Contr. Tert. Quatern. Geol. 32(4) 97-132 43 figs Leiden, December 1995

An outline of cassoidean phylogeny

(Mollusca, Gastropoda)

Frank Riedel

Berlin, Germany

Riedel, Frank. An outline of cassoidean phylogeny (Mollusca, Gastropoda). — Contr. Tert. Quatern. Geo!., 32(4): 97-132, 43 figs.

Leiden, December 1995.

The phylogeny of cassoidean gastropods is reviewed, incorporating most of the biological and palaeontological data from the literature.

Several characters have been checked personally and some new data are presented and included in the cladistic analysis. The

Laubierinioidea, Calyptraeoidea and Capuloidea are used as outgroups. Twenty-three apomorphies are discussed and used to define

cassoid relations at the subfamily level. A classification is presented in which only three families are recognised. The Ranellidae

contains the subfamilies Bursinae, Cymatiinae and Ranellinae. The Pisanianurinae is removed from the Ranellidae and attributed to

the Laubierinioidea.The Cassidae include the Cassinae, Oocorythinae, Phaliinae and Tonninae. The Ranellinae and Oocorythinae are

and considered the of their families. The third the both paraphyletic taxa are to represent stem-groups family, Personidae, cannot be

subdivided and for anatomical evolved from Cretaceous into subfamilies reasons probably the same Early gastropod ancestor as the

Ranellidae. have from Ranellidae the Late Cretaceous. The Cassidae (Oocorythinae) appears to branched off the (Ranellinae) during

The first significant radiation of the Ranellidae/Cassidaebranch took place in the Eocene. The Tonninae represents the youngest branch

of the phylogenetic tree.

Key words — Neomesogastropoda, Cassoidea, ecology, morphology, fossil evidence, systematics.

Dr F. Riedei, Freie Universitat Berlin, Institut fiir Palaontologie, MalteserstraBe 74-100, Haus D, D-12249 Berlin, Germany.

Contents superfamily, some of them presenting a complete classifi-

cation. Modern prosobranch classification schemes (e.g.

Ponder & Warén, 1988; Vaught, 1989) recognise five Introduction p. 97 cassoid families: Bursidae, Cassidae, Ficidae, Ranellidae Material and methods p. 97 and Tonnidae. Beu (1988a; see also Beu & Maxwell, Classification p. 98 1990) convincingly showed that Distorsio and its allies, habitat and diet 99 Distribution, p. which previously had been attributed to the Ranellidae, masses and 102 Egg intracapsular development .... p. distinct This is represent a family. strongly supported Larval development p. 105 here and was adopted in the latest comprehensive publi- Protoconch and teleoconch p. 107 cation on the Cassoidea by Warén & Bouchet (1990), Anatomical features p. Ill who also introduced the Laubierinidae as a new cassoid Fossil record p. 120 family. The classification has been subsequently revised Phylogenetic cladogram and discussion p. 124 by Riedel (1994), who removed the Ficidae from the Acknowledgements p. 125 Cassoidea and by Bandel & Riedel (1994), who excluded References 125 p. the Laubierinidae. The remaining five families are the

subject of this paper, which is an attempt at a clearly

defined phylogenetic analysis on which a classification

Introduction can be based.

The Cassoidea (= Doliacea = Tonnoidea), containing such

MATERIAL METHODS spectacular gastropods as the tritons and helmet-shells, AND

have been the subject of research for a long time. Many authors have contributed the of to knowledge the Living animals were observed and collected by the author 98

Fig. 1. A: Initial whorl of shell with indication how to measure the maximum diameter (MD) and method of counting whorls; N -

of I - initial axis B: of the last nonspiral shell, defining zero point. height protoconch whorl has to be correlated to get a better

understanding of height/width ratio.

(by snorkeling, diving and dredging) at several places in specimens were photographed using a reflex camera. The

New Zealand (including Leigh Laboratory, Portobello method of counting whorls and measuring dimensions of

Marine Station), Australia (Long Reef/Sydney, Heron protoconchs is illustrated in Fig. 1.

Island, Lizard Island) and Bali (Indonesia). The snails

usually were relaxed using a magnesium chloride solution CLASSIFICATION isotonic with seawater (3.5 %) and then fixed in 70 %

ethanol or 5 to 10 % formalin.

Klaus Bandel shells of The of the is of based (Hamburg, Germany) provided arrangement taxa course on

Recent cassoids from the Caribbean, the Mediterranean phylogenetic considerations. However, in contrast to the

and the Red Sea. Alan Beu (Lower Hutt, New Zealand) phylogenetic cladogram, it appears preferable to introduce

and Bob Penniket (deceased, formerly Warkworth, New the classification as understood by the author, before the

donated characters of certain discussed. Zealand) generously several juvenile specimens taxa are This procedure with preserved protoconchs for further comparison. will hopefully avoid possible nomenclatural confusion.

Additional and material borrowed from the The main other dry wet was changes or rearrangements, compared to

Australian Museum (Sydney), the Institut fiir Hydro- classifications (e.g. Thiele, 1929; D.W. Taylor & Sohl,

biologie und Fischereiwissenschaft (Hamburg), the Musee 1962; Golikov & Starobogatov, 1975; Beu, 1981, 1985,

royal de l'Afrique Centrale (Tervuren), the National 1988a; Ponder & Waren, 1988; Beu & Maxwell, 1990;

Museum of & New Zealand (Wellington) and the Zoo- Waren Bouchet, 1990) are briefly introduced, but will

logisches Institut und Museum (Hamburg). be discussed in more detail elsewhere in the present

Fossil material originating from the Campanian of paper.

Mississippi (Coon Creek) and the Paleocene of Alabama The Ficidae Meek, 1864 have already been removed

from the (Matthews Landing) was collected by Klaus Bandel in Cassoidea and been recognised as a superfamily co-operation with David T. Dockery (Jackson). Klaus (Riedel, 1994). The inclusion of the Laubierinidae Waren

Bandel and colleagues provided Eocene gastropods from & Bouchet, 1990 and Pisanianurinae Waren & Bouchet,

1990 in the the Brazos River (Texas) and Oligocene gastropods from Cassoidea presents difficulties on account of

Glimmerode which cassoids the absence of larval which unites (northern Germany) among a pallial appendage, with the other families Bandel & well-preserved protoconchs were examined. Several (compare Riedel, 1994). No excellently preserved gastropod faunas from various secondary periostracum on the larval shells is documented

European Paleocene, Eocene and Miocene localities were in the literature (Waren & Bouchet, 1990). Moreover, the borrowed from the Institut royal des Sciences naturelles Laubierinidae and Pisanianurinae do not show the de Belgique (Brussels), the Nationaal Natuurhistorisch episodic growth of the teleoconch which is characteristic

Museum (Leiden) and the Naturhistorisches Museum for cassoids. It is difficult to imagine that this character

lost. other (Vienna). was Many gaps in knowledge have to be

and Protoconchs, shells of juveniles and radulae were bridged other anatomical features are of uncertain mounted coated with the in the on stubs, gold and examined with interpretation, e.g. monopectinate osphradium the Laubierinidae aid of an electron microscope (CamScan). Larger or the asymmetrical osphradium in the 99

Pisanianurinae have not been recorded in the Cassoidea. 3a - Subfamily Cassinae Latreille, 1825

The Laubierinidae had already been removed from the Cassis Scopoli, 1777

Cassoidea (Bandel & Riedel, 1994) and the Pi- Cypraecassis Stutchbury, 1837

sanianurinae, which, in addition to conchological charac-

3b - Oocorythinae Fischer, 1885 also show anatomical features which Subfamily ters are very similar Dalium Dall, 1889 to those of laubierinids, should be provisionally placed Galeodea Link, 1807 within the superfamily Laubierinioidea.Alan Beu (pers. Oocorys Fischer, 1883 comm.) concurs with this view. Sconsia Gray, 1847 The tun-shells and its (Tonna relatives) are recognised as a subfamily of the Cassidae and the frog-shells (Bursa 3c - Subfamily Phaliinae Beu, 1981 and its the sister of the relatives) possibly represent group Casmaria H. & A. Adams, 1853

Cymatiinae. Sassia and Charonia have been removed Phalium Link, 1807

from the Cymatiinae and are attributed to the Ranellinae Semicassis Morch, 1852 but perhaps represent an independent ranellid branch

3d - Tonninae Suter, 1913 (subfamily). Dalium, Galeodea and Sconsia have usually Subfamily Eudolium Dali, 1889 been assigned to the Cassinae, but are here considered to Malea Valenciennes, 1883 be better placed in the Oocorythinae. Subgenera and Tonna Briinnich, 1772 extinct taxa not included in the classification are pre- sented below. Personopsis currently contains but a single

Recent species, which, however, will be reassigned by DISTRIBUTION, HABITAT AND DIET Beu (pers. comm.) to a new personid genus. This means that Personopsis will have to be removed from the list of Personidae live in most warm oceans and are mainly when that Recent genera new name is coined. represented by the genus Distorsio (compare Clench &

Turner, 1957; Lewis, 1972; Beu, 1985). Personopsis (?)

is known exclusively from the West Pacific (Beu, 1985)

Subclass Caenogastropoda Cox, 1959 and Distorsionella appears to be restricted to the southern Order 1991 Neomesogastropoda Bandel, West Pacific (Beu, 1978b, 1985). Suborder Troschelina Bandel & Riedel, 1994 Common species such as Distorsio anus (Linne, 1758) Superfamily Cassoidea Latreille, 1825 and Distorsio reticulata Roding, 1798, can be found in

shallow-water habitats (Orr Maes, 1967; Cernohorsky,

1 - Personidae 1854 Family Gray, 1971; Lewis, 1972), usually under coral rubble (Cerno- Distorsio Rdding, 1798 horsky, 1971). Distorsio kurzi Petuch & Harasewych, Distorsionella Beu, 1978 1980 was reported from 120-130 m depth in the Sulu Sea ? Personopsis Beu, 1988 (Petuch & Harasewych, 1980), which however is easily

surpassed by Distorsionella lewisi Beu, 1978, which was 2 - Family Ranellidae Gray, 1854 dredged in more than 600 m depth (Beu, 1978b). Shasky 2a - Subfamily Bursinae Thiele, 1925 (1992) (?) Bufonaria Schumacher, 1817 reported Personopsis pusilla (Pease, 1861)

from about 10-30 stones and of Bursa Roding, 1798 m depth living on pieces

corals. Crossata Jousseaume, 1881

There observations of Tutufa Jousseaume, 1881 are no on the feeding strategy

the Personidae. However, J.D. Taylor (1980) found

2b - Iredale, 1913 Subfamily Cymatiinae remains of a chaetopterid (Polychaeta) in the alimentary

Cabestana 1798 Roding, tract of Distorsio reticulata. Personal examination of the

Cymatium Roding, 1798 stomach of Distorsio sp. has revealed the cuticle of a

'worm'.

2c - Subfamily Ranellinae Gray, 1854 Ranellidae are distributed worldwide and at present Argobuccinum Hcrrmannsen, 1846 are missing only from Arctic and Antarctic seas. Some ? Charonia Gistel, 1848 species occur at places across the world, Cy- Fusitriton Cossmann, 1903 many e.g.

matium parthenopeum (von Salis, 1793), which was Gyrineum Link, 1807

recorded from the western Atlantic Ranella Lamarck, 1816 (see Beu, 1985)

the Atlantic ? Sassia Bellardi, 1872 (Bermudas to Uruguay), eastern (Spain to

Angola), the Mediterranean, South Africa, the northern

3 - Family Cassidae Latreille, 1825 Indian Ocean, Japan, Hawaii, Australia and New Zealand 100

Fusitriton (Laxton, 1968; Beu, 1970c). Other species of Cymatium hard substrates (Beu, 1978a). aurora Hedley, which usually do not exceed these latitudes. Cabestana has been 1916 was reported from 3,300 m (Hedley, 1916),

would have been the record for the reported as far north as Great Britain (Fretter & Graham, greatest depth on

this is buccinid 1981; Graham, 1988) and as far south as New Zealand ranellids, but species a (compare Dell,

be found (Powell, 1979) and Argentina but does not occur in the 1972). Sassia kampyla (Watson, 1855) can tropics (Beu, 1985). Charonia, with the two species C. associated with F. magellanicus and is reported from as tritonis (Linne, 1758) and C. lampas (Linne, 1758), has deep as 900 m (Beu, 1978a). Argobuccinum pustulosum a similarly wide distribution as does Cymatium (Laxton, (Lightfoot, 1786) was (personally) found at depths

rocks 1968; Beu, 1970a, 1985; Fretter & Graham, 1981; between 40 to 100 m but also occurs on intertidal

Charonia, Graham, 1988). Ranella, also with two species attributed in southern New Zealand (Beu, pers. comm.).

in shallow lives but also to it, occurs from the western Atlantic (via the Indian fairly water, on rocky bottoms,

sand seaweeds Ocean) to New Zealand (Dell & Dance, 1963; Arnaud & on amongst (Laxton, 1968; Bandel,

both be found Beurois, 1972; Beu, 1985). Fusitriton has been reported 1976a). The two species of Charonia can

of the northern in Fretter & from the temperate and cool waters deeper water (Beu, 1970a; Graham, 1981; hemisphere (J.T. Smith, 1970; Stickle & Mrozek, 1973; Graham, 1988), which probably is the more common

Bursinae live in shallow Somerton, 1981; Beu, 1985) as well as from the southern habitat for the genus. Most or

hard of coral hemisphere (J.T. Smith, 1970; Penchaszadeh & De fairly deep water. They prefer substrates

Mahieu, 1975; Beu, 1978a, 1985) but not from the tropics reefs and surrounding areas (Orr Maes, 1967; Cer-

Bandel & (see discussion). Argobuccinum is restricted to the nohorsky, 1971; Bandel, 1976a; Wedler, 1987; southern hemisphere, being absent from the tropics (J.T. pers. obs.).

ranellids consists of dead animals Smith, 1970; Beu, 1985). Sassia is a predominantly Indo- The diet of living or

of Caribbean wide of taxa. Several have Pacific genus with the exception a single representing a range genera which habit is species (Sassia lewisi Harasewych & Petuch, 1980). been observed to prey on echinoderms,

has 'worldwide' in otherwise known from Cassidae. Gyrineum a distribution, mainly exclusively Cymatium has been found hunt other tropical, but also in temperate seas (Beu, 1985). The nicobaricum to gastropods,

Indo-Pacific Conus 'worms' and Bursinae occur mainly in the (Beu, 1981, including (see Kohn, 1959), bivalves,

1985) but also in the Atlantic Ocean, including the barnacles (Houbrick & Fretter, 1969; Bandel, 1976a).

secretes Mediterranean (Sabelli & Spada, 1982). The group is Cymatium parthenopeum probably a relaxing

their unknown from cool waters at high latitudes. substance, which induces bivalve prey to open

valves has been ob- The species of Cymatium can be found most com- (Loch, 1982a). Cymatium pileare monly in calm (Bandel & Wedler, 1987) and shallow served to prey on mangrove oysters (Littlewood, 1989). waters, usually under stones, broken platform corals, etc. C. (Linatella) caudatum has 'a catholic diet' (Morton, and (Orr Maes, 1967; Houbrick & Fretter, 1969; Bandel, 1990, p. 478), preying on gastropods, cirripedes

such C. nicobaricum ascidians and is of farms in southern 1976a; pers. obs.). Some species, as a major pest oyster

Cabestana to (Roding, 1798), enter open (unsheltered) areas relatively India (Beu, pers. comm.). spengleri seems

ascidians often. Cymatium pileare (Linne, 1758) prefers the prefer (Tunicata) (Laxton, 1968; Riedel, 1992), seaward side of reefs where currents are strong (J.D. but was also observed feeding on polychaetes (Laxton,

The Charonia became famous Taylor, 1978), while C. parthenopeum is known to dig 1968). genus through

into soft substrates (Kilburn, 1970). Cymatium (Linatella) Copland (1966), who reported that the triton preyed on caudatum (Gmelin, 1791) has been reported from coral the reef coral-killing asteroid Acanthaster planci (Linne,

rubble and sandy bottoms in fairly deep water 1758). Charonia has since been observed to feed on other

well echinoids and cucumbers (Cernohorsky, 1971; Morton, 1990) but also occurs sea stars, as as on sea

in Bandel commonly on intertidal rocks Hong Kong (Morton, (Laxton, 1968; Beu, 1970a; Bandel, 1976a, 1984;

and bivalves In 1990). The genus Cabestana, with C. spengleri (Perry, & Wedler, 1987), on (Percharde, 1972).

Laboratoire 1811), is known from intertidal rocky areas (Riedel, the aquarium tanks of the Arago (southern

triton fed for several fish and 1992) and C. cutacea (Linne, 1767) from soft bottoms France), a was years on

alone Kohn (Fretter & Graham, 1981; Graham, 1988). The Ranellinae squid (Bandel, pers. comm., 1992). (1983)

live Fusitriton the ranellid with the widest contain genera or even single species which usually designated as range

in of water and because of this can of food discussion). This feeds on echinoids, a wide range depth (see genus bivalves, also be found on different substrates. A good example is asteroids, ophiuroids, tunicates, gastropods,

Fusitriton magellanicus (Roding, 1798), which is known chitons and polychaetes. Fusitriton is able to paralyse its

down and substance based on tetra- from 20 m (pers. obs.) to 1,500 m depth (Beu, prey possibly uses a

but also & Itoh, on 1978a), living on sandy or muddy bottoms, on methylammonium (Asano 1960). Day (1969), 101

the basis of aquarium experiments, proposed that eastern Africa to western America (Kilias, 1962).

Argobuccinum pustulosum preferred polychaetes. Gyri- Most Cassidae seem to prefer soft substrates, from

neum natator found to shallow water. However, there are few (Roding, 1798) was consume to very deep very algae (J.D. Taylor et al., 1980), which habit is unknown records. Cassis cornuta (Linne, 1758) is regularly found in other Cassoidea (except for larvae). Among the on sandy bottoms at depths of 5 to 10 m (pers. obs.).

Bursinae, several species ofBursa have been observed to Bandel & Wedler (1987) described C. tuberosa (Linne,

1822 also from consume 'starfish' (Orr Maes, 1967), ophiuroids and 1758) and C. madagascariensis Lamarck, echinoids (J.D. Taylor, 1978, 1984; Kohn, 1983), anne- sandy bottoms and observed the juveniles to dig them- lids and other 'worms' (Houbrick & Fretter, 1969; J.D. selves in. This applies also to Cypraecassis testiculus

Taylor, 1978, 1984; Bandel & Wedler, 1987), molluscs (Linne, 1758), but digging is not restricted to early and cirripedes (Bandel, 1976a; Bandel & Wedler, 1987) ontogeny (Bandel, 1976a). Cypraecassis coarctata (So- and The alimen- observed the vertical sides even sponges (J.D. Taylor, 1978, 1984). werby, 1825) was to move on

of in addition of tary tract Bufonaria rana (Linne, 1758), to of rocks (Abbott, 1968). Both species Cypraecassis ophiuroids and polychaetes, contained remains of fish were found in comparatively shallow water. Phalium

(J.D. Taylor, 1980). I observed how a specimen of Tutufa glaucum (Linne, 1758) prefers sandy areas of the unshel- rubeta (Linne, 1758) swallowed a comparatively large tered seaward side of reefs (Abbott, 1968). Semicassis

whole. ophiuroid labiata was found in the intertidal, covered by sand (pers.

Species richness in the Cassidae is greatest in tropical obs.). One species of Semicassis, S. microstoma (von

Some into and from soft bottoms in seas. species, however, range temperate Martens, 1903) was dredged water cooler The that make the waters. two genera up Cassinae, depths of 200 to c 1,100 m (Abbott, 1968). Members of

Cassis and be confined Cypraecassis, appear to to warm the Oocorythinae reach the greatest depth amongst oceans, and probably both genera are longitudinally representatives of the superfamily. Fischer (1883) re- distributed worldwide. Among the Phaliinae, the genus corded Oocorys from depth in excess of 3,600 m.

Semicassis with distribution. predominates, a near-global Galeodea rugosa (Linne, 1771) has been found living on

Semicassis labiata be found in the (Perry, 1811) can a muddy soft bottom at depths down to about 700 m southern hemisphere and like S. semigranosa (Lamarck, (Fretter & Graham, 1981; Graham, 1988). Tonna galea,

1822) occurs in subantarctic seas (Abbott, 1968; Powell, when resting, has usually been found subtidally, hidden

1979). Phalium is restricted to the Indo-Pacific and the in sand (Bandel & Wedler, 1987). Riedl (1983) character- third genus, Casmaria, occurs in the tropical Indo-Pacific, ised the habitat of the same species as a sublittoral soft with vibexmexicana the exception of C. (Stearns, 1894), bottom which may be largely covered by empty shells. which is found off tropical west America, and of C. Wilson & Gillet (1971) recorded other species of Tonna ponderosa (Gmelin, 1791), which is also known from the burying themselves in soft substrates. Eudolium has been

Caribbean & (Emerson Old, 1963; Abbott, 1968; Bradner dredged from soft bottoms, down to 800 m (Turner,

& DuShane, 1982). Among the Oocorythinae the genus 1948; Marshall, 1992). Kilias (1962) maintained that

Galeodea is distributed. Its in Malea most widely species occur might occur even in the deep sea. Bandel (pers. the and eastern Atlantic & warm temperate (Fretter comm., 1992) found Malea pomum (Linne, 1758) in

the Mediterranean Graham, 1981; Graham, 1988), shallow water, on a sandy bottom.

(Erlanger, 1893; Fioroni, 1966a; Sabclli & Spada, 1978) There are a few reports on cassid diets. Schenck and in the southwest Pacific (Dell, 1953; Ponder, 1984; (1926) assumed bivalves to be the main food item. pers. obs.). Oocorys has been recorded in the literature However, several species of Cassis obviously prefer only from the (sub)tropical Atlantic (Fischer, 1883; regular (Lyman, 1937; D.R. Moore, 1956; Schroeder,

Turner, 1948; Quinn, 1980; Waren & Bouchet, 1990; 1962; Hughes & Hughes, 1971, 1981) or irregular

Bouchet & Waren, but O. sulcata 1883 is echinoids obs. for 1993), Fischer, (Foster, 1947; D.R. Moore, 1956; pers. abundant at bathyal depths all around New Zealand (Beu, Cassis cornuta). Edmunds & Edmunds (1973) observed pers. comm.). Thus the genus may have a similarly wide (under aquarium conditions) Cassis tuberosa to feed on distribution as Galeodea. Dalium is known from the asteroids. Cypraecassis testiculus has been reported to

Caribbean and Sconsia lives in only the western Atlantic prey on regular (Hughes & Hughes, 1981) as well as on

(Clench & Abbott, 1943; Clench, 1959). The Tonninae irregular echinoids (Bandel & Wedler, 1987). Among the

and which is that comprises two genera (Tonn a Eudolium) can Oocorythinae there one report (under aquarium be found in the Atlantic well in the Indo-Pacific. Galeodea as as conditions) echinophora (Linne, 1758) preys on

Tonna galea (Linne, 1758) represents the species with the the irregular echinoid Echinocardium cordatum (Pennant, widest east-west distribution(Turner, 1948; Kilias, 1962; 1777) (Hughes, 1986a). Quinn (1980), on the basis of the

Abbott, The third Malea, is known from feed 1974). genus, environment, speculated that Oocorys could on 102

'worms'. Species of Phalium and Semicassis have been Feeding experiments under aquarium conditions have described eating echinoids (D.R. Moore, 1956; J.D. clearly demonstrated that a hungry cassid loses its food

bivalves The and this also under natural circum- Taylor, 1980) or (Abbott, 1968; Day, 1974). preference may occur

it is offer Tonninae are recorded to feed on fish, crustaceans stances. However, not very convincing to only

(Cernohorsky, 1972), echinoderms in general (Cerno- holothurians (see Morton, 1991) as food item and con-

these echino- horsky, 1972; Powell, 1979) and bivalves (Powell, 1979). clude that the genus Tonna specialises on

of in the derms. In addition Weber offered Weber (1927) found a species Natica oesopha- to holothurians, (1927)

naticids and these under gus of Tonna galea. Species of Tonna have several times to Tonna, were preferred (also been observed to swallow holothurians whole (Grange, aquarium conditions). However, observations under

stomach 1974; Bandel, 1976a, 1984; Sarver, 1977; Kohn, 1983; natural conditions and based on examination of

also & indicate Morton, 1991; Fig. 2). contents (see Bentivegna Toscano, 1991)

for that the Cassidae actually have a strong preference

echinoderms and that several genera or species prefer

of echinoderms. In particular groups or species contrast,

the Ranellidae are not specialised echinoderm feeders, but

the fact that they may eat these animals unites them with

the Cassidae. It is assumed that the Personidae do not

basis of two prey on echinoderms, purely on the negative

these reports and judging also from the anatomy of

gastropods (see below).

EGG MASSES AND INTRACAPSULAR

DEVELOPMENT

Fig. 2. Tonna olearia feeding on a holothurian (after Morton, of the Personidae is based Knowledge of the masses 1991). Not to scale. egg three different of Distorsio on reports on species (d'As-

Kasinathan aro, 1969a; Cernohorsky, 1971; et al., 1974).

Marshall found sclerites of holothurian in the about 1 (1992) a The egg capsules are flattened spheres (Fig. 4e),

tract of a Eudolium alimentary species. mm in diameter, each containing 20-40 embryos

— the distributionof Summarising remarks naturally, (d'Asaro, 1969a; Kasinathan et al., 1974). The egg

the Cassoidea is correlated their in more-or-less to postembryonic capsules are usually arranged regular arcs;

Below, the almost 'worldwide' occurrence not development. however, with many capsules produced, they form

is linked to of species such as Cymatium parthenopeum one but several masses (d'Asaro, 1969a; Cernohorsky,

larval The of two of the development. separation species 1971). The largest known spawn (d'Asaro, 1969a)

Fusitriton by the tropical ocean acting as a barrier contained about 50,000 embryos. The embryos of

appears strange. However, temperature changes of ocean Distorsio clathrata (Lamarck, 1816) and D. reticularis

surface occurred several times since the evolution waters (Linne, 1758) were observed to hatch as free-swimming

A of Fusitriton (at least since the Oligocene). temperature veligers (d'Asaro, 1969a; Kasinathan et al., 1974). The

decrease of about 5° C Plio-Pleistocene d'Asaro during glaciations only detailed embryology was presented by

(Krommelbein, 1986) may have been sufficient for the (1969a).

larvae to cross the tropics (compare Beu, 1970d) and a of most of the Ranellidae have The egg masses genera since second species of Fusitriton may have evolved been described in the literature. Among the Cymatiinae

then. 1928; the spawn of several species of Cymatium (Lamy,

Genera reaching great water depths are obviously Bouillaire, 1953; Arakawa, 1960; Laxton, 1969; Bandel,

adapted to cold temperatures. Consequently the same 1976a; Ramon, 1991) and Cabestana (Simroth, 1907;

be found closest to the genera, such as Fusitriton, can Lamy, 1928; D.T. Anderson, 1959; Laxton, 1968, 1969; such polar oceans. Vice versa, shallow-water species as Penchaszadeh & De Mahieu, 1975; Douglas, 1985;

known extend latitudes. most Bursinae, are not to to high Ramon, 1991; Riedel, 1992) have been figured and/or

The substrate preferred certain cassoids usually can be of the masses is hemi- by described. The shape egg always

correlated their A wide flattened base. The hemi- directly to potential prey. range spherical, with a gelatinous

of food also wide of different sub- usually means a range sphere contains about 80 to 350 egg capsules, arranged in

strates to live on. a three-dimensional spiral (Fig. 3). 103

The diameters of the egg masses match the maximum

width of the teleoconch aperture and sizes of egg masses

and capsules are correlated to the sizes of the spawning

females. The females were usually found to incubate the

egg masses. The embryos of these recorded egg masses

hatched as free-swimming veligers (except D.T. Ander-

but The number of son, 1959, see Riedel, 1992). embryos

varies from about 100,000 to 900,000 (pers. obs.) per

spawn (see remarks).

The egg capsules in Fusitriton are also arranged in a

spiral, which is, however, two-dimensional (Philpott,

1925; Howard, 1962; Penchaszadeh & De Mahieu, 1975).

Kilburn 1982; Argobuccinum (see & Rippey, pers. obs.),

Ranella (see Laxton, 1968, 1969; Latigan, 1976),

3. view of Fig. Apical semi-hemispherical egg mass character- Charonia (see Lo Bianco, 1888; Bougis, 1950;

istic of and Bursinae. Not to scale. Cymatiinae McCamley, 1965; Berg, 1971; Cross, 1973; Amoroux,

1974; Bandel, 1976a; Marler & Marler, 1982) and The shape of the capsules is more-or-less finger-shaped Gyrineum (see Petit & Risbec, 1929) arrange their egg- (Fig. 4d). capsules in more-or-less regular rows.

Fig. 4. Egg capsule types of Cassoidea in lateral view and apical view in d and h:

a - Sassia (after B.J. Smith et al., 1989); b - Fusitriton, Ranella; c - Argobuccinum; d - Cymatiinae and Bursinae; e - Distorsio

- Tonna - Marler & h - Cassis (after d’Asaro, 1969a); f (after Bandel, 1976a); g Charonia (after Marler, 1982); (after d’Asaro,

1969b); i - Cypraecassis (after Bandel, 1976a). Not to scale. 104

well The are Cassis Anonymous, 1961; d'Asaro, 1969b) as The shapes of the egg masses vary. egg capsules (see

for roughly finger-shaped in Charonia, Ranella, Argo- as Cypraecassis (see Work, 1969; Bandel, 1976a;

The in buccinum and Fusitriton (Fig. 4) and may be somewhat Hughes & Hughes, 1987). egg capsules

are column-like (Fig. 4i), with a rounded angulate (Latigan, 1976; pers. obs.). Cypraecassis

and in Cassis vasiform with The capsules of Charonia can be easily distinguished cross section, are a more-or-

section The of the by their circular cross section and the bulbous apex (Fig. less rectangular cross (Fig. 4h). apices

subdistorta in both are somewhat collar-shaped, the 4g). The egg capsules of Sassia (Austrotriton) capsules genera

wine- central of the surrounded (Lamarck, 1822) have been described as being plate apex being by a gelatinous glass-shaped (cited by Beu, 1988b from Watson, 1971; wall. Cassis cornuta has been reported to secrete about

and Cassis B.J. Smith et al., 1989; Fig. 4a). Sassia bassi (Angas, 230 capsules per spawn (Anonymous, 1961)

about 260 1869) produces similar stalked egg capsules (B.J. Smith madagascariensis (d'Asaro, 1969b).

than 800 et al., 1989). The capsules of Gyrineum gyrinum (Linne, Cypraecassis testiculus may spawn more egg

flattened & In- & Cassis 1758) are ellipsoids (Petit Risbec, 1929). capsules (Hughes Hughes, 1987). Reports on

cubation has been reported in Ranella australasia (Perry, cite about 600,000 embryos per spawn all of which hatch

than 1811) (see Laxton, 1969; Latigan, 1976), Charonia lam- as free-swimming veligers. In Cypraecassis more a

and Sassia subdistorta Beu, million hatch also Bandel, 1976a). pas (see Bandel, 1976a) (see veligers may (see

varies Incubation is known in the Cassinae. 1988b). The number of egg capsules per spawn not

of three greatly, usually from 100 to 300. An egg mass of Amongst the Phaliinae, the egg masses

Argobuccinum pustulosum contained 130 capsules, with Semicassis species have been described (D.T. Anderson,

Wilson & The altogether nearly 500,000 embryos that all hatched as 1965; Abbott, 1968; Gillet, 1971). egg

and for free-swimming veligers (pers. obs.). Free-swimming capsules are connected apically basally (except

veligers have also been recorded for Charonia, Ranella the 'ground floor') to each other (D.T. Anderson, 1965),

In and Fusitriton. The spawn of Gyrineum gyrinum was forming a tower-shaped egg mass. cases were commu-

Wilson & figured (Petit & Risbec, 1929) to consist of 18 capsules, nal spawning occurs (D.T. Anderson, 1965;

direction secretion is horizon- containing a total of about 250 embryos (see remarks). Gillet, 1971) the of capsule

tal rather than vertical Abbott, The The embryos of Sassia subdistorta feed on nurse eggs (see 1968). egg

more-or-less and lack and only a single crawling juvenile emerges from each capsules are finger-shaped an apical

but capsule (Watson, 1971; B.J. Smith et al., 1989). This collar. The number of developing embryos is vague,

applies also to Sassia bassi (see B.J. Smith et al., 1989). probably amounts to hundreds of thousands (compare

of not Many Bursinae have egg masses and capsules which D.T. Anderson, 1965). Hatching embryos was incubated 3, observed. masses are commonly are very similar to those of the Cymatiinae (Figs 4d). Huge egg

Anderson, 1965; Wilson & Gillet, smaller A more-or-less hemispherical or cup-shaped spawn was (D.T. 1971),

individuals described for Bursa granularis (Roding, 1798) (see ones by single (Abbott, 1968).

is also assumed for Risbec, 1931; McGinty, 1962; Orr Maes, 1967; Ganapati Communal spawning large egg

of Galeodea in- & Sastry, 1973; Bandel, 1976a), Bursa corrugata (Perry, masses (see Erlanger, 1893). However,

cubation has been observed. The 1811) (see d'Asaro, 1969a), Bursa rhodostoma (Sowerby, not egg capsules are wide 1835) (see Loch, 1982b), Bufonaria echinata (Link, more-or-less irregularly (pea-) shaped, nearly as as

1807) (see Thorson, 1940) and Tutufa bufo (Roding, high (Erlanger, 1893; Fioroni, 1966a; Hughes, 1986a).

(1893) and Fioroni (1966a) observed nurse 1798) (see Coleman, 1975). The egg masses were Erlanger eggs while incubated by most species. The finger-shaped capsules in the egg capsules of Galeodea echinophora,

in number from about 100 to 150 Hughes (1986a) reported intracapsular cannibalism. About vary per spawn.

the of Bursa 10 % of the few hundred capsule (Erlanger, According to d'Asaro (1969a) egg mass eggs per

and hatch as crawling juveniles. corrugata contains more than 100,000 embryos, which all 1893) finally develop

The of Galeodea (see Fioroni. hatch as free-swimming veligers. Thorson (1940) counted early development rugosa

of The about 50-60 developing embryos in each capsule of 1966a) is very similar to that G. echinophora.

of other have not been Bufonaria echinata spawn. There is one report of nurse embryogenies Oocorythinae

observed. eggs in Bursa (see Fioroni, 1966b, but compare remarks).

Bursa The of Tonninae is restricted A complete embryogeny of corrugata was pre- knowledge egg masses

sented by d'Asaro (1969a). to the genus Tonna. The egg capsules always form a flat,

three of tongue-shaped (or more-or-less irregular) (Thorson, The Cassidae exhibit at least types egg spawn

more-or-less Panikkar & 1950; masses. Most Cassinae form an irregular, 1940, 1942; Tampi, 1949; Ostergaard,

two-dimensional with the bases Amio, 1963; Gohar & Eisawy, 1967; Bandel, 1976a; spawn, egg-capsule (in

hard substrate. This holds for Penchaszadeh, The are in rows) attached to a true 1981). egg capsules arranged 105

arcs and have the shape of a flattened sphere (Fig. 4f), Cassis, whereas Distorsio produces the lowest number of

similar to those of the Personidae. The largest spawn was eggs in the superfamily. It must be emphasised that it is

reported by Bandel (1976a) to have a length of 43 cm important to compare the number of eggs in the early

and width of 8 The number of the number is reduced cm. egg capsules embryogeny, as greatly when

amounted to about 7,000 in that The nurse particular egg mass. egg feeding occurs. The low number of eggs

number of in each Tonna eggs egg capsule varies from reported from capsules of Gyrineum gyrinum (see Petit &

about 20 to 100 1940; 1950; Risbec, results from (Thorson, Ostergaard, 1929) probably nurse egg feeding or

Bandel, 1976a). Large egg masses therefore contain intracapsular cannibalism (compare Sassia subdistorta).

several hundred thousand embryos, which hatch Fioroni nurse from Bursa usually (1966b) reported eggs egg

as free-swimming & 1949; However, the veligers (Panikkar Tampi, capsules. capsules are too large for any

Bandel, 1976a). Thorson (1940) and Penchaszadeh (1981) species of Bursinae known from the Mediterranean.

both described in which egg masses, they suspected that Moreover, the spawn is of a different shape and was

two different modes of different development (in cap- probably produced by a neogastropod.

sules) occurred. Except for one report (Gohar & Eisawy, It is commonly accepted, and it will also be shown

the of Tonna all collected from 1967), egg masses were here, that the protoconchs of species in most cases can be

soft bottoms. Incubation is unknown, though Ostergaard used to distinguish non-planktotrophic (nurse eggs,

(1950) observed individuals of Tonna remaining near intracapsular cannibalism, lecithotrophic) from plank- their spawn, which could imply a kind of protection. totrophic development, without actual observations on the

remarks — Summarising the shape of an egg capsule early ontogeny.

is of correlated course with anatomical features. The Estimated egg contents of capsules in the literature,

of the be viewed in the such 'a few hundred' shape spawn, however, must as or 'several hundred', in some

context of ecology and behaviour. In cases where an cases need to be checked. Several tests I did demon-

individual of the Distorsio personid forms several egg strated that the actual number may be ten times the

masses (which appears to be the rule), it is probably not estimated total. Numbers of eggs and capsules have been

possible to protect or incubate the spawn. The hemispher- confused in the literature by Amio (1963), who quoted

ical which unites the and Arakawa's egg mass (Fig. 3), Cymatiinae (1960) count of the number of eggs per

be Bursinae, can protected perfectly, while the two- capsule in Cymatium parthenopeum, which, however, was

dimensional of Charonia is too to be the number of for that spawn e.g. large actually capsules egg mass. Kilias

fully incubated and several at the usually egg capsules (1962), citing Ostcrgaard (1950), reported 3,000 eggs in

eaten periphery are by predators (e.g. neogastropods; a spawn [Tonna perdix (Linne, 1758)], which was

Bandel, pers. comm.). Communal spawning and incuba- actually the number of capsules.

tion in Ranella and Semicassis have to be seen as Embryogenies usually have to be observed under

The of Semicassis characteristic conditions. convergences. spawn is aquarium Tank conditions in some instances

and cannot be confused. Tonna is the known only genus (e.g. D.T. Anderson, 1959; Gohar & Eisawy, 1967;

in the Cassoidea that places the egg mass directly on Latigan, 1976) may have led to abnormal development.

sand. The flat, broad spawn is well adapted to that However, it must be taken into consideration that these

habitat. However, in naticids, does not conditions also under camouflage, as possibly may occur natural circum-

occur. stances, and therefore such results should be included in

There three are main types of egg capsules in the discussions on macro-evolution.

Cassoidea (Fig. 4). The more-or-less finger-shaped A final remark on egg masses might seem trivial, but

is for with it capsule typical most Ranellidae, the exception was a surprise to me to discover that Fusitriton of Sassia and similar Gyrineum. A egg capsule type as in oregonensis (Redfield, 1846), occurring in the northern

Sassia be found also in the the counter- can Calyptraeoidea (see hemisphere, arranges egg capsule spiral

and the basal condi- and Fusitriton Bandel, 1976b) possibly represents clockwise, magellanicus, which is re-

tion in the Cassoidea. An collar is found in stricted the apical only to southern hemisphere, secretes a clockwise

the Cassinae. The flattenedegg capsules of Distorsio and spiral (the Caribbean Bursa granularis: counter-clock-

Tonna are superficially similar, but differ in details wise; the Tasman Sea Cabestana spengleri: clockwise).

1969a; Bandel, This These (d'Asaro, 1976a). capsule type gastropods appear to react to the Coriolis force, but

the of data (comparing same strategy development) usually more are needed.

contains significantly fewer embryos than the finger-

However, Tonna LARVAL shaped type. secretes many more egg DEVELOPMENT

than all other cassoid capsules genera and therefore reaches a similar veliger output as e.g. Cymatium or Larval development is here understood to be directly 106

indentations limitation useful Charonia, or (Argo- correlated to planktotrophy. This appears Bursa) triangular

some ?) are in the context of embryonic and larval shell characters buccinum, Fusitriton, Sassia, Cymatium

all whorls of the larval (see next section). Intracapsular larval stages are attrib- secreted and potentially fixed on and uted to the embryonic development. Lecitotrophic free- shell. This is accomplished by a pallial appendage

has been observed in Charonia 1895; swimming veligers are somewhat intermediate, but a (see Simroth,

and et distinct larval shell is not built, which (also in the context Bandel et al., 1994) Cymatium (Bandel al., 1994). of interpretation of fossil shells) is the main point here The mode of secretion and formationof periostracum was

described in detail Bandel al. Bursine larvae (see also next section). by et (1994). differentiated from other ranellids the Specific attributions of larvae call for careful consid- can be by

of the which remains thin in the late larval eration because in most cases only aspects periostracum, stage,

known Distorsio clathrata vice the is thicker in the ontogenies are (compare e.g. or versa, periostracum usually of d'Asaro, 1969a and Laursen, 1981). However, larval larvae of Cymatiinae and Ranellinae (see Bandel et al., features of the Cassoidea are often characteristic at the 1994).

and the velum is said to be generic level. Cassoid larvae are usually very large In all descriptions early

have been adult bilobate Riedel, and the late velum some of them even interpreted as gastro- (see e.g. 1992)

Larval food could pods (e.g. MacDonald, 1855; Pilsbry, 1945). It is very quadrilobate. The foot is usually large.

identified in the tracts of (see difficult to rear veligers in an aquarium. Consequently be alimentary Cymatium

which contained nearly all the literature data reflect only stages of larval Richter, 1987; Bandel et al., 1994),

be remains of life, from which, however, some strategies can Radiolaria, Foraminifera, tintinnids, extrapolated. Reports listed below all provide information dinoflagellates, unidentified phytoplankton, and of Bursa

in which Radiolaria, on at least characteristic periostracum sculptures (which (see d'Asaro, 1969a, Richter, 1987),

corroded after Foraminifera and found. are usually metamorphosis). Living dinoflagellates were

observed in the The larvae of the Cassidae other than Tonninae veligers of the Personidae were genus are

Distorsio. D'Asaro (1969a) provided information of the scarcely known. Two larval shells of Cypraecassis (Laur-

and of early veliger of D. clathrata. The veliger uses a pallial sen, 1981; Waren & Bouchet, 1990) one tentacle to secrete and form periostracal, carina-like, Semicassis (see Laursen, 1981) have been figured, which chambers. The velum is bilobate, but already shows show the characteristic large size, but no special pe- lateral indentations, foreshadowing the quadrilobate stage. riostracum sculpture.

the 'late' of The of the Tonna has been Bandel et al. (1994) described veliger a veliger genus figured or

species of Distorsio from the Red Sea. The larva has a described by several authors (MacDonald, 1855; Fischer,

there four velar lobes 1940; Lebour, 1945; comparatively large foot, are long 1887; Simroth, 1895; Thorson,

and form 1981; and a pallial tentacle is used to secrete a Ostergaard, 1950; Scheltema, 1971; Laursen,

Bandel The velum of the periostracal carina (without chambers). Two larval shells Bandel, 1991; et al., 1994). and in late of Distorsio species were figured by Laursen (1981), both early veliger is already slightly quadrilobate,

is different the four velar lobes bearing a periostracal carina, which, however, veligers are comparatively very long.

formed in the hatched in the two specimens. An undetermined larva figured by A proboscis is already newly

The larvae Lebour (1945) also shows this characteristic periostracal veliger (Thorson, 1940; Ostergaard, 1950). which sculpture. bear an operculum (Simroth, 1895; Bandel, 1991),

The larval shells bear Veligers of the Ranellidae have been observed or is lost during metamorphosis.

1895; as well as spirally arranged, tiny, examined in the genera Cymatium (see Simroth, periostracal spines

Lebour, 1945; Pilsbry, 1945, 1949; Scheltema, 1966; triangular indentations, which resemble homogeneous

Laursen, 1981; Richter, 1984; Waren & Bouchet, 1990; lirae under the light microscope (Bandel et al., 1994). A

observed in Tonna Bandel, 1991; Bandel et al., 1994), Cabestana (see larval pallial appendage was by

Riedel, 1992), Argobuccinum (sec Pilkington, 1974; pers. Simroth (1895) and Bandel et al. (1994).

Cassidae the stomach of obs.), Charonia (see Simroth, 1895; Lebour, 1945; Among the contents Cy-

Scheltema, 1966; Laursen, 1981; Waren & Bouchet, praecassis larvae have been examined (Richter, 1987),

1990), Fusitriton (see Pilkington, 1976; Waren & which comprised fragments of dinoflagellates.

Scheltema coined and defined the term 'te- Bouchet, 1990; pers. obs.), Sassia (sec Pilkington, 1976) (1971)

and Bursa (see d'Asaro 1969a; Scheltema, 1972; Laursen, leplanic' for long-term veligers. Simroth (1895) was the

that larvae of the Cassoidea drift 1981; Richter, 1984, 1987; Waren & Bouchet, 1990; first to recognise may

Scheltema Bandel et al., 1994). for several months. This was taken up by (e.g.

certain who demonstrated that certain All larvae have in common that, during a 1966, 1971, 1972, 1986),

larval in the for to a stage, periostracal spines (some Cymatium, veligers may stay plankton up year. 107

remarks — there is evidence that Summarising strong pronounced on comparatively small embryonic shells (e.g.

the larval life of the Cassoidea reveals a synapomorphic in Cabestana spengleri), while it is scarcely developed

character, allowing the three families to be united within in species with large initial whorls [e.g. Ranella olearia

the same superfamily. In some genera (Distorsio, ( Linne, 1758)].

of all three Cymatium, Charonia, Tonna) families, a The diameters of embryonic shells (of planktotrophic

larval tentacle has pallial actually been observed. More- species) are given here (alphabetically) at the generic

certain over, periostracal sculptural elements could not be level:

produced by any means other than the pallial appendage.

In this the existence of the be way, appendage may - Argobuccinum: 400 /ym (pers. obs.)

demonstrated without direct observation details - 330 & (for see Bufonaria: /rm (Waren Bouchet, 1990)

Bandel et al., 1994). - Bursa : 150-320 fj.m (d'Asaro, 1969a; Bandel, 1975;

The of small larvae periostracal spines (young) are Laursen, 1981; pers. obs.)

relatively larger than those of large ones of the same - Cabestana: 240 (Ramon, 1991; Riedel, 1992)

species (compare Simroth, 1895; Laursen, 1981; Bandel - Charonia: 500-850 /OTI (Berg, 1971; pers. obs.)

et al., The the diameter of a larval 1994). spines enlarge - Cymatium: 210-430 /vm (Arakawa, 1960; Bandel,

shell, which is a from small possibly protection predators 1975; Beu, 1986; Ramon, 1991; pers. obs.)

A thick and the must 220 (e.g. copepods). periostracum spines - Cypraecassis: pm (Laursen, 1981) be interpreted also as from - protection boring organisms Distorsio: 210-350 ftm (d'Asaro, 1969a; Beu, 1986;

or Moreover, the epibionts. periostracal sculptures have pers. obs.)

effects on the swimming activities (or buoyancy) of the - Eudolium: 290-300 /rm (Waren & Bouchet, 1990;

veligers (compare Richter, 1973; Bandel et al., 1994). Marshall, 1992)

Short, indentations such as in Fusi- simple periostracal - Fusitriton: 400 pm (pers. obs.)

triton and Sassia considered magellanicus kampyla, are - Gyrineum (Biplex): 500 pm (pers. obs)

to be the more plesiomorphic condition. The possible Malea: - 450 //m (pers. obs.) absence of periostracal in certain appendages species - Oocorys: 420 pm (Bouchet & Waren, 1993)

must be considered lost (apomorphy). - ’Personopsis’: 360 pm (pers. obs.)

Dawydoff (1940) illustrated a veliger that was attrib- Ranella: 540-710 & Bouchet, - /im (Waren 1990; pers.

uted by Waren & Bouchet (1990) to the genus Distorsio. obs.)

However, from a similar periostracal this apart sculpture, - Sassia: 330-350 /rm (pers. obs.)

larva has little in common with a personid gastropod, - Semicassis: 330 /rm (pers. obs.) since the are on of the tentacles, as in Tonna: 400-560 eyes top cephalic - wm (Thorson, 1940; Ostergaard, The of this Strombimorpha. systematic position peculiar 1950; Amio, 1963; Gohar & Eisawy, 1967; Bandcl, veliger remains unclear. 1975; pers. obs.)

- Tutufa : 330 «m (pers. obs.).

PROTOCONCH AND TELEOCONCH

The embryonic shells of non-planktotrophic cassoids The embryonic shell (or protoconch 1 of authors) is usually consist of 1-2 whorls (pers. obs.). However, the

defined here that of the conch that as part precedes a diameters are relatively much larger than in

larval shell. This or a juvenile/adult means that the planktotrophic species. A few examples: in Cassis nana

of all protoconch consists of - non-planktotrophic species Tenison-Woods, 1874 2.7 mm, Galeodea triganceae the shell whereas embryonic only, in - - planktotrophic Dell, 1953 2 mm, Gyrineum roseum (Reeve, 1844) 1.2

the be divided into species protoconch may an embryonic mm (Fig. 6). The typical embryonic shell ornament of and a larval shell (or protoconch 2 of authors). The planktotrophic species is lost. However, sometimes

teleoconch includes all shell material that does not belong sculptural elements such as lirae, may be present, but the to protoconch. these can be interpreted as having been derived

The shells of cassoids embryonic planktotrophic are (heterochronically) from the former larval shell sculptural

with the initial whorl of the conch. of usually synonymous pattern a hypothetical ancestor.

The shells of the Personidae far The larval shells embryonic are always (as of cassoids may reach huge sizes,

as smooth, whereas those of the Ranellidae and known) which cannot be found in many other gastropod taxa. The

Cassidae bear a more-or-less regular pattern of tiny number of whorls, maximum diameters (D) and heights

hollows with sharp projecting rims (compare Ramon, (H) of larval shells of 86 Recent species have been taken

1991; Riedel, 1992; Fig. 5). This sculpture is well (see Material and methods) from the literature (Cotton, 108 109

1945; Clench & Turner, 1957; McGinty, 1962; Barnard, Larval shell ornament consists of axial and spiral lirae (or

1963; Abbott, 1968; Lewis, 1972; Beu, 1978a, b, 1986; ribs) forming a reticulate pattern. Reticulation may be

Pctuch Gibson-Smith reduced to all to total absence in Sassia Finlay, 1978; & Harasewych, 1980; degrees up (e.g.

& Gibson-Smith, 1981; Beu & Knudsen, 1987; Beu & parkinsonia (Perry, 1811) [Figs 7, 8] or species of

Kay, 1988; Waren & Bouchet, 1990; Marshall, 1992; Distorsio). Usually the number of sculptural elements is

Abbott, 1993; Bouchet & Waren, 1993) or personally reduced and/or the reticulation has vanished from parts of

is measured. The largest larval shell (as far as known) the larval whorls (Figs 9-13). that of Tonna allium (Dillwyn, 1817) with H = 5.7 mm The teleoconchs are not described in detail and and D = 5.45 mm (compare Bandel et al., 1994). Ap- readers are referred to the pertinent literature (e.g. Fi- proaching that size are the larval shells of Ranella, scher, 1887; Simroth, 1907; Vredenburg, 1919; Thiele,

Charonia and some species of Cymatium. The smallest 1929; Clench & Abbott, 1943; Clench, 1944; Wenz, larval shell is found in Distorsio euconstricta Beu, 1986 1944; Turner, 1948; Clench & Turner, 1957; Kilias, 1962,

with H = 1.0 and = 1.05 At this size mm D mm. present, 1973; Dell, 1963; Cernohorsky, 1967, 1971, 1972; defines cassoid 1970c, 1977, the lowest limit among species. Abbott, 1968, 1974; Beu, 1970a, 1970b,

Putting the H and D measurements together leads to 1978a, b, 1981, 1985, 1986, 1988a, b; Bayer, 1971; Keen, the following H/D ratios, and six groups of larval shell 1971; Wilson & Gillet, 1971; Sabelli & Spada, 1978, shape are defined: 1982; Powell, 1979; Fretter & Graham, 1981; Beu &

Cernohorsky, 1986; Beu & Knudsen, 1987; Beu & Kay,

1988; Graham, 1988; Lai, 1990; Warén & Bouchet, 1990;

I globular: H/D = 0.91-1.10: Personidae, Cassidae Emerson, 1991; Marshall, 1992; Bouchet & Warén,

for except some Phaliinae, some Bufonaria, some 1993).

Characters considered be for Sassia, Gyrineum sp., Fusitriton magellanicus. to important (sub)family

phylogeny are the more-or-less distorted teleoconch of all II ovoid: H/D = 1.11-1.30: Bursinae except for some Personidae and among the Ranellidae only the Bursinae Bufonaria, Ranella, some Sassia, Argobuccinum pus- (probably all) have an anal canal present in the outer lip tulosum, some Phaliinae. the end of the at posterior aperture (Beu, 1981). conical: Ill low H/D = 1.31-1.50: Cabestana, Charonia, Summarising remarks — Embryonic shell sculpture

some Cymatium. (Fig. 5) similar to that of the Ranellidae and Cassidae

occurs in other such as Cerithiimorpha and IV conical: H/D = 1.51-1.70: some Cymatium. many groups, also in other neomesogastropod families (see Bandel, V high conical: H/D = 1.71-1.90: some Cymatium. to 1975). Therefore, the ornament has be seen as sym-

VI turriform: H/D = 1.91: some Cymatium. plesiomorphic for the Cassoidea, and lost in the

Personidae. The typical sculpture of ranellids and cassids

is not known from the Neogastropoda (which in other

The number of larval whorls varies from almost 2 to shell characters may be somewhat convergent [?] on nearly 6. More than 3.5 whorls are found only in the certain cassoids), except for a few species such as

Ranellidae, the highest numbers occurring in the genus Melongena melongena (Linne, 1758), which, however,

Cymatium. has non-planktotrophic development (Bandel, 1975).

Figs 5-13. Photomicrographs illustrating morphology of cassoidean gastropod protoconchs.

5 - Sculptural pattern of embryonic shell (of Argobuccinum), which is symplesiomorphic for the Cassoidea. Scale bar equals 10

µm.

6 - Apical whorls of Gyrineum roseum, scale bar represents 1 mm.

7 - Juvenile shell of Sassia parkinsonia, scale bar equals 3 mm.

8 - Apical whorls of the same specimen as in Fig. 7, showing diameter and ornament of the embryonic shell and part of larval

larval whorl. shell, which is smooth, apart from few remains of axial ribs on the first Scale bar equals 0.3 mm.

9 - Apical whorls of Sassia kampyla. Scale bar represents 2 mm.

10- The same specimen as in Fig. 9, lateral view of transition from embryonic to first larval whorl, which is partly spirally lirate.

Scale bar equals 0.2 mm.

11- Juvenile the Scale bar is 2 shell of Gyrineum (Biplex) sp., showing c 180° alignment of varices. mm.

12- in Scale bar Apical whorls of the same specimen as Fig. 11. equals 1 mm.

13- Apical whorls of Tutufa rubeta,showing cancellate ornament on first larval whorl. Scale bar represents 2 mm 110 111

the of The embryonic shell sculpture is possibly a useful the Cymatiinae.The larval shells with highest number

character to distinguish higher taxa (compare Bandel & whorls are found in this subfamily. The larval shell sculp-

Riedel, 1994). ture is always based on a pattern of rectangles. How-ever,

different The large size of the initial whorl allows the this can be used as a character at several taxo-

Cassoidea to be differentiated from most other gastro- nomic levels. No larval shells in the Cymatiinae and

pod taxa {i.e. their early ontogenetic stages) with Bursinae are totally covered by axial and spiral ribs (form-

similar sculpture of the embryonic shell. The diameter ing the rectangles). However, complete reticulate ornament

in the Cerithiimorpha for example seldom exceeds 100 occurs in the Ranellinae and Oocorythinae. The Tonninae fim (Bandel, 1975). Among the Cypraeoidea some show a reduced pattern of rectangles, which is characteristic

this has occurred in all species are known (Ostergaard, 1950; Bandel, 1975; of subfamily. Loss of sculpture

in which the diameterof the Tanaka, 1980; pers. obs.) families.

embryonic shell lies within the range of the Bursinae. The plesiomorphic teleoconch type in the Cassoidea is

of the assumed be that of the for However, postembryonic stages Cypraeoidea to Ranellinae, example Gyrineum

Fossil The canal certainly cannot be confused with those of any cassoid. (see record). pronounced posterior siphonal

the Bursinae The correlation of embryonic shell size to teleo- in represents an apomorphy.

conch size is difficult. In several cases species with The embryonic, larval and adult shell characters are not

large adult teleoconchs have a large initial whorl, while repeated in the discussion on the fossil record (see below).

with adult have fossil taxa is of based the species a small teleoconch a compara- The recognition of course on

tively small embryonic shell. However, this cannot be modern fauna and plesiomorphy or apomorphy of characters

generalised at all, as too many exceptions are known. is explained further in the context of the chronology of

The size of the embryonic shell must be interpreted in fossils.

individual. the context of the entire ontogeny of an

Comparing two species with a similar teleoconch size and with the same number of larval shell whorls, it can ANATOMICAL FEATURES be expected that the embryonic shells are also about

in diameter. if for instance the number the of the Personidae Distorsio has equal However, Among genera only of larval whorls is comparatively lower, the embryonic been examined, but most references deal only or mainly

shell is larger in most taxa. This has to be seen as a with the radula (Troschel, 1863; Clench & Turner, 1957;

derived character, because in early ontogenies, where Cernohorsky, 1967; Beu, 1978b; Bandel, 1984; Waren &

free larval is the initial whorl in The radula of Distorsio is no phase present, Bouchet, 1990). taenioglossan

small. For the central teeth of D. almost all taxa is largest (with a few exceptions, see comparatively example, e.g. Wilson, 1985). clathrata are about 60-70 /um wide (see Bandel, 1984;

Planktotrophy, and therefore the existence of a Waren & Bouchet, 1990), and the central teeth of

larval shell, is a plesiomorphic character for the three Distorsionella are only about 20 «m wide (see Beu, 1978b).

of Distorsionella show the cassoid families. A more-or-less globular shape and a The radulae Distorsio and same

comparatively small protoconch size (Personidae and characteristic features in having small but strong, serrated,

some Ranellinae) are also 'primitive' characters within central (with down-curved ends) and lateral teeth and

the Cassoidea (compare Bandel & Riedel, 1994). smooth, simple marginal teeth (Fig. 22). The radula of ’Per-

in Conical or even turriform protoconchs occur only sonopsis ’is not known, neither is the animal.

Figs 14-21. Photomicrographs of cassoidean gastropod radulae.

14- Section of radula of Fusitriton magellanicus. Scale bar equals 0.3 mm.

15- Fusitriton magellanicus, magnification of central and lateral radular teeth. Scale bar represents 0.2 mm

0.2 16- Section of radula of Cabestana spengleri. Scale bar equals mm.

17- Cabestana spengleri, magnification of central and lateral radular teeth. Scale bar represents 0.1 mm.

18- Central radular teeth of Bursa Scale bar 0.2 mm. sp., showing basal interlocking processes. equals

of lateral and radular teeth. Scale bar 0.5 19- Bursa sp., magnification marginal equals mm.

of central and lateral radular teeth. Scale bar 0.2 20- Cassis cornuta, magnification represents mm.

21- Radula of Semicassis labiata. Scale bar equals 1 mm. 112

Fig. 22. Radulae (half row) of Cassoidea and members of The soft parts of Distorsio have been examined by Lewis

outgroups (not to scale): (1972) and by myself. The animal of Distorsio sp. is

about 1 cm long (Fig. 25). The foot is medium sized, the

for the The operculum typical genus. head is compara-

small. The of the tively bases cephalic tentacles are

- danieli a Capulus (after Orr, 1962); thickened to lateral swellings carrying the eyes. Below

b - Gyrineum gyrinum (after Cernohorsky, 1967); the base of the right tentacle arises a relatively small

c - Ranella australasia; penis which is constricted twice to a smaller diameter.

d - Akibumia orientalis (after Warén & Bouchet, 1990); The seminal groove is open and runs along the right body e - Pisanianura grimaldii (after Warén & Bouchet, 1990); wall. The mantle is and edge smooth regular except for f - Cymatium muricinum (after Bandel, 1984);

the fold of the inhalant The brownish is mo- Distorsio siphon. gill g - clathrata (after Bandel, 1984); nopectinate, and rounded anteriorly. The bipectinate h - Charonia lampas; is The is i - Cymatium (Linatella) caudatum (after Morton, 1990); osphradium yellowish. hypobranchial gland

- The is j Oocorys sulcata (after Bayer, 1971); inconspicuous. rectum comparatively short,

k - Galeodea maccamleyi (after Ponder, 1984); terminating some distance from the mantle edge. l - Casmaria vibexmexicana (after Bradner & DuShane, 1982); The body cavity is dominatedby the anterior alimen-

m - Eudolium canal. The is coiled crosseanum (after Marshall, 1992); tary proboscis in several loops when

n - Tonna Bandel, galea (after 1984). retracted and can be extended to about three times the 113

length of the animal. The buccal mass, radula and jaws

are all relatively small. The oesophageal gland is brown and appears to be homogeneous, but sectioning might reveal a complex of glands. Salivary ducts could not be observed.

Distorsio perdistorta Fulton, 1938 (see Lewis, 1972)

is much larger and differs in having a finger-shaped, late- rally flattened penis, similar to that of Fusitrition magel-

that the is lanicus (Fig. 26). Apart from gross anatomy

very similar to that of Distorsio sp. Beu (1978b) pre-

sented some anatomical features of Distorsionella lewisi

characteristic anterior revealing the same very elongate

alimentary tract.

The gross anatomies of most genera of the Ranellidae

have been described in the literature. Argobuccinum pustulosum, Bursa sp., Cabestana spengleri, Charonia

lampas, Fusitriton magellanicus, Ranella australasia and

Tutufa rubeta have been examined personally, but only

here because brief descriptions of some features are given

most have already been described. General characters of

the family or superfamily are pointed out (see also

summarising remarks).

The radulae of the Ranellinae (see Troschel, 1863;

Clench & Turner, 1957; Barnard, 1963; Dell, 1963; Dell

& Dance, 1963; Cernohorsky, 1967, 1970; Arnaud &

Beurois, 1972; Rang, 1976; Beu, 1978a; Bandel, 1984;

Waren & Bouchet, 1990; Habe, 1992; Figs 14, 15, 22)

central and 23. Gross of Argobuccinum pustulosum. Scale are all comparatively large, having powerful Fig. anatomy

bar equals 2 cm. Abbreviations are as follows (also lateral teeth with a varying number of cusps. The mar- applicable to Figs 25-32): ginal teeth are always smooth. The lateral and marginal

teeth of Charonia are significantly longer compared to

- - B - C columellar CT buccal mass; muscle; cephalic those of the other genera of the Ranellinae. The radulae

E - F - G - HG - tentacle; eye; foot; gill; hypobranchial of the Cymatiinae (see Troschel, 1863; Clench & Turner, gland; IS - inhalant siphon; M - mantle; N - nerve 1957; Barnard, 1963; Cernohorsky, 1967; Kilburn, 1970; - OP - collar; O - oesophagus; OG oesophageal gland; Kang, 1976; Bandel, 1984; Beu & Kay, 1988; Morton, operculum; OS - osphradium; OV - oviduct; P - probos-

1990; Waren & Bouchet, are all similar, 1990) very cis; PE - penis; PR - prostate; PRD - prostate duct; R -

central and lateral teeth set with - RAS - radula S - seminal having strong very sharp rectum; RA radula; sac;

teeth be indented, and the SD - SG - V - cusps. The inner marginal may duct; salivary duct; salivary glands;

smooth visceral outer marginal teeth are always (Figs 16, 17, 22). mass.

The Bursinae (see Troschel, 1863; Cooke, 1917; Thiele,

1929; Thorson, 1940; Cernohorsky, 1967; Habe, 1973;

The animal of has Melone, 1975; Beu, 1981; Bandel, 1984; Waren & Argobuccinum pustulosum (Fig. 23) a

radulae that which is filled Bouchet, 1990; Figs 18, 19) have are easily comparatively long body cavity, widely by

distinguished from those of Ranellinae and Cymatiinae by the glandular complex of the anterior alimentary tract

the of two from the base of the (compare Day, 1969). The salivary ducts through the presence cusps projecting pass which is the in all Ranellidae and central tooth. The central and lateral teeth are powerful nerve collar, case

or The Cassidae that have been examined anatomically. The and the cutting edges may be set with cusps not.

seminal is where the is intercalated inner marginal teeth may bear denticles, the outer mar- groove open prostate

distance from but closed where it ginal teeth are always smooth. (and at some it) runs the flattened The bodies of two Ranellinae are figured here but along long, laterally penis, where, however,

in of adhesion be J.T. Smith only some characteristic features are described briefly a seam can recognised. (1970),

the context of literature data (Bouvier, 1887; Amaudrut, however, described a genital duct open over its whole

1898; Barnard, 1963; Dell & Dance, 1963; Cemohorsky, length. A short prostate duct opens into the posterior

1967; Laxton, 1968, 1969; Day, 1969; J.T. Smith, 1970). mantle cavity. 114

24. Fig. Nervous system of Tonna galea (after Haller,

1893), not to scale. Abbreviations: BG - buccal

ganglia; CPG - cerebral and pleural ganglia; PG -

pedal ganglia; SB - suboesophageal ganglion; SP -

supraoesophageal ganglion.

Fig. 25. Gross anatomy of Distorsio sp. Scale bar equals 5

mm.

The hypobranchial gland a row of thick, 26. Penis Fusitriton comprises Fig. of magellanicus. Scale bar repre-

whitish leaflets. The is and brown as gill monopectinate sents 1 cm.

in in all other Cassoidea. The which gill tapers to a point,

appears to be the case in all Ranellidaeand Cassidae. The

is is usual in the Casssoidea. osphradium bipectinate as The animals of Gyrineum and Sassia are poorly known.

The of Fusitriton differs from that of body magellanicus Cernohorsky (1967) provided a short description of

in few the Argobuccinum pustulosum characters, mainly Gyrineum gyrinum, in which the 'very thick' proboscis seminal open groove, the lack of an additional prostate and the 'club-shaped' penis might be of interest. Beu

duct and the shape and muscular structure of the penis (1978a) figured and briefly described the animal of Sassia

The (Fig. 26; compare Beu, 1978a). animals of the two kampyla, which externally closely resembles that of

species of Ranella (see Bouvier, 1887; Amaudrut, 1898; Argobuccinum species.

Barnard, 1963; Dell & 1963; Laxton, 1968, 1969; Dance, As an example of Cymatiinae, the animal of

differ in such the of pers. obs.) only details, as shape the Cabestana spengleri is figured (Fig. 28) and briefly

The in Ranella similar penis. gross anatomy is very to described here. Conspicuous features are the long gill

that of and Fusitriton The seminal Argobuccinum species. which terminates close to the mantle edge, the short duct is in Ranella open species. The animal of Charonia rectum which terminates far back in the mantle cavity,

lampas differs in some Overall differ- (Fig. 27) aspects. the simple but comparatively long penis (with an open

the broader the ences are head, hypobranchial gland being seminal duct), the stripe-like lamellae of the hy- formed by numerous lamellae covered by a regular pobranchial gland, the salivary ducts fused with the wall

epithelium, and the salivary glands being clearly demar- of the anterior oesophagus and the salivary glands

cated both functionally and physically, and distinguished restricted to the left-hand side of the body cavity. the by orange anterior glands and white posterior glands. Cymatium nicobaricum, described by Houbrick & Fretter

The of Charonia tritonis differs hypobranchial gland (see (1969), mainly with respect to the glandular

Bouvier, 1887) to be similar to that of is appears complex, but otherwise very similar to Cabestana

Argobuccinum. in most anatomical features. spengleri gross 115

Fig. 27. Gross anatomy of Charonia lampas. Scale bar equals 2 cm.

28. Gross of Cabestana Scale bar 1 Fig. anatomy spengleri. equals cm.

Fig. 29. Anterior alimentary tract of Tutufa rubeta. Scale bar equals 1 cm. 116

1963; Abbott, 1968; Hughes & Hughes, 1981; Bradner &

DuShane, 1982; Bandel, 1984; Waren & Bouchet, 1990;

Figs 20-22) than in Oocorythinae or Tonninae. The

central and lateral teeth closely resemble those of the

Oocorythinae. The tip of each marginal tooth is serrated

in the most taxa. Data on same species (e.g. Abbott,

1968; Waren & Bouchet, 1990 on Semicassis granulata)

be but indicate considerable appear to contradictory may

intraspecific variation as is the case in Galeodea (Ooco-

rythinae).

The Tonninae defined their are well by radular teeth.

The radulae of Tonna, Malea and Eudolium species

(Troschel, 1863; Fischer, 1887; Turner, 1948; Barnard,

1963; Kang, 1976; Bandel, 1984; Waren & Bouchet,

1990; Marshall, 1992) are closely similar (Fig. 22). The

marginal teeth of all three genera are reported to be

smooth. The broad central tooth bears two denticles

Fig. 30. Anterior body of Bursa sp. Scale bar represents 1 projecting from the left and right flanks of the base in the

cm. direction of the main cusp. The cutting edges of the

central and lateral teeth of most Tonna species are

smooth while those of Malea and Eudolium are serrated

Morton (1990) provided some anatomical data on However, the radula of Tonna maculosa Dillwyn, 1817

caudatum which the attribu- Cymatium (Linatella) justify (see Turner, 1948) is intermediate, having few serrations. tion of Linatella as a subgenus to Cymatium (Beu, pers. Turner (1948) and Waren & Bouchet (1990) figured comm.). radulae of the veliger of Tonna sp., showing serrated

The of the Bursinae Houbrick central and lateral teeth. This genera (Risbec, 1955; suggests that the smooth

& Frettcr, 1969; Beu, 1981; Figs 29, 30) all have a condition of the cutting edges is apomorphic within the comparatively broad and short proboscis, a large buccal Tonninae. mass without in Beu, There of cassid in the (minute Tutufa) jaws (see 1981), are some descriptions anatomy and a well-developed mantle fold forming the exhalant literature, but only Tonna galea has been described in siphon. The arrangement of the salivary gland complex detail (Bouvier, 1887; Haller, 1893; Simroth, 1896-1907; varies and Some of the focus the greatly (inter- intragenerically; compare Figs Weber, 1927). publications on

29, 30). The seminal ducts of Bursa species are usually salivary glands (Troschel, 1854; Panceri, 1869; Schonlein, open, and in Tutufa or Bufonaria open genital grooves 1898), which produce sulphuric acid. The complex of

well closed ducts occur as as (Beu, 1981; pers. obs.). salivary glands is extremely large, in full contact with the

The radulae of the Cassidae allow the taxa to be body wall, which by muscular contraction is able to

in three units. The radula grouped larger systematic type squeeze the glands and force the salivary secretions

of the Oocorythinae is considered to be basic within the through the salivary ducts to perform an ejaculation (see

The four attributed this The family (Fig. 22). genera to Troschel, 1854). salivary ducts as in all cassoids run

all have central and lateral teeth with subfamily strong a through the nerve collar. The nervous system (Fig. 24) varying number of cusps on the cutting edge (Troschel, has been described by Bouvier (1887) and Haller (1893).

1863; Fischer, 1883; 1905; Thiele, 1929; Turner, Reynell, The gross anatomy reveals the typical cassoidean arrange-

1948; Barnard, 1963; 1971; Quinn, 1980; Bandel, Bayer, ment of the organs of the pallial cavity, with a large mo-

Waren & The 1984; Ponder, 1984; Bouchet, 1990). nopectinate gill and a bipectinate osphradium. The

marginal teeth are more-or-less hooked and scarcely hypobranchial gland is inconspicuous. The rectum

than the laterals. The of each inner longer cutting edge terminates close to the mantle edge where a siphonal fold

marginal tooth is serrated in most taxa. Variation in the is The seminal duct is the is developed. open, penis

number of cusps on the central and lateral teeth occurs approximately finger shaped but laterally flattened. The

intraspecifically (Ponder, 1984) as well as interspec- proboscis is pleurembolic as is the case in all cassoids

which that ifically, means using numbers of cusps to and characteristic for the suborder Troschelina Bandel &

needs statistical confirmation. distinguish species Riedel, 1994. The jaws have developed a terminal hook

The of the Cassinae and Phaliinae united species are and are extremely powerful (Weber, 1927; Waren & by having significantly longer marginal teeth (Troschel, Bouchet, 1990). A characteristic feature is the large foot

1863; Fischer, 1887; Thiele, 1929; Cotton, 1945; Barnard, lacking an operculum. The foot spreads out very wide 117

soft and its anterior when crawling on a substrate, edge

hook each bears a lateral fleshy at extremity (see e.g.

Wilson & Gillct, 1971). Marshall (1992) presented some anatomical features of the genus Eudolium, which are not significantly different from Tonna. The body of Malea has not been described.

with Among the Oocorythinae the genus Galeodea the species G. echinophora (see Simroth, 1896-1907; Niiske,

1973; Fange & Lidman, 1976; Hughes, 1986a) and G. rugosa (see Reynell, 1905, 1906) has been examined to

extent. of is some The complex salivary glands very

have been shown large. The huge accessory glands to produce sulphuric acid (Fange & Lidman, 1976). The nervous system has been described by Reynell (1905,

similar 1906) and is concentrated in a style as that of

Tonna. The gross anatomy does not show any marked differences from the 'bauplan' of Tonna. The body in

Oocorys species (Fischer, 1883; Bayer, 1971; Quinn,

1980; Waren & Bouchet, 1990) is very similar to that of

Tonna and Galeodea species, except for the head, which has thick tentacles. The developed very long, eyes are

There the of Dalium missing. are no reports on anatomy and Sconsia.

The shells of Cassinae have been examined exten-

there few the Gross of Cassis bar sively, but are very reports on anatomy Fig. 31. anatomy cornuta. Scale repre-

(e.g. Amau-drut, 1898; Hughes & Hughes, 1981). sents 2 cm.

Fig. 32. Gross anatomy of Semicassis labiata. The oesophageal gland is not shown. Scale bars equal 2 cm. 118

A sketchy figure (Fig. 31) of Cassis cornuta is provided Summarising remarks — Waren & Bouchct (1990, pp.

here to give a rough idea of the spatial arrangement of 69, 73) placed emphasis on 'typical salivary glands'

the body, which is significantly flattened due to the ('normal tonnoid type') and indirectly (?) made this construction of the shell. The important feature is the anatomical feature the superfamily key character, with the

large complex of salivary glands filling most of the body intention of including the Laubierinioidea(see Bandel & cavity. The anterior salivary glands are comparatively Riedel, 1994) within the Cassoidea. Bandel & Riedcl

coloured The the Laubierinidae small, pea-shaped and beige. two accessory (1994) showed that Waren & Bouchet,

acid glands are huge and slightly greenish. The buccal 1990 is best treated as a separate superfamily - to which

is is also known Pisanianuridae added herein - is mass remarkably large which, however, the are which, however,

from some species of other cassoid subfamilies. Among close to the Cassoidea. The complex of salivary glands is

the other organs the penis is conspicuous. It is very small difficult to interpret in the systematic context of the

and constricted to form The seminal is the of the of a tiny tip. groove Neomesogastropoda as knowledge anatomy

open. the group is very incomplete. Within the Cassoidea, the

Beu (1981) provided anatomical data on the Phaliinae Personidae have no accessory salivary glands, which based Semicassis the refutes the idea character; the on pyra (Lamarck, 1822) (for penis as a superfamily key

see also Abbott, 1968), and Hughes (1986b) presented an Ranellidae have large accessory salivary glands, but with

outline of the anatomy of S. granulata (Born, 1778). quite different arrangements (and contents) among the

Gross anatomical features of S. labiata are figured here, taxa; and the Cassidae have very large accessory salivary

and a few key characters are commented upon (Fig. 32). glands, which contain sulphuric acid in possibly all taxa.

The animal is orange except for the visceral mass, which The acid produced by ranellid and cassid species is

columellar which is associated with echinoderms. is brown, and the muscle, yellowish. apparently feeding on

of the the is Among the organs pallial cavity osphradium The nervous system of Tonna (Fig. 24), with minor

remarkably large and hypobranchial gland is well devel- differences, is representative of the Cassoidea. Within the

oped, which is in contrast to the smaller ones in species Neomesogastropoda a similar concentration of the ganglia

of the other cassid subfamilies. Again a very large can be found e.g. in the Calyptraeoidea, and a more complex of salivary glands is present, with a pair of concentrated nervous system occurs in the Lamellarioidea

anterior, beige, small glands and a pair of white accessory (Bouvier, 1887).

(acid-secreting) glands that are relatively as huge as in Shape and size of the penis is of limited value in

Tonna galea. The salivary ducts are brown, pass through arranging systematic units. In the Ranellidae, however, a

the nerve collar (cerebral and pleural ganglia fused) and long tapering penis predominates, while a finger-shaped,

in Cassidae. run laterally along the oesophagus. However, the left duct laterally flattened one occurs mainly the joins the buccal cavity dorsally while the right duct There are several exceptions in each family and the

terminates from two of anatomical differences be between of ventrally. Apart figures a penis may greater species

there is the of the than between of different families. (Abbott, 1968), no report on anatomy same genus genera

Casmaria in the literature.

Figs 33-41. Photomicrographs (except Fig. 41) of apical whorls of cassoidean gastropods.

33- Apical whorls of Sassia septemdentata (Eocene, Texas). Scale bar equals 1 mm.

34- Initial whorls of Gyrineum gwinae (Campanian, Mississippi). Scale bar equals 0.2 mm

35- Apical whorls of Distorsio tortuosus (Miocene, The Netherlands). Scale bar represents 1 mm.

36- Apical whorls of Sassia delafossei (Eocene, Paris Basin). Scale bar represents 1 mm.

37- 0.1 The same specimen as in Fig. 36, magnification of initial shell. Scale bar equals mm.

38- shell of Sassia Scale bar 0.1 Embryonic sp. (Paleocene, Alabama). represents mm.

39- as in 38, reduction of cancellate ornament of larval shell and first varix of teleo- The same specimen Fig. showing juvenile

conch. Scale bar equals 1 mm.

40- Early juvenile shell of Sassia flandrica (Oligocene, Germany), showing the transition from protoconch to teleoconch. Scale

bar represents 1 mm.

41- Galeodea cf. echinophora (Miocene, northern Germany). Scale bar equals 3 cm. 119 120

The seminal duct of most cassoid taxa is open and where not a full review of all the literature dealing with fossil

is closed of cassoids, but brief of it (e.g. Argobuccinum pustulosum) a seam a summary important data to adhesion is visible. In comparison with the other super- produce evidence for the chronological order of at least families of the Neomesogastropoda the open genital the subfamily origin.

be Personidae have distorted groove must the plesiomorphic state. The all a more-or-less

is Other basal neomesogastropod characters include the teleoconch, which a character to recognise even in monopectinate gill, the bipectinate osphradium with large poorly preserved fossils. A species called Eutritonium

leaflets, and for the Troschelina, the Cottreau, 1922 from the 'Senonian' pleurembolic probos- praegranosum (Late cis. A coiled proboscis (when retracted) is found only in Cretaceous) of Madagascar (see Cottreau, 1922) is the

the Personidae and represents one good character to oldest known personid. Beu (1988a) attributed this

that The radulae of to justify family (see Beu, 1988a). species the genus Distorsio on the basis of its strong

reduced personid species appear to be in size. However, teleoconch distortion and coarsely cancellate sculpture. the down-curved shape of the central teeth is a better From the Paleocene of Poland, Personopsis (= Eutri- character to this from the Ranellidaeand described separate family tonium) rutoti was (Krach, 1963; compare Beu,

Cassidae. In comparison with the outgroups (Lau- 1988a) and Beu (1988a) erected the new genus and bierinioidea, Calyptraeoidea, Capuloidea), the basic species Kotakaia simplex from the Paleocene of the Chat- cassoid radula in in Sassia type occurs the Ranellinae, i.e. ham Islands (New Zealand). Distorsio septemdentata and Gyrineum (see discussion on phylogenetic (Gabb, 1860) (see Pilsbry, 1922; Palmer & Brann, 1966; cladogram). The subfamilies of the Ranellidae and Maxwell, 1968; Beu, 1978b) was described from the

Cassidae are each homogeneous in respect to their Eocene of Texas (Fig. 33). Beu (1988a) recognised that radulae. The Cassinae and Phaliinae share the same this species actually belonged to the genus Sassia (Ra- radula type, which is apomorphic to that of the nellidae) and that the slight distorsion of the whorls was

The characteristic Oocorythinae (simple marginals). due to convergence. A new species of genuine Eocene

the bases of the central radular interlocking processes at personid of New Zealand, Distorsio beui, was recorded teeth have developed convergently in the Bursinae and in by Maxwell (1968) and attributed by Beu (1988a) to the the Tonninae. the genus Personopsis. From Oligocene Distorsio

interposita (Tate, 1894) from Victoria (Australia), D.

crassidens (Conrad, 1848) from Mississippi (USA) (see

FOSSIL RECORD MacNeil & Dockery, 1984) and D. tortuosum

subclathrata (d'Orbigny, 1852) from the Adour Basin

have been described. The preceding chapters provide the basis for tracing the (France) (see Lozouet, 1987) of Miocene material Cassoidea back through earth history. Convergences exist Descriptions are more numerous

of and D. between adult shells, larval shells or embryonic shells include, among many others, gatunensis Toula, 1909 illustrated the cassoids and other families of the Neomesogastropoda (Panama; Woodring (1959) pro-

D. (e.g. Trichotropidae) or Neogastropoda (e.g. Cancellarioi- toconch), tortuosus (Borson, 1822) (The Netherlands)

these when for which species Janssen (1984) documented the dea). However, convergences operate only

D. constrictus simillimus comparing the same ontogenetic stage and not when protoconch (Fig. 35), (Sowerby, from Santo using all different ontogenetic stages as a complex of 1849), a large species Domingo (see Pilsbry,

and D. reticularis from Burma and characters (see protoconch and teleoconch chapter). 1922), (Daiton, 1908) India (Dey, 1961). In the literature, very few authors provide data on the The oldest documented member of the Ranellidae is early ontogenetic shell of cassoids, which only partly results from the diagenesis of fossils. The characters of probably a specimen from the Aptian (Early Cretaceous)

of northern Schroder The the protoconchs of only sixteen species could be ex- Germany figured by (1992). shows tracted from the literature and another fourteen specimen is a damaged juvenile, which, however,

could be a cancellate larval shell and some protoconchs examined personally. This seems to densely postlarval be whorls with varices. Other Cretaceous shells which quite a poor basis to trace back a systematic unit ranellids the Turonian of Utah which had its origin sometime during the late Early appear to be come from

Cretaceous. 1893, Tritonium kanabense; see Beu, However, there are a fair number of descrip- (USA) (Stanton, from the Campanian of Mississippi (USA) tions of teleoconchs in the literature and many of them 1988a),

1993: Sassia with are diagnostic without the reconfirmation of the (Dockery, carlea, Gyrineum gwinae,

Thus few difficult protoconch, Fig. 34), from Lybia (Wanner, 1902, Triton protoconch characters. very taxa are to

attribute to the next higher taxon and the protoconchs in tuberculosum); Madagascar (Basse, 1932, Triton sauryi)

and Tennessee 1926, Tritonium some cases can resolve these problems. The following is (Wade, univaricosum). 121

Fig. 42. Chronology of the evolution of Cassoidea at the subfamily level (see text). 122

Fig. 43. Phylogenetic cladogram of the Cassoidea (see text). 123

Malea Cossmann & 1923, Malea In the Paleocene of Belgium occur species which Glibert 1922, sp.; Peyrot,

orbiculatum 1986, Eudolium The (1973) assigned to the genus Charonia, subgenus Sassia Yokes, subfasciatum).

radiated (see remarks). Kollmann & Peel (1983) recorded a other three subfamilies of the Cassidae have

ranellid from the Paleocene of Greenland, while worldwide (see Ravn, 1907; Dall, 1909; Cossmann &

Zinsmeister (1983, Gyrineum judithi) and Beu (1988a, Peyrot, 1923; Gardner, 1948; Parker, 1948; Woodring,

Ranella louellae) from California and Traub (1979) 1959; Dey, 1961; E.J. Moore, 1963; Jung, 1971; Kanno,

and their described some ranellids from the Paleocene of Austria. 1973; Janssen, 1984; Beu & Maxwell, 1990)

relation with the modern faunas is obvious All these gastropods are best referred to the subfamily (Fig. 41).

remarks — the evolution of Personidae Ranellinae. A protoconch of a species of Sassia from the Summarising

Paleocene of Matthew's Landing (Alabama) is illustrated and Ranellidae has been analysed and discussed by Beu

in and there is need here (Figs 38, 39). numerous papers (see References) no

revision. additionalremarks have In the Eocene, the ranellid fossil record increases in for a However, some to

and the be made in the of classification of quantity as well as in quality (Figs 36, 37) context a partly new

branch the Ranellidae and the of the Cassidae needs Bursinae and the Cymatiinae can be recognised to story some off and diverge (see Cossmann, 1903; Weaver, 1912; reevaluation. The author fully supports Beu's (e.g. 1988a)

certain Olsson, 1930; Squires, 1983; Beu, 1988a). Several assignment of fossils to genera, though Sassia (see

protoconchs have been documented by Beu (1988a) and classification) is now placed in the Ranellinae (but see

teleoconchs have become more characteristic (see chapter final discussion), which means that this subfamily has the

record and the of the on protoconch and teleoconch). The radiation of the three longest fossil represents stem-group

The ranellid subfamilies is very well reflected in the fossil Ranellidae and also of the superfamily (Fig. 42).

Ranellinae the record of the Oligocene (see Beyrich, 1854; von Koenen, possibly originated during Aptian (Schro-

1866; Speyer, 1866; Ravn, 1907; Hickman, 1969; J.T. der, 1992) and are definitely recognised from the

Smith, 1970; Kuster-Wendenburg, 1973; Giirs, 1983; Turonian onwards (Beu, 1988a).

MacNeil & Dockery, 1984; Lozouet, 1987; Fig. 40) and Personidae (see final discussion) and Cassidae

the latest. In the Miocene(see Cossmann, 1903; Dall, 1909; Pilsbry, 1922; branched off during the Campanian at

Cossmann & Peyrot, 1923; Sorgenfrei, 1958; Woodring, context of the phylogenetic analysis (see following

1959; Dey, 1961; Beu, 1970c; Jung, 1971; Beets, 1984; section), the Personidae must have branched off earlier

Janssen, 1984). than the Cassidae. A significant radiation of the

Paleocene- The origin of the Cassidae has been discussed, Ranellidae and Cassidae occurred at the

Schenck Eocene The subfamilies Bursinae and amongst other authors, by Cossmann (1903), boundary. Cyma-

and tiinae in several Eocene faunas. (1926) and Abbott (1968). Cossmann Abbott pro- are easily recognised

while Schenck assumed the Kase attributed Hainabursa posed an Eocene origin (1984) aquilana (Parana,

to the origin to be in the Late Cretaceous. Schenck's view 1909) from the late Early Cretaceous of Japan

record of few Bursinae. This is cassoid but (1926) is accepted here based on the a species not a closely

certain cerithioidean Cerithium fossils of Late Cretaceous age (Wanner, 1902, Cassidaria resembles gastropods (e.g. 1

1958, columna 1834). sp.; F.M. Anderson, Haydenia impressa; Poyarkova Sowerby,

Dzhalilov, 1985, Cassidaria Abdel-Gawad, 1986, The oldest cassid fossils have to be attributed to the & sp.;

the which the of the Cassidaria truncata) appear to be close to genus Oocorythinae, represent stem-group

radiation of the Cassidae occurred Galeodea and represent members of the Oocorythinae. A family (Fig. 42). A at

the where the subfamilies Cassinae Paleocene Galeodea species from New Zealand was the start of Eocene

Tonninae recorded by Bcu & Maxwell (1990). The Cassidae and Phaliinae can be differentiated. The cannot

radiated during the Eocene, which is well documented in be recognised with certainty prior to the Early Miocene,

of others and the branch of the Cassoidea. a number papers (see among Deshayes, 1866; they represent youngest

is record Cossmann, 1903; Oppenheim, 1906; Clark & Woodford, As for the Bursinae, there a single (see Wenz,

The Proto- 1927; Wrigley, 1934), and in addition to the 1938-44) of a Cretaceous specimen. genus

Oocorythinae, the subfamilies Cassinae and Phaliinae can dolium Wilckens, 1922is comparatively small, has a very

columellar and the be recognised. Cassids from Oligocene deposits have thick shell, no siphonal canal, no plait

Stilwell been described by (among others) Beyrich (1854), Speyer inner lip of the aperture is pronounced. (1994)

(1862, 1866), Cossmann (1903), Ravn (1907), Kuster- has recently re-assigned this genus to the Neritopsidae

Wendenburg (1973), MacNeil & Dockery (1984) and Beu (Neritimorpha).

of the Cassoidea in the of & Maxwell (1990). The fossil record context discussed Miocene strata yield the first shells attributable to the the Neomesogastropoda has recently been by

Malea Bandel & Riedel Cassoid such the Tonninae (Dalton, 1908, Dolium sp.; Toula, 1909, (1994). outgroups as

and can be traced back to the camura [see also Woodring, 1959; Jung, 1971]; Pilsbry, Calyptraeoidea Capuloidea 124

Mesozoic and probably evolved during the Early Creta- morphic. Sassia is not only closely related to Gyrineum, ceous. but also to Charonia and Cabestana (Beu, pers. comm.).

Sassia and Charonia are somewhat intermediate between

Ranellinae sensu Beu (e.g. 1988a) from which they

originated and Cymatiinae/Bursinae to which they gave PHYLOGENETIC CLADOGRAM AND DISCUSSION rise. Sassia and Charonia should be considered to

constitute an independent subfamily.

Neomesogastropod phylogeny has been discussed by The Cymatiinae and Bursinae both have semi-globular

Bandel & Riedel with the introduction of two with the in three- (1994), egg masses egg capsules arranged a

Simrothina new suborders, (comprising the superfamilies dimensional spiral (10). The Bursinae have developed a

Naticoidea, Cypraeoidea and Lamellarioidea), and prominent posterior siphonal canal (11), the central

Troschelina Lau- radular teeth bear (comprising superfamilies Cassoidea, basal interlocking processes (12) and bierinioidea, and The Calyptraeoidea Capuloidea). the whorls of the larval shell do not have an open suture

used define characters to these systematic units are not and are covered only by a comparatively thin perio- repeated here apart from those of the cassoid branch, stracum (13). The Cymatiinae have developed conical to which are presented in a more detailed phylogenetic tree turriform protoconchs (14).

The (Fig. 43). following characters have already been The stem-group of the Cassidae is represented by the introduced in the preceding chapters and therefore are Oocorythinae, which shows the plesiomorphic conditions

summarised here. only (teleoconch, protoconch, radula, egg capsules, etc.) within

The Cassoidea have developed a larval pallial tentacle the family. The Cassinae and Phaliinae are united by two

(1) which allows the veliger to secrete, form, fix and synapomorphies: the cancellation of the larval shell is lost dissolve elements. It and periostracal sculptural represents a (15) the marginal radular teeth are extended, while special adaptation to a long-term planktonic life (see the central and lateral teeth are widened (16). Despite

Bandel et and the for the these other characters Beu's al., 1994) only synapomorphy two synapomorphies support

Cassoidea that could be found the available data among (1981) introductionof the subfamily Phaliinae. The egg

this on superfamily. Alan Beu (pers. comm., 1994) is of capsules of the Cassinae have collar-like apices (17), the opinion that of the teleoconch is while the Phaliinae retain basal and episodic growth a egg capsule type another for the Cassoidea. tower-like which is synapomorphy produce a egg mass (18), usually

In subdividing the Cassoidea into families, the brooded (19).

Personidae can be defined by at least four apomorphies. The fourth subfamily of the Cassidae, the Tonninae,

The whorls of the teleoconch distorted the larval are (2), is very well defined by at least four apomorphies: the

carina periostracal sculpture forms a (3), the egg capsules operculum is lost during metamorphosis (20), the egg

more-or-less and the is flattenend solid are globular (4) proboscis ex- capsules are spheres forming a tongue- tremely long and has to be coiled when retracted (5). The shaped mass (21), the ornament of the larval shell is

Ranellidae and Cassidae both include echinoderms in largely reduced (22) and the bases of the central radular their diet (6) and are also united by the possession of teeth bear interlocking processes (23), as in the Bursinae.

acid. The large accessory salivary glands (7) containing In the past there have been some classificatory

Cassidae can be separated from the Ranellidae by two problems with the tun-shells, the helmet-shells and features: they are specialised for feeding on echinoderms Oocorys and relatives. Fischer (1887) introduced the

(8) and the accessory salivary glands are extremely family Oocorythidae, while Dall (1909) placed Oocorys

the wall to exert enlarged (9), allowing body pressure by in the Cassidae, Thiele (1929) and Turner (1948) in the muscular contraction. Tonnidae. Quinn (1980) again recognised a family

The Ranellidae show the plesiomorphic conditions and Oocorythidae. Waren & Bouchet (1990) preferred assign- cannot be defined by an apomorphy. The Ranellinae ment of the Oocorythinae to the Cassidae. Beu (pers. represent the basal line from which the other two subfam- comm.) synonymises the Oocorythinae with the Cassinae, ilies have branched off. The genera Charonia and Sassia which together with the Phaliinaeconstitute the Cassidae. are placed in the Ranellinae on the basis of their radulae, The Tonnidae are considered to be closely related to egg capsules and protoconchs, although the shape of the Cassidae. The classification presented here collects these larval shell of Charonia is similar to those of some taxa at the subfamily level within a single family and

Cymatiinae. Sassia and Gyrineum are probably the oldest therefore incorporates former proposals, which initially genera of the Ranellidae and Sassia may have derived appeared to be controversial, but in fact combined from Gyrineum. The alignment of teleoconch varices is observations on different taxonomic levels. about 180° in Gyrineum and about 270° in Sassia (com- The Ranellinae and Oocorythinae are both para-

180° considered is of pare Beu, 1981, 1985), being plesio- phyletic taxa which, however, a good indication 125

stem-groups. The chronology of the fossil record coin- edition). New York (Van Nostrand Reinhold), 663 pp

cides largely with the cladistic analysis and therefore Abbott, R.T., 1993. Phalium (Semicassis) vector, a new deep-

from central — The to Tonninae water species the Indian Ocean. appears be reliable. The cannot be older than Nautilus, 107(3): 94-96. the Oocorythinae, the Cassinae and Phaliinae are mono- Abdel-Gawad, G.I., 1986. Maastrichtian non-cephalopod phyletic and must have evolved at the same time, the mollusks (Scaphopoda, Gastropoda and Bivalvia) of the Personidae do not have large accessory salivary glands

middle Vistula Valley, central Poland. — Acta Geologica and therefore must have branched off from the stem-line Polonica, 36(1): 69-224, 48 pis. earlier than the Cassidae, etc. The latter, however, implies Amaudrut, M.A., 1898. La partie anterieure du tube digestif et that all ranellids and cassids are descended from a single la torsion chez les Mollusques gasteropodes. — Annales des species, which had developed accessory salivary glands. Sciences naturelles, Zoologie, 7: 1-290, 10 pis. This would that the other Ranellinae without mean early Amio, M., 1963. A comparative embryology of marine gastro-

such glands became extinct. A second possibility could pods, with ecological considerations. — Contributions of

the Shimonoseki of 392: 15-139. be the loss of accessory salivary glands in the Personidae. University Fisheries,

Amoroux, J.M., 1974. Observations sur la biologie du In my opinion it is more likely that the Personidae are as

Charonia nodiferus — old as the Ranellidae and did not branch off from the mollusque gasteropode (Lamarck).

Vie et Milieu, 24: 365-368. Ranellinae, both taxa having evolved from a common Anderson, D.T., 1959. The reproduction and early life history ancestor. In this case the fossil record must be interpreted of the gastropod Cymatilesta spengleri (Perry) (Fam. Cy- as incomplete and the Personidae must have evolved

matidae). — Proceedings of the Linnean Society of New during the Aptian at the latest. South Wales, 84(2): 232-238.

Anderson, D.T., 1965. Further observations on the life histories

of littoral gastropods in New South Wales. — Proceedings

ACKNOWLEDGEMENTS of the Linnean Society of New South Wales, 90(3): 242-

252.

1958. Cretaceous of the Pacific I am very grateful to Klaus Bandel (Hamburg) and Alan Anderson, F.M., Upper coast.

Beu (Lower Hutt, New Zealand), who both contributed — Memoirs of the Geological Society of America, 71: 1-

378. greatly to my knowledge of the Cassoidea. Alan Beu

1961. Cassis cluster. — Anonymous, cornuta (helmet shell) egg reviewed the manuscript and proposed many important Hawaiian Shell News, 10(1): 1. alterations, including those of a linguistic nature. 1 wish Arakawa, K.Y., 1960. Miscellaneous notes on Mollusca (2). to extend thanks to the many following colleagues: Mating and spawning habits of some marine molluscs. — Australian Museum (Sydney: Phil Colman, Ian Loch, Venus, 21(1): 72-78. Winston Ponder and Janet Waterhouse), Institut fur Hy- Arnaud, P.M., & J. Beurois, 1972. Premiere signalisation des und Fischereiwissenschaft Horst drobiologie (Hamburg: Charonia Ranella iles Saint-Paul genre et aux et Amsterdam Institut des Sciences naturelles de Weikert), royal revision du (Ocean Indien) et genre Argobuccinum (Gas-

Belgique (Brussels: Annie Dhondt), Leigh Laboratory tropoda, Cymatiidae). — Tethys, 3(4) (1971): 865-874.

(New Zealand: Bill Ballantine), Lizard Island Research Asano, M., & M. Itoh, 1960. Salivary poison of a marine

Station (Queensland: Anne Hoggett, Lyle Vail, Marianne gastropod Neptunea athritica Bernardi, and the seasonal

variations — and Lance Pearce), Mississippi Office of Geology of its toxicity. Annals of the New York Academy of Sciences, 90: 674-688. (Jackson: David T. Dockery), Nationaal Natuurhistorisch d'Asaro, C.N., 1969a. The comparative embryogenesis and Museum (Leiden: Arie Janssen), National Museum of early organogenesis of Bursa corrugata Perry and Distorsio New Zealand (Wellington: Bruce Marshall),

clathrata Lamarck (Gastropoda: Prosobranchia). — Ma- Naturhistorisches Museum (Vienna: Ortwin Schultz), lacologia, 9(2): 349-389. Portobello Marine Station (New Zealand: John Jillet) and d'Asaro, C.N., 1969b. The spawn of the emperor helmet shell, Bob Penniket (Warkworth, New Zealand). I dedicate this Cassis madagascariensis Lamarck, from South Florida. — to Bob Penniket, who died in 1993. paper January Bulletin of marine Science, 19(4): 905-910.

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Manuscript received 1 September 1995, revised version accepted 11 October 1995