FAU Institutional Repository http://purl.fcla.edu/fau/fauir This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute. Notice: ©1990 The Bailey-Matthews Shell Museum. This author manuscript appears courtesy of The Nautilus, a peer-reviewed, not-for-profit quarterly published by the non-profit organization The Bailey-Matthews Shell Museum. The published version is available at http://shellmuseum.org/nautilus/index.html and may be cited as: Harasewyeh, M. G. (1990). Studies on bathyal and abyssal buccinidae (Gastropoda: Neogastropoda): 1. Metula fusiformis Clench and Aguayo, 1941. The Nautilus, 104(4), 120-129. o THE NAUTI LUS 104(4):120-129, 1990 Page 120 Studies on Bathyal and Abyssal Buccinidae (Gastropoda: Neogastropoda): 1. Metula fusiformis Clench and Aguayo, 1941 M. G. Harasewych Department of Invertebrate Zoology National Museum of Natura l History Smithsonian Institution Washington , DC 20560, USA ABSTRACT fact that the vast majority of taxa are based exclusively on features of the shell and operculum, supplemented Based on the morphology of the radu la and shell, Metula [u­ occasionally by observations on radu lar morphology. sifo rmis Clench & Aguayo, 1941 is transferred to the predom­ inantl y Indo-w estern Pacific genus Manaria . This species occurs Shells of Buccinid ae tend to be simple, and offer few in upper continental slope communities (183- 578 m) of the readily discernible morphological characters. These are Caribbean Sea and the northern coast of South America . The subject to convergence, especia lly in polar regions and holotype was collected dead in 2,633 rn, well below the depth the deep sea, where effects of habitat on shell form are inhabited by this species. The large well-developed gland of most prono unce d (Graus, 1974). Leiblein, a sepa rate sperm ingesting gland between the capsule Detailed ana tomical data are available for compara­ gland and albumin gland , and three-cusped rachidian teeth are tively few, mainly shallow-water taxa (e.g., Dakin, 1912; features that Manaria shares with other fusiform buccinids Golikov, 1963, 1980; Kosuge, 1967; Ponder, 1973; Lus, (e.g., Penion, Serratifusus ) as well as with primitive members 1981; Kantor, 1990). The lack of well-defined, synapo­ of other fami lies within Muricacea. These features are inter­ preted as being symplesiomorphic, and suggest that the fusi­ mor phic anatomical features (other than radular mor­ form buccinids are among the more primitive members of the phology), even between the fami lies Buccinid ae, Nas­ Buccinidae. sariidae, Fasciolariidae, and Melongenidae have led Ponder (1973a) to suggest these groups might all be con­ Key words: Buccinidae ; Caribbean ; bathyal; Manaria; Me­ sidered subfamilies of Buccinidae. This arra ngemen t was tula . subsequently adopted by Ponder and Waren (1988). Bouchet and Waren (1985) revised the deep-water Buccinid ae (sensu Wenz, 1943) of the northeastern At­ INTRODUCTION lant ic Ocean , and later (Bouchet & Waren, 1986) re­ viewed many of the tropical deep-water species. Despite The family Buccinidae comprises one of the most diverse these significant contributions, most of the nearl y 200 and dominant groups of predatory prosobranch gastro­ supraspecific taxa within Buccinidae (sensu Wenz, 1943) pods at high latitudes and at bath yal, abyssal and hadal are poorly defined, and the assignme nt of many species depths. It is represented in the fossil record of the Lower to genera remain tentative. Cretaceous (Albian), and ranks among the oldest of the Among the taxa listed by Bouchet and Waren (1986) neogastropo d families (Taylor et al., 1980). Like most as "insuffi ciently known" is Metula f usiform is Clench predatory prosobranch fami lies, it is believed to have and Aguayo, 1941. The placement of this species in Met­ evolved in tempera te climatic zones at higher latitudes ula was disput ed by Olsson and Bayer (1972) who sug­ (SohI, 1987). Although the majority of these families be­ gested that it had affinities with Fusin us or a fusiform came predominantly tropical during the Cenozoic, most buccinid. Abbott (1974) referred this species to the genus Buccinidae remained in temperate and polar regions , Bartschia. Bouchet and Waren (1986) considered it to were the family dive rsified since the late Miocene (Taylor be a buccinid, and commented on its conchological re­ et al., 1980). The success of Buccinidae at high latitudes semblance to Euthriostoma. and in the deep sea has been attri buted to their broad During a recent dive aboa rd the research submersible habitats and diets, which are considered to be adaptations Johnson-Sea-Link I off Navassa Island, situa ted off the to unpredictable resources (Ta ylor, 1978). southwestern peni nsula of Ha iti, the author had the op­ Despite the high diversity and abundance of Buccin­ port unit y to observe and collect several living specime ns idae, the systema tics of this group is poorly understood of "Met ula" fu siformis. These observations, together with at all taxonomic levels. This is due, in large par t, to the data from additional material discovered in the USNM \ M. G. Harasewych , 1990 Page 121 collections, form the basis of this report , the first in a Tabl e 1. Shell characters used for phenetic analysis. Char­ series on enig matic deep- water buccinid taxa. acters 1 through 8 describe the geometry of the generalized shell form (Harasewych, 1982). MATERIALS AND METHODS 1. Shape of the gene rating curve of the body cavity (Sbc). 2. Shape of the generating curve of the siphonal canal (Sse). Five speci me ns of "Me tula" fu siformis Clench and 3. Relative siphonal length (RsI). Aguayo, 1941 were observed, record ed on videotape and 4. Siphonal angle (fl). collected either in (1 spec ime n) or within 2 meters (4 5. Angle of the gene rating curve (0). specime ns) of a bucket baited with decomposing octopus 6. Rate of whorl expansion (W). and set on an ooze-covered area (slope about 20°) off the 7. Position of the genera ting curve relative to the axis (D). 8. Hate of whorl translation (T). west coast of Navassa Island (l8°24'42"N, 75°03'OO"W ) 9. Spire angle (a ). at a depth of 578 m for 50 hours. Th e spec ime ns, which 10. Number of axial ribs on four th teleoconch whorl (no. rib). were moribund upon reaching the surface, were fixed in 11. Number of spiral cords on fourth teleoconch whorl (no. 10% neutral buffered formalin and stored in 70%ethanol cord). until dissection. Phenetic analyses wer e used to assess the relationships of three conchologically similar taxa, each proposed on the basis of a single specimen . All specim ens listed in the conch gradual, marked by formation of axial ribs, fol­ " ma terial examined" section, as well as the holotype of lowed within 112 whorl by the formati on of six fine spiral Buccinofusus suriname nsis Okutani, 1982 and two spec­ cords. Teleoconch of up to 8% conve x whorls, rounded imens of a southern variant of Buccinu m cane tae Clench at first, becoming sharply shouldered by the fifth post­ and Aguayo, 1944, described as Plicijusus jamarci Oku­ nucl ear whorl. Suture broadly adpressed. Axial sculpture tani , 1982, were scored for the 11 shell charac ters listed of broad , rounded, regularl y-spaced, axial to slightly pro­ in table 1. Th ese data were standardized (mean = 0, socline ribs that do not extend onto the an teriormost standa rd deviation = 1), a Euclidean distance matrix portions of the body whorl or the siphonal canal. Axial calculated, and a phenogram based on the UPGMA clus­ ribs number 11-12 on the first and 11-16 on the penul­ tering algorithm was produced using SYSTAT version tim ate whorl. Spiral sculpture of strong cords, as broad 4.0 (Wilkinson, 1988). A Principle Component Analysis or broader than intervening spaces, that overlay axial using the same data matrix (25 spec imens x 11 char­ ribs. Cords number 12-13 between suture and shoulder, acters) was performed , also using SYSTAT, and the in­ 19-21 betw een should er and siphonal canal, 16-18 on dividu als plott ed using the first two pri ncipal components siphonal canal. Sixteen to 21 cords rem ain exposed on as axes. penultimate whorl. Aperture elliptica l, tap ering poste­ Repositories of examined spec ime ns are indi cated by riorly beneath suture to form anal sulcus. Outer lip with the following abbreviations: 18-23 thin spiral Iirae pronounced beneath axial ribs and weak or absent between. Inner lip smooth, with thin, MCZ-Museum of Comparative Zoology, Harv ard Uni ­ porcellaneous inductura. Columella solid, sinuate, lack­ versity ing folds. Siphonal canal broad , slightly shorter than ap­ NSMT-National Science Museum, Tok yo erture, crossing coiling axis. Siphonal fasciole weak, ad­ USNM-Nationa l Museum of Natural History, Smith­ jacent to colume llar edge of siph on. Exterior surface of sonian Institution shell dull ivory to light amber, ape rture and columella whit e. Periostra cum (figure 9) thi ck, straw-colored to SYSTEMATICS brown, consisting of thin, axial blad es that are broadest be tween spiral cords and abrad ed along their surfaces. Family Buccinidae Rafinesque, 1815 Operculum (figures 2, 13, op) thick, elongate, with ter­ Genus Manaria E. A. Smith, 1906 minal nucl eus (usually abrade d), attached along slightl y Manaria f usiform is (Clenc h & Aguayo , 1941) less that lh of its inner surface , glazed along posterior (figures 1-5, 7- 17) and left inner margins, fills aperture V4 whorl from out er lip. Met ula f usif ormis Clenc h & Aguayo, 1941:179, pl. 14, fig. 1; Bouchet & Waren, 1986:485, fig. 116. Shell ultrastructure: (figure 10) She ll composed of thr ee " Me tula" fusifo rmis Clench & Aguayo.-Olsson & Bayer, 1972: layers of crossed-lame llar crystals and an outermost pris­ 925.