Elephant snail Scutus ambiguus, copyright Leon Altoff.

Belongs within: Vetigastropoda.
Contains: Diodora, Fissurella, Puncturella, Tugali, Emarginula.

The Fissurellidae, keyhole and slit limpets, are a group of limpet-like gastropods in which the shell bears a slit or perforation for the exhalant current from the gills. Members of the Diodorinae and Fissurellinae have an apical aperture on the shell bordered by a callus whereas Hemitoma and Emarginulinae have a more-or-less developed longitudinal slit running back from the front of the shell. In some cases the slit is only developed on the shell ventrally with only an anterior emargination or notch visible dorsally. Diodorinae and Fissurellinae may be distinguished by the shape of the callus around the aperture which is rounded in Fissurellinae but posteriorly truncate in Diodorinae. Members of the genus Scutus have a depressed shell that in life is largely covered by the expanded mantle. In Hemitoma, the shell may be moderately elevated but not conical, with a distinct internal groove and a short slit or notch anteriorly (Knight et al. 1960).

Characters (from Knight et al. 1960): Shell conical, porcellaneous; protoconch spiral; with perforation, slit, notch or emargination for passage of exhalant current; muscle scar horseshoe-shaped, open anteriorly.

<==Fissurellidae [Cervicobranchiata, Fissonellidae, Fissurellacea, Fissurelladae, Fissurelloidea, Larvadae, Patelloideae]
    |  i. s.: Leurolepas roseola McLean 1970 PP78
    |         Incisura Hedley 1904 P61
    |           `--*I. lytteltonensis (Smith 1894) P61 [=Scissurella lyttletonensis KC60]
    |         Laevinesta atlantica M84
    |--Hemitoma Swainson 1840 BR05 (see below for synonymy)
    |    |  i. s.: H. bella (Gabb 1865) [=Emarginula bella] O27
    |    |         ‘Emarginula’ clathrata Ad. & Reeve 1850 [=Subemarginula clathrata] H09
    |    |         ‘Emarginula’ cumingii Sowerby 1863 [=Subemarginula cumingii] H09
    |    |         H. golischae Dall 1916 O27
    |    |         H. yatesii Dall 1901 O27
    |    |--H. (Hemitoma) octoradiata (Gmelin 1791) KC60 (see below for synonymy)
    |    |--H. (Montfortia Récluz 1843) KC60
    |    |    `--H. (*M.) australis (Quoy & Gaimard 1834) [=Emarginula australis] KC60
    |    `--H. (Montfortista Iredale 1929) KC60
    |         `--H. (*M.) excentrica (Iredale 1929) [=*Montfortista excentrica] KC60
    |--Diodorinae [Diodorini] BR17
    |    |--Diodora BR05
    |    |--Megathura Pilsbry 1890 [=Macrochasma Dall 1915] KC60
    |    |    `--M. crenulata (Sowerby 1825) KC60 (see below for synonymy)
    |    |--Pseudofissurella Haber 1932 KC60
    |    |    `--*P. corallensis (Buvignier 1852) [=Fissurella corallensis] KC60
    |    |--Stromboli Berry 1954 KC60
    |    |    `--*S. beebei (Hertlein & Strong 1951) [=Fissurella beebei] KC60
    |    `--Monodilepas Finlay 1927 KC60
    |         |--*M. monilifera (Hutton 1873) P61 [=Lucapina monilifera P61, Fissuridea monilifera F27]
    |         |    |--M. m. monilifera P61
    |         |    `--M. m. cookiana Dell 1952 P61
    |         |--M. carnleyensis Powell 1955 P61
    |         |--M. diemenensis Finlay 1930 P61
    |         |--M. otagoensis Finlay 1930 P61
    |         `--M. skinneri Finlay 1928 P61
    |--Fissurellinae BR05
    |    |--Fissurella S11
    |    |--Atractotrema Cossmann 1888 KC60
    |    |    `--*A. grata (Deshayes 1861) [=Fissurella grata] KC60
    |    |--Cosmetalepas Iredale 1924 [incl. Profissurellidea Wenz 1938] KC60
    |    |    `--*C. concatenata (Crosse & Fischer 1864) KC60 [=Fissurella concatenata KC60, Fissuridea concatenata F27]
    |    |--Megatebennus Pilsbry 1890 KC60
    |    |    `--*M. bimaculatus (Dall 1872) [=Fissurellidea bimaculata] KC60
    |    |--Lucapina Gray ex Sowerby 1835 [incl. Chlamydoglyphis Pilsbry 1890, Foraminella Guilding ex Sowerby 1835] KC60
    |    |    |--L. sowerbii (Guilding ex Sowerby 1835) (see below for synonymy) KC60
    |    |    `--L. elisae Costa & Simone 2006 S11
    |    |--Amblychilepas Pilsbry 1890 [incl. Sophismalepas Iredale 1924] KC60
    |    |    |--A. javanicensis (Lamarck 1822) KC60 (see below for synonymy)
    |    |    |--‘Fissuridea’ nigrita [=*Sophismalepas nigrita] F27
    |    |    `--‘Sophismalepas’ oblonga M54
    |    |--Macroschisma Gray ex Sowerby 1839 [=Machrochisma Swainson 1840, Macrochisma Gray 1840] KC60
    |    |    |  i. s.: M. madreporaria Hedley 1907 H09
    |    |    |--*M. (Macroschisma) macroschisma (Humphrey ex Solander 1786) [=Patella macroschisma] KC60
    |    |    |--M. (Dolichoschisma Iredale 1940) KC60
    |    |    |    `--M. (*D.) producta Adams 1854 KC60
    |    |    `--M. (Forolepas Iredale 1940) KC60
    |    |         `--M. (*F.) tasmaniae Sowerby 1862 KC60
    |    `--Lucapinella Pilsbry 1890 KC60
    |         |--*L. callomarginata (Carpenter ex Dall 1872) [=Clypidella callomarginata] KC60
    |         |--L. gigas Usticke 1959 BC01
    |         |    |--L. g. gigas BC01
    |         |    `--L. g. oblonga Usticke 1959 BC01
    |         |--L. limatula BC01
    |         |    |--L. l. limatula BC01
    |         |    |--L. l. delicata Usticke 1969 BC01
    |         |    `--L. l. hassleri Pérez Farfante 1943 BC01
    |         `--L. nigrita (Sowerby 1834) [=Fissurella nigrita] H09
    `--Emarginulinae [Emarginulidae, Emarginulini] BR05
         |--Puncturella KC60
         |--Tugali KC60
         |--Emarginula BR05
         |--Emarginella Pilsbry 1891 KC60
         |    `--*E. cuvieri (Audouin 1826) [=Emarginula cuvieri] KC60
         |--Emersonia Haber 1932 KC60
         |    `--*E. costata (Emerson 1870) [=Cemoria costata] KC60
         |--Loxotoma Fischer 1885 KC60
         |    `--*L. neocomiensis (d’Orbigny 1843) [=Emarginula neocomiensis] KC60
         |--Vacerrena Iredale 1958 P61 [=Vacerra Iredale 1924 non Godman 1900 KC60]
         |    `--*V. demissa (Hedley 1904) [=Puncturella demissa] P61
         |--Clypidina Gray 1847 [Clypidinidae, Clypidinoidei] BR05
         |    `--*C. notata (Linnaeus 1758) [=Patella notata] BR17
         |--Rimulopsis Haber 1932 KC60
         |    |--*R. (Rimulopsis) goldfussi (Roemer 1836) [=Emarginula goldfussi] KC60
         |    `--R. (Koniakaua Haber ex Dacqué 1933) KC60
         |         `--R. (*K.) multistriata (Zittel 1873) [=Rimula multistriata] KC60
         |--Fissurellidea d’Orbigny 1839 [Fissurellideinae, Fissurellideini, Fissurellidinae] BR05
         |    |--F. (Fissurellidea) hiantula (Lamarck 1822) (see below for synonymy) KC60
         |    `--F. (Pupillaea Sowerby 1835) [=Papillaea (l. c.), Pupilia (l. c.)] KC60
         |         `--F. (*P.) aperta (Sowerby 1825) [=Fissurella aperta] KC60
         |--Rimula Defrance 1827 BR05 [=Rimularia Waldheim 1834 KC60; Rimulacea, Rimulidae]
         |    |--*R. blainvillii (Defrance 1827) BR17 [=Emarginula blainvillii BR17, *Rimularia blainvilli KC60]
         |    |--R. exquisita Adams 1853 H09
         |    |--R. mazatlanica C64
         |    `--R. mexicana Berry 1969 PP78
         |--Austriacopsis Haber 1932 KC60
         |    |--*A. (Austriacopsis) austriaca (Hörnes 1853) [=Rimula austriaca] KC60
         |    |--A. (Balinula Haber ex Dacqué 1933) KC60
         |    |    `--A. (*B.) triontina (Greco 1899) [=Emarginula triontina] KC60
         |    `--A. (Puncturellopsis Haber 1932) KC60
         |         `--A. (*P.) acuta (Eudes-Deslongchamps 1842) [=Fissurella acuta] KC60
         |--Zeidora Adams 1860 BR05 (see below for synonymy)
         |    |  i. s.: Z. lodderae Tate & May 1900 H09
         |    |         Z. maoria Powell 1937 P61
         |    |--*Z. (Zeidora) calceolina Adams 1860 P61
         |    `--Z. (Nesta Adams 1870) KC60
         |         `--Z. (*N.) candida (Adams 1870) [=*Nesta candida] KC60
         |--Montfortula Iredale 1915 [incl. Plagiorhytis Fischer 1885 non Chaudoir 1848] KC60
         |    |--*M. rugosa (Quoy & Gaimard 1834) KC60 (see below for synonymy)
         |    |--M. chathamensis Finlay 1928 P61
         |    |--M. conoidea (Reeve 1842) [=Hemitoma conoidea] F27
         |    |--M. kaawaensis (Bartrum 1919) [=Tugalia kaawaensis] F27
         |    |--M. lyallensis Mestayer 1928 P61
         |    |--M. pulchra P95
         |    `--M. variegata Adams 1852 W78
         `--Scutus Montfort 1810 BR05 (see below for synonymy)
              |  i. s.: S. anatinus (Donovan 1820) (see below for synonymy) S79
              |         S. breviculus (Blainville 1817) P61 [=Parmophorus breviculus S79, Emarginula brevicula S79]
              |         S. elongatus [=Patella elongata,; incl. Parmophorus laevis] S79
              |         ‘Parmophorus’ fragilis S79
              |         ‘Parmophorus’ fissurella S79
              |         ‘Parmophorus’ gibbosus S79
              |         ‘Parmophorus’ sinensis S79
              |         S. unguis (Linnaeus 1758) H09 (see below for synonymy)
              |--S. (Scutus) ambiguus (Chemnitz 1795) F27 (see below for synonymy)
              |--S. (Nannoscutum Iredale 1937) KC60
              |    `--S. (*N.) forsythi (Iredale 1937) [=*Nannoscutum forsythi] KC60
              `--S. (Proscutum Fischer 1885) KC60
                   `--S. (*P.) compressus (Deshayes 1861) [=Parmophorus compressus] KC60

Amblychilepas javanicensis (Lamarck 1822) KC60 [=Fissurella javanicensis KC60, Fissuridea javanicensis F27, Megatebennus javanicensis H09; incl. Fissurella trapezina Sowerby 1835 KC60, *Amblychilepas trapezina KC60]

Fissurellidea (Fissurellidea) hiantula (Lamarck 1822) [=Fissurella hiantula; incl. *Fissurellidea megatrema d’Orbigny 1841] KC60

Hemitoma Swainson 1840 BR05 [=Subemarginula Gray 1847 KC60; incl. Siphonella Issel 1869 non Hagenow 1851 KC60; Hemitomidae, Hemitominae BR17]

Hemitoma (Hemitoma) octoradiata (Gmelin 1791) KC60 [=Patella octoradiata KC60, Emarginula (Hemitoma) octoradiata O27; incl. E. tricostata Sowerby 1823 BR17, *Hemitoma tricostata KC60, Patella tricostata non Gmelin 1791 KC60, *Subemarginula tricostata KC60]

Lucapina sowerbii (Guilding ex Sowerby 1835) [=*Foraminella sowerbii; incl. Fissurella cancellata Sowerby 1835 non Gray 1825, *Lucapina cancellata] KC60

Megathura crenulata (Sowerby 1825) KC60 [=Fissurella crenulata KC60, Lucapina crenulata O27; incl. *Megathura californica Pilsbry 1890 KC60, *Macrochasma californicum KC60]

*Montfortula rugosa (Quoy & Gaimard 1834) KC60 [=Emarginula rugosa P61, Clypidina (*Montfortula) rugosa KC60, Hemitoma rugosa F27, Subemarginula rugosa H09]

Scutus Montfort 1810 BR05 [=Parmaphora Bowdich 1822 KC60, Parmophora Desmarest 1859 KC60, Parmophorus deBlainville 1817 KC60, Scutum Sowerby 1842 non Schumacher 1817 KC60; incl. Aviscutum Iredale 1940 KC60; Scutinae, Scutini]

Scutus anatinus (Donovan 1820) [=Patella anatina; incl. Parmophorus australis Lamarck 1822, Par. convexus, Par. elongatus Blainville 1817 non Patella elongata, Emarginula elongata, Scutus elongatus, S. tumidus Ad. 1851] S79

Scutus unguis (Linnaeus 1758) H09 [=Patella unguis H09, Mytilus lingua Lightfoot 1786 K65; incl. Scutus angustatus Adams 1851 S79, Parmophorus corrugatus S79, S. corrugatus S79, Par. elegans Gray 1825 S79, Par. emarginatus Philippi 1851 S79, Par. granulatus Blainv. 1817 S79, Scutus granulatus S79, Par. imbricatus S79, S. imbricatus S79, Par. rueppeli Philippi 1851 S79]

Scutus (Scutus) ambiguus (Chemnitz 1795) F27 [=Patella ambigua S79, *S. antipodes Montfort 1810 P61, S79, P61]

Zeidora Adams 1860 BR05 [=Zidora Fischer 1885 BR05; incl. Crepiemarginula Seguenza 1880 KC60, Legrandia Beddome 1883 non Hanley 1872 KC60; Zeidoridae, Zidoridae]

*Type species of generic name indicated


[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[BC01] Boyko, C. B., & J. R. Cordeiro. 2001. Catalog of Recent type specimens in the Division of Invertebrate Zoology, American Museum of Natural History. V. Mollusca, part 2 (class Gastropoda [exclusive of Opisthobranchia and Pulmonata], with supplements to Gastropoda [Opisthobranchia], and Bivalvia). Bulletin of the American Museum of Natural History 262: 1–170.

[C64] Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 33: 517–686.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[K65] Kay, E. A. 1965. The Reverend John Lightfoot, Daniel Solander, and the Portland Catalogue. Nautilus 79 (1): 10–19.

[KC60] Knight, J. B., L. R. Cox, A. M. Keen, R. L. Batten, E. L. Yochelson & R. Robertson. 1960. Gastropoda: systematic descriptions. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I169–I331. Geological Society of America, and University of Kansas Press.

[M54] Macpherson, J. H. 1954. Molluscs (sea shells and snails). In: Willis, J. M. (ed.) The Archipelago of the Recherche pp. 55–63. Australian Geographical Society: Melbourne.

[M84] McLean, J. H. 1984. A case for derivation of the Fissurellidae from the Bellerophontacea. Malacologia 25 (1): 3–20.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

[P95] Ponder, W. F. 1995. Book review: Australian Marine Shells. Prosobranch Gastropods. Parts 1 and 2. Molluscan Research 16: 97–102.

[PP78] Poorman, F. L., & L. H. Poorman. 1978. Additional molluscan records from Bahía de Los Angeles, Baja California Norte. Veliger 20 (4): 369–374.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[S11] Simone, L. R. L. 2011. Phylogeny of the Caenogastropoda (Mollusca), based on comparative morphology. Arquivos de Zoologia 42 (4): 161–323.

[S79] Smith, E. A. 1879. Notes on the species of the genus Scutus. Journal of Conchology 2: 252–264.

[W78] Wells, F. E. 1978. Zonation of marine gastropods on a rocky intertidal shore in the Admiralty Gulf, Western Australia, with emphasis on the genus Nerita. Veliger 20 (3): 279–287.

Last updated: 17 October 2021.

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