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On the morphology and relationships of some oliviform gastropods
ARTICLE · JANUARY 1991
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Yu I Kantor Severtsov Institute of Ecology and Evo…
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Available from: Yu I Kantor Retrieved on: 31 December 2015 16 A.A.lliHJJelli Illi1JIE:tiKO ¡, A 1986. CwcreMa H <]lWIOreHHH Vltrlnldae SOLEM A. 1966. Sorne non-marine moJiuscs fr()m Thalland, Ruthenica (1991), 1(1-2): 17-52. © Ruthenica, 1991 (Gaslropoda Pu1monata). B KHHre: Mopt/JoJWzwrecKue u with notes on classificatlon of the Helicarionidae. Spolia 3KOJlOlUIWCKUC OCII06bl CUCnteMQtnUKU MOIVliOCKOS, Tpy Z{J(J/oglca Musei haurliensis, 24: 7-11 O. iJbl300JWZUtWCI(.ot0t uncmumyma AH CCCP, 148: 124- SOLEMA. 1976. Endodontoid landsnaiisjromPacific Islands 157. (Mollusca: Pulmonata: Sigmurethra). Part l. Family En BAKER H.B. 1928. Minute American Zonltldae. Proceedings o/ dodoniidae. Chlcago, 508 p. the Academy of natural Sciences, Philadelphia, 80: 1-44. SOLEMA.1982. Endodonioid /aJul snai/sjrom Pael/le!slands BLANFORD W T., GODWlN-AUSTEN H.H.1908. The fmma (Mollusca: Pulnwnata: Sigmurethra). Part JI. Families of British India, including Ceylon and Burma. Mollusca. Punetidae ami Charopidae, Zoogeography. Chlcago, 336 p. Testacellidae and Zmtitidae. London, 311 p. Van GOETHEM J.L. 1977. Revislon systematique des Uro On the morphology and relationships of sorne HOFFMANN H. 1941. Anatomische und systematlsche Unter cycJinae (Mollusca, Pulmona1a, Urocyclldae). Annales suchungen uber die Parmarioninen (Gastr. Putm.). Zoolo Musee Royal de l'Afrique Centra1e, Tervuren, Belgique, olivifonn gastropods gischen Jahrbucher. Abteilung jür Systematik, i:Jkalogie serlelN-8, 218: !-355. und GeographiederTiere, 74(1-2): 1-156. Van MOL J.-J. 1970. Revlslon des Urocyclldae (Mollusca, Ga RIEDEL A. 1980. Genera Zonitidarum. Rotterdam, 197 S. stropoda, Pulmonata). Annales Musee Royal de t Afrique YURI I. KANTOR SCHILEYKO A.A. 1989a. Taxonmnlc status and phylogenetic Centra/e, Tervuren, Belgi K MOp ID.H.KAHTOP Hncmumym OOOJ!IO!iUOHHOU .owp,PoJUJzuu u aJw;weuu :«ueomnh!X u.u. A.H.Ceeep~oea AH CCCP, MocJCea 117071, Jfenunc~euu npocneiCm 33 PE3IDME. Ha pyqHoM ncKpnmm: 11 cepHiíHhlx cpeaax HCCJICJIOBaHa Mop The family Olividae Latreille, 1825 (sensu la Pseudolivinae Cossmann, 1901 [Ponder, Warén, to) is a rather complex and heterogeneous group, 1988]. which includes five subfamilies: Olivinae Latreil The taxonomic status of severa! subfamilies is le, 1825; Olivellinae Troschel, 1869; Ancillinae still unclear and sometimes they are considered Swainson, 1853; Agaroniinae Olsson, 1956, and to be separate families. For instance, Marcus and Marcus [1959] studied the anatomy of dif- 18 Yu.I.Kantor Morphology and relationships of oliviform gastropods 19 ferent olivids and found clear differences betwe Oliva sayana Ravenel, 1834 [morphology of en Oliva and Olivel/a. Bandel [1984], following the oesophagus only ] Marcus and Marcus [ 1959], considered it Subfamily Olivelliñae necessary to separate Olivella and Oliva at least Olivel/a borealis Golikov, 1967 at the subfamily level. Golikov and Starobogatov Subfamily Ancillinae [1975] erected the new family Olivancillariidae Amalda montrouzieri (Souverbie, 1860) (type-genus 0/ivancillaria d'Orbigny, 1841, by Subfamily Pseudolivinae ' original designation) on the basis of morpholo Pseudoliva zebrina A.Adams, 1853 gical features of the radula and female reproduc Pseudoliva ancilla Hanley, 1859 tive system. Final! y, Golikov and Starobogatov Melapium lineatum (l.amarck, 1822) [1988] established the separate family Olivelli Benthobia tryoni Dall, 1889 dae (as Olivellidae Golikov et Starobogatov, 1988 In studying anatomy both dissections and = Olivellidae Troschel, 1869 = Olivellidae Olsson, sectioning have been u sed. I usually tried to com 1956, as subfamily) and referred it to new sub bine both methods for each species 'but in the order Olivelloidei Golikov et Starobogatov. The cases of very small animals (e.g. Benthobia try position of Pseudolivinae is also indefinite. Ols~ oni) reconstruction of the anatomy was made on son [1956] stated that the systematic position of ly on the basis of serial sections. Sections 8-1 O Pseudoliva and Zemira is questionable and that pm thick were obtained after routine techníque most authors listed Pseudoliva among Buccina and stained with Mallory or Masson triple stains. cea. Golikov and Starobogatov [1975] erected a family Pseudolividae and placed it in the super For clarífying phylogenetic interrelationships family Buccinoidea. a cladistic analysis was conducted. The proce The suprageneríc classification of olivids is dure is descríbed in detail in the corresponding usually based on conchological and radular fea section of the paper. turcs, and on some characters of the gross mor phology of the soft body. Only a few authors des OB03HA'IEHMSI HA PHCYHKAX. cribed the anatomy of severa! representatives of the family in detail [Küttler, 1913; Marcus, Mar cus, 1959; Marcus, Marcus, 1968, and some ag - auan&HaSI :>Keneaa; amt - nepe.uHec MRHTHlfnoe others ]. llzyflaJlbl(Cj aog - )lOITOJlHHTeJlhHRSl flH~CBOJt.Hast >KCJle3Rj 1 had the opporiunity of examining the ana asg - .nononmnenhuan cJttoHHaH >Keneaa; be - 6yKKaJihHaR tomy of different olivids belonging to the genera flO/IOCTh; btn - 6yKK3JlbHast Macea; bt - 6yKKaJibl!Ril Tpy6- Oliva, Olivella, Pseudoliva, Melapium, Benthobia Ka; C -- U..CKyM; CID - KOJIYMCIDH1pHbH1 M)'CKYJJ; Ct - KTC and Amalda. The comparative study of these ge HH)lHii; CU - K)'TMKyna; daS _;._ OpOTOK ,l\OflOJHHfTCJJbHOii nera allowed meto shed light on the relationships CJHOUHOif >KCJ1e3bi; dd - npoTOK nl1lll,CBapHTeJlbHOfi >KeJIC3bt; between different olivid subfamilies. dg - flHUI,CBBpHTeJibHaSI >I FIG. 1 (fronting page). 4.-F- Shells of the species s1udied: A-B- MelapitJm l(neatum• (Lamarck, 1822} (A- female with 2 egg capsules, shelllength 24.0 mm; B ~ male, shelllength 25.5 mm); C- 0/lvella borealis Golikov, 1967, shelt Jength 9.0 mm; D Oliva bulbosa (ROding, 1798), shelt length 22.7 mm; E- Pseudo/iva zebrinaA.Adams, 1853, shelllength 10.2·mm; F-G Benthobia tryoni Dall, 1889, shelllength 8.6 mm. PMC. l. A-F- PaKoamJbl HCCJICAOnat-mhlx »H~oa: A-B- Melapium lineatum (Lamarclt, 1822) (A- caMKa CJ1BYMR nHu.enwMH KancynaMH, 8biCOTa paKOBHHbl 24.0 MM; B- caMCII., DblCOTa paKOBHUbT 25.5 MM); e- Olivella borea/is Golikov, 1967. BbiCOTa paKODHHbl 9.0 MM; D- Oliva bulbosa (ROding, 1798), nr.1coTa paKonuuht 22.7 MM; E- Pseudo/iva zebrina A.Adams, 1853, nMcora pa~ pl :amt A ··: ' .. : ·.': . ' .. ': ::·:>...... ' ... ··.· ...... ' .. . ·.::' ' ...... '. ' ... . . : . .. . -:>· ...... '. B FIG.3. A- Crawllng 0/ivel/a borealis Golikov; B- positlon of the mollusc into the sediments PHC. 3. A- nonayu.tan: Oliveila boreaUs Golikov; B- nonomeuue MOJlJHOCKa n rpynTe. FIG. '2 (frontirtg page). A- longitudinal section through the muscular bu lb of the accessory salivary gland of Belltlwbia tryoni, scale B - 0.5 mm; B- sectiou through the oesophagus and accessory ocsophageal gland of Melapium lineatum, scale- 1 mm. PMC. 2. A- npo¡¡:OJIDHMH cpca •tepea M)'CKYJlhHhlH: MClliOK J{OHOJ1HHTCJlbHofi CJlfOHHOft me.ne3hJ JJenthobia tryoni, r.tacuna6 0.5 MM; B- Cpe3ttCpe3 liHU~CUOl{ H f{OUOJJHt-fTCJibliYIO mmtCBOAIIY[{) >K8JIC3Y Me/apium fineatum, MUCtuTaÓ- 1 MM. 22 Yu.I.Kantor Morphology and relationships of oliviform gastropods 23 FIG 4. Morphology of the soft body of 0/ivella horealis Gollkov. A,B- body, removed from the shell; C- removed visceral mass; D - scheme of the positlon and attachment of the columellar muscle, the Jnner volume of the shell dotted, the remnants of the inner walls white; E- mantle. Scales 1 mm. PJ{C. 4. Mop4JonornSI MSU'KOOO Tena 0/ive/la borealis Golikov. A,B - TeJIO, M3BJIClJCHHOC H3 p8KOBHHbli e - om:eneHHbU~:f BHCL(C pMbHbtli MCIUOKj D - CXCM8 pacnoJIO>KCHlUI H npHKpefiJICHJUt KOJIYMM'JJHpHOI'O M)'CK)'JIR, BH)'tpCHHHSJ nOJIOCTh paKOBHIIbl 3aTOlJKomma, ocraswaHCst tJaCT& BH)'TJWHHHX CTeHOK paKO»HHbt 6ena~I; E- MI\HTHH. Macurra61 MM. )KeJie3aj Sh ~ raCTpH'lecKHli UI,HTOKj St - >KeJ!Y,ll;OKi t - probaseis; prr - proboscis retractors; r ~ rectum; rd - nujmooon«; tf- nonepettHafl CKJJap;Ka; tn - ronoott&ie uzy radula; rdv - radular divertlculum; rn - rhynchostome; nanbu,a; tr - cne.n.bl CmipMbHOÜ CKYJJbnT)'pbl¡ vl - KJJanau ro - renal openlng; rr - radular retractar; rs - radular Jiel16neifna; vs - vesicula semlnalls. sac; rw - remnant of hmer wall of the pénultlmate whorl; s - slphon; sa - sorting arca; se - subradular cartllages; KEY TO ABBREVIATIONS IN FIGURES sd - sallvary duct; sg- salivary gland¡ sh - gastlic shield; st - stomach; t - typhlosoles; tf - transverse fold; tn - ag- anal gland; amt- anterior mantle tentacle; aog - cephallc tentacles; tr - traces of the spiral sculpture; vt - valve of Lelblein; vs - vesicula semlna/is. accessory oesophageal gland; asg- acccssory salivary gland; be - buccal cavlty; bm - buccal mass¡ bt - buccal tube; e - caecum; cm ~ coiumellar muscle¡ ct - ctenldlum; cu - DESCRIPTIONS OF MORPHOLOGY cuticle; das - duct of the accessory salivary gland; dd - duct of the dlgestlve gland; dg - dlgestive gland¡ di - duct Olivella borealis Golikov, 1967 of the gland of Leibfeln; e - eye; el - eye-lohe; f - IliG. S. MoYphology of the digeslive system of 0/ivella borealls Golikov. A-scmidiagrammatic longitudinal section of the anterior p~rt ofthe dlgestive system, nervous rlnz not shown (scale 0.5 mm); B-scmidiagrammatic transvcrse section ofthc anterior part of the cephalic flap; g - gona~; gr ~ glandular foJds of the oeso saHdy bottom. Most of the time these molluscs borealis never feed on Bivalvia [Kantor, Kiyash are buried in sand and are able to move rather ko, 1990 ]. rapidly through the sediments. In aquarium Thus rather different modes of feeding and observa1ions the mean velocity of movement in diet was found amongst Olivella species. Accor the sediments is 20 cm/h (range 4-40 cm/h). ding to Marcus and Marcus [1959] O. verreauxii Sometimes molluscs without apparent cause feed on bivalves, swallowing the prcy entirely. appeared on the sediment surface, where they Thc strange feeding mechanism was described can move more rapidly. for O. columellaris [Seilacher, 1959]. The latter The foot is broad, thin and semitransparent. . species projects the head above the surface of se The propodium is of crescent form and divided by diments. Mucous "bags" are spun from the pro a longitudinal cleft. The pedallobes are well de podium and billow out in the swash of receding veloped and embrace the shelllaterally (Fig. 3). waves. Periodically, the nets are drawn forwards When the mollusc withdraws the pedallobes are and probaseis reaches from behind to ingest greatly reduced in size due to the outflow of the them. liquid from broad !acunes of the lobes. The sip ANATOMY. The shell is characterized by the hon during crawling is directed forward. The wac resorption of the columella, and the inner walls of ter currents through the siphon and above the the spire. Only the inner wall of about half of the propodium inside the mantle cavity and outside penultimate whorl remains (Fig. 4D). The col u the mantle cavity above the posterior part of the mellar muscle has the form of a thick, slightly foot can be seen clearly. concave plate, which passes along the outer In aquarium observalions, molluscs with a surface of the visceral mass and is attached lo the shell length of less than a centimeter are usually remnants of the inner wall of !he penultimate buried in sand at a depth of 1-1.5 cm. The siphon whorl. The only partly spirally eoiled visceral is usually protruded through the sediment, al mass in its upper part above the stomach loses its though it can be pulled entirely under the se spiral morphology and eorresponds in shape to diment for long periods. Molluscs are usually ori the inner volume of the shell thatlacks columella entated nearly parallel to the surface with the an and inner walls (Fig. 4 A-C). terior part of the shell slightly raised (Fig. 3B). The head is absent. Two flaps triangular While crawling under the sediment a track in the when contracted stage and pigmented at their form of a shallow groove is seen on the surface. tips, are placed at !he anterior end of the body. When the mollusc is buried under sand, the pe The rhynchostome opens below the right flap. dallobes are greatly expanded and embrace most The operculum is large, very thin, light yel of the shell. There is a moderately large cavity in low and transparent. It fills the entire aperture. the sand below the spire. The water currents out It bears few growth lines, which are thread-like side the mantle cavity above the posterior part of and slightly raised; in a specimen with a shell the foot are clearly seen. Sediment particles length of 10.6 mm, there are only three growth which have passed through the mantle cavity ap lines. pear to be entangled with mucus to form long Mantle complex (Fig. 4E): the mantle almost threads. During crawling the propodium plays entirely covers the propodium when the mollusc the role of a wedge in moving the sediment par withdraws in~o t~e shell. The dark gray osph lides. radium is clearly seen through the mantle wall. Gut content analysis revealed that the mol The siphon is highly muscular with the wall for lusc swallows food and sediments particles with med of circular muscle.fibers and numerous bnn out sorting. Thus sand grains and other inor dles of longitudinal muscles which are separated ganic particles are usually present in the sto by radial fibers connecting the walls of the mach. Sometimes the size of the particles is sur siphon. prisingly large. For instancc, the stomach of a The mantle forms anterior and posterior ma mollusc with a shelllength of 9.0 mm contained ntle tentacles and a small posterior mantle lobe. a piece of bivalve shell 1.6 x 0.8 mm large, which The posterior mantle tentacle is derived from the occupied nearly the whole stomach volume. Rem right side of the mantle; it passes through the nants of different Arthropoda (Cladocera, Ostra upper corner of the aperture and is situated in the F!G. 6. Morphology of 1he sort body of Ollvella bor~alis Gollkov. A,B - stomach; C-stomach, opened mid-dorsally; D - transverso coda, Cirripedia larvae, water mites), Foramini channeled suture. It covsists of epithelium, a !a sectlon ofthestomach at the entran ce ofthe duct of the digestlve gland; E-transverse section of the stomach in thearea of muscular fera, Nematoda, and gastropods (Fartulum sp., yer of longitudinal muscles, parenchymal and l.a ring; F- transversc sectlon of the caecum of the stomach¡ G ~ transveroo section of the posterior mantle Iobe; H- radula~ I Caecidae) were identified in the stomach cont rge vacuolated ce!!¡; aqd bunqles of longitudinal diagrammatJc transverse ooctlon of the radula in the radularsac; J ~ penls; K- vesicula semlnalis. Scale 0.05 mm for Flg. H and ents. No remains of living bivalves were ever re muscle fibers. The thicll:nerve passes through the · O.S mm for aU others. middle part of the tentacle, closer to the ventral corded. PHC. 6. Moptfxmomu MiJfi(Qf() 'feM Oli-vella borealis Golikov. A-B ~ >KCJlYJWK; e - :>KCJty):{OK, BCJ functiop is the sccretion of the columellar callus probaseis. There is neither valve nor gland of [Marcus, Marcus, 1959]. The mantle lobe has a Leiblein, nor aceessory salivary glands. shallow channel seen in the transverse section At the battom of thc bucea! eavity there is a (Fig. 6G). I ts wall beside the muscle fibers is for large odontophore fonncd of two large subradular med of large supporting vacuolated cells, which cartilages connected at their anterior parts by are histologically similar to the cells fonning the transverse muscle. The radula (Fig. 6H) is ty mantlc wall. pical for the genus, short and consisting of broad The ctenidium is large, occupies up to 4'3 of rachidian teeth with numerous cusps, large, cur the mantle length and is fonned of tal!, irregu ved and sharply pointed laterals, and small rec larly triangular lamcllae. The osphradium is ne tangular plates, which are probably the rudi arly equal to the ctenidium in length and is slig ments of marginal teeth. The radula is folded in htly wider than the latter. The hypobranchial the radular sae along two pairs of longitudinal gland is moderately developed and forms low folds, situated between the rachidian and lateral folds near the rectum. The rectum is thin-walled teeth, and the lateral and marginal teeth respec and lined with very high epithclium. The rectal tively (Fig. 61). In a specimen with a shelllength gland is situated ncar !he anal opening. It is for of 9.0 mm, the radula is formed of 24 rows of med of a fcw large connecting chambers filled by teeth and the basal radula membrane has a width parenchyma. The gland opens near the anal opc of 0.37 mm. The percentage ratio of the radula ning by a broad duct. memhrane width and the shelllcngth expressed Digestive system (Fig. 5,6): thc probaseis in as a percentage is 4.1 %. its contracted state is narrow, with smooth walls. The ocsophagus opens into a moderate sized The probaseis and rhynchodacum are lined with stomach (Fig. 6A-C). The openings of the oeso low, nearly cubic epithelium with large oval nuc phagus, intestine and unpaired duct of the diges lei. The cpithelium is covered by a thin cuticle. tive gland are dos e to each other. The stomach is cm Below the epithelium there is a thin ]ayer of lon lined with a cuticle which is especial! y thick in the gitudinal muscles and a thicker layer of circular middle part of thc stomach, where the wall form musclc fibers. The oral opening has the fonn of a thick muscular ring and where the epithelium FIG. 7. Morphology of the soft body of Oliva bulbosa (ROding). A,B - body, removed from 1he shell; C- anterior view of the hcad. narrow slit and lacks a distinct sphíncter. The lacks definite folds. At the hind end of the sto Scates O.S cm. bucea! tube leads from the mouth to the bucea! mach, the cuticle becomes thinner and the epi PMC. 7. MoJXI>oROr.mJ MSH'KOI'O TeJia Oliva bulbosa (ROding). A,B- TeRO J.13BJJC'-ICHIIOC H3 paKOBHHbl; e~ llHJ~ fOJ100hl cncpe;-lH. cavity. The bucea] eavity is situated at the base of thelium forms numerous equal folds. The duct of Macurra6 0.5 CM, the probaseis in its contracted state. The wall of the digestive gland is broad, and is lined with the bucea! tube is highly foldcd and lined with cuticle at the opening into the stomach. It rami of the penis and opens at the tip of a long papilla In fixcd specimens, the anterior and posterior tal! epithelium and has a thin !ayer of circular fies into numerous channcls at a short distance . FIG. 9.0esophagus of Oliva sayanaRavenel, 1834, dissected dorsally. PHC. 9. TIHu~eBo.n. Oliva sayana Ravenel, 1834, ncKphlrhJtí: .n.opcanbHO. and minor typhlosoles border the intestinal groo Amalda montrouzieri (Souverbie, ve. There is a concavity at the base of the major 1860) typhlosole. The large·unpaired duct of the diges tive gland is situated near the entrance of the (Fig. 10 F,G; 11) oesophagus. The radula is small (Fig. 8E), its length about MATERIAL STUDIED: 2 specimens dredged 2.35 mm in a specimen with a proboscis 6 mm in in the outer lagoon of Noumea, New Caledonia, length. Lateralteeth are curved, with a broad ba 10.IX.l989, coll. J. Taylor; one female (shell FIG. 8. Morphology of the digestivc system of Oliva bulbosa (ROding), A- organs of the body haemocoel (scale 0.5 cm), salival)' length 27.4 mm) sectioned. glands removed; B ~longitudinal section of the valve ofi..eiblein (seale- 0.5 mm) C-slomach {scale O. S cm); D ~ stomach, se and pointed tip. The broad rachidians bear opened dorsally (scale0.5 cm); E- radula (scale 0.1 mm). three cusps. In the anterior par! of the radula, the ANA TOMY: The body consists of approxi teeth are damaged. The radula in a specimen mately 3 whorls (Fig. lOF). Mantle short, ex PH:C. 8. MopíjJoJIOrJHI mm~erutp»TeJJbtiOM CHCTCMbt 0/ivabu/bosa (RQding). A- opraHhfT)'JIODmU,HOI'O reMOlleJHI (MRClUTaÓ0.5 CM); wíth shelllength 22.7 mm consists of 100 tran lending up to \IÍ of last whorl, with a well deve B - flpú]:{OJlbHbltf Cpe3 KJJ8nana JieM6nei1Ha (MaCIUTR6 0.5 MM); C- )KCJIY,AOK (MRCUITa6 0.5 CM) j D - )KeJtYflOK 1 DCKpbrrbJli: loped posterior mantle lobe. Foot broad with a .nopcan&no (MaCmTaÓ 0.5 cM); E~ pa.rcyna (MaCmTaó 0.1 MM). sverse rows of teeth, 9 of which are incompletely formed. Thc ratio between radula membrane wi crescent-shaped propodium divided by a deep dth and shelllength expressed as a percentage is longitudinal deft. Pedal lobes well developed. formed of circular muscles. Bucea! tube short, li portion of the oesophagus between the opening'of 1.09%. The body lacks pigmentation and is creamy ned with tal! epithelium and a thick cuticular !a the duct and the nerve ring are lined with a glan In general, the morphology of O. bulbosa is white in color. The poorly developed anal gland is yer. The wall of the tube is formed of circular dular epithelium, which is so high that it restricts very similar to that described by Marcus and seen through the mantle as a ramified black line. · muscles. the oesophageallumen to a dorsal slit. The valve Marcus [1959] as O. sayana (the correct identi The columellar muscle forms about 2 whorls, Salivary glands large, formed of ramified tu of Leiblein is large, pear-shaped with a broad fication of the species should be O. circinata ieaving a wide cavity. Ovary yellow and bes. The wall of each tube consists of a single funnel-like cavity (Fig. 8B). The walls of the val Marra! ~ B.Tursch, personal communication) occupying the inner side of the whorls of the !ayer of angled cells. Glands are situated at both ve consist of very long and narrow cells. The lu and differs mainly in details of stomach visceral mass. Operculum large, very thin and sides of the rhynchodaeum, and cover the bend of men is almost completely filled with the folds for morphology. transparent (Fig. lOG). It is yellow with the oesophagus; they are attached to the oeso med by small oval cells, the outer !ayer of which indistinct growth lines and subcentral nudeus. Head small, poorly distinguished from the. phagus by numerous connective tissue fibers. bears long cilia. Oliva sayana Ravenel, 1834 The right salivary gland is larger than the left The small accessory salivary gland is situated body and formed of small dorso-ventrally flat one. Branches of the proboscis retractors pene in front of the nerve ring. Its lumen is lined with FIG. 11; Amalda nwntrouzieri Souverble. A~ semtdlagraminatic longitudinal sectlon of the nnteriorpart of the dlgestlvesystem (sea le pi O.S cm), salivary gtandswith the ductsafe notshown; B- mantle (scale 1 mm); C- stomach {scale 1 mm). PHC. 11. Amalda montrouzierl Souverble. A- cxeMaTH3Hpommu&tH nponon&Hblií cpe3 qepea nepe,nmoiD 'lacr& nHll(esapmenbliOH CHC1"CMht Amalda montrou.zleri Souverbie (MacwTa6 0.5 CM), cnrouu~>~e >KeJie3bl e npoTOKRMH ue H306pa»eeu&t; B - MannHI (uaC1111'a6l MM); C - >KeJIYAOK (M8Cli1Ufi 1 MM). diverticulum is lined with a thick cuticular !ayer. of slightly lesser diameter than the gland itself. Cartilages paired, not fusing anteriorly. Radular The glandular part of the oesophageal epithe sac only slightly longer than the odontophore. lium is absent. The radular refractor does not pass through the Radula was illustrated by Kilburn and Bou nerve ring and is attached to the floor of the bady chet [1988, Fig. 51, 53] and described by them pr haemoroel. · as follows: "Rachidian with side cusps barely Salivary glands large, formed of ramifying stronger !han median cusp; tiny, irregular den FIG. 10. Morphology of the soft body of Pseudoliva zebrina A.Adams (A-E) and Amalda nwntrouzieri Souverbie (F,G). A,B,F tubes. The barders of the cells which form the ticles devcloped on bases of cusps". body, re~ove.d from the shell (scale 0.5 cm); C- partof the visceral mass (scale0.25 cm); D- penis (scale 1 mm); E- viewof jo Stomach (Fig. llC) U-shaped, without cae the crawlmg hve snail (from the sketch of S.Gofas); G- operculum (scate 0.25 cm). tube walls are not visible. The glands ad in the oesophagus and valve of Leiblein. The barders cum. Dueto the fixation I was unable to study its PHC. 10. MoJXPonornH MSJf'Koro te.n.a Pseudoliva zebrina A.Adarns (A-E) »Ama/da montrouzieri Souverbie (F-G) A,B,F- TCJIO, between the salivary glands could not be seen interna! morphology. H3BJIClJCHHOC lt3 paKOBHHbl (M8CWT8Ó 0.5 CM); C- YlfRCTOK BHCl{Cpa:JlbHOI'O MCWKR (MRCUlTa6 0.25 CM); D- nemtC (MaCuiTa6 even in sections and it can be deduced that there In general, anatomy of A. montrouzieri is 1 MM); E- BH)l nonayll(ero MOJJJIIOCKa (e uaópocu S. Gofas); G- KpwruelfJ hg FIG. 12. Morphology of the soft body of PsJudo/iva anc/1/a Hanley. A- body, removed lrom the shell; B -anterior view of the head (scale 1 cm}; C- stomach; D ·- manthnA,C,D- in the same scale, scale 1 cm) PHC. 12. MopclKlnomsr MSITKoro Tena Pseudo/iva ancilla Hantcy. A - reno MOJUJIOCKa 6e3 paKonHHM; B - BH~ OOJIOBbl cnepe¡pt (MaCunafil CM); C- >KMYJ{OK; D- MftHTHil (A,C,D.- B O.AHOM M8CLUta6e, MRClliTI\61 CM), .Museum Nl! E 2094; one bady dissected and part ated at the base of the probaseis in the lower of the digestive system sectioned (female). right side of the rhynchodaeum. Subradular ear ANATOMY: Body consists of 2.25 whorls in tilages paired, thick and fused anteriorly. Radu a specimen with a bady length of abaut 48 mm lar diverticulum opens into a poorly differentia (Fig. l2A). The columellar muscle is very .thick ted bucea! cavity at the probaseis base. 'l'he very and wide with very broad eavity. Its upper edge thick-walled long bucea! tube leads from the is turned outside. The muscle comprises abaut mouth opening to the bucea! eavity; the walls of 1.5 whorls. The foot is broad, pedallobes absent. the bucea! tube are formc:d of circular muscle fi Propodium very narrow. The foot is pigmented bers. This tube occupies almos! the whole inner with black spots. Mantle thick, ctenidium and volume of the probaseis and is lined with cuticle, osphradium are poorly seen through it. Siphon is which in the posterior part is. formed of many very long and thick walled. layers of up to 0.2 mm thick. Rhynchodaeum very Head small (Fig. 12B), well differentiated thick with numerous muscles attached to its an from the bady. Tentacles in fixed specimens very terior part by one end and to the skin by the short, cone shaped, with eyes near the tips. In other. Oesophagus with thick muscular walls and the live mollusc, it is seen, that the tentacles are forms a S-shaped curve after leaving the probas elongate and the eyes are situated at the small eis. The valve of Leiblein large, cone-shaped. lobes approximately in the middle of the tentacle Small and compact acinous salivary glands [Kilburn, 1989: Fig. 2]. are tightly attached to the oesophagus on both Mantle complex: Mantle (Fig. 12D) long, 1.5 sides of the valve of Leiblein. Numerous muscle times longer than its width. Siphon very long, fibers, which are attached to the rhynchodaeum without distributive valve. Ctenidium very long, by one end, and to the oesophagus and valve of occupying abaut 213 of the mantle length, and for Leiblein by the other, penetrate the salivary med of high triangular lamellae. Osphradium na glauds. The right salivary gland is situated below rrow, symmetrical, equaling 1/z of the ctenidium the odontophore, the left mainly above it. Ducts FIG.13.Pseudo/lvaancil/aHanley. A -semidlagramrnaticlongltudinalsectlon ofthe proboscis (scale 1 cm), sallvary ducts notshm;n; Jength. Hypobranchial glaud well developed, for of the glands are short, and immediately after B~D- organsof the body haemocoel (scale 0.5 cm): B- from the ventral side. C ~ from the dorsal slde, D- from the rights1de ming distinct folds in the posterior part of the leaving the glands enter the walls of the oeso (proboscis ls CUl'Ved to the lert to show the val ve of Lelblein). mantle. Rectum and pallial oviduct growing fused phagus abave the valve of Leiblein aud pass iu PHC. 13. Pseudoliva ancil/a Hanley. A-cxeMaru3wposaHublif npononr.uwtl epea xoúma (Macurra61 CM), npoTOKH CJIIOHHblX men~ without distinct border. Anal opening and gono side the walls. They open into the bucea! tube He Hao6pa» chambers. The glandular epithelium differs from Somali, depth 4750-4420 m, Galathea trawl, the rest of the oesophagus in taking up more ex mud, 8.V.l967 (1 dried specimen - radu1a tensive staining. extracted). The radula was illustrated by Barnard [1959, ANATOMY: Body rounded, in the specime11 Fig. 15 a). It consists of 57 rows of teeth. Ra with the shelllength of 8.3 mm consisting of 1.75 chidian broad, with three large acute cusps. La whorls (Fig. 14 A,B). It lacks pigmentation terals with two sharply pointed cusps. except to the visceral mass. The digestive gland Stomach small, sac-like (Fig. 12C), with a containing numerous black granules, which can large eaecum and wide unpaired duct of the di be seen through the skin. Visceral mass gestive gland, which is situated at the opening to occupying 1.25 of a whorl. Columellar muscle the intestine. Externally the stomach is similar to consists of about one whorl and leaving a wide that of Oliva. I was unable to study its interna! cavity. Propodium narrow, with a moderately morphology. wide and shallow cleft. Head poorly differentiated from the body, with very long, Pseudoliva zebrina A.Adams, 1853 broadly spaced tentacles (Fig. 14D). Mantle complex (Fig. 14E): Mantle 1ength (Fig. lE, 10 A-E) equals its width. Ctenidium long, equal to 3/,¡ of the mantle length, narrow. Posteriorly near the nr MATERIAL EXAMINED: Two spedmens: nephridium, the ctenidium is formed of mode Praia Amelía, prov. M~medes, Angola, depth rately high lamellae; this lamellae bear long fla 60-80 m, coll. S.Gofas; both sectioned. gellae which adhere to their inner sides, but are ANA TOMY: The body of a specimen with absent from the anterior part of the ctenidium. shell length of 10.2 mm consists of 3.5 whorls Osphradium is very large, nearly twice as broad (Fig. 1OA,B). Col u mellar muscle forming a as the ctenidium and equals the length of cte funnel with a wide lumen. The foot Iacks pedal nidium 's inner side. The osphradium is greenish lobes and has a very narrow propodium. Head, gray, i1s right portion broader than the left. The siphon and mantle edge pigmented, brown right portian of the osphradium consists of 26 la (S.Gofas, personal communication). Head well mellae, theJeft of 21 lamellae. differentiated from the body, ·with moderately Siphon broad and shallow. On the right side, long tentacles, subdivided at their tips. Eyes are the mantle edge forms a rather narrow posteríor situated on small lobes, which have a small mantle lobe. The rectum is thin~walled and creamy-white spot (Fig. lO E). Operculum large, narrow. thin and transparent, with a terminal nucleus. The specimen was a mature female with large Mantle long, occupying about a whorl and, in pallial oviduct. The gonopore has the form of a fixed specimens, covering the base of the head. long slit with slightly frilled borders. The mantle edge forms a deep notch near the Digestive system (Fig. 15, 16): The rhyncho siphon. Nephridium very narrow, brown. stome has the form of a flattened slit about 0.5 A second specimen with a shelllength of 10.2 mm in width. A distinct sphincter is absent. Pro mm is an immature maJe with a very short, smo boscis of medium length, gradually narrowing to oth, flattened penis with a small papilla at its tip the oral opening. Oral opening small, rounded, (Fig. lOD). without sphincter. The proboscis, rhynchodaeum The mantle complex is similar to that of P. and skin are lined with epithelium covered with a H ancilla but differs in the shorter siphon. moderately t:hick cuticle. A proboscis wall and The digestive system is also very similar to. rhynchodaeum are slightly folded ¡¡nd lined with · · A bod removed from the shell (sea le-0.5 cm); C -longitudinal that of P. ancil/a and differs. only in the larger FIG. 14. Morphology of the soft body of Bentlwbw. tryom Dan. ,B -· Y¡ th head (mantle removed) (sea le - 0.5 cm); E - mantle epithelium formed of tal! c.ells with large oval section of the val ve of Leiblein (sea le- 0.5 mm); D - anterlodr re(~ le e O OS mm). H -lateral tooth in the different projection. accessory saliv~ry gland and poorly developed elongated cells. Pro~tis walls very thin: formed (sca1e ~ o.s cm); F-form of the ctenldium lamellae; 0 ~ ra u a - · • glandular folds 'of the oesophagus between the of an outer thin !ayer ,pf circular muscles and an . D n A B TeJIO H3.Meqe¡:moe H3 paKOBHHbl (Macurra6 0.5 CM); e - opening of the duct of gland of Leiblein and the inner !ayer of longitudil\al ,muscle fibers. The PHC. 14. Mop$onornSJ MYrKOro Te11R Benthobw. tryoni a · • .--m; roJIOBhlcrtCpe]Vf,MaHTH$1YAMeua (r.tacurra60.5ct>ü; nervous ring. flpüAOJihHNÜ epea 1.fCpe3 KJianau Jieütineliua (Macurra6 0.5 MM) • D , G ~ mlrrvJia (Macurra6 0.05 MM); H - naTepanbH&tt:i ayO proboscis lumen is filleil with friable, probably E _ MaHTHSI (Macmn6 0.5 cM); F - rs aog FIG. 16. Rcconstruction of the position of the organs of thc body hacmocoel of Bentlwbia tryoní Dall from the serial seclions. Nervous ring not shown. A- from the right si de; B - from the left si de. Columellar muscle indicated by dotled arca. Scal,e 0.5 mm. PHC. 16. PeKoHCTPYKU.HH noJfO>KCJHfH oprauou T)'JlOUHll.(noro reMOU.CJut y Bentlwbia tryoni Dall, nony•ICI:IHI'» e cepHt111biX cpcaou. Hepnuoe KOJtbU.O ue J.I306pa}l(euo. A-e npanoH cTopoHbl; B -e nenojf croponM. KoJryMeJmuptlb!H MYCKY Jf3aTOliKOnau. M a cm ra6 O.SMM FIG. 15. A--: Semidlagrammatlc longitudinal section of the anterior part of the digestive system of Benthobia tryoni Dall (scale O.S covered with a very thin connective tissue layer of Leiblein posteriorly. The lumen of the gland is mm)· Sahvary glands not shown, arrow indica tes the place of the opening of the salivary duct; B ~ mid oesophagus of Nucella and consist of orange granulated tissue. Borders lined with a single !ayer of low cubic epithelium heyseana (Dunker). opened ventrally (scale 0.5 cm). Arrows Jndlcate the direction of the food movement (drawlng of A.I.Medinskaya). of the cells were not seen. Salivary ducts pass and filled with a finely granulated secretion stai within the oesophagus walls and open into the ning grayish-blue. The duct has a diameter of PHC. 15. A- cxeMaTHaHponamthtfi npoJ,\OJibHbl~ cpea ttepea nepe~HIOIO li8CTb nMll{euapJffeJlbHOit CMCTeMbf Benthobia tryoni Dall oesophagus immediately behind the bucea! cavity about 0.1 mm, slightly coiled and opening into (Macrura(} 0.5 MM)· CJUOHHbte >KeJle3bt He uao6pa>KCHbl, CTpeJIKO~ OÓ the wall of the gland is formed by 3 layers: two folds. Rectum of medium· diameter, opening at epithelial with thin circular muscle !ayer between the middle of the mantle length. Anal gland them. In Benthobia the outer !ayer practieally is absent. not developed. Moreover, the sharp dif Digestive system (Fig. 18): Probaseis highly ferentiation into the muscular sac and the long, extensible, short in the contracted position, thick moderately thick duct is not present in other and gradua!ly narrowing to its tip. The probaseis neogastropods. Nevertheless, the opening of the wall is formed of a thick !ayer of circular muscle duct into anterior part of the bucea! tube probably fibers with few longitudinal ones. Almost entire indieates the homology of the "typieal" accessory probaseis lumen is filled with numerous muscle salivary gland of neogastropods and the gland of bands. Oral opening broad-oval, nearly rectan Benthobia. gular. Rhynchodaeum thick-walled at its dorsal Radula of the specimen with the shelllength part and very thin, nearly transparent at the ven of 8.3 mm (Fig. 14G) formed by 45 rows of tran tral side. Probaseis retractors are attached to the sparent and yellowish teeth. Rachidians broad rhynchodaeum dorsal! y on its anterior third and with 7 cusps, the outer pair of which are shorter to the cover of the bady haemocoel. The probaseis than the other. Lateral teeth have the form of is highly innervatcd. The oesophagus is attached curved plate. to the probascis walls with numerous radial mus ele fibers. Odontophore very large, díameter of REMARKS. The genus Benthobia Dall was the bucea! mass exceeds that of the oesophagus recently attributed to the Pseudolivinae by Bou by 3-4 times. Bucea! mass protrudes backward chet and Warén [1985]. Anatomieal data confirm from the probaseis base. The radular diverticu this attrlbution. One of the most importan! fea lum opens into the bucea! eavity near the oral tures (in my opinion) is the position of the radular opening. The odontophore is probably very mo sac at the probascis base. This feature is otherwise bile and may be protracted through the mouth found only in the order Toxoglossa. opening. The short bucea! tnbe leads from the mouth to the poorly differentiated bucea! mass Melapium lineatum (Lamarck, 1822) and is lined with a moderate! y thick (abaut 0.025 mm) cuticular layer. Subradular cartilages (Fig. 1 A-B, 2 B, 17, 18) paired, fusing anteriorly to form a deep groove and extending to the end of the bucea! mass. MATERIAL EXAMINED: 3 specimens, South The oesophagus after leaving probaseis, pas Afriea, off East London, 33o04,6'S, 27°54,8'E, ses abave the organs of the bady haemocoel and depth 70 m, gray sandymud, 16.VII.1984. makes a long curve directed forward. Behind the nervous ring, the oesophagus follows below the ANATOMY: Body consists of approximately organs of the bady haemocoel and then abruptly 2 whorls (Fig. 17 A,B). Foot broad, with a very widens and forms a ring, positioned in the frontal narrow propodium. In live specimen the foot has surface. unusual coloration in the form of 3 narrow con The val ve of Leiblein is moderately small and centric colored bands [Kilburn, 1989; Kilburn, oval. Gland of Leiblein very Jarge, dark-gray, personal communication ]. Operculum absent. occupying half of the volume of the bady haemp Head well differentiated from the bady, with long coel and opening into the oesophagus on Hs right tentacles, which are gradually narrowing to their side by a moderately thin duct. The very large tips. Eyes small, positioned at the base of the accessory gland is situated between the valve of tentacles. Rhynchostome has the form of a broad Leiblein and the gland of Leiblein (Fig. l8A-D flattened dorso-ventrally slit. Mantle thick, os aog). It is formed of numerous large glandular phradium and ctenidium being searcely visible sacs of the oesophagus (Fig. 2B). through it. Border between nephridium and Paired salivary glands are formed by rami mantle eavity barely apparent. Columellar muscle fied tubes. The left salivary gland is situated be leaving a broad cavity and formed by 1-1.5 low the probaseis, in front of the nerve ring, at whorls. the leve! of the probaseis tip. The ríght salivary Penis short, thickening to the end and flat gland is positioned on the ríght side of the pro tened at the front edge (Fig. 17C). The gonopore baseis and joins the left salivary gland. Salivary · papilla is very small and has the form of a small FIG. 17. Morphology of1he sort body or Melapium /ineatum (Lamarck). A,B- body, removed from 1he shell (scaÍe 1 cm); e-penls ducts are of small diameter and lie on both sides (scate _1 cm); D - vesicula seminalis (In the same sea le as C); E - mantle (scale 1 cm); F- form -of the ctenidlum Jamettae; G,H bump at the front edge of the penis. of the oesophagus. ~ egg capsule: G- from the dorsal slde, H-from the ventralside (scate O.S cm); 1-radula; J-margtnal1ooth in the dlfferent Mantle comp/ex (Fig. 17E): Mantle short, its Radula (Fig. 17 I,J) is formed by 70 rows of projectlon (sca1e 0.25 mm); K- stomach (sea le 0.25 cm). length equaling its width. Maulle edge thickened. teeth. Rachidian tooth broad but !hin, with three Siphon long, thick-walled. Ctenidium very wíde; acute cusps. Bases of the laterals are dark and PHC. 17. MoP<}>onomR MRrKOro TeJJa Melapium lineatum (Laruarck). A,B - Te.,IO, H3BJICtJCHHOO Ha paKOBHHbi (Macnmu5 1 CM); e its width is half that of the mantle and equal to nemtc btacnrra61 CM); D ~ veslcula seminalis (MacmTa6 1 cM); E- MftHTHR (Macurra6 1 cM); F- 4JopMa nenecTKa KTCHHJ\Hst; highly sclerolized. G,H- Bfi~en.as:~ Kancyna, G- e ,«opcanbHOfi CTopoHbt, H-e BCHTpanbuoü cropouw; I-pa.eyna; J- MaprnHaJJbHbltf ay6 u the mantle length. It is formed of moderately tall, Stomach (Fig. 17K, 18E) sack-shaped wlth a J1l)yroti npoeKU,HH (M8ClliT86 0.25 MM); K- >KeJIYJ(OK (Macurra6 0.25 CM), triangular lamellae with small flagellae at their ventral oesophagus opening. Caecum very short. tips. Osphradium narrow, equals ;¡/3 of the cte Ducts of the digestive gland are paired; one of nidium length, nearly symmetrieal with lamellae them opens at the entrauce of the oesophagns, that are wider in the right portion of the osph another near the entrance to the intesline. A lar- radium. Hypobranchial gland without distinct 40 Yu.I.Kantor Morphology and relationships of olivifonn gastropods 41 membrane. The surface of the capsule is smooth, the propodium functions as a wedge, moving sd aog measurements 10.3 x 5.4 mm and 11.5 x 5.0 mm. sediment particles aside. The capsule wall is comprised of a single layered wall about O. 1 mm thick. Each capsule is filled 5. Morpho/ogy of the head. Morphology of thc wíth rounded eggs, whose diameter is about 0.2 head is variable in thc groups studied. 4 statcs mm. can be distinguished: the plcsiomorphic Barnard [1959] and Kilburn and Rippey condition, found among Pseudolivinae: hcad {1982) described the similar egg capsules posi, more or lcss differentiated from the body with tioned in the same way. The former author men, long tentad es (state 0); head, formed of vertical tioned, that his capsules were calcified externally flaps which bear long tcntacles (found in Oliva, (the feature not found in studied specimen). state 1); head poorly differentiatcd from the body and formed of rounded horizontal flaps (found in Ama/da, state 2); and the reduction of PHYLOGENETIC ANALYSIS the head, found in 0/ive/la (state 3). Small triangular flaps in the latter species eannot be In arder to clarify the systcmatic position of regarded as a true head for two reasons: l. thc differcnt oliviform groups, a phylogcnetic analy asymmetrical position of the rhynchostomc, sis was conducted. The following characters were which opens below the right flap and 2. absencc used for the analysis. of innervation of the flaps. For comparison, in lintricula and 0/ivancil/aria, also without l. Form of the she/1 suture. The most marked tentacles and eyes, the flaps are supplicd with character of Oliva and Olivella is the channelled, nerves [Marcus, Marcus, 1959]. narrow suture, which contains the posterior man tle tentacle. Such a suture is unique among neo 6. Operculum. An operculum is absent in gastropods and thus may be considered to be a Oliva and Melapium, the state being considered derived state. In Ancilla, the suture is covered by as apomorphic. The loss of the operculum is the callus and is not seen from the surface. widcly distributed among gastropods and Finally, Pseudolivinae has a slightly impressed probably took place indepcndently in both genera suture, usual for gastropods. This type may be mentioned. considered as an ancestral stage of the ingroup. Since the sequence of transformation of character 7. Presence of the posterior mantle /obe. The states is :mclear, the character was unordered posterior mantle lobe which produces the colu during the analysis. mellar callus is well developed in Olivinae, Oli vellinae and Ancillinae and in Benthobia tryoni. 2. Resorption o/ inner wal/s of the she/L As As it is absent in other neogastropods, the feature can be seen from sections of the shell in Oliva, the may be considered apomorphic. inner walls of the shell are partially resorbed and very thin. In 0/ive/la the columella and inner 8. Presence of the anterior mant/e tentacle. wa1i: in the upper psrt of the shell have been Anterior mantle tentacle was found in Olivinae cor:.pletely resorbed and only part of the inner and Olivellinae. Its functional significance is un wa!I of the penultimate wall remains. This may be clear. Since it is absent in other neogastropods, regarded asan advljnced (apomorphic) state. the feature may be considered apomorphic. 3. Presence of pedallobes. AII the subfamilies 9. Presence of the posterior manlle tentacle. lt except Pseudolivinae have well developed pedal is well developed in genera with a channelled su lobes which embrace the shell. The feature is uní• ture, that is in Oliva and 0/ivella. As a posterior que among Neogastropoda and may be mantle tentacle is absent in other neogastropods, considered apomorphic. The function of the pedal the feature may be considered apomorphic. Iobes is to probably a protect the shell from the FIG. 18. M~rp~ology of t~e di~estivc system of Me/apium lineatum (Lamarck). A~D - organs of.the body haemocoel (sea le 0.5 cm), abrasive action of sediments and they are thus 10. Position of the radula and place of Arrows mdt~ate thc dtrecuon of thc food movemcnt in the oesophagus: A - from thc right si de, B- from the left si de, e- from connected wíth a burrowíng mode of life. opening of the radular diverticulum. The radular thc dorsal stde, D - frorn !he verllral si de; E - stornach, opcned dorsally (sea le 0.25 cm). sac in N eogastropoda (except in the ord'er 4. Toxoglossa) is usually situated at the proboscis PHC. 18. MopclJononni mmtenapmcJthHOfi cuneMol Melapium lineatum (Lamarck). A-D ~ opraubl ryJlonwntuoro reMol(eJm Form of the propodium. AII the subfamilies (Macmra60.5 CM). Crpe.nKaMH noKa3auo uanpaMCUHCADH>KCHmt HHIIVI n mmteno.nc: A- en pasa; B-cnena; C- cnopcaJJhHOii except Pseudolivinae have a very mobile tip in its contracted position. Such a condition CTOpOHbl; D - C HCHTpaJtMtOft CTOpOnbl~ E-- mCJJYAOK, DCKph!ThJM nopcaJlbHO (MUCIJJT36 0.25 CM), crescent-shaped propodium, subdivided by a among studied species was found in Oliva. deep longitudinal cleft. Such a propodium can be Toxoglossa have a specialized inlraembolic type found only among the Olividae and thus it may of the probaseis, in which the radular be considered to be an apomorphic feature. It is diverticulum opens at the proboscis base. Origin ge longitudinal fold, originating near the groove. A moderatcly large gastric shield is situ' highly innervated in Oliva {Marcus, Marcus, of this proboscis type is probably connected with entrancc of the ocsophagus, dirccts food into a ated atthe base of the typhlosoles. 1959] and is used for seizing the food ítem and their specialized feeding mechanism [Sysoev, Kantor, 1987; Kantor, 1990 ]. Nevertheless, the posterior mixing atea. which is mainly lined by Egg capsu/es (Fig. lA, 17 G,H): Egg probably also for food location [Tursch, 1991 ]. radular sac is situated at the proboscis base in transvcrsc folds and also with somc irrcgularly capsules ate altached on thc inner Iip of the Such a propodium may also be an adaptalion toa sorne olivids, that is Pseudoliva, Benthobia, oricntatcd folds. Distinct major and minor aperture of thc female shell. They have the form borrowíng mode of life. Thus, during aquarium Olivella and Ama/da. Two types of proboscis typhlosolcs ate short and rcstnct thc intestinal of an orange segment and are attached by a thin observations on Olivel/a borealis, it was seen that 42 Yu.I.Kantor Morphology and relationships of oliviform gastropods 43 TABLE l. Llst of characters, ured in the cJadistivc anatysis. TABLE 2. Character states in 1he species, used in the ctadístic analysis l. The form of tlte suture of the sheiJ: O- depressed, 1 - channeled, 2-covered by the caUus NSZ - Pseudo/hu zebrlna; MEL ~ Melapium lineatum; OLB ~ 0/ivella boreal{-s,· OLV - Olivella verreauxii,· OLI - Oliva bulbosa; AMA ~ Ama/da nwntrow:ieri; AMD - Amalda dimidinta. morphology were found among the mentioned servations). In neogastropods and highér meso genera. In Melapium and Amalda the gastropods, such a conneclion is lost with the ap odontophore seems to lie at the proboscis base, pearance of a long proboscis and the retractar as through the nerve ring and that is why this con ingroup: acinous (Pseudoliva and Benthobia) and but as seen from th.e sections, the radular well as various bucea! muscles attached to the pro diverticulum opens near the proboscis tip, nection cannot be considcred to be homologous formed of ramifíed tu bes (the others). The baseis wall [Graham, 1973; personal obser with the connection of the retractar with the functional significance and transformation whereas in Benthobia and Olivella the vations ). Nevertheless, such a connection was fo col u mellar muse! e (se e fea tu re 11). The sequence of the character states are not known diverticulum opens at the probaseis base. In und in a few neogastropods, e.g. in sorne primitive Benthobia the position of the radula at the appearance of this feature is unclear and the and therefore the character was unordered. Turridae - species of Splendrillia and Clavus character was unordered during the analysis. proboscis base is correlated with a very prlmitive (subfamily Clavinae) [Sysoev, Kantor, 1989; 15. Length of the salivary ducts. The salivary feature - the connection of the radular retractar Kantor and Taylor, unpublished observalions] 13. Number of radular teeth per traJtsverse ducts may pass freely along the oesophagus (011- with the columellar muscle (see below) and this and inBentJwbia tryoni. In all these neogastropods row. Typically N eogastropoda have three teeth or vella, Oliva, Melapium) or immediately after lea confinns the plesiomorphic nature of the position the radular refractor al so pass es through the nerve a single one in each row. Only some primitive ving the gland enter the oesophageal walls and of the radula at the proboscis base in Benthobia. ring. Therefore this feature may be considered ¡¡s Turridae (subfamily Clavinae) and Olivella have follow within the walls (other genera). The fun Nevertheless, in other olivids we cannot state for very prlmitive and plesiomorphic. 5 teeth in a row. The small rectangular plates in ctional significance of the feature is not known certain, whether it is a prlmitive feature or not. The presence of such a connection to my the Iatter genus should be considcred as marginal and therefore it was unordered. As the possibility of independent appearance mind indicates the position of the radula at the teeth. This is obvious from the manner of folding of this feature in different olivifonn gastropods proboscis base in Benthobia to be plcsiomorphic. of the radular membrane in the radular sac (see 16. Accessory salivary glaJtds. Accessory sali cannot be rejected, the feature was unordered The alternative, i.e. a secondary shift of the description of the species). The same type of vary glands have the same morphology in all the during the analysis. radular sac from the proboscis tip to the base, folding is found among molluscs with a neogastropods. They are usually paired (plesio would be to assume the restora1ion of a previously morphic condition) but unpaired in studied 11. Connection of the radular refractor with the taenioglossan radula, from which neogastropods lost connection, which scarcely feasible. probably originated [Kantor, 1990 ]. So, the olivids except Olivella and Melapium, in which columellar musde. In archaegastropods and these giands are absent (apomorphic condítion). mesogastropods Iacldng a long proboscis the ra presence of marginal teeth in Olivella is a 12. Connection of the radular retractor wilh plesiomorphic state. dular retractar passes through the nerve ring and the body walL In Amalda there is a connection 17. The gland of Leiblein. The gland of is connected with the columellar muscle (e.g. in between the radular retractor and the flóor of the 14. Morphology of the salivary glaJtds. Two Leiblein is a synapomorphic feature of Trochidae, Uttorinidae, Naticidae, personal ob- body haemocoel. The retractar does not pass types of salivary glands were found in the neogastropods and among studied olivids is Yu.I.Kantor Morphology and relationships of olivifonn gastropods 45 absent only in Olivella . So, reduction of the gland is very large and represented by numerous gland may be cousidered to be an apomorphic sacs opening independently into the oesophagus condition. (Fig. 2C), it is not connected with the duct of the 18. Accessory oesophageal gland. The acces gland of Leiblein. The latter gland opens inde sory oesophageal gland, known also as "glande pendently backward from the accessory framboisee" was first recorded in thc Muricacea oesophageal gland. Nothing definite can be said and usual! y has the form of dorsal glandular folds on the origin and development of the accessory (Nace/la - Fretter, Graham, 1962) which pass oesophageal gland, and consequently the char backward from the nervous ring to the duct of the acter was not ordered during the analysis. gland of Leiblein, where they are spread far iuto 22. The position of the columellar muse/e. the ramifications of the gland of Leiblein. In Usually the spirally coiled columellar muscle sorne M uricidae and Thaididae the gland is passes along the inner side of the whorls of the subdivided internally into small lobes (e.g. body and attaches to the columella. The only Ocenebra japonica, Nucella heyseana known exccption was found in Olivelia in which A.Medinskaya, Kantor, unpublishe(l the muscle has the form of a thick, slightly observations) (Fig. l5B). The similar glandular concave plate, which passes along the outer epithelium was found in Oliva peruviana surface of the visceral mass and is attached to the [Kuttler, 1913], "0. sayana" <~ O. circinata) remnants of the inner wall of the penultimate [Marcus, Marcus, 1959], O. bulbosa and O. saya whorl. This feature is an autapomorphy for na (this study). lt is al so present in Pseudoliva Olivella. and Benthobia. The glandular tissuc of thc gland 23. The morphology of the visceral mass. The in these taxa also passes into the duct of the gland visceral mass in al! prosobranchiate gastropods of Leiblein as in Muricacca. The diffcrent except Olivella is spiral coiled. On the contrary, situation is in M elapium (see bclow, character in Olivella the partially spirally coiled visceral 21). The functional significan ce of the feature is mass in its upper part above the stomach has lost nuclear and it was unordered during the analysis. its spiral morphology and corresponds in its shape to the inner volume of the shell that lacks 19. Morphology of the stomach. The externa! col u mella and inner walls (Fig. 3 A-C). This and interna! morphology of the stomach is rather feature is an autapomorphy for Olivella. variable in the species studied. Three main types As described above, 23 characters coded as 211 141 can be distinguished: l. oesophagus opens ven 57 states were chosen for the analysis 4. Form of the propodium. Appearance of Moreover, although this marginal teeth are flat, are brackets), the character states are presented PHYLOGENY AND CLASSIFICATION OF crescent-shaped propodium is a synapomorphy they are well defined and there is a longitudinal in round brackets. THE OLIVIFORM GASTROPODS. for the clade 14. fold of the radular membrane between the marginal and the lateral teeth. As such folds Clade 14: Amalda, Oliva, Olivella [3(1), The cladistic analysis presented here could S. Morphology of the head. The clearly define the marginal teeth from the lateral 4(1) 5(1), 7(1), 11(!), 14(1) J. The monophyly not confirm the monophyly of oliviforms. This is reconstruction of the character evolution shows in taenioglossan radulas, I think that the plates in of the clade is supported by 6 apomorphies. The the evidence of the fact, that Olividae sensu lato the change of its states from plesiomorphic to 1 Olivella are true marginals. plesiomorphic state of the character 1 is unclear. is a polyphyletic group, including severa! fami (clade 14) and independently from 1 to 3 In the cladograms produced with P A UP program, lies. (autapomorphy of Olivella) and from 1 to 2 14. Morphology of the salivary glands. Accor this character synapomorphic state for the dad e The well documented clade 14 corresponds to (autapomorphy of the clade 12). ding to the analysis the tubular salivary glands is 2 and in the cladograms produced by the volume of the family Olividae adopted by Ols appeared independently from acinous in Mela HENNIG86 program the plesiomorphic state son [19S6]. As it was said in introduction, sorne 6. Operculum. Operculum is lost indepen pium and clade 14. may be either O, or 1 or 2. The channeled suture dently in Oliva and Melapium. authors expressed their opinion on the isolation (state 1) is correlated with the presence of the of the subfamily Olivellinae as a separate family. 1 S. The length of the salivary tjucts. The posterior mantle tentacle (the apomorphic Golikov and Starobogatov [1988] established the 7. Presence of the posterior mantle lobe. The réconstruction of the character evolution suggest condition). So, it seems more likely that the family Olivellidae and placed it into a new mono character is synapomorphic for clade 14 an4 the independent origin of the shortened salivary plesiomorphic state of the character is Oor 2. typical suhordcr Olivelloidei Golikov et Starobo appears independently in Benthobia. ducts in clade 13 and 12. Clade 11: Oliva, Olivella [2(1), 8(1), 9(1) ]. gatov, 1988, thus opposing it to ncarly al! other 8. Presence of the anterior mantle tentacle.. 16. Accessory salivary glands. The Oliva [6(1), 13(1), 18(1) ]. The appearance stenoglossan Neogastropoda. Their brief diagno The character is synapomorphic for clade 11. reconstruction suggests the plesiomorphic of the glandular epithelium of the oesophagus is sis of the subordcr lacks a lot of importan! fea condition of the state 1 of the character in remarkab1e, but occurs also in clade 13 and Me Jures: "Bucciniformes with smooth tun-eted shell. 9. Presence o! the posterior mantle tentacle. Head Jacks tentacles and eyes. Radula has outer The character is synapomorphic for cladc 11. comparison with the state 2. The Jast one is the lapium. apomorphy for Melapium and Olivella. pair of marginal teeth lacking thc bend and ha Reduction of one of the accessory salivary glands Clade 15: Olivella [2(2), S(3), 10(1), 16(2), ving the fonn of rectangular plates. Family Oli 1 O. Position of the radula. Although the vellidae Golikov et Starobogatov fam.nov. (Shell reconstruction suggests the position of the radula is most probably synapomorphic for the ingroup. 17(1), 19(1), 22(!), 23 (1) ]. The genus is well defined with 8 apomorphies, 6 of which are aut small, turreted-cylindrical. Foot large with broad a t the proboscis tip (state 0) to be plesiomorphic, lateral lobes. Central radular tooth broad and this is in contradiction with our data on Benthobia apomorphies. Shift of the radula to the probaseis 17. The gland of Leiblein. Reduction of the base is probably secondary and therefore apo with many cusps)." (op.cit., p. 73). anatomy (see description and discussion of the This point of view is disproved by my data. character 11). Thus, the radular position at the gland of Leiblein is an autapomorphy for morphic in comparison with Oliva. The interme Olivella. diate morphological stage is found in Amalda and Dueto the numbcr of synapomorphies Olivella is probaseis base should be considered as closely related to Oliva, although differs greatly plesiomorphic for the clade 13. Nevertheless the Melapium, in which the radula position is at the 18. Accessory oesophageal gland. Reconstruc in possessing many autapomorphic chara~ters. In cladogram shows a possibility of reversion of the probaseis base, but the radular diverticulum tion suggests the possibilities of either indepen opens near the probaseis tip. Characters 2, 22 m y opinion, the genus Olivella should be ¡solated character in Olivella. This reversion should be dent origin of the character in Oliva, Melapium as a separate family but its p1acement ínto a considered as an autapomorphy for 0/ivella. and 23 are connected with each other. and clade 13, or reduction of the gland in clades scparate from other Olividae subor~er (as Position of the radula at the probaseis base in 12 and Olivella. Clade 12: Amalda montrouzieri, A. dimidiata proposed by Golikov and Starohogatov) ~s. dou~ Olivella may be connected with the feeding me [1(2), S(2), 12(0), 13(1), lS(l), 19(?2) ]. The tful. I clearly realize, that the rest of Olmdae·Is chanism (see Discussion of the morphology). 19. Morphology of the stomac'h. monophyly of this clade is supported with 5 apo morphologically heterogeneous and complex gro Reconstruction suggests the independent origin 11. Connection of the radular retractar wíth morphies one of which - reduction of the u p. Much more species should be studied, before of the tubular U-shaped stomach (type 3, marginal teeth [13(1)] occurs independently in the final decisions on the phylogeny and rela the columellar muscle. According to the formal autapomorphy for Amalda) and the type 2 analysis the state 1 of the character should be other clades. (No information is available on the tionships of all olivid genera can be made. In this stomach (autapomorphy for Olivella) from the morphology of stomach of A. dimidiata.) So, the connection although I do not propose)he system considered as plesiomorphic. Nevertheless, the plesiomorphic type 1 stomach. plesiomorphic condition is the state O (see state · 2 of the character 19 may he either an of the whole group, it is worth to re.. establish the description of the character). So, to m y mind, 20. Presence of the anal gland. The analysis apomorphy for the whole clade or autapomorphy supcrfamily Olivoidea [as proposed by Olsson, losses of the connection occurred independently shows a possibility of numerous reductions of the of Ama/da montrouzieri. Only one character 19S6] to include Olivellidae and Olividae. It sho in clade 14, Pseudoliva and Melapium. anal gland (in Olivella verreauxii, Amalda dimi distinguishes the two genera, the absence of the uld be emphasized, that the latter family proba diata, Melapium and clade 13). anal gland in A. dimidiata. bly is a paraphyletic group. 12. Connection of the radular refractor with The clade 13 corresponds to the subfamily the body wall. Reconstruction of the character As it was said in the description of the char Clade 13: Benthobia, Pseudoliva [13(1), Pseudolivinae, which was already isolated as a evolution suggests that the presence of this acters, state l of the character 21 is an autapo IS (1), 18 (1), 20(1) ]. The monophy~y of the separate family [Golikov, Starobogatov, 1975]. connection is an apomorphic condition. So, the morphy for Melapium, while states 1 of the char clade is supported by only 4 apomorph1es, all of In note 34 of the cited paper, the authors wrote: state O is an autapomorphy for the clade 12. acters 22 and 23 are the autapomorphies for which appear to be parallel in the other clades. "The subfamily Pseudolivinae, being rather far Pseudoliva [11 (1) J is characterized by only Olivella. from other Olividae asto shell characters and dif 13. The number of radular teeth per transverse one weak apomorphy. fering greatly from them in radula, is elevated to Benthobia [7 (1) also is characterized by a row. According to formal analysis the presence of CLADE ANALYSfS. J family rank... On the basis of shell structure and marginal teeth in Olivella should be considered single apomorphy which appears independently radula we assign the family Pseudolividae lo the as an apomorphic condition. Pondcr [ 1973 J also During the discussion of the principal clades in ciad e 14. superfamily Buccinoidea." 1 agree with the ne stated, that additional plates appeared the character-state reconstruction advocated in Clade S: Melapium [6(1), 11(1), 13(1), ccssity of the isolation of Pseudolivinae into se secondarily in Olive/la. Nevertheless, it seems to the preceding section is used. Each clade is fol paraJe family. Cladistic analysis presented h~re 14(1), 16(2), 20(1), 21 (l) J. The genus is charac me, that the inGependent reduction of the lowed by included taxa and by a list of synapo did not finally resolve the problem of evaluatmg terized by 7 apomorphies with only one marginal teeth in other clades probably more morphies. Discussion is based on the consensus autapomorphy [21 (1) J. its position in the system of Neogastropo~a. As adequately describes the situation. Indeed, the cladogram with Nucella as the outgroup. Each Pseudolividae and Olivoidea do not cons1ltute a general evolutionary trend in Neogastropoda is clade is characterized by its apomorphies (in squ- monophylctic taxon, it is necessary to include the reduction of the number of teeth in a row. more differcut groups of Neogastropoda iuto the ::!:48~-----~------Yu.I.Kantor Morphology and relationships af oliviform gastropods 49 analysis. Genus Benthobia Dall, 1889 was only the radular sac at the proboscís base in Benthobia partial resorption of the inner walls of the shell, uali Macea pacnonaraeTCll y ocnonaHHll xOÓOTa rccently included in Pseudolívinae [Bouchet, and Zemira is a plesiomorphic condition. The sa resulting in their extreme thinness. This resor nepe11 OKOJIOrJIOTO'IHhiM nepnnhiM KOJiñ"OM. Pe Warén, 1985], and my morphological data me situation occurs ín Pseudolíva. ption of the walls does not result in any mor TpaKTOP pa;llyJihl MOJKeT COC;lii!Hl!TbC>I C KOJIYMM confirm thís point of view. The Pseudolividae is Ponder [1973] dístinguished 3 basic patterns phological changes of the body. In 0/ivel/a the JiliPHI>IM MYCKYJIOM. :lKeneaa Jieft6neií:na xopomo characterized by few apomorphíes and possesses of organization of the foregut ín the Neogastropo columella and inner walls in the upper part of the pa3BHTa. many primitivo features. The most importan! of da, probably evolved separately and derived from shell are completely resorbed (Fig. 4D) and this Tia¡¡;OTp>IJ.I OTJIH'laeTCll OT Conoidei nanH'liieM them are the plesiomorphic posítion of the radula a early neogastropod forerunner before the elon cat:ses consequent morphological changes. These JKCJie3hl H KJiarrana Jieií:6neií:na, OT Muricoidei at the proboscis base and the connection of the gation of the proboscis. These groupings are: l. are: the Ioss of the spiral morphology of the vis OTJlU'laCTC>I CUOCOÓOM Suborder Muricoidei Rafinesque, 1815 IIOJIHHTCJJbHOÜ CJHOHHOÜ %e.TIC3hl, pacnOJIO)KC... concave, plate which passes along the outer raCTC.H C BHCUIHC:if CTOpúllbi :BUCIJ;CpaJibHOI'O Hl!eM MCCTa OTKpb!TUll pa¡:¡yJillpHOro ):\HBCpTHKy sulface of the visceral mass and is attached to the Ka, liaCTitl.JHO norpy:>KcH B ceMeHHHK. K¡>hliiic·q¡ Family Melapiidae fam. nov. Jia B 6yKK3JibHYIO nOJIOCTh y BCplliHHbl XOÓOTa H B reinnants of the inner wall of the penultimate KpynHa5I lf OliCHb TOHKa5l, MJIH OTCyTCTByeT. O'lieHb 3H3l.J:HTeJlbHOM p33BHTHH )(OITOJIHHTeirbHO:Ü whorl. Vesicula seminalis large, situated at the JIOBa HC AH¡jJ¡j¡epeHI\IIPYCTCll OT TeJia, 6e3 Shell suboval with vcry large and convex body flHII(CBO.ll.HO:Ü: JKeJIC3bi, outer side of the visceral mass, partially embed JICII u rJiaa. ITporro¡:¡nil noru cepnoBUJIHhlil, whorl and small spire. Spiral sulcus on the body 3AME'IAHH51': OKOH'!aTeJibHOC IIOJIO>KCHHC ded into testis. Operculum large and very thin, or no¡:¡11n xpynm>le. MauTnll e nepe¡:¡nnMf~~E:z~ whorl absent. Siphonal canal short. Operculum Melapiidae cpe¡:¡n Muricoidei ocraercll ue>ICHhlM. absent. Head not differentiated from the body, MaHTllllHh!MH II1YnaJlhl\3MH H C 33JIHCH absent. Head tentacles long and slcndcr without without tentacles and eyes. Propodium crescent noil nonacrbro. PMynnpnhlil Memox eye-lobes. Eyes present, or absent. Propodium Superfamily Olivoidea Latreille, 1825 shaped. Parapodia large. Mantle with anterior CTCll y ocuonanu¡¡ xOÓOTa. )J(eJieaa JieM.nelína narrow. Parapodia absent. Radula sac positions and posterior mantle tentacles and with posterior ):{OITOJIHHTCJtbHaSI CJIIOHHa.SI )KeJIC33 OTCyTCTBYIOT. at the probaseis base in the probaseis contracted Shell usually glossy. Sutures usually narrow mantle lobe. Radula sac positions at the probaseis Pa¡:¡yJia o6paaonana xpynuh!M l\CHTpaJibHh!M state but the radular diverticulum opens into the and channeled, but may be covered by the callus. base. Gland of Leiblein and accessory salivary MHOr03Y6l\OBh!M 3YOOM, 0):\H03Y6110Bh!MH Jia'fe buccal cavity near the probaseis tip. Oesophagus Foot with parapodia and well developed crescent gland absent. Radula formcd of broad central paJihHh!MH lf ne60JiblliHMH Ilp!IMOyrOJibllb!MH betwecn the opening of the gland of Leiblein and shapcd propodium, subdivided by longitudinal multicuspidate tooth, unicuspidate laterals and MaprnHaJibHhiMII. the valve of Leiblein forms nearly a complete ring. cleft. Salivary glands formed of ramifying tubes. small rectangular marginals. CeMeilcrno BKJIIO' Tipe¡~Ba¡m:reJibHbiM aHaJIM3 COOTHoweHHll TYPPHJ{ (Gastropoda, Toxoglossa, Turridae) e pa3JIH'IHbiM THUOM paJ{yJillpHoro annapaTa 11 COBpeMeHHblX H HCKOnaeMblX Q>ayHaX A.B.CbiCOEB Hucmumym napa3umo;zozuu AH CCCP, JieuurtcKuii. npocne~em 33, 117071 Moc~eea PaccMaTpnnaercR COOTHomeHne "HeroKcorJIOCCHñiX" H "roKcorJIOCCHDix" (T.e. e npH MHTnnnoii: HJIH npo)IBHHYTOH crpyKTypoii: PMYJIS!plllo!X 3yOOB) BH.t\OBTurridae B COBpe MCHHhiX 1i HCKOHaCMNX 4Jaynax. lloKa3aHO, 'ITO B HOCJICJl;OBaTCJibliO CMCHliiOI!IHX ,n;pyr ¡q¡yra Kalinoaoii:cKnx cj¡aynax onpe,n;Me11noro pemo11a 11a6.mo,n;aeTCJI nocrenennoe cMe meune YK338HHOI'O COOTHOillCHlHI B flOJU>3y "TOKCOrJIOCCHhiXu BH!(OB. ?Jro, Bepo.SITHO, CBH.t\CTeJThC'l'BYCT 00 SBOJIIO"HOHHOM BNTCCHeHHH OOJieC HpHMHTRBH"'X BH.t\OB OOJieC npo• ,n;nnnyTI>IMn. Haó.nro)laercl! ana'lHTCJihHall reorpa4Jn'lecKaSI neo)lnOpoi\HOCTh KaK ncKO· 11 naeMhiX, TaK H COBpCMCHHDlX cPayH IIO COÚTHOilleHHIO "HeTOKCOrJIOCCHhiXn li TOKCOrJIOC- . cHnrx" BlfAOB. B conpeMeHHHIX epeHl.I,HpyeTC51. B.l{OJlb BCTBCIÍ npHBC,UCHh1 anOMOp type was found in Olivella. 3. The stomach has unordered (nonadditive) (characters 1, 10, 12, 1 the form of U-shaped tube, caecum absent. This 14, 15, 18, 19, 20). Either representative of the ci>HH1 llOJiy>KHpHbiM mpHtP'rOM Bb1J(MCH HOMCp np»3HRKR, flpocThiM - ero COCTOmnte. type was found in Amalda. family Volutidae (Alcithoe arabica Gmelin, 1791 The pathways of the changes of stomach mor - the data obtaincd from Ponder, 1970) or rep phology is nuclear. That is why the feature was resentative of Thaididae (Nucella heyseana Analysis of the parsimonious trees revealed that i:HSCUSSION OF THE CLADOGRAMS unordered during the analysis. (Dunker, 1882) - unpublished data of Kantor only two basic topologies exist. The differenee in 20. Presence of the anal gland. The anal and A.Medinskaya) were used as the outgroup. the topologies concerns the position of Melapium. CHARACTER ANALYSIS gland was found in Olivella borealís, Ama/da and Analysis wíth partially unordered characters A.nalysis with Nucella as an outgroup produced Oliva. lt is absent in all other genera studied as and with all characters unordered produeed the trees in which Melapium was included either in l. Form of the shell suture. The reconstruc well as in 0/ivella verreauxii [Marcus, Marcus, same resu1ts. When Nucel/a was used as the out the same clade with Pseudoliva or in the clade tion of the character evolution shows either the 1959]. The presence of the anal gland is group, the PAUP program produced 6 equally with Oliva, Olivella, and Amalda. This results in possibility of the origin of the channeled suture eonsidered to be a primitive feature among parsimonious trees (consisten e y index = O. 763 the consensus tree, in which Melapium is placed (state 1) from the suture, covered with the eallus Neogastropoda [Ponder, 1973 ], so its reduction with al! characters unordered and 0.744 with only in the separate clade (Fig. 19). Analysis with Al (state 2) in clade 12 or both states 1 and 2 inde is considered to be apomorphic. sorne unordered). When Alcithoe was used as !he cithoe as an outgroup produced a single topo pendently from the plesiomorphic state. The lat 21. Connection o[ the accessory oesophageal outgroup, the PAUP program produced 3 equally logy, the same as the second topology, obtained ter seems more probable to me. gland with the gland of Leiblein. As was said parsimonious trees (consistency index ~ O. 7 44 with Nucella as the outgroup. Below is the dis above, the glandular folds of the oesophageal with all characters unordered and O. 725 with only cussion of both cladograms. Discussion of the 2. Resorption o! inner walls of the shell is gland are usually continuous with the glandular sorne unordered). The HENNIG86 program pro characters and clades is based on the consensus synapomorphic for the clade 11 while the tissue of the gland of Leiblein. On the contrary, duced 2 trees with Nucel/a as the outgroup and cladogram with Nucella asan outgroup. complete resorption of the inner walls is the in Melapíum, in which the accessory oesophageal the single tree with Alcithoe as the outgroup. It is importan! to mention, that in al! the cla autapomorphic state for Olivel/a. dograms the taxa of the ingroup do not form a monophyletic taxon. 3. Presence o[ the pedallobes. Appearance of pedallobes is a synapomorphy of the clade 14. 46 Yu.J.Kantor Morphology and relationships of oliviform gastropods 47