The superfamily Cavolinioidea is a taxonomic group of small floating sea snails, pelagic marine opisthobranch gastropod mollusks.

The Cavoliniodea are part of a larger group which is commonly known as the sea butterflies because they swim by flapping what appear to be small "wings".

This superfamily contains the greatest number of different species of sea butterflies.


These sea butterflies are circumglobal, carried by the sea currents to all the seas of the world.


Cavoliniids prefer deep waters, from 100 m up to 2,000 m. They do best in warm oceanic water.

Shell description

Species in this superfamily have a calcareous, bilaterally symmetrical conical or globular shell with a completely different form than those in the family Limacinidae. This shell is twisted in a flat manner or slightly curved.

Identification of juveniles is difficult as the juvenile shells differ greatly from adults.

Life habits

Towards the anterior end of the animal, two parapodia (winglike flat lobules) protrude between each half of the shell. The parapodia enable these sea butterflies to float along in the water currents, using slow flapping movements. The parapodia are also covered with cilia, which produce a minute water current that pushes the planktonic food to the mouth of the animal.


Cavoliniids have a strange sexual life. They develop from males as juveniles into hermaphrodites and then later convert into females. More than one male stage can occur. This bizarre-seeming, but not very uncommon phenomenon is called protandry. (This is also common among many species of fish, some may all start as females, others species may start as males.)


Recent changes

In 2003, the family Cavoliniidae was raised to the rank of superfamily Cavolinioidea. At the same time, the subfamilies were given the new status of families : Cavoliniidae, Cliidae, Creseidae and Cuvierinidae (Cainozoic Research, 2(1-2): 163-170, 2003). In 2006 a new family Praecuvierinidae was created on evolutionary grounds. In the new taxonomy of Bouchet & Rocroi (2005) several families have been categorized as subfamilies of the family Cavoliniidae.

Family Cavoliniidae

Genus Cavolinia Abildgaard, 1791

A very distinctive shape of shell with a marked bulge on the ventral plate. The species are protandric hermaphrodites.

Genus Diacavolinia van der Spoel, 1987

Twenty two species of Diacavolinia. Diacavolinia species are characterised by the absence of a caudal spine

  • Diacavolina angulosa J.E. Gray, 1850
    • Distribution : Indo-Pacific, Atlantic.
    • Length : 4 mm
  • Diacavolinia bicornis van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Indo-Pacific, Atlatic Ocean.
    • Length : 8 mm
  • Diacavolina constricta van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Bermuda, Venezuela.
  • Diacavolinia deblainvillei van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Caribbean, Western Atlantic
    • Length : 7 mm
  • Diacavolinia deshayesi van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution ; Bermuda, Panama, French Guyana
    • Length : 8 mm
  • Diacavolinia elegans van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : New Yersey
    • Length : 6 mm
  • Diacavolinia flexipes van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : SE Asia
    • Length : 5 mm
  • Diacavolinia limbata van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Brazil, southern Indo-Pacific
    • Length : 13 mm
  • Diacavolinia longirostris (de Blainville, 1821) Long-snout Cavoline
    • Distribution : circumglobal, Red Sea, Madagascar, West Pacific, Australian; Gulf of Mexico
    • Length : 7 mm; width : 4.9 to 6.8 mm
    • Description : globulous brownish shell with two distinct lateral spines and a long rostrum on the dorsal rim. Dorsal side of the shell is relatively flat whereas the ventral side is deeply rounded.
  • Diacavolinia mcgowani van der Spoel, Bleeker and Kobayashi, 1993
  • Diacavolinia ovalis van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Caribbean, West Atlantic
    • Length : 6 mm
  • Diacavolinia robusta van der Spoel, Bleeker and Kobayashi, 1993
    • Distribution : Caribbean, West Atlantic
    • Length : 5.4 mm
  • Diacavolinia strangulata (G.P. Deshayes, 1823)
    • Distribution : Panama, Brazil, Cuba
    • Length : 4 mm

Genus Diacria J. E. Gray, 1847

The genus comprises two species groups and a total of ten species. The species may be globular, with both dorsal and ventral sides rounded, or bilaterally symmetrical with a long caudal spine. The species are protandric hermaphrodites. They are the largest of the Cavoliniids.

  • Diacria atlantica L. Dupont, 1979
    • Distribution : Massachusetts
    • Length : 9 mm
  • Diacria costata G. Pfeffer, 1879
    • Distribution: Indo-Pacific
  • Diacria danae van Leyen and van der Spoel, 1982
    • Distribution : circumglobal in warm seas
    • Length : 9 mm
  • Diacria maculata Bleeker and van der Spoel, 1988
  • Diacria major (Boas, 1886)
    • Distribution : Florida, Bermuda, Atlantic Ocean, Indian Ocean.
    • Length : 13 mm
  • Diacria quadridentata (de Blainville, 1821) -- Four-tooth Cavoline
    • Distribution : circumglobal, Red Sea, Madagascar, Gulf of Mexico, Japan
    • Length : 3 mm; width : 1.8 to 2.5 mm
    • Description : a small, globular shell, with curved spinal and ventral sides. There are no caudal or lateral spines. The dorsal side extends further than the ventral side.
    • Subspecies:
  • Diacria rampali Dupont, 1979
    • Distribution : Florida, Brazil
    • Length: 10 mm
  • Diacria rubecula Bontes & van der Spoel, 1998
    • Distribution : warmer regions of North Atlantic
    • Length: 11 mm
  • Diacria schmidti Leyen & van der Spoel, 1982
    • Distribution : Pacific
  • Diacria trispinosa (de Blainville, 1821) -- Three-spine Cavoline
    • Distribution: circumglobal, Gulf of Mexico, Madagascar.
    • Length: 13 mm; width : 10 mm
    • Description: The slightly transparent, brownish shell is bilaterally symmetrical and is darker on the ribbed sections. Very long caudal spine and strong lateral spines. There are five ribs on the dorsal side and three ribs on the ventral side.

Family Clioidae Jeffreys, 1869

This family has been named for a long time Clioidae and Cleodoridae with type genus Clio, confusingly the same name as another molluscan family Clionidae with type genus Clione. The International Commission on Zoological Nomenclature (ICZN) has therefore changed the first name into the original spelling Cliidae Jeffreys, 1869, type genus Clio Linnaeus, 1767

Genus Clio Linnaeus, 1767

All species in this genus are characterised by a bilaterally symmetric, straight or adapically dorso-ventrally slightly curved shell, with an elliptical to triangular transverse section; protoconch clearly separated, globular or elliptical, frequently with a spine at the tip. Subgenera are used for some species (e.g. Clio s.str., Balantium Bellardi, 1872, Bellardiclio Janssen, 2004), but most species cannot yet be assigned to one of these. Numerous fossil species have been described. Recognised extant species are:


  • forma excisa van der Spoel, 1963
  • forma lanceolata Lesueur, 1813 (tropical/subtropical, circumglobal)
  • forma martensi (Pfeiffer, 1880) ??
  • forma sulcata (Pfeffer, 1879)

The true status of these formae has to be evaluated, they might be real formae, subspecies, or even species.

  • Clio recurva (Children, 1823)
    • Distribution : tropical-subtropical, circumglobal (bathypelagic species)**
      • Height: to over 30 mm

Family Creseidae Curry, 1982

Genus Creseis Rang, 1828

The shells of the species in this genus have the form of a more or less narrow, conically widening tube.

  • Creseis chierchiae Boas, 1886
    • Distribution : tropical-subtropical, circumglobal.
    • Length: shell up to 9 mm high.
    • Description: shell straight or slightly curved, initially slowly, later hardly increasing in diameter, with clear annulations, protoconch with rounded tip, followed by a distinct swelling. A form with lacking annulations, also known as a Pliocene fossil, is described as C. chierchiae forma constricta Chen & Bé, 1964.
  • Creseis clava (Rang, 1828) (synonym: Creseis acicula (Rang, 1828)
    • Distribution : tropical-subtropical, circumglobal.
    • Length: shell is up to 35 mm high.
    • Description : shell, long and straight or slightly irregular, with small apical angle, circular in cross section; smooth shell surface; protoconch without swelling; there is a characteristic tentacular lobe on the
  • Creseis conica Eschscholtz, 1825
    • Distribution : tropical-subtropical, circumglobal.
    • Length: shell height up to 20 mm.
    • Description : shell straight or slightly curved, with a wider apical angle than C. clava, transverse section circular, protoconch slightly swollen.
  • Creseis virgula (Rang, 1828)
    • Distribution : tropical-subtropical, circumglobal.
    • Length: shell height up to 12 mm
    • Description : shell with circular transverse section, curved in its basal part.

Genus Hyalocylis Fol, 1875

  • Hyalocylis striata (Rang, 1828)
    • Distribution : tropical-suntropical, circumglobal.
    • Length : 10 mm
    • Description : shell slightly curved dorsally, with distinct annulations, transverse section initially circular, later slightly dorso-ventrally compressed. The animal is often easily recognised by the very large fins.

Genus Styliola Gray, 1850

  • Styliola subula (Quoy & Gaimard, 1827)
    • Distribution : tropical-suntropical, circumglobal, absent in the Red Sea.
    • Length : 13 mm
    • Description : needle-like shell, transparent and round in cross-section. A prominent oblique dorsal furrow runs from a short distance above the protoconch to the aperture, building a toothlike process. There is no tentacular lobe on the anterior margin of the fins.

Family Cuvierinidae

Extinct genera:

Extant genera:

The genus Cuvierina developed from the Ireneia lineage during the Early Miocene and splits up in two subgenera:

  • Subgenus Cuvierina s. str.

Extant species:

Extinct species:

  • Subgenus Urceolarica Janssen, 2006
  • Extant species:

    Extinct species:

    Family Praecuvierinidae

    Janssen, 2005 (extinct genera)

    Bouchet & Rocroi

    In the new taxonomy of Bouchet & Rocroi (2005) the superfamily Cavolinioidea is treated differently :

    • Family Cavoliniidae Gray, 1850 (1815)
      • Subfamily Cavoliinae Gray, 1850 (1815) (formerly Hyalaeidae Rafinesque, 1815 )
      • Subfamily Clioinae Jeffreys, 1869 (formerly Cleodoridae Gray, 1840 - nomen oblitum)
      • Subfamily Cuvierininae van der Spoel, 1967 (formerly : Cuvieriidae Gray, 1840 (nom. inv.); Tripteridae Gray, 1850 )
      • Subfamily Creseinae Curry, 1982
    • Family Limacinidae Gray, 1840 (formerly : Spirialidae Chenu, 1859 ; Spiratellidae Dall, 1921 )
    • † Family Sphaerocinidae A. Janssen & Maxwell, 1995



    • Rampal, J., 2002. "Biodiversité et biogéographie chez les Cavoliniidae (Mollusca, Gastropoda, Opisthobranchia, Euthecosomata). Régions faunistiques marines" Zoosystema, 24(2):209-258.
    • Janssen, A.W., 2003. Notes on the systematics, morphology and biostratigraphy of fossil holoplanktonic Mollusca, 13. "Considerations on a subdivision of Thecosomata, with the emphasis on genus group classification of Limacinidae" Cainozoic Research, 2(1-2): 163-170.
    • Janssen, A.W., 2005. Development of Cuvierinidae (Mollusca, Euthecosomata, Cavolinioidea) during the Cainozoic: a non-cladistic approach with a re-interpretation of Recent taxa
    • Janssen, A.W., 2006. Notes on the systematics, morphology and biostratigraphy of fossil holoplanktonic Mollusca, 16. Some additional notes and amendments on Cuvierinidae and on classification of Thecosomata (Mollusca, Euthecosomata. -- Basteria, 70(1-3): 67-70.

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