Bela africana shows a larger and differently sculptured protoconch as compared to B. brachystoma (Tab. 1) and a teleoconch sculpture with less pronounced spiral ribs. Bela africana is also smaller and its colour different, being brownish-reddish between the whitish axial costae and on the inner lip, with a reddish band running in the middle of the spire. From B. exilis it differs in the less angulated outline, the larger protoconch, and different sculpture (Tab. 1). For differences with B. ardovinii n. sp., see below under this species. Bela exilis Ardovini, 2004 (Figs 94-96, 101, 111-118, 147-148, 165) Bela exilis Ardovini, 2004 (Malacologia 43: 8, figs [unnumbered]) Original description. “Bela exilis sp. n. è specie di piccole dimensioni, vivente in ambiente circa litorale 7 su fondali sabbiosi. La protoconca è paucispirale, composta da tre giri, i primi due apparentemente lisci, ma che, se osservati al microscopio, a forte ingrandimento, rivelano leggerissime pliche longitudinali che cedono il posto nel terzo giro a quattro file di tubercoli microscopici. La conchiglia nel complesso presenta una superficie percorsa da strie granulose più o meno evidenti, la sutura è poco marcata, la spalla leggermente marcata le coste poco numerose, ma evidenti in numero di sei circa per ogni giro. Il colore è brunoarancio uniforme più intenso presso la sutura e nella parte mediana.” Types depositories. Holotype in MMP; paratypes nos. 1-3 in RA. Type locality. Eastern Atlantic Ocean, Senegal, South of Dakar, - 45 m. Material examined Holotype, 3 paratypes and 18 exx (topotypes), Senegal, 30 miles South of Dakar, - 45 m (taken with soft parts from the stomach of Astropecten sp.), CS-PM (topotypes). Description Shell very small for the genus, height 3.4-3.6 mm, width 1.3-1.5 mm, biconical, turriculate elongate, solid. Protoconch multispiral, dome shaped, of 2.8-2.9 convex whorls. Protoconch-I (embryonic shell) of 0.70.8 whorls, separated by a demarcation from protoconch-II (larval shell). First 1.6-1.7 apical whorls apparently smooth, covered with microgranules, the remaining with reticulated sculpture of 4 granulose spirals (3 major, 1 smaller subsutural), crossed by oblique axial riblets. Maximum diameter of protoconch 710-720 µm. Protoconch-teleoconch transition not well marked. Teleoconch of 2.5-3 whorls, rounded, sutural ramp convex, whorl sides gently convex. Last whorl about 3/5 of shell length. Axial sculpture of 10-11 prominent, flexuous and narrowly rounded axial ribs, regularly spaced, with broader interspaces. Spiral sculpture of 2 major granulose cords, with 25-36 irregularly alternating smaller granulose cordlets and interspaces of variable size. Each cordlet actually consisting of a rows of densely packed rounded granules. Aperture narrow, ovate, about 2/5 of the shell height. Siphonal canal moderately long, broad and open, deviating on the left. Inner lip with a moderately developed parietal callus. Outer lip not varicose. Anal sinus marked, arcuate on shoulder slope. Colour yellowish-brownish with two dark brown bands, one subsutural and the second on the middle of the last whorl; parietal callus brownish with siphonal canal white. 8 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO Distribution So far known only from the type locality. specimens having been found mostly with the soft parts in the stomach of Astropecten sp. Remarks The teleoconch of B. exilis is smaller than that of B. brachystoma. The species differs from B. africana in the less angulated outline, the smaller protoconch (but with more whorls) and the different sculpture (Tab. 1). For differences with B. ardovinii n. sp., see below under this species. Remarks Variation of the paratypes: length 3.0-3.9 mm, width 1.0-1.6 mm, protoconch of 3.2-3.3 whorls, maximum diameter of protoconch 690-710 µm; teleoconch of 2.5-2.7 whorls, axial sculpture of 8-10 ribs, spiral sculpture with 20-28 cordlets. Bela ardovinii n. sp. differs from B. brachystoma by the smaller size of the adult, larger protoconch (in size and number of whorls) and different sculpture of the teleoconch (weaker spiral cords) and protoconch (with strong spiral cords). From B. africana it differs by the slightly smaller size, more angulated outline, uniformly reddish-brown colour (instead of brownishreddish between the whitish axial costae, with a spiral reddish band), and smaller protoconch (but with more whorls) with a different sculpture (Tab. 1). From B. exilis it mainly differs by the totally different colour pattern and smaller protoconch (but with more whorls) with different sculpture (Tab. 1). Bela ardovinii Mariottini & Oliverio n. sp. (Figs 97-99, 102, 119-126, 149-150, 166) Type material Holotype. Eastern Atlantic Ocean, Senegal, 30 miles South of Dakar, - 45 m, MNHN (no. 21321). Paratypes. 14 exx (labelled as A to N), same locality as the holotype, RA (A-F), CS-PM (G-J, L), MNHN (K, no. 21322) and MO (M-N). Description Shell very small for the genus, height 3.7 mm, width 1.4 mm, biconical, turriculate elongate, solid. Protoconch multispiral, dome shaped, of 3.2-3.3 convex whorls. Protoconch-I (embryonic shell) of 0.8 whorls, separated by a demarcation from protoconch-II (larval shell). First 1.0-1.1 apical whorls densely covered by microgranules, next 0.7-0.8 whorl apparently smooth, the remaining with reticulated sculpture of 5 spiral series of tubercles (3 major, 1 smaller subsutural, 1 smaller suprasutural), crossed by weak opisthocline axial riblets, more evident subsuturally. Spiral cords corresponding to each spiral series of tubercles gradually appearing on the last rotoconch whorl. Maximum diameter of protoconch 690-710 µm. Protoconch-teleoconch transition not well marked. Teleoconch of 2.5-2.7 whorls, rounded, sutural ramp convex, whorl sides very gently convex. Last whorl about 3/5 of shell length. Axial sculpture of 8-10 rounded axial ribs fading at base, regularly spaced, with narrower interspaces. Spiral sculpture of one major granulose cord, and 20-28 granulose cordlets, irregularly spaced. Spiral granulose threads in most interspaces. Entire surface covered by microgranules. Aperture narrow, ovate, about 2/5 of shell height. Siphonal canal short, broad and open, slightly deviating to the left. Inner lip with a weak parietal callus. Outer lip not varicose. Anal sinus marked, arcuate on shoulder slope. Colour uniformly reddish-brown. Distribution Only known from the type locality, the original CONCLUSION This study focused nearly exclusively on shell characters. We provisionally include Pleurotoma brachystomum Philippi, 1844 within the genus Bela s.l. The overall morphological analyses resulted in the recognition of several mangeliine species within the Recent and fossil material, that we consider as related to Bela brachystoma: 1) a Pliocene to Recent Mediterranean species, corresponding to the currently recognised B. brachystoma; 2) Bela sp. from the Pliocene of Guidonia (Latium, Central Italy), that may deserve a deeper study; and the Recent eastern Atlantic 3) B. exilis Ardovini, 2004; 4) B. africana Ardovini, 2004; 5) B. ardovinii Mariottini & Oliverio n. sp. ACKNOWLEDGMENTS We would like to express our gratitude to Alberto Zilli (Museo Civico di Zoologia di Roma), for the examination of the Pleurotomum granuliferum material kept in the Monterosato (ex Brugnone) collection and helpful advices; to Virginie Héros and Philippe Maestrati (MNHN) for the help in the study of the samples under their care; to Roberto Ardovini (Rome) for the generous gift of specimens of B. exilis, B. africana and B. ardovinii; to Angelo Vazzana (Museo di Biologia Marina e Paleontologia di Reggio Calabria) for sending fossil specimens of B. brachystoma, to Jakov Prkic (Split) for sending B. brachystoma from Croatia. Work partly funded by PNRA (2004/1.9 POLARTOX). 9 BELA BRACHYSTOMA COMPLEX 1 2 3 4 5a 5b 6a 6b 7a 7b 8a 8b 9a 9b 10a 10b 11a 11b 12a 12b 13a 13b Figs 1-13 Bela brachystoma. (1) Original drawing by Philippi (1844: pl. 26, fig. 10). (2) Drawing of B. brachystoma brachystoma by Nordsieck (1977: pl. 11, fig. 85). (3) Drawing of B. brachystoma apicalis by Nordsieck (1977: pl. 11, fig. 86). (4) Original label in Monterosato’s handwriting (no. 202.b.2, MCZR). (5a-b) Italy, Viareggio (PI), 43°51’N 10°11’E, 5.6 x 1.9 mm, Monterosato collection (no. 202.b.2, MCZR). (6a-b) San Vincenzo (LI), 43°05’N 10°24’E, - 34 m, 5.5 x 2.1 mm, CS-PM. (7a-b) Idem, 6.0 x 2.2 mm, CS-PM. (8a-b) Fiumicino (RM), 41°38’N 12°11’W, - 140 m, 7.3 x 2.5 mm, CS-PM. (9a-b) Idem, 41°43’N 12°06’W, - 80 m, 4.1 x 1.7 mm, CS-PM. (10a-b) Pescara, 42°31’N 14°12’E, 50 m, 3.3 x 1.4 mm, CS-PM. (11a-b) Libya, - 110/150 m, 4.0 x 1.7 mm, CS-PM. (12a-b) Spain, Estepona, 36°25’N 5°09’W, - 150 m, 5.1 x 2.1 mm, SR. (13a-b) Croatia, Brac Is., 43°24’N 16°30’E, - 50 m, 4.4 x 1.9 mm, CS-PM. 10 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO 14a 14b 15 16 14c 17 21a 21b 22 23 21c 24 27 20 18 25 19 26 30 29 28a 28c 28b 28d 31 32 33 34 Figs 14-34 Bela brachystoma. (14a-20) Italy, San Vincenzo (LI), 43°05’N 10°24’E, - 34 m, 5.2 x 1.8 mm, CS-PM. (21a-27) Idem, 4.1 x 1.7 mm, CS-PM. (28a-32) Ponza (LT), 40°51’N 12°55’W, - 40 m, 3.2 x 1.4 mm, CS-PM. (33-34) Linosa (AG), - 85 m, bioclastic muddy sand (living animal). BELA BRACHYSTOMA COMPLEX 11 35d 36 35a 35b 35c 37 38 39d 40 39a 39b 39c 41 42 43d 44 43a 43b 43c 45 46 47d 48 47a 47b 47c 49 50 Figs 35-50 Bela brachystoma. (35a-38). Libya, - 110/150 m, 3.3 x 1.5 mm, CS-PM. (39a-42) Italy, San Vincenzo (LI), 43°05’N 10°24’E, - 34 m, 3.2 x 1.6 mm, CS-PM. (43a-46) Pescara, 42°31’N 14°12’E, - 50 m, 3.6 x 1.3 mm, CS-PM. (47a-50) Civitanova Marche (MC), 43°18’N 13°46’E, - 50 m, 3.6 x 1.4 mm, CS-PM. 12 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO 52 51 53a 54 55 56 57a 53b 57b 59 58a 58b 58c 58d 60 61 62 63a 63b Figs 51-63 Bela brachystoma (fossil, = Pleurotomum granuliferum). (51) Original drawing by Brugnone (1862: pl. 1, fig. 13). (52) “Pl. granuliferum var. parva Ficarazzi”, original label in Brugnone’s handwriting (ex Brugnone, Monterosato collection, no. 16980, MCZR). (53a-b) Pleurotomum granuliferum, lectotype, Italy Ficarazzi (PA), 38°04’N 13°29’E, Pliocene, 5.7 x 2.2 mm, MCZR. (54) Idem, paralectotype, ex. A, 4.5 x 1.9 mm. (55) Idem, ex. B, 4.6 x 2.0 mm. (56) Idem, “var. striiselevatiorus. Ficar.”, Brugnone’s autographic label. (57a-b) Idem, ex. C, 4.5 x 1.8 mm. (58a-61) Idem, ex. D, 4.9 x 1.9 mm. (62) “Pl. granuliferum m. Pelle.”, Brugnone’s autographic label. (63a-b) Monte Pellegrino (PA), 38°13’N 13°19’E, Pliocene, ex. E, 4.8 x 1.9 mm, MCZR. 13 BELA BRACHYSTOMA COMPLEX 65 64a 64b 64c 64d 66 67 68 69 70 71 72 73 75a 76a 75b 74 77a 76b 77b Figs 64-77 Bela brachystoma (fossil, = Pleurotomum granuliferum). (64-74) Italy, Ficarazzi (PA), 38°04’N 13°29’E, Pliocene, ex. F, 4.6 x 1.9 mm, MCZR. (75a-b) Gallina (RC), 38°05’N 15°41’E, Pliocene, 2.8 x 1.3 mm, CS-PM. (76a-b) Cava Tacconi-Pomezia (RM), 41°38’N 12°28’E, Upper Pliocene-Lower Pleistocene, 5.5 x 1.8 mm, CS-PM. (77a-b) Terre Rosse (SI), 43°19’N 11°23’E, Pliocene, 4.5 x 1.6 mm, CS-PM. 14 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO 78a 81a 78b 79b 81b 82a 79a 82b 80a 80b 82c 82d 83 86 84 85 87 88 89 90 Figs 78-90 Bela spp. (78a-b) Bela sp. (prope brachystoma), Italy, Guidonia (RM), 42°00’N 12°43’E, Pliocene, ex. A, 5.1 x 1.8 mm, CS-PM. (79a-b) Idem, ex. B, 6.1 x 2.3 mm, CS-PM. (80a-b) Idem, ex. C, 5.5 x 1.8 mm, CS-PM. (81a-b) Bela nitida, Italy, Guidonia (RM), 42°00’N 12°43’E, Pliocene, 4.8 x 1.9 mm, CS-PM. (82a-90) Bela sp. (prope brachystoma), Italy, Guidonia (RM), 42°00’N 12°43’E, Pliocene, ex. D, 4.8 x 1.8 mm, CS-PM. BELA BRACHYSTOMA COMPLEX 15 91a 91b 92a 92b 93a 93b 94a 94b 95a 95b 96a 96b 97a 97b 98a 98b 99a 99b 100a 100b 101a 101b 102a 102b Figs 91-102 Bela spp. (91a-b) Bela africana, West Africa, Senegal, 30 miles south of Dakar, - 45 m, ex. A, 3.8 x 1.6 mm, CS-PM. (92a-b) Idem, ex. B, 3.7 x 1.6 mm, CS-PM. (93a-b) Idem, ex. C, - 45 m, 3.4 x 1.5 mm, CS-PM. (94a-b) Bela exilis, West Africa, Senegal, 30 miles south of Dakar, - 45 m, ex. A, 3.6 x 1.4 mm, CS-PM. (95a-b) Idem, ex. B, 3.4 x 1.3 mm, CS-PM. (96a-b) Idem, ex. C, 2.8 x 1.3 mm, CS-PM. (97a-b) Bela ardovinii, holotype, West Africa, Senegal, 30 miles south of Dakar, 3.7 x 1.4 mm, MNHN. (98a-b) Idem, paratype G, 3.5 x 1.3 mm, CS-PM. (99a-b) Idem, paratype J, 3.2 x 1.2 mm, CS-PM. (100a-b) Same as figs 93a-b. (101a-b) Same as figs 95a-b. (102a-b) Same as figs 99a-b. 16 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO 103 104 105 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 107 106 Figs 103-126 Sculpture details of the protoconch of Bela spp. (103-110) B. africana, same as figs 93a-b. (111-118) B. exilis, same as figs 96a-b. (119-126) B. ardovinii, same as figs 99a-b. BELA BRACHYSTOMA COMPLEX 17 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 Figs 127-150 Sculpture details of the protoconch of Bela spp. (127-128) B. brachystoma, same as figs 35a-d. (129-130) Idem, same as figs 39a-d. (131-132) Idem, same as figs 43a-d. (133-134) Idem, same as figs 47a-d. (135-136) Idem, same as figs 28a-d. (137-138) P. granuliferum, same as figs 58a-d. (139-140) Idem, same as figs 64a-d. (141-142) Bela sp. (prope brachystoma), same as figs 82a-d. (143-144) Bela menkhorsti, Tunisia, coast off of Sfax, - 90 m, 2.6 x 1.4 mm, CS-PM. (145-146) B. africana, same as figs 93a-b. (147-148) B. exilis, same as figs 96a-b. (149-150) B. ardovinii, same as figs 99a-b. 18 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO 153 154 152 151 155 156 157 158 159 160 161 162 163 164 165 166 Figs 151-166 Outline of the teleoconch of Bela spp. (151) B. brachystoma, same as figs 14a-c. (152) Idem, same as figs 21a-c. (153) Idem, same as figs 32a-d. (154) Idem, same as figs 28a-d. (155) Idem, same as figs 35a-d. (156) Idem, same as figs 43a-d. (157) Idem, same as figs 47a-d. (158) P. granuliferum, same as fig. 54. (159) Idem, same as fig. 55. (160) Idem, same as figs 58a-d. (161) Idem, same as figs 64a-d. (162) Bela sp. (prope brachystoma), same as figs 82a-d. (163) Bela menkhorsti, same as fig. 143. (164) B. africana, same as figs 93a-b. (165) B. exilis, same as figs 96a-b. (166) B. ardovinii, same as figs 99a-b. BELA BRACHYSTOMA COMPLEX Variables Bela brachystoma Recent “P. granuliferum” Protoconch diameter (mm) 510-650 Protoconch number of whorls 19 Bela sp. (prope brachystoma) Bela africana Bela exilis Bela ardovinii 620-650 610-620 760-780 710-720 670-680 2.3-2.4 2.3-2.6 2.4-2.5 2.6-2.7 2.8-2.9 3.2-3.3 1.3-1.4 apical smooth, remaining reticulated consisting of 4-5 granulose spirals (3 major in the middle of whorl, 1 smaller subsutural and 1 smaller above the teleoconch suture) crossed by oblique axial costae 1.4-1.5 apical smooth, remaining reticulated consisting of 4-5 granulose spirals (3 major in the middle of whorl, 1 smaller subsutural and 1 smaller above the teleoconch suture) crossed by oblique axial costae 1.60-1.70 apical smooth, remaining reticulated consisting of 4 major granulose spirals and a smaller subsutural granulose spiral crossed by oblique axial costae 1.7-1.8 apical smooth (nucleus with fine striae visible at high magnification), remaining reticulated consisting of 5-6 granulose spirals (3 major in the middle of whorl, 1-2 smaller subsutural and 1 smaller above the teleoconch suture) crossed by oblique axial costae 0.7-0.8 apical showing strong microgranules, next 0.3-04 whorl partially smooth with few scattered microgranules, remaining reticulated consisting of 4 granulose spirals (3 major, 1 smaller subsutural) making a sort of cordelets in the last whorl, crossed by oblique axial costae, microgranules well ordered in row on the spirals, scattered on the rest of protoconch Teleoconch axial folds (last whorl) 1.0-1.1 apical showing strong microgranules, next 0.7-08 whorl smooth, remaining reticulated consisting of 4 strong spiral cords (3 major, 1 smaller subsutural) crossed by oblique axial costae that are slightly aculeated, thinner and more numerous reaching the suture, microgranules scattered, densely grouped and ordered in cordelets on the spirals 8-9 8-9 8-9 8-9 10-11 7-8 Shell length range (mm) 4.5-7.0 4.5-6.0 4.6-6.1 3.4-3.8 3.4-3.6 3.2-3.7 Shell length/width ratio (range, mean and SD) 2.4-2.9 2.61 (0.05) 2.2-2.8 2.48 (0.04) 2.6-3.0 2.74 (0.05) 2.3-2.4 2.31 (0.03) 2.5-2.6 2.57 (0.02) 2.6- 2.7 2.66 (0.02) reddish-brown, yellowish-brown, whitish ? ? yellowish with white axial ribs, darker brown band in the middle of teleoconh whorl and remaining basal part milk white, peristome brown yellowish-brownish with two dark brown bands, one covering the suture and the second in the middle of the last whorl, peristome brownish with siphonal canal white Protoconch sculpture Shell colour reddish-brown Tab. 1 Shell morphological features of Bela spp. REFERENCES ARDOVINI R., 2004. Due nuove specie e una nuova sottospecie di Turridae dal Senegal, West Africa. Malacologia 43: 7-9. ARDOVINI R., 2008. Bela africana n. sp. (Gastropoda Turridae) West Africa, Senegal. Malacologia 58: 12-13. BOUCHET P., ROCROI J.-P., 2005. Classification and Nomenclature of Gastropod Families. Malacologia 47: 1-397. BRUGNONE G.A., 1862. Memoria sopra alcuni pleurotomi fossili dei dintorni di Palermo. Stabilimento Tipografico F. Lao, Palermo, 42 pp. CAVALLO O., REPETTO G., 1992. Conchiglie fossili del Roero. Associazione Naturalistica Piemontese Amici del Museo “F. Eusebio”, Alba, 129 pp. CHIRLI C., 1997. Malacofauna Pliocenica Toscana. Superfamiglia Conoidea. Vol. 1. B.M.B., Firenze, 92 pp. CHIRLI C., RICHARD C., 2008. Les mollusques plaisanciens de la Côte d’Azur. Devaye Imprimeurs, Cannes, 128 pp. GOFAS S., LE RENARD J. (eds), [2007]. Check List of European Marine Mollusca. http://www.somali.asso.fr/clemam/index. clemam.html [downloaded III.2007]. GRAY J.E., 1847. A list of the genera of Recent Mollusca, their synonyma and types. Proceedings of the zoological Society of London 15: 129-219. ICZN (International Commission on Zoological Nomenclature), 1999. International Code of Zoological Nomenclature. 4th edn. The International Trust for Zoological Nomenclature, London, XXIX + 306 pp. KOHN A., 1998. Superfamily Conoidea. In: Beesley P.L., Ross G.J.B., Wells A. (eds), Mollusca: the Southern Synthesys. Fauna of Australia 5 (B). CSIRO, Melbourne, pp. 846-854. MARIOTTINI P., SMRIGLIO C., DI GIULIO A., OLIVERIO M. (in press). A new fossil conoidean from the Pliocene of Italy, with comments on the Bela menkhorsti complex (Gastropoda: Conidae). Journal of Conchology. MONTEROSATO T.A., 1872. Notizie intorno alle conchiglie fossili di monte Pellegrino e Ficarazzi. Tip. Amenta, Palermo, 44 pp. MONTEROSATO T.A., 1890. Conchiglie delle profondità del Mare di Palermo. Il Naturalista siciliano 9: 157-166. NORDSIECK F., 1977. The Turridae of the European Seas. La Conchiglia, Roma, 131 pp. 20 PAOLO MARIOTTINI, ANDREA DI GIULIO, CARLO SMRIGLIO, MARCO OLIVERIO PAVIA G., 1975. I molluschi del Pliocene inferiore di Monteu Roero (Alba, Italia NW). Bollettino della Società di Paleontologia italiana 14: 205-226. PHILIPPI R.A., 1844. Enumeratio Molluscorum Siciliae. Vol. II. Tipografia Saxorum, 298 pp. POWELL A.W.B., 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Museum 5: 1-157. PUILLANDRE N., SAMADI S., BOISSELIER M.-C., SYSOEV A.V., KANTOR Y.I., CRUAD C., COULOUX A., BOUCHET P., 2008. Starting to unravel the toxoglossan knot: molecular phylogeny of the ”turrids” (Neogastropoda: Conoidea). Molecular Phylogenetics and Evolution 47: 1122–1134. SABELLI B., GIANNUZZI-SAVELLI R., BEDULLI D., 1992. Catalogo annotato dei Molluschi marini del Mediterraneo 2. Libreria Naturalistica Bolognese, Bologna, 150 pp. TAYLOR J.D., KANTOR Y.I., SYSOEV A.V., 1993. Foregut anatomy, feeding mechanisms, relationships and classification of the Conoidea (= Toxoglossa) (Gastropods). Bulletin of the natural History Museum of London (Zoology) 59: 125-170. TUCKER J.K., 2004. Catalog of Recent and fossil turrids (Mollusca: Gastropoda). Zootaxa 682: 1-1295. AUTHORS’ ADDRESSES Paolo Mariottini Andrea Di Giulio Dipartimento di Biologia, Università degli Studi “Roma Tre” Viale Marconi 446, I-00146 Rome, Italy e-mails: mariotpa@uniroma3.it; digiulio@uniroma3.it Carlo Smriglio Via di Valle Aurelia 134, I-00167 Rome, Italy e-mail: carlo.smriglio@deagostini.it Marco Oliverio Dipartimento di Biologia Animale e dell’Uomo, Università di Roma “La Sapienza” Viale dell’Università 32, I-00185 Rome, Italy e-mail: marco.oliverio@uniroma1.it