CHECKLIST OF GASTROPOD SPECIES REFERRED TO THE ORDER COCCULINIFORMIA HASZPRUNAR, 1987 (GASTROPODA: COCCULINOIDEA ET LEPETELLOIDEA) WITH SOME ...
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CHECKLIST OF GASTROPOD SPECIES REFERRED TO
THE ORDER COCCULINIFORMIA HASZPRUNAR, 1987
(GASTROPODA: COCCULINOIDEA ET LEPETELLOIDEA)
WITH SOME REMARKS ON THEIR FOOD PREFERENCES
ANDRZEJ LESICKI
Department of Animal Physiology, Institute of Experimental Biology, A. Mickiewicz University,
Fredry 10, 61–701 Poznañ, Poland (e-mail: alesicki@main.amu.edu.pl)
ABSTRACT: Cocculiniform limpets live mostly in aphotic zones of deep-sea areas. In this environment, rather
poor in organic nutritients, they have specialized in utilizing different odd food sources. The paper presents
the list of all species included in the superfamilies Cocculinoidea Dall, 1882 and Lepetelloidea Dall, 1882.
Data on their habitat, food preferences and distribution are also included. Adaptive features in cocculiniform
limpets that permit them to assimilate organic nutritients of various origin are also discussed. It is suggested
that some expanded parts in their alimentary tracts may be used as “fermentative chambers” in which sym-
biotic bacteria could change the unassimilable food particles into simple compounds absorbed by limpets.
KEY WORDS: limpet, food preferences, alimentary tract adaptations, Cocculiniformia, Cocculinoidea, Lepetel-
loidea, checklist
INTRODUCTION: COCCULINIFORM LIMPET FAMILIES
Sunlight does not reach bathyal and abyssal zones 1987(a), but recently only Cocculinoidea have been re-
of seas and oceans, and photosynthesis cannot take tained in Cocculiniformia (BIELER 1992, LINDBERG &
place there. Thus, apart from the exceptional animal PONDER 1991, PONDER & LINDBERG 1997). This super-
communities connected with chemoautotrophic pri- family includes the families Cocculinidae Dall, 1882(a)
mary production near hydrothermal vents and cold and Bathysciadiidae Dautzenberg et Fischer, 1899.
seeps (C HILDRESS & F ISHER 1992, T UNNICLIFFE Lepetelloidea (with 8 families: Lepetellidae Dall,
1991), animal life at the sea bottom can be fuelled 1882(a), Addisoniidae Dall, 1882(a), Bathyphytophil-
only by organic particles falling down from the
idae Moskalev, 1978, Choristellidae Bouchet et Warén,
euphotic zone (GOODAY & TURLEY 1990, STOCKTON
1979, Cocculinellidae Moskalev, 1971, Osteopeltidae
& DELACA 1982). Such organic falls reaching the
Marshall, 1987, Pseudococculinidae Hickman, 1983
deep-sea floor are rather scarce, therefore deep-sea
animals often specialize in consuming very odd food. and Pyropeltidae McLean et Haszprunar, 1987) have
Among such specialized consumers, there is a group been shifted to Vetigastropoda Salvini-Plawen, 1980
of snails, most of them with limpet-shaped shells, (PONDER & LINDBERG 1997). This recent change in the
which is especially noteworthy. They are included taxonomic position of cocculinoid and lepetelloid lim-
into two superfamilies: Cocculinoidea Dall, 1882(a) pets is recognized, however in this paper they all are
and Lepetelloidea Dall, 1882(a). Not long ago were called cocculiniform limpets according to the former
they joined in the order Cocculiniformia Haszprunar, classification.48 Lesicki A.
FOOD PREFERENCES IN COCCULINIFORM LIMPETS
Cocculinidae and Pseudococculinidae feed on substrata, rather than directly on the substrata. It is an
wood which has sunk to the ocean floor (HASZ- open question if the cocculiniform limpets consume
PRUNAR 1988ab, MARSHALL 1986, MOSKALEV 1976, bacteria free living on the substrata or if they harbour
WOLFF 1979). Although it is not excluded that they symbiotic bacteria in their alimentary tracts. Al-
eat directly wood (maybe due to contamination with though particular families specialize in characteristic
symbiotic bacteria which digest cellulose), probably nutrition sources, there are some exceptions. Teuthi-
their food consists of different microorganisms which rostria cancellata feeds on cephalopod beaks (MOSKA-
decompose plant remains (MARSHALL 1986). Bathy- LEV 1976) (the other cocculinids feed on wood), so
phytophilidae also eat plants, such as algal holdfasts does Helicopelta rostricola (MARSHALL 1996) (the other
and seagrass (MOSKALEV 1978, WOLFF 1976). Lepetel- addisoniid limpets prefer elasmobranch egg cases).
lidae are found on empty polychaete tubes (DALL Pyropelta corymba and P. musaica probably feed on che-
1882a, 1889a, MOSKALEV 1976, 1978, VERRILL 1880). moautotrophic bacteria living on sulphide crust at hy-
It is again disputable whether the tubes themselves, or drothermal vents (MCLEAN & HASZPRUNAR 1987) but
microorganisms living on them are the nutrition for they have also been collected from whale skulls
lepetellids. Bathysciadiidae feed on chitinous beaks of (MCLEAN 1992a). Coccopigya spinigera usually con-
dead cephalopods (D ALL 1908, D AUTZENBERG & sumes wood, like the other cocculinid limpets, but it
FISCHER 1899, MOSKALEV 1973). The same source of was found on a whale skull, too (WARÉN 1991). An-
food is used by cocculinid Teuthirostria cancellata other cocculinid limpet, Paracocculina cervae, was usu-
(MOSKALEV 1976) and addisoniid Helicopelta rostricola ally collected from deep-sunken wood or algal hold-
(MARSHALL 1996). Furthermore, Cocculinellidae de- fasts (HASZPRUNAR 1987a, MARSHALL 1986) but MAR-
rive their nutrition from decaying fish bones (HASZ- SHALL (1994) noted its occurrence on whale bones.
PRUNAR 1988c, MARSHALL 1983). On the other hand, Such exceptional cases have led WARÉN (1996b) to a
whale skeletons have become food for Osteopeltidae conclusion that many deep-sea limpets can live on a
(MARSHALL 1987) although some representatives of “second choice” substratum when the first is not avail-
Cocculinidae (e.g. Cocculina craigsmithi) and Pyropel- able. Such a conclusion would be in agreement with
tidae (e. g. Pyropelta wakefieldi) may be also found on the hypothesis that these limpets feed on free living
decaying whale bones and vertebrae (MARSHALL bacteria. However, it seems that the hypothesis of
1994, MCLEAN 1992a). This latter family is included symbiotic bacteria in the alimentary tract of cocculini-
in the hydrothermal vent fauna, i.e. animal communi- form gastropods is more probable. Several authors
ties connected with chemoautotrophic primary pro- noted substratum debris observed in the digestive
duction of this unique ecosystem (LESICKI 1998). Ad- tracts (HASZPRUNAR 1987ab, 1988b, HASZPRUNAR &
disoniidae and Choristellidae (as well as pseudococ- MCLEAN 1996, SIMONE 1996). The actual consump-
culinid Tentaoculus balantiophaga) have been found tion of the odd substrata may be also evidenced by the
exclusively in empty egg cases of sharks and skates very diverse radular structures in particular families,
(MARSHALL 1996, MCLEAN 1985, 1992b, VERRILL suggesting adaptive differentiation (HICKMAN 1983)
1882, VILLA 1985). Two other pseudococculinids of and by differences of the alimentary tract anatomy in
the genus Tentaoculus (T. lithodicola and T. neolithodi- these families (H ASZPRUNAR 1987ab, 1988abc,
cola) live on carapaces of living deep-sea crabs of the 1992ab, HASZPRUNAR & MCLEAN 1996). HASZPRUNAR
family Lithodidae (MARSHALL 1986). A bathysciadiid- (1988a) suggested that the alimentary tract of the
like limpet feeds on the periostracum of a gastropod families feeding on wood, that is Cocculinidae and
of the genus Capulus (WARÉN 1993). Pyropelta bohlei Pseudococculinidae, represents the most primitive or-
was also found attached to the shell of Bathyacmaea ganization. Strongly cuticularized epithelium of some
jonassoni (BECK 1996). stomach part, creating a so called gastric shield, may
The substrata used by cocculinoid and lepetelloid be used for mechanical crushing of hard food before
gastropods as sources of their nutrition are built of it is digested by bacteria in the long intestine coiled in
unassimilable substances. Wood is built of cellulose, three loops (a long intestine is usually characteristic
cephalopod beaks are composed of chitin. Both are for plant consumers). The specialization of the other
large, filamentous polysaccharides. Egg cases of elas- cocculiniform families to other food sources is re-
mobranch fish comprise a fibrous protein, collagen. flected in serious modifications of their alimentary
Gastropod periostracum is also built of fibrous pro- tracts. Polysaccharide (chitin) consumers, such as
teins. Animals usually do not secrete their own en- bathysciadiids, have a very large stomach but they
zymes which could digest such molecules. Only bac- have lost their midgut gland, which has been replaced
teria can produce appropriate enzymes. MARSHALL with a greatly expanded oesophageal gland. A very
(1986) suggested that the cocculiniform limpets feed large stomach is also characteristic for bathyphyto-
on bacteria which are associated with their biogenic philids (HASZPRUNAR & MCLEAN 1996). On the otherCocculiniformia, Checklist of species 49
hand, fibrous protein diet resulted in progressive re- bers” in which symbiotic bacteria could change the
duction of the stomach, observed in cocculinellids unassimilable food particles into simple compounds
and addisoniids (members of the latter family have no absorbed by limpets. However, further physiological
stomach at all) (HASZPRUNAR 1987b, 1988ac). It ap- studies are desperately needed to explain the details
pears that, instead of stomach, addisoniid limpets of digestive processes in cocculinoid and lepetelloid
have an extremely large intestinal sac which occupies gastropods.
about two thirds of the animal’s body (HASZPRUNAR MARSHALL (1996) noticed that decaying beaks,
1987b). Although Haszprunar initially reported a lack egg cases, bones, wood and algal holdfasts are gener-
of stomach in choristellids and a similarity of their ali- ally rare at the sea-floor. This raises a further unan-
mentary tract anatomy to that of addisoniids (HASZ- swered question: how can the limpets find their food?
PRUNAR 1988a), later he described a very large, highly The development of these gastropods is unknown.
modified stomach for this family (H ASZPRUNAR Their larvae are probably lecithotrophic and may be
1992ab). Nevertheless it may be said that in all coccu- dispersed by bottom currents (MARSHALL 1986). The
liniform gastropods there is some expanded chamber chemoreception-based recognition of the food sub-
in their alimentary tracts. It is suggested here that strata may be only speculated upon.
such chambers may be used as “fermentative cham-
RECENT KNOWLEDGE OF COCCULINIFORM LIMPETS
During the last 25 years our knowledge of the ution (HASZPRUNAR 1988ab, 1992ab, HASZPRUNAR &
snails of the superfamilies Cocculinoidea and Lepe- MCLEAN 1996, HICKMAN 1983, MARSHALL 1996). The
telloidea has been seriously increased. Studies initi- papers published during the last three years (BECK
ated by MOSKALEV in the seventies (1971, 1973, 1976, 1996, HASEGAWA 1997, HASZPRUNAR & MCLEAN 1996,
1978) and continued in the eighties and nineties by LEAL 1996, MARSHALL 1996, MCLEAN & HARASEWYCH
HASZPRUNAR (1987ab, 1988abc, 1992ab, 1996), MAR- 1995, SIMONE 1996, WARÉN 1996ab) contain descrip-
SHALL (1983, 1986, 1987, 1994, 1996), M C L EAN tions of 17 new species (further 8 have been presented
(1985, 1988, 1991, 1992ab) and WARÉN (1989, 1991, but not named yet), redescription and reclassification
1993, 1996ab) have brought not only the discovery of another 22 species, establishment of a new genus
and description of several new species. As a result of and a new subfamily. Descriptions of further species
the detailed comparative anatomical studies, they are being prepared (MARSHALL, LEAL, MCLEAN – per-
have revealed relationships between them. It has also sonal communication). This is a proof that this gastro-
been possible to revise descriptions of earlier dis- pod group still inspires new research. Therefore it
covered species as well as to improve the system intro- seems reasonable to compile in one paper all informa-
duced on the turn of the 19th century (DALL 1882ab, tion on the species included in the superfamilies Coc-
1889ab, 1896, DAUTZENBERG 1886, 1889, THIELE culinoidea and Lepetelloidea, as well as all data on
1908, 1909, 1925, 1929, VERRILL 1880, 1882, 1884). their habitat, food preferences and distribution areas.
The recent discoveries throw some new light on the The checklist should also give a detailed bibliography
problems of the gastropod origin, specially on the of both superfamilies.
role of limpet-shelled forms in the gastropod evol-
CHECKLIST OF COCCULINIFORM LIMPET SPECIES
Superfamily: C o c c u l i n o i d e a Dall, 1882(a) Cocculina angulata Watson, 1886
(Thiele, 1909 emend.) (western Pacific: Philippines)
(MCLEAN & HARASEWYCH 1995)
Family: COCCULINIDAE Dall, 1882(a)
“Cocculina” alta Smith, 1894
Genus: Cocculina Dall, 1882(a)
(southwest Pacific: off New South Wales, 750 m)
(type species: Cocculina rathbuni Dall, 1882)
“Cocculina” alveolata Schepman, 1908
Several species have been referred to the genus
(Indo-Pacific)
Cocculina Dall, 1882(a), however their generic
Cocculina baxteri McLean, 1987
(and family) status has not been recently con- (northeastern Pacific: Prince William Sound,
firmed by the studies on their anatomies and radu- Alaska; 424–430 m; on wood)
lae, these species are listed as “Cocculina”. (HASZPRUNAR 1987a)50 Lesicki A.
Fig. 1–8. Shells of cocculiniform limpets. Cocculinidae: 1 – Cocculina surugaensis Hasegawa, 1997; 2 – Cocculina tenuitesta
Hasegawa, 1997; 3 – Coccopigya okutanii Hasegawa, 1997; 4 – Coccopigya punctoradiata (Kuroda et Habe, 1949); Pseudo-
cocculinidae: 5, 6 – Pseudococculina subcingulata (Kuroda et Habe, 1949); 7 – Notocrater pustulosa (Thiele, 1925); 8 – Copu-
labyssia similaris Hasegawa, 1997. Scale bars: 500 µm (1, 2, 5–8) or 1 mm (3, 4). Published by the courtesy of KAZUNORI
HASEGAWA (National Science Museum, Tsukuba, Japan) after HASEGAWA (1997)Cocculiniformia, Checklist of species 51
(northeastern Pacific: Queen Charlotte Sound, (off Southwest Reef, New Providence Island Baha-
Vancouver Isl., British Colombia; 265 m) mas; 518 m; on palmetto fronds)
(DALL 1921, MCLEAN 1987) (MCLEAN & HARASEWYCH 1995)
Cocculina cowani McLean, 1987 “Cocculina” striata Schepman, 1908
(northeastern Pacific: off Moresby Isl., Queen (Indo-Pacific)
Charlotte Islands, British Columbia; 1370 m; on “Cocculina” subcompressa Schepman, 1908
wood) =? “Cocculina” nipponica Kuroda et Habe, 1949
(HASZPRUNAR 1987a) (northwestern Pacific: off Japan)
Cocculina craigsmithi McLean, 1992(a) “Cocculina” subquadrata Schepman, 1908
(northeastern Pacific: Santa Catalina Basin be- (Indo-Pacific)
tween Santa Catalina Isl. and San Clemente Isl.; Cocculina surugaensis Hasegawa, 1997 (Fig. 1)
1240 m; on whale bones) (northwestern Pacific: Suruga Bay, Honshu, Ja-
“Cocculina” diomedae Dall, 1908 pan, 345–920 m; Sagami Bay, Honshu, Japan,
(east Pacific: off West America) 750–870 m; on wood)
“Cocculina” dofleini Thiele, 1925 Cocculina tenuitesta Hasegawa, 1997 (Fig. 2)
(Indo-Pacific) (northwestern Pacific: Suruga Bay, Honshu, Ja-
Cocculina emsoni McLean et Harasewych, 1995 pan, 275–500 m; on wood)
(northwestern Atlantic: off Southwest Reef, New Cocculina n. sp. (1) (MOSKALEV 1976)
Providence Island, Bahamas; 518 m; on palmetto Cocculina n. sp. (2) (HASEGAWA 1997)
fronds) (northwestern Pacific: Suruga Bay, Honshu, Japan,
“Cocculina” fragilis Thiele, 1925 180–710 m; on wood)
(western Indian Ocean: Zanzibar Channel,
E. Africa) Genus: Coccorater Haszprunar, 1987(a)
“Cocculina” japonica Dall, 1908 (type species: Cocculina radiata Thiele, 1904)
(northwestern Pacific: off Japan) Coccocrater agassizii (Dall, 1908) (Cocculina)
“Cocculina” japonica uncinata Kuroda et Habe, 1949 (eastern Pacific: Gulf of Panama; 1015 m)
(HASZPRUNAR 1987a, MCLEAN 1987)
(northwestern Pacific: off Japan)
Coccocrater pocillum (Dall, 1890) (Cocculina /Coccopygia/)
“Cocculina” leptoglypta Dautzenberg et Fischer, 1897
(western Atlantic: off Tobago; 1600 m)
(Atlantic: off Azores; 1550 m)
(MCLEAN & HARASEWYCH 1995)
(DANTART & LUQUE 1994)
Coccocrater portoricensis (Dall et Simpson, 1901) (Coccu-
Cocculina messingi McLean et Harasewych, 1995
lina)
(northwestern Atlantic: south of Settlement Point,
(western Atlantic: off San Juan Harbor, Porto
Grand Bahama Island; 412 m; on wood)
Rico; 566 m) (MCLEAN & HARASEWYCH 1995)
“Cocculina” nassa Dall, 1908 Coccocrater radiata (Thiele, 1904) (Cocculina)
(equatorial eastern Pacific) (eastern Indian Ocean: off Sumatra; 614 m)
“Cocculina” oblonga Schepman, 1908 (HASZPRUNAR 1987a)
(Indo-Pacific)
Cocculina ovata Schepman, 1908 Genus: Coccopigya Marshall, 1986
(Indo-Pacific: Saleh Bay, north coast of Sumbawa (nomen novum pro Coccopygia Dall, 1889/a/)
Island, Indonesia; 274 m; Philippines; 187–210 m) (type species: Cocculina spinigera Jeffreys, 1883)
(HASZPRUNAR 1987a, MCLEAN 1987) Coccopigya barbatula Marshall, 1986
“Cocculina” pacifica Kuroda et Habe, 1949 (southwestern Pacific: off New South Wales;
(northwestern Pacific: off Japan) 384–457 m; on wood)
Cocculina sp. cf. pacifica Kuroda et Habe, 1949 (HASE- Coccopigya compunctum (Marwick, 1931) (Tectisumen)
GAWA 1997) (early or middle Miocene, NE of Tekaraka, Gis-
(northwestern Pacific: Suruga Bay, Honshu, Ja- borne District, New Zealand) (MARSHALL 1986)
pan, 180–680 m; on wood) Coccopigya crebrilamina Marshall, 1986
Cocculina pristina Marshall, 1986 (southwestern Pacific: Whale Island to Tetara
(Otaian, early Miocene, 1.6 km NW of Pakaurangi Head, New Zealand; 55–146 m; on large log)
Point, Kaipara, New Zealand) Coccopigya crinita Marshall, 1986
Cocculina rathbuni Dall, 1882(a) (southwestern Pacific: New Zealand; New South
(western Atlantic: Massachusetts, Barbados, Mar- Wales; 203–914 m; on wood)
tinique; 730–919 m) Coccopigya hispida Marshall, 1986
(264 km SE off Martha's Vineyard Isl.; 925 m; off (southwestern Pacific: White Island to off Timaru,
Ford Pierce, Florida, 124 m) New Zealand; 833–1514 m; on wood)
(HASZPRUNAR 1987a, MCLEAN 1987) (HASZPRUNAR 1987a)52 Lesicki A.
Coccopigya komitica Marshall, 1986 (western Atlantic: Puerto Rico Trench; 8595 m)
(Otaian, early Miocene, 1.6 km NW of Pakaurangi (LEAL 1996)
Point, Kaipara, New Zealand) Fedikovella n. sp. (1) (MOSKALEV 1976)
Coccopigya lata Warén, 1996(a) (western Atlantic: off Puerto Rico; 7950–8100 m)
(north Atlantic: off southwestern Iceland; on Fedikovella n. sp. (2) (MOSKALEV 1976)
sunken driftwood) (western Atlantic: off Puerto Rico; 7950–8100 m)
Coccopigya mikkelsenae McLean et Harasewych, 1995 Fedikovella n. sp. (3) (MOSKALEV 1976)
(western Atlantic: off Chateau Belair Bay, St. Vin- (western Atlantic: off Puerto Rico; 8330 m)
cent, Lesser Antilles; 421 m; on wood)
Coccopigya oculifera Marshall, 1986 Genus: Paracocculina Haszprunar, 1987(a)
(southwestern Pacific: White Island to off Timaru, (type species: Cocculina laevis Thiele, 1904)
New Zealand; 248–962 m; on wood) Paracocculina cervae (Fleming, 1948) (Cocculina)
Coccopigya okutanii Hasegawa, 1997 (Fig. 3) (southwestern Pacific: North Cape to Long Sound,
(northwestern Pacific: Suruga Bay, Honshu, Ja- New Zealand; 18–891 m; on whale bones, algal
pan; 205–740 m; on wood) holdfast, deep-sunken wood) (H ASZPRUNAR
Coccopigya otaiana Marshall, 1986 1987a, MARSHALL 1986, 1994)
(Otaian, early Miocene, 1.6 km NW of Pakaurangi Paracocculina laevis (Thiele, 1904) (Cocculina)
Point, Kaipara, New Zealand) (eastern Indian Ocean: off Nias Island, Sumatra)
Coccopigya punctoradiata (Kuroda et Habe, 1949) (Coc- (HASZPRUNAR 1987a)
culina) (Fig. 4)
(northwestern Pacific: off Tosa, Shikoku; Suruga Genus: Teuthirostria Moskalev, 1976
Bay, Honshu; 120–1708 m; on wood) (type species: Teuthirostria cancellata Moskalev, 1976)
(HASEGAWA 1997) Teuthirostria cancellata Moskalev, 1976
Coccopigya spinigera (Jeffreys, 1883/c/) (Cocculina) (eastern Pacific: off Peru; 5200–5540 m; on cepha-
= Cocculina conspersa Dautzenberg et Fischer, 1897 lopod beaks)
(northwestern Atlantic: off the northeastern
United States to western and southern Iceland,
Gen. n., sp. n. (LEAL 1996)
and to north of the Hebrides; also Mediterranean:
(western Atlantic: Puerto Rico Trench; 8595 m)
between Catalonia and Mallorca, and Chafarinas
Island, N. Marocco; 200–1534 m; on submersed
wood and whale skeletons, also on tubes of xy- Family: BATHYSCIADIIDAE Dautzenberg et Fischer,
lophagid and teredinid ship-worms) 1899
(DALL 1889a; DANTART & LUQUE 1994; MCLEAN &
HARASEWYCH 1995; WARÉN 1991) Genus: Bathysciadium Dautzenberg and Fischer, 1899
Coccopigya viminensis (Rocchini, 1990) (Cocculina) (type species: Bathysciadium conicum Dautzenberg
(Mediterranean: between Barcelona, Tarragona et Fischer, 1899)
and Mallorca, Tusean Archipelago; 450–1883 m; = Bathypelta Moskalev, 1971 (? Family: Bathypelti-
on wood) (DANTART & LUQUE 1994) dae Moskalev, 1971; Superfamily: Bathypeltoidea
Coccopigya sp. (HASEGAWA 1997) Moskalev, 1971) (type species: Bathysciadium pacifi-
(northwestern Pacific: Suruga Bay, Honshu, Ja- cum Dall, 1908) (WARÉN 1996b)
pan; 180–680 m; on wood) = Bonus Moskalev, 1973
(type species: Bonus petrochenkoi Moskalev, 1973)
Genus: Fedikovella Moskalev, 1976 (type species: (WARÉN 1996b)
Fedikovella caymanensis Moskalev, 1976) Bathysciadium concentricum Dall, 1927
Fedikovella beanii (Dall, 1882/a/) (Cocculina) (northwestern Atlantic: off Georgia; 800 m)
(northwestern Atlantic: Martha's Vineyar Island, Bathysciadium costulatum (Locard, 1897) (Lepeta)
Massachusetts; Martinique; 183–1846 m) (Atlantic: south of the Azores; 3175 m; on a cepha-
(off Chateau Belair Bay, St. Vincent, Lesser Antil- lopod jaw)
les; 421 m; on wood) (MCLEAN & HARASEWYCH = Bathysciadium conicum Dautzenberg et Fischer,
1995, MOSKALEV 1976) 1899
Fedikovella capulus (Thiele, 1925) (Cocculina) (Atlantic: off Azores, off Spanish Sahara;
(western Indian Ocean: Zanzibar Channel; 463 m) 1000–2000 m; on cephalopod beaks) (PELSENEER
doubtfully included in Fedikovella (MCLEAN & 1899, 1940; WARÉN 1996b)
HARASEWYCH 1995, MOSKALEV 1976) Bathysciadium pacificum Dall, 1908
Fedikovella caymanensis Moskalev, 1976 (eastern Pacific: off Peru; 4115 m; on cephalopod
(western Atlantic: Cayman Trench; 6740–6800 m; beaks) (WARÉN 1996b)
on sunken wood) (MCLEAN & HARASEWYCH 1995) Bathysciadium petrochenkoi (Moskalev, 1973) (Bonus)Cocculiniformia, Checklist of species 53
(northwestern Pacific: Kurile-Kamchatka Trench; Cocculina dalli Verrill, 1884
9130–9430 m) (WARÉN 1996b) close to Iothia rugosa (Jeffreys, 1883/b/) Lepetidae
Bathysciadium rotunda (Dall, 1927) (Cocculina) Dall, 1869 (MCLEAN & HARASEWYCH 1995)
(northwestern Atlantic: off Fernandina, Florida) Cocculina lissocona Dall, 1927
(tentatively referred to Bathysciadium by MCLEAN & referred to Propilidium Forbes et Hanley, 1849,
HARASEWYCH 1995) Lepetidae Dall, 1869 (MCLEAN & HARASEWYCH
Bathysciadium xylophagum Warén et Carrozza, 1996 (in 1995)
WARÉN 1996b) Cocculina maxima Dautzenberg, 1925
(Mediterranean: off Sardinia; 630 m; in holes belongs to patellogastropod genus Pectinodonta
made by ship-worms in a piece of sunken wood) Dall, 1882(a) (MARSHALL 1985)
Bathysciadium sp. cf. xylophagum Warén et Carrozza, Cocculina obtusa Thiele, 1925
1996 (WARÉN 1996b) probably belongs to patellogastropod genus Pecti-
(Atlantic: off southwestern Portugal, Josephine nodonta Dall, 1882(a) (MARSHALL 1985)
Bank; 200 m; on a cephalopod beak) Cocculina petasus Thiele, 1925
Bathysciadium n. sp. (1) (HASZPRUNAR 1988a) is a basal plate of a barnacle – Crustacea: Cirripe-
(southwestern Pacific: off New Zealand) dia (WARÉN 1985)
Bathysciadium n. sp. (2) (HASZPRUNAR 1988a) Cocculina reticulata Verrill, 1885
(southwestern Pacific: off New Zealand) referred to Propilidium Forbes et Hanley, 1849,
Bathysciadium n. sp. (3) (WARÉN 1996b) Lepetidae Dall, 1869 (MCLEAN & HARASEWYCH
(southwestern Indian Ocean: off Reunion Island; 1995)
2830–2850 m) Cocculina rhyssa Dall, 1925
Bathysciadium n. sp. (4) (WARÉN 1996b) belongs to patellogastropod genus Pectinodonta
(Mediterranean: off Capraia Island; 150 m) Dall, 1882(a) (HASEGAWA 1997, MARSHALL 1985)
Cocculina scabra Kuroda et Habe, 1949
Genus: Pilus Warén, 1991 (tentatively placed in the referred to Iothia Forbes, 1849, Lepetidae Dall,
Bathysciadiidae by WARÉN 1993) 1869 (HASEGAWA 1997, INABA & OYAMA 1977)
(type species: Cocculina conica Verrill, 1884) Cocculina superba Clarke, 1960
Pilus conica (Verrill, 1884) (Cocculina) referred to Lepetidae Dall, 1869 (M C L EAN &
(northwestern Atlantic: northeastern coasts of HARASEWYCH 1995)
United States and southwestern coasts of Iceland; Cocculina teramachii Kuroda et Habe, 1949
900–1000 m) referred to Iothia Forbes, 1849, Lepetidae Dall,
(MCLEAN & HARASEWYCH 1995; WARÉN 1993) 1869 (HASEGAWA 1997, INABA & OYAMA 1977)
Genus: Xenodonta Warén, 1993 Genus: Dallia Jeffreys, 1883 (a) (type species: Tectura
(tentatively placed in the Bathysciadiidae) (Dallia) galeola Jeffreys, 1883/a/) with the species:
(type species: Xenodonta bogasoni Warén, 1993) D. galeola (Jeffreys, 1883/a/), D. pusilla (Jeffreys,
Xenodonta bogasoni Warén, 1993 1883/a/) and D. adunca (Jeffreys, 1883/a/)
(north Atlantic: off western and southwestern Ice- probably belongs to Lepetidae Dall, 1869 (MAR-
land; 260–770 m) SHALL 1986) but their specific, generic and famil-
Xenodonta n. sp. (WARÉN 1993) iar status is unclear (DANTART & LUQUE 1994)
(eastern Pacific: off Galapagos Islands; on the shell
periostracum of Capulus sp.) Genus: Maoricrater Dell, 1956 (type species: Notoacmea
explorata Dell, 1953) belongs to Lepetidae Dall,
species excluded from Cocculinoidea Dall, 1882(a) 1869 (HICKMAN 1983, MOSKALEV 1977)
Acmaea parva var. tasmanica Pilsbry, 1895
= Cocculina meridionalis Hedley, 1903 Genus: Propilidium Forbes et Hanley, 1849 (type spe-
= Acmaea excentrica Test, 1945 cies: Patella ancyloides Forbes, 1840) belongs to
sometimes referred in Cocculina or Notocrater be- Lepetidae Dall, 1869 (DANTART & LUQUE 1994,
longs to Propilidium Forbes et Hanley, 1849, MARSHALL 1986)
Lepetidae Dall, 1869 (MARSHALL 1986)
Cocculina aethiopica Thiele, 1925 Superfamily: L e p e t e l l o i d e a Dall, 1882(a)
is a basal plate of a barnacle – Crustacea: Cirripe- (Thiele, 1908 emend.)
dia (WARÉN 1985)
Cocculina casanica Dall, 1919 Family: LEPETELLIDAE Dall, 1882(a)
is a junior synonym of Lepeta caeca (Müller, 1776) Genus: Lepetella Verrill, 1880 (type species: Lepetella tu-
and belongs to Lepetidae Dall, 1869 (MCLEAN bicola Verrill et Smith in Verrill, 1880)
1987) Lepetella barrajoni Dantart et Luque, 199454 Lesicki A.
(Atlantic: off Iberian Peninsula, 82–86 m; on poly- Genus: Bogia Dantart et Luque, 1994
chaete Hyalinoecia tubicola tubes) (type species: Cocculina labronica Bogi, 1984)
Lepetella espinosae Dantart et Luque, 1994 (family uncertain, should be excluded from Lepe-
?= Lepetella laterocompressa auct. non Patella latero- tellidae according to DANTART & LUQUE 1994)
compressa De Rayneval et Ponzi, 1854 (partim) Bogia labronica (Bogi, 1984) (Cocculina)
(Mediterranean: off Iberian Peninsula, off Capraia (Mediterranean: Tyrrhenian Sea; 80–220 m)
Island (Italy); 58–272 m; on polychaete Hyalinoecia
tubicola tubes) Genus: Sablea Allen, 1970
Lepetella ionica Nordsieck, 1973 (type species: Sablea minuta Allen, 1970)
= Cocculina mamilla Di Geronimo, 1974 Sablea minuta Allen, 1970
(Mediterranean: Ionian Sea; off Sardinia; off Ibe- (Eocene-Oligocene)
rian Peninsula between Catalonia and Mallorca; conchologically similar to Lepetella Verrill, 1880
900–4210 m) (DANTART & LUQUE 1994, JANSSEN (DANTART & LUQUE 1994; STEARNS & DOCKERY
1989) 1984)
Lepetella laterocompressa (De Rayneval et Ponzi, 1854)
(Patella) Genus: Tectisumen Finlay,1927 (type species: Cocculina
(Pleistocene, Monte Mario, Italy) clypidellaeformis Suter, 1908) (synonymised with
?= Gadinia compressa Tiberi in Jeffreys, 1883(a) Lepetella Verrill, 1880 by WARÉN (1972) and HICK-
MAN (1983) but not by HASZPRUNAR (1988a))
(recent, Mediterranean: off Palermo) (MONTERO-
SATO 1890) Tectisumen clypidellaeformis (Suter, 1908) (Cocculina)
?= Lepetella tubicola Jeffreys, 1883(a) (southwestern Pacific: deep water off New Zea-
land, on polychaete Hyalinoecia tubes) (DELL 1956,
?= Patella tricornis Turton, 1821
MOSKALEV 1978)
(recent, Mediterranean: Bay of Naples; North Sea:
Tectisumen compunctum (Fleming, 1966) (Lepetella
off Norway, 30–60 m) (WARÉN 1972)
/Tectisumen/)
?= Cocculina clipeus Thiele, 1925
(Cenozoic, New Zealand) (MOSKALEV 1978)
(off Cape Bojador, W. Africa) (WARÉN 1972) may
Tectisumen mayi Finlay, 1927
be a distinct Lepetella species (DANTART & LUQUE
(southwestern Pacific: off New Zealand)
1994)
(MOSKALEV 1978)
?= Propilidium aquitanense Locard, 1886
Tectisumen parallela (Fleming, 1966) (Lepetella /Tecti-
(Gulf of Biscay) may be a distinct Lepetella species sumen/)
(DANTART & LUQUE 1994) (Cenozoic, New Zealand) (MOSKALEV 1978)
(according to DANTART & LUQUE (1994) name L. Tectisumen tasmanica (May, 1920)
laterocompressa should be restricted to the fossil spe- (southwestern Pacific: off Australia)
cies, recent Mediterranean specimens, identified
as L. laterocompressa, probably were in fact L. espino-
Genus: Tecticrater Dell, 1956 (type species: Cocculina
sae or L. sierrai; status of North Sea specimens is
compressa Suter, 1908) (synonymised with Lepetella
unclear)
Verrill, 1880 by WARÉN (1972) and HICKMAN
Lepetella postapicula Dell, 1990 (1983) but not by HASZPRUNAR (1988a))
(Antarctica) Tecticrater compressa (Suter, 1908) (Cocculina)
Lepetella sierrai Dantart et Luque, 1994 = Tectisumen subcompressa Powell, 1937
?= Lepetella laterocompressa auct. non Patella latero- = Tectisumen finlayi Powell, 1937
compressa De Rayneval et Ponzi, 1854 (partim) (southwestern Pacific: off New Zealand; 260 m)
(eastern Atlantic: Bay of Biscay, 116–120 m; on (DELL 1956, MOSKALEV 1978)
empty polychaete Hyalinoecia tubicola tubes) “Tecticrater” grandis Crozier, 1966
Lepetella tosaensis (Kuroda et Habe, 1949) (Cocculina) (southwestern Pacific: off New Zealand)
(northwestern Pacific: off Japan) this species should be probably referred to Lepeti-
tentatively attributed to Lepetella Verrill, 1880 dae Dall, 1869 (MARSHALL, personal information)
(HASEGAWA 1997)
Lepetella tubicola Verrill et Smith in Verrill, 1880 Gen. nov. Warén in prep. (HASZPRUNAR 1988a)
(northwestern Atlantic: off New England; on poly-
chaete Hyalinoecia tubes, 351–651 m) (D ALL Family: ADDISONIIDAE Dall, 1882(a)
1882ab, 1889a, HICKMAN 1983, MOSKALEV 1978)
Lepetella n. sp. (?) (DANTART & LUQUE 1994) Subfamily: Addisoniinae Dall, 1882(a)
(eastern Atlantic: Bisagos Archipelago, Guinea- Genus: Addisonia Dall, 1882(a)
Bissau, W. Africa; 216–378 m) (type species: Addisonia paradoxa Dall, 1882)
Addisonia brophyi McLean, 1985Cocculiniformia, Checklist of species 55
(northeastern Pacific: Santa Barbara Basin to
Santa Catalina Basins; 155–174 m; in shark egg Family: CHORISTELLIDAE Bouchet et Warén, 1979
cases) = CHORISTIDAE auct.
Addisonia enodis Simone, 1996
(southwestern Atlantic: off Ubatuba, Sao Paulo St. Genus: Bichoristes McLean, 1992(b)
Brazil, 184 m) (type species: Bichoristes wareni McLean, 1992)
Addisonia excentrica (Tiberi, 1855) (Gadinia) Bichoristes wareni McLean, 1992(b)
= Addisonia lateralis auct., non Gadinia lateralis (southwestern Pacific: Norfolk Ridge, south of
Réquien, 1848 New Caledonia; 505–515 m)
= Addisonia eccentros Jeffreys, 1883(a)
(Mediterranean: Sicily to Corsica; Eastern Atlan- Genus: Choristella Bush, 1897 (type species: Choristella
tic: Bay of Biscay to Guinea Bissau) leptalea Bush, 1897)
(DAUTZENBERG 1886, 1889, GUBBIOLI & NOFRONI Choristella hickmanae McLean, 1992(b)
1986, HASZPRUNAR 1987b, LOCARD 1897, MCLEAN (northeastern Pacific: Northern Cascadia Abyssal
1985, TIBERI 1857, VILLA 1985) Plain, Strait of Juan de Fuca; 2176 m)
= Addisonia paradoxa Dall, 1882(a) (HASZPRUNAR 1992b)
(western Atlantic: Nova Scotia to Jamaica; in Choristella leptalea Bush, 1897
empty egg cases of sharks and skates) = Choristella brychia Bush, 1897 (MCLEAN 1992b)
(DALL 1882bc, 1889b, MCLEAN 1985, DANTART &
(northwestern Atlantic: off Martha's Vineyard Is-
LUQUE 1994, WARÉN 1996b)
land, Massachussetts; 713–1481 m)
Choristella marshalli McLean, 1992(b)
Subfamily: Helicopeltinae Marshall, 1996
(southwestern Pacific: off New Zealand; 376–1116
m; in empty skate egg case) (HASZPRUNAR 1992b)
Genus: Helicopelta Marshall, 1996
Choristella nofronii McLean, 1992(b)
(type species: Helicopelta rostricola Marshall, 1996)
= Cintha naticiformis auct., non Cintha naticiformis
Helicopelta rostricola Marshall, 1996
Jeffreys, 1883(b)
(southwestern Pacific: Chesterfield Plateau, Coral
Sea; 685–700 m; on a detrital cephalopod beak) (Alboran Sea, westernmost Mediterranean near
Spain; 50–100 m; in Raia egg cases)
Helicopelta n. sp. (MARSHALL 1996)
(southwestern Pacific: southeast of New Caledo- (GUBBIOLI & NOFRONI 1986, HASZPRUNAR 1992b)
nia; 750 m; on a detrital cephalopod beak) Choristella ponderi McLean, 1992(b)
(southwestern Pacific: off Australia; 91–552 m; in
species excluded from Addisoniidae Dall, 1882(a) skate egg case)
Gadinia lateralis Réquien, 1848 Choristella tenera (Verrill, 1882) (Choristes)
is a junior synonym of Trimusculus mammilaris (northwestern Atlantic: off Martha's Vineyard Is-
(Linnaeus, 1758) (Pulmonata) land, Massachussetts and off Cape Hatteras, North
(DANTART & LUQUE 1994, WARÉN 1996b) Carolina; 353–580 m; inside an old egg-case of
skate Raia) (MCLEAN 1992b)
Family: BATHYPHYTOPHILIDAE Moskalev, 1978 Choristella vitrea (Kuroda et Habe in Kuroda et al.,
1971) (Choristes)
Genus: Bathyphytophilus Moskalev, 1978 (type species: (northwestern Pacific: Sagami Bay, Japan; on egg
Bathyphytophilus caribaeus Moskalev, 1978) capsules of shark) (MCLEAN 1992b)
Bathyphytophilus caribaeus Moskalev, 1978 Choristella n. sp. (1) (MCLEAN 1992b)
(western Atlantic: Cayman Trench; 5800–6500 m; (southwestern Indian Ocean: Mozambique Chan-
on turtlegrass Thalassia testudinarum) nel; 3716 m)
Bathyphytophilus diegensis Haszprunar et McLean, 1996 Choristella n. sp. (2) (MCLEAN 1992b)
(northeastern Pacific: San Diego Trough; 1224 m; (southwestern Pacific: Norfolk Ridge; 503 m)
on surfgrass Phyllospadix scouleri) Choristella n. sp. (3) (MCLEAN 1992b)
Bathyphytophilus n. sp. (HASZPRUNAR & MCLEAN 1996) (Indo-Pacific: Banda Sea, off Tanimbar Island, In-
(northeastern Pacific: San Diego Trough; donesia; 356–368 m)
1207–1234 m; on surfgrass Phyllospadix torreyi) Choristella n. sp. (4) (MCLEAN 1992b)
(Indo-Pacific: Arafura Sea, off Kai Islands, Indone-
Genus: Aenigmabonus Moskalev, 1978 (type species: sia; 390–502 m)
Aenigmabonus kurilokamtschaticus Moskalev, 1978)
Aenigmabonus kurilokamtschaticus Moskalev, 1978 species excluded from Choristellidae Bouchet et
(northwestern Pacific: Kurile-Kamchatka Trench; Warén, 1979
6120–6160 m) Choristes agulhasae Clarke, 196156 Lesicki A.
probably in Trenchia Knudsen, 1964, Skeneidae
Clark, 1851 (MCLEAN 1992b) Genus: Osteopelta Marshall, 1987
Choristes agulhasae argentinae Clarke, 1961 (type species: Osteopelta mirabilis Marshall, 1987)
probably in Trenchia Knudsen, 1964, Skeneidae Osteopelta ceticola Warén, 1989
Clark, 1851 (MCLEAN 1992b) (north Atlantic: off southwestern Iceland between
Choristes carpenteri Dall, 1896 Vestmannaeyjar and Reykjanesridge; on whale
should be placed in Naticidae Forbes, 1838 bone)
(MCLEAN 1992b) Osteopelta mirabilis Marshall, 1987
Choristes coani Marincovich, 1975 (southwestern Pacific: Challenger Plataeu, north-
should be placed in Naticidae Forbes, 1838 east of Chatham Islands and Chatham Rise, New
(MCLEAN 1992b) Zealand; 800–955 m; on whale skulls and bones)
Choristes elegans Carpenter in Dawson, 1872 (HASZPRUNAR 1988c)
is synonym of naticid Amauropsis islandica (Gmelin, Osteopelta sp. cf. mirabilis Marshall, 1987 (MARSHALL
1791) (MCLEAN 1992b) 1994)
Choristes mollis Okutani, 1964 (middle Eocene, Waihao Greensand, New Zea-
probably in Granigyra Dall, 1889, Skeneidae Clark, land; associated with bones of fossil turtle)
1851 (MCLEAN 1992b) Osteopelta praeceps Marshall, 1994
Choristes nipponica Okutani, 1964 (southwestern Pacific: Chatham Rise and Chal-
should be excluded from Choristellidae (MCLEAN lenger Plataeu, New Zealand; 372–912 m; on
1992b) whale bones)
Cintha naticiformis Jeffreys, 1883(b)
probably in Trenchia Knudsen, 1964, Skeneidae Family: PSEUDOCOCCULINIDAE Hickman, 1983
Clark, 1851 (MCLEAN 1992b) (the subfamiliar attribution of particular genera
recently to Xyloskenea Marshall, 1988, Skeneidae may be premature according to MCLEAN 1991)
Clark, 1851 (WARÉN 1996a)
Cyclostrema pompholyx Dall, 1889(a) Subfamily: Pseudococculininae Hickman, 1983
should be not referred to Choristellidae (MCLEAN
1992b) Genus: Pseudococculina Schepman, 1908 (type species:
Cyclostrema valvatoides Jeffreys, 1883(b) Pseudococculina rugosoplicata Schepman, 1908)
probably in Skeneidae Clark, 1851 (M C L EAN Pseudococculina cingulata Schepman, 1908
1992b) (Indo-Pacific: off Indonesia)
Pseudococculina granulata Schepman, 1908
Family: COCCULINELLIDAE Moskalev, 1971 (Indian Ocean)
Pseudococculina gregaria Marshall, 1986
Genus: Cocculinella Thiele, 1909 (southwestern Pacific: off Southern New Zealand
(type species: Acmaea minutissima E. A. Smith, 1904) and New South Wales; 384–891 m; on wood)
Cocculinella coercita (Hedley, 1907) (Cocculina) (HASZPRUNAR 1988b)
(southwestern Pacific: off New South Wales and ”Pseudococculina” rosea Habe, 1952
eastern Victoria; 146–393 m; off Cape Jaffa, S. Aus- (northwestern Pacific: off Japan)
tralia; 238 m; on fish bones) (MARSHALL 1986) status not confirmed by recent anatomical studies
Cocculinella kopua Marshall, 1983 Pseudococculina rugosoplicata Schepman, 1908
(southwestern Pacific: off North Cape, New Zea- (eastern Indian Ocean: Sunda Sea, Indonesia,
land; 257–327 m) (species of uncertain validity, 2798 m)
MARSHALL 1986) Pseudococculina subcingulata (Kuroda et Habe, 1949)
Cocculinella minutissima (E. A. Smith, 1904) (Acmaea) (Cocculina) (Fig. 5, 6)
(northwestern Indian Ocean: Arabian Sea; on tele- (northwestern Pacific: off Tosa, Shikoku; Suruga
ost fish bones) (HASZPRUNAR 1988c) Bay, Honshu, Japan; 140–490; on wood)
Cocculinella osteophila Marshall, 1983 (HASEGAWA 1997)
(southwestern Pacific: Whangaroa Harbour, New Pseudococculina n. sp. (1) (MOSKALEV 1976)
Zealand; 13 m; on bone) Pseudococculina n. sp. (2) (MOSKALEV 1976)
Cocculinella salisburyensis Ludbrook, 1956 Pseudococculina n. sp. (3) (MOSKALEV 1976)
(Pliocene, South Australia) Pseudococculina n. sp. (4) (MOSKALEV 1976)
doubtfully referable to the genus, probably be- Pseudococculina n. sp. (5) (MOSKALEV 1976)
longs to Lepetellidae Dall, 1882(a) or to Acmaei-
dae Forbes, 1849 (MARSHALL 1983) Genus: Bandabyssia Moskalev, 1976 (type species:
Bandabyssia costoconcentrica Moskalev, 1976)
Family: OSTEOPELTIDAE Marshall, 1987 Bandabyssia costoconcentrica Moskalev, 1976Cocculiniformia, Checklist of species 57
(Pacific: Banda Trench; 5700 m) (northwestern Pacific: off Japan) (H ASEGAWA
1997, MARSHALL 1986)
Genus: Kaiparapelta Marshall, 1986 (type species: Notocrater pustulosa (Woodring, 1928 non Thiele,
Kaiparapelta singularis Marshall, 1986) 1925) (Cocculina)
(a subgenus in Notocrater Finlay, 1927 according to (Miocene, Jamaica) (MCLEAN & HARASEWYCH 1995)
HASZPRUNAR 1988ab) the species should probably receive a replacement
Kaiparapelta askewi McLean et Harasewych, 1995 name as the name pustulosa appears to be preoccu-
(northwestern Atlantic: 165 km E of Charleston, pied
South Carolina; 194 m) Notocrater youngi McLean et Harasewych, 1995
(spongivorous) (WARÉN & GOFAS 1996) (western Atlantic: off Southwest Reef, New Provi-
Kaiparapelta singularis Marshall, 1986 dence Island, Bahamas; 518 m; on palmetto
(early Miocene, Pakaurangi Point, Kaipara, New fronds)
Zealand)
Genus: Tentaoculus Moskalev, 1976
Genus: Kurilabyssia Moskalev, 1976 (type species: (type species: Tentaoculus perlucida Moskalev, 1976)
Kurilabyssia squamosa Moskalev, 1976) Kurilabyssia Tentaoculus balantiophaga Marshall, 1996
antipodensis Marshall, 1986 (southwestern Pacific: off Castlepoint and western
(southwestern Pacific: off New South Wales and Chatham Rise, New Zealand; 1065–1335 m; on
southern New Zealand; 823–962 m; on wood) spent skate egg cases)
(HASZPRUNAR 1988b) Tentaoculus eritmeta (Verrill, 1884)
Kurilabyssia squamosa Moskalev, 1976 (Puncturella /Fissurisepta/)
(northwestern Pacific: Kurile-Kamchatka Trench; (western Atlantic) (MCLEAN & HARASEWYCH 1995)
5220 m) Tentaoculus georgiana (Dall, 1927) (Cocculina)
(western Atlantic: off Georgia; 805 m) (MCLEAN &
HARASEWYCH 1995)
Genus: Mesopelex Marshall, 1986
Tentaoculus haptricola Marshall, 1986
(type species: Mesopelex zelandica Marshall, 1986)
(southwestern Pacific: off Castlepoint, New Zea-
Mesopelex zelandica Marshall, 1986
land; 1070–1200 m; on Macrocystis holdfast)
(southwestern Pacific: off Aldermen Islands, Bay
Tentaoculus lithodicola Marshall, 1986
of Plenty, New Zealand; 443–872 m)
(southwestern Pacific: off Kaikoura, New Zealand;
Mesopelex n. sp. (to be described by B. A. Marshall ac-
400 m; on dorsal surface of carapace of stone crab
cording to HASZPRUNAR 1988ab)
Lithodes murrayi Henderson, 1888)
(southwestern Pacific: off Timaru, New Zealand;
Tentaoculus neolithodicola Marshall, 1986
852–876 m)
(southwestern Pacific: off Jackson Bay, New Zea-
land; 1027–1041 m; on dorsal surface of carapace
Genus: Notocrater Finlay, 1927 of stone crab Neolithodes brodiei Dawson et Yaldwyn,
(type species: Cocculina craticulata Suter, 1908) 1970) (HASZPRUNAR 1988b)
= Punctolepeta Habe, 1958 Tentaoculus perlucida Moskalev, 1976
(type species: Punctolepeta minuta Habe, 1958) (western Pacific: off New Guinea; 300–450 m)
Notocrater craticulata (Suter, 1908) (Cocculina)
(southwestern Pacific: off Timaru and Dusky Subfamily: Caymanabyssiinae Marshall, 1986
Sound, New Zealand; 37–738 m; on wood)
(MARSHALL 1986) Genus: Caymanabyssia Moskalev, 1976
Notocrater gracilis Marshall, 1986 Subgenus: Caymanabyssia s. str. Moskalev, 1976
(southwestern Pacific: off Hicks Bay, New Zealand, (type species: Caymanabyssia spina Moskalev, 1976)
55–73 m; in terenid tubes in wood) Caymanabyssia (Caymanabyssia) rhina Marshall, 1986
Notocrater houbricki McLean et Harasewych, 1995 (southwestern Pacific: off White Island, New Zea-
(western Atlantic: south of Settlement Point, land; 1075–1100 m; on wood)
Grand Bahama Island, Bahama Islands; 412 m) Caymanabyssia (Caymanabyssia) spina Moskalev, 1976
Notocrater maxwelli Marshall, 1986 (western Atlantic: Cayman Trench; 6740–6800 m;
(early Miocene, Pakaurangi Point, Kaipara, New on wood) (MCLEAN & HARASEWYCH 1995)
Zealand) (western Atlantic: Puerto Rico Trench; 8595 m)
Notocrater ponderi Marshall, 1986 (LEAL 1996)
(southwestern Pacific: off New South Wales; Caymanabyssia (Caymanabyssia) vandoverae McLean,
165–84 m; on wood) (HASZPRUNAR 1988b) 1991
Notocrater pustulosa (Thiele, 1925) (Cocculina) (Fig. 7) (northeastern Pacific: Escanaba Trough, Gorda
= Punctolepeta minuta Habe, 1958 Ridge; 3362 m; on wood)58 Lesicki A.
Subgenus: Dictyabyssia McLean, 1991 (type species: (northwestern Pacific: Suruga Bay, Honshu, Ja-
Caymanabyssia sinespina Marshall, 1986) pan; 180–740 m; on wood)
Caymanabyssia (Dictyabyssia) fosteri McLean, 1991
o
(East Pacific Rise 11 N; 2700 m; on wood) Genus: Punctabyssia McLean, 1991
Caymanabyssia (Dictyabyssia) sinespina Marshall, 1986 (subfamiliar status unclear)
(southwestern Pacific: White Island to Jackson (type species: Punctabyssia tibbettsi McLean, 1991)
Bay, New Zealand; 800–1147 m) Punctabyssia tibbettsi McLean, 1991
(HASZPRUNAR 1988b) o
(East Pacific Rise 11 N; 2700 m; on wood)
Genus: Amphiplica Haszprunar, 1988(b) Genus: Yaquinabyssia Haszprunar, 1988(b)
Subgenus: Amphiplica s. str. Haszprunar, 1988(b) (type species: Yaquinabyssia careyi McLean, 1988)
(type species: Amphiplica venezuelensis McLean, 1988) Yaquinabyssia careyi McLean, 1988
Amphiplica (Amphiplica) concentrica (Thiele, 1909) (northeastern Pacific: Cascadia Abyssal Plain, 171
(Pseudococculina) km west of Cape Foulweather, Oregon; 2774 m)
(Atlantic: north of Azores) (MCLEAN 1988) (HASZPRUNAR 1988b)
Amphiplica (Amphiplica) knudseni McLean, 1988
(southwestern Pacific: Tasman Basin, New Zea- Family: PYROPELTIDAE McLean et Haszprunar,
land; 3610 m) (HASZPRUNAR 1988b) 1987
Amphiplica (Amphiplica) venezuelensis McLean, 1988
(western Atlantic: Venezuelan Basin; Genus: Pyropelta McLean et Haszprunar, 1987 (type
3476–5057 m) (HASZPRUNAR 1988b) species: Pyropelta musaica McLean et Haszprunar,
Amphiplica n. sp. (LEAL 1996) 1987)
(western Atlantic: Puerto Rico Trench; 8595 m)
Pyropelta bohlei Beck, 1996
Subgenus: Gordabyssia McLean, 1991 (type species:
(western Pacific: Edison Seamount, off Lihir Is-
Amphiplica (Gordabyssia) gordensis McLean, 1991)
land; 1483 m; hydrothermal vents, on shell of
Amphiplica (Gordabyssia) gordensis McLean, 1991
Bathyacmaea jonassoni Beck, 1996)
(northeastern Pacific: Escanaba Trough, Gorda
Pyropelta corymba McLean et Haszprunar, 1987
Ridge; 3305 m; on sulfide crust)
(northeastern Pacific: Guaymas Basin; 2022 m; hy-
drothermal vents)
Genus: Colotrachelus Marshall, 1986 (type species:
(northeastern Pacific: Santa Barbara Basin; 1240
Colotrachelus hestica Marshall, 1986)
m and Monterey Bay, off Point Sur, California; 940
Colotrachelus hestica Marshall, 1986
m; on whale bone or skull) (MCLEAN 1992a)
(southwestern Pacific: off Castlepoint and off
Pyropelta musaica McLean et Haszprunar, 1987
Kaikoura, New Zealand; 1174–1723 m; on wood)
(HASZPRUNAR 1988b) (northeastern Pacific: Axial Seamount, Juan de
Fuca Ridge; 1575 m; hydrothermal vents)
Genus: Copulabyssia Haszprunar, 1988(b) (northeastern Pacific; 940–1400 m; on whale
(type species: Cocculina corrugata Jeffeys, 1883) skulls) (MCLEAN 1992a)
Copulabyssia corrugata (Jeffreys, 1883/c/) (Cocculina) Pyropelta wakefieldi McLean, 1992(a)
(northeastern Atlantic; Tyrrhenian Sea; 100–1778 (northeastern Pacific: Monterey Bay, off Point Sur,
m; from submerged wood) California; 940 m; on whale skull)
(DI GERONIMO & BELLAGAMBA 1985, HASZPRUNAR
1988b, NORDSIECK 1973, WARÉN 1991) ACKNOWLEDGEMENTS:
Copulabyssia gradata (Marshall, 1986) (Pseudococcu- I would like to thank Dr. BRUCE MARSHALL (Museum Te
lina) Papa Tongarewa, Wellington NZ), Dr. JAMES H. MCLEAN
(southwestern Pacific: White Island and East Cape, (Museum of Natural History, Los Angeles) and Dr.
New Zealand; 1075–1280 m; on wood) ANDREW MCARTHUR (Smithsonian Institution, Washing-
(HASZPRUNAR 1988b) ton) for their kind remarks that helped me in the prepara-
Copulabyssia leptalea (Verrill, 1884) (Cocculina) tion of this list. Dr. KAZUNORI HASEGAWA (National Science
(northern Atlantic: Iceland, north of the Hebri- Museum, Tsukuba) kindly supplied me with the photo-
des; from wood bored by shipworms) (MCLEAN & graphs of cocculiniform gastropods. I am also indebted to
HARASEWYCH 1995) Mrs. JOLANTA MAÆKOWIAK for her help in searching for
Copulabyssia similaris Hasegawa, 1997 (Fig. 8) bibliographical sources in libraries.Cocculiniformia, Checklist of species 59
REFERENCES
ALLEN J. E. 1970. New species of Eocene Mollusca from Gulf DALL W. H. 1890. Scientific results of explorations by the
coast. Tulane Stud. Geol. Paleont. 8: 69–78. U.S. Fish Commission steamer ”Albatross”, 7: Prelimi-
BECK L.A. 1996. Morphology and anatomy of new species of nary report of the collection of Mollusca and Brachio-
neolepetopsid, acmaeid, fissurellid and pyropeltid lim- poda obtained in 1887–1888. Proc. U. S. natn. Mus. 12:
pets from Edison Seamount off Lihir Islands (West Pa- 219–362.
cific) (Gastropoda: Prosobranchia: Neolepetopsidae, Ac- DALL W. H. 1896. Diagnoses of new species of mollusks from
maeidae, Fissurellidae, Pyropeltidae). Arch. Moll. 125 the west coast of America. Proc. U. S. natn. Mus. 18
(1/2): 87–103. (1034): 7–20.
BIELER R. 1992. Gastropod phylogeny and systematics. DALL W. H. 1908. Reports on the dredging operations off
Annu. Rev. Ecol. Syst. 23: 311–338. the west coast of Central America to the Galapagos, to
BOGI C. 1984. Cocculina labronica n. sp. Boll. Malacologico 20 the west coast of Mexico, and in the Gulf of California...
(5–8): 155–160. XIV. The Mollusca and Brachiopoda. Bull. Mus. comp.
BOUCHET P., WARÉN A. 1979. The abyssal molluscan fauna of Zool. Harv. Univ. 43: 205–487.
the Norwegian Sea and its relation to other faunas. Sar- DALL W. H. 1919. Descriptions of new species of Mollusca
sia 64: 211–243. from the north Pacific Ocean in the collection of the
BUSH K. J. 1897. Revision of the marine gastropods referred United States National Museum. Proc. U. S. natn. Mus.
to Cyclostrema, Adeaorbis, Vitrinella and related genera: 56: 293–371.
with description of some new genera and species belong- DALL W. H. 1921. Summary of the marine shell bearing mol-
ing to the Atlantic fauna of America. Trans. Connecticut lusks of the northwest coast of America, from San Diego
Acad. Sci., New Haven 10: 97–143. to Polar Sea. U. S. Nat. Mus. Bull. 112: 1–217.
CHILDRESS J. J., FISHER C. R. 1992. The biology of hydrother- DALL W. H. 1925. Illustrations of unfigured types of shells in
mal vent animals: physiology, biochemistry, and auto- the collections of the United States National Museum.
trophic symbioses. Oceanogr. Mar. Biol. Annu. Rev. 30: Proc. U. S. natn. Mus. 66(17): 1–41.
337–441.
DALL W. H. 1927. Small shells from dredgings off the south-
CLARK W. 1851. On the classification of the British marine east coast of the United States by the United States Fish-
testaceous Mollusca. Ann. Mag. Nat. Hist. s. 2, 7: eries steamer “Albatross” in 1885 and 1886. Proc. U. S.
469–482. natn. Mus. 70: 1–134.
CLARKE A. H. JR. 1960. A giant ultraabyssal Cocculina (C. su- DALL W. H., SIMPSON T. 1901. The Mollusca of Porto Rico.
perba n. sp.) from the Argentina basin. Natur. History Pa- United States Fish Commission Bulletin for 1900, vol. 1,
per, Nat. Mus. Canada 7: 1–4. 351–524.
CLARKE A. H. JR. 1961. Abyssal mollusks from the South At-
DANTART L., LUQUE A. 1994. Cocculiniformia and Lepeti-
lantic Ocean. Bull. Mus. Comp. Zool. Harv. Univ. 125
dae (Gastropoda: Archaeogastropoda) from Iberian wa-
(12): 343–387.
ters. J. Moll. Stud. 60: 277–313.
CROZIER M. A. 1966. New species and records of Mollusca
from off Three Kings Islands, New Zealand. Trans. R. D AUTZENBERG P. 1886. Note sur l’Addisonia lateralis
Soc. N. Zealand 8: 39–49. Requien. J. Conchyl. 34: 203–208.
DALL W. H. 1869. Materials for the monograph of the family DAUTZENBERG P. 1889. Contribution a la faune malacolo-
Lepetidae. Am. J. Conch. 5: 140–150. gique des iles Açores. Révision des mollusques marins
des Açores. Resultats des Campagnes Scientifiques ac-
DALL W. H. 1882a. On certain limpets and chitons from the complies sur son yacht par le Prince Albert I. 1: 1–113.
deep waters off the eastern coast of the United States.
Proc. U. S. natn. Mus. 4: 400–414. DAUTZENBERG P. 1925. Mollusques nouveaux provenant des
croisiéres du Prince Albert 1er de Monaco. Bull. Inst.
DALL W. H. 1882b. On certain limpets and chitons from the
Océanogr. Monaco 457: 1–12.
deep waters off the eastern coast of the United States.
Ann. Mag. Nat. Hist. ser. 5, 10: 11–26. DAUTZENBERG P., FISCHER H. 1897. Dragages effectues par l'
DALL W. H. 1882c. Note on Gadinia excentrica Tiberi. Amer. “Hirondelle” et par la “Princesse Alice”. Mem. Soc. Zool.
Naturalist: 737. France 10: 139–234.
DALL W. H. 1889a. Reports on the results of dredging under DAUTZENBERG P., FISCHER H. 1899. Description d'un mol-
the supervision of Alexander Agassiz, in the Gulf of Mex- lusque nouveau. Bull. Soc. Zool. Frances 24: 207–209.
ico (1877–78) and the Caribbean Sea (1879–80), by the DAWSON J. W. 1872. The post-Pliocene geology of Canada.
U.S. coast survey steamer “Blake“, Lieut. Commander Can. Naturalist Quart. J. Science NS 6 (3): 369–416.
C.D. Sigsbee, U.S.N., and Commander J.R. Bartlett,
DELL R. K. 1953. A molluscan fauna from the Chatham Rice,
U.S.N., commanding. Bull. Mus. comp. Zool. Harv. Coll.
New Zealand. Rec. Domin. Mus. 2.
18: 1–492.
DALL W. H. 1889b. A preliminary catalogue of the shell- DELL R. K. 1956. The archibenthal mollusca of New Zea-
bearing marine mollusks and brachiopods of the south- land. Dominion Mus. Bull. 18: 1–235.
eastern coast of the United States, with illustrations of DELL R. K. 1990. Antarctic Mollusca, with special reference
many species. U. S. Nat. Mus. Bull. 37: 1–221. to the fauna of the Ross Sea. R. Soc. N. Z. Bull. 27: 1–311.60 Lesicki A.
DI GERONIMO I. 1974. Molluschi bentonici in sedimento re- HASZPRUNAR G. 1992b. On the anatomy and relationships of
centi batiale e abissale dello Jonio. Conchiglie 10: the Choristellidae (Archaeogastropoda: Lepetelloidea).
133–172. Veliger 35 (4): 295–307.
DI GERONIMO I., BELLAGAMBA M. 1985. Malacofaune dei dra- HASZPRUNAR G., MCLEAN J. H. 1996. Anatomy and system-
gaggi BS 77–1 e BS 77–2 (Sardegna Nord Orientale). atics on bathyphytophilid limpets (Mollusca, Archaeo-
Boll. Soc. Paleontol. Ital. 24: 111–129. gastropoda) from the northeastern Pacific. Zool. Scripta
25 (1): 35–49.
FINLAY H. J. 1927. A further commentary on New Zealand
molluscan systematics. Trans. New Zealand Inst. 57: HEDLEY C. 1903. Scientific results of the trawling expedition
320–485. of H.M.C.S. Thetis off the coast of New South Wales. 6.
Mollusca 2. Austr. Mus. Sydney Mem. 4: 327–402.
FLEMING C. A. 1948. New species and genera of marine Mol-
HEDLEY C. 1907. The results of deep sea investigations in the
lusca from the Southland fiords. Trans. R. Soc. N. Z. 77:
Tasman Sea. 3. Mollusca from eighty fanthoms off Narra-
72–92.
been. Rec. Austr. Mus. 6(4): 283–304.
FLEMING C. A. 1966. Marwick's illustrations of New Zealand HICKMAN R. S. 1983. Radula patterns, systematics, diversity
shells, with a checklist of New Zealand Cenozoic mol- and ecology of deep-sea limpets. Veliger 26(2): 73–92.
lusca. Bull. N. Z. Dept. Sci. Industr. Res. 173:
INABA T., OYAMA K. 1977. Catalogue of molluscan taxa de-
FORBES E. 1840. On some new and rare British Mollusca. scribed by Tadashige Habe during 1939–1975, with illus-
Ann. Nat. Hist. 5: 102–108. trations of hitherto unfigured species (for commemor-
FORBES E. 1849. On the genera of British Patellacea. The ation of his sixtieth birthday). 185 pp. Okinaebisu-no-
Athenaeum 1145: 1018. kai. Tokyo.
FORBES E., HANLEY S. 1849. A history of British Mollusca and JANSSEN R. 1989. Benthos-Mollusken aus dem Tiefwasser des
their shells. 2. 557 pp. J. Van Voorst, London. östlichen Mittelmeeres, gesammelt während der “Me-
teor” - Fahrt 5 (1987). Seckenbergiana Maritima 20:
GMELIN J. F. 1791. Systema Naturae. Editio decima tertia. 265–276.
Vol. 1, part 6. pp. 3021–3910. G. Beer, Lipsiae.
JEFFREYS J. G. 1883a. On the mollusca procured during the
GOODAY A. J., TURLEY C. M. 1990. Responses by benthic or- “Lightning” and “Porcupine” expeditions 1868–70. Part
ganisms to inputs of organic material to the ocean floor: V. Proc. Zool. Soc. London 1882: 656–687.
a review. Phil. Trans. R. Soc. Lond. A 331: 119–138. JEFFREYS J. G. 1883b. On the mollusca procured during the
GUBBIOLI F., NOFRONI I. 1986. Prima segnalazione di Cintha “Lightning” and “Porcupine” expeditions 1868–70. Part
naticiformis Jeffreys, 1883 in Mar Mediterraneo: note VI. Proc. Zool. Soc. London 1883: 88–115.
ecologiche e sistematiche. La Conchiglia 18: 6–7. JEFFREYS J. G. 1883c. On the mollusca procured during the
HABE T. 1952. Pseudococculina rosea, new species (Cocculini- cruise of H.M.S. Triton, between the Hebrides and
dae). Ill. Cat. Jap. Shells (20): 152, pl. 30. Faroes in 1882. Proc. Zool. Soc. London 1883: 389–399.
HABE T. 1958. Descriptions of ten new gastropod species. KNUDSEN J. 1964. Scaphopoda and Gastropoda from depths
Venus 20(1): 32–42. exceeding 6000 meters. Galathea Rep. 7: 125–136.
KURODA T., HABE T. 1949. On the gastropod genus Coccu-
HASEGAWA K. 1997. Sunken wood-associated gastropods col-
lina from Japan. Venus (Jap. J. Malac.) 15(1/4): 58–67.
lected from Suruga Bay, Pacific side of the Central Hon-
shu, Japan, with descriptions of 12 new species. Nat. Sci. KURODA T., HABE T., OYAMA K. 1971. The sea shells of Sa-
Mus. Monogr. 12: 59–123. gami Bay. 489 pp., Tokyo.
HASZPRUNAR G. 1987a. Anatomy and affinities of cocculinid LEAL J. H. 1996. How deep is deep? or museum-hunting for
limpets (Mollusca, Archaeogastropoda). Zool. Scripta 16 the deepest Atlantic mollusks. Am. Conchologist 24(4):
(4): 305–324. 10–11.
LESICKI A. 1998. Molluscs in ecosystems fuelled by chemoau-
HASZPRUNAR G. 1987b. The anatomy of Addisonia (Mol-
totrophy (Fauna of ecosystems connected with chemo-
lusca, Gastropoda). Zoomorphology 106: 269–278.
autotrophy, part 3). (in Polish). Przegl. zool. 42(1/2):
HASZPRUNAR G. 1988a. Comparative anatomy of cocculini- 5–32.
form gastropods and its bearing on archaeogastropod LINDBERG R., PONDER W. H. 1991. A phylogenetic analysis of
systematics. Malacol. Rev. Suppl. 4: 64–84. the Gastropoda. Geol. Soc. Am., Abstr.: A160–A161.
HASZPRUNAR G. 1988b. Anatomy and affinities of pseudo- LOCARD A. 1886. Prodrome de Malacolgie française. Cata-
cocculinid limpets (Mollusca: Archaeogastropoda). logue général des Mollusques vivants de France. Mol-
Zool. Scripta 17 (2): 161–180. lusques marines. H. Georg, Lyon and J. B. Baillière,
HASZPRUNAR G. 1988c. Anatomy and relationships of the Paris.
bone-feeding limpets, Cocculinella minutissima (Smith) LOCARD A. 1897. Mollusques Testaces 2. In: Expeditions Sci-
and Osteopelta mirabilis Marshall (Archaeogastropoda). entifiques du Travailleur et du Talisman pendant les
J. Moll. Stud. 54: 1–20. années 1880, 1881, 1882, 1883. 4: 1–515.
HASZPRUNAR G. 1992a. Anatomy and relationships of the LUDBROOK N. H. 1956. The molluscan fauna of the Pliocene
Choristellidae (Archaeogastropoda, Lepetelloidea). strata underlying the Adelaide Plains. Part III. Scapho-
Proc. Tenth Intern. Malacol. Congress (Tübingen 1989): poda, Polyplacophora, Gastropoda (Haliotidae to Torni-
509–510. dae). Trans. R. Soc. South Austr. 79: 1–36.You can also read
