Daphnella rissoides, copyright G. & Ph. Poppe.

Belongs within: Conoidea.
Contains: Asperdaphne (Asperdaphne), Pseudodaphnella, Eubela, Rugobela, Buccinaria, Nepotilla, Philbertia, Hemilienardia, Raphitoma, Veprecula, Eucyclotoma, Kermia, Pleurotomella (Pleurotomella), Taranis, Gymnobela, Daphnella (Daphnella).

The Raphitomidae is a very diverse group of conoid gastropods, many species of which show a reduction in the radular apparatus or other structures of the foregut (Bouchet et al. 2011).

Characters (from Bouchet et al. 2011): Shell very variable in shape and size, from buccinoid to ovate or to elongate-fusiform or high-cylindrical and from about 2 to over 140 mm high. Sculpture variously developed, from nearly smooth shell to well-developed spiral and axial elements; subsutural ramp, when separated from rest of the whorl, usually smooth or with traces of anal sinus growth. Apertural armature rarely well expressed, inner lip usually smooth. Anal sinus either subsutural, shaped as a ‘reversed-L’, or on subsutural ramp, asymmetric, very shallow to rather deep, typically not constrained but sometimes even almost tubular. Siphon short to rather long. Protoconch typically planktotrophic, multispiral, of 2.5–6.5 whorls, protoconch I often spirally striated, protoconch II with diagonally cancellated sculpture. Paucispiral protoconch with predominantly spiral striation. Operculum absent. Radula of hypodermic marginal teeth of very variable morphology. Teeth relatively very long to very short and reduced. Solid base of teeth from very small to rather large, side projections around base sometimes present. Barb(s) present or absent; cockscomb-shaped structure present near tooth tip in at least one species. Tooth canal opening laterally or subapically. Reduction in radular apparatus and other structures of anterior foregut frequent, grading to complete absence.

<==Raphitomidae [Daphnellinae, Daphnellini, Pleurotomellinae, Raphitominae, Thatcheriidae, Thatcheriinae]
    |--+--Hemilienardia PK11
    |  `--Raphitoma PK11
    `--+--+--Veprecula PK11
       |  `--Rimosodaphnella Cossmann 1915 PK11, P66 [=Rimosodalphnella (l. c.) P66]
       |       |--*R. textile (Brocchi 1814) [=Murex textilis] P66
       |       |--R. ciprianii (Hornung 1920) [=Daphnella (Rimosodaphnella) ciprianii] P66
       |       `--R. salinasi (Calcara 1841) [=Pleurotoma salinasi] P66
       `--+--Glyphostomoides Shuto 1983 PK11, BK11
          |--Tritonoturris Dall 1924 PK11, P66
          |    |--*T. robillardi (Adams 1869) [=Clathurella robillardi] P66
          |    |--T. amabilis (Hinds 1843) P66 [=Clavatula amabilis P66, Clathurella amabilis H09]
          |    |--T. capensis (Smith 1882) [=Pleurotoma (Defrancia) capensis] P66
          |    |--T. cumingii (Powys in Powys & Sowerby 1835) BK11 [=Buccinum cumingii P66]
          |    |--T. elegans (Pease 1860) [=Clathurella elegans] P66
          |    |--T. menecharmes (Melvill 1923) [=Veprecula menecharmes] P66
          |    `--T. paucicostata (Adams 1869) [=Clathurella paucicostata] P66
          `--+--+--Eucyclotoma PK11
             |  `--Kermia PK11
             `--+--Pleurotomella Verrill 1873 PK11, BK11 (see below for synonymy)
                |    |  i. s.: ‘Daphnella’ aculeola Hedley 1915 [=Tasmadaphne aculeola] P66
                |    |         P. aguayoi (Carcelles 1953) S11
                |    |         *Fusidaphne’ bullata Laseron 1954 P66
                |    |         *Tasmadaphne’ spicula Laseron 1954 P66
                |    |         P. thalassica Dall 1919 O27
                |    |--P. (Pleurotomella) P66
                |    |--P. (Anomalotomella Powell 1966) P66
                |    |    |--P. (*A.) anomalapex Powell 1951 P66
                |    |    |--P. (A.) annulata Thiele 1912 P66
                |    |    |--P. (A.) enderbyensis Powell 1958 P66
                |    |    |--P. (A.) frigida Thiele 1912 P66
                |    |    `--P. (A.) ohlini (Strebel 1905) [=Thesbia ohlini] P66
                |    |--P. (Cyrtoides Nordsieck 1968) BK11
                |    `--P. (Systenope Cossmann 1889) P66
                |         |--P. (*S.) polycolpa (Cossmann 1889) [=Raphitoma (*S.) polycolpa] P66
                |         |--P. (S.) goniocolpa (Cossmann 1889) [=Raphitoma goniocolpa] P66
                |         |--P. (S.) guepellensis (Cossmann 1889) [=Raphitoma guepellensis] P66
                |         `--P. (S.) linophora (Cossmann 1889) [=Raphitoma linophora] P66
                `--+--Taranis BK11
                   `--+--+--Teretiopsis Kanto & Sysoev 1989 PK11,BK11
                      |  |    `--T. hyalina Sysoev & Bouchet 2001 BK11
                      |  `--+--Gymnobela PK11
                      |     `--Thatcheria Angas 1877 BK11, BR05 [incl. Cochlioconus Yokoyama 1928 P66]
                      |          |--T. (Thatcheria) P66
                      |          |    |--*T. (T.) mirabilis Angas 1877 P66
                      |          |    |--T. (T.) carinata (Martin 1933) [=Cryptoconus carinatus] P66
                      |          |    |--T. (T.) gradata (Yokoyama 1928) [=*Cochlioconus gradatus] P66
                      |          |    |--T. (T.) liratula (Powell 1942) [=Waitara liratula] P66
                      |          |    |--T. (T.) pagodula (Powell 1942) [=Waitara pagodula] P66
                      |          |    `--T. (T.) vitiensis Charig 1963 P66
                      |          `--T. (Waitara Marwick 1931) P66
                      |               `--T. (*W.) waitaraensis (Marwick 1926) [=Turricula waitaraensis] P66
                      `--Daphnella Hinds 1844 PK11, BR05 (see below for synonymy)
                           |  i. s.: D. cassandra Hedley 1904 H09
                           |         D. cladara Sysoev & Bouchet 2001 BK11
                           |         D. clathrata Gabb 1865 O27
                           |         D. effusa D19
                           |         D. electra D19
                           |         D. hayesi Usticke 1959 BC01
                           |         D. lymnaeiformis (Kiener 1840) S11
                           |         D. mitrellaformis (Nomura 1940) BK11
                           |         D. pluricarinata (Reeve 1845) [=Pleurotoma pluricarinata] H09
                           |         D. pulvisculus Chino 2006 BK11
                           |         D. stegeri BC01
                           |           |--D. s. stegeri BC01
                           |           `--D. s. monocincta Usticke 1969 BC01
                           |         D. vitrea (Reeve 1845) P66
                           |--D. (Daphnella) P66
                           |--D. (Diaugasma Melvill 1917) P66
                           |    |--D. (*D.) epicharta Melvill & Standen 1903 P66
                           |    `--D. (D.) olyra (Reeve 1845) [=Pleurotoma olyra] P66
                           `--D. (Hemidaphne Hedley 1918) P66
                                |--D. (*H.) souverbiei (Smith 1882) [=Pleurotoma (Daphnella) souverbiei] P66
                                |--D. (H.) axis (Reeve 1846) [=Pleurotoma axis] P66
                                |--D. (H.) cyclophora (Deshayes 1863) [=Pleurotoma cyclophora] P66
                                |--D. (H.) lirata (Reeve 1845) [=Pleurotoma lirata] P66
                                |--D. (H.) reeveana (Deshayes 1863) [=Pleurotoma reeveana] P66
                                |--D. (H.) rissoides (Reeve 1843) [=Pleurotoma rissoides] P66
                                |--D. (H.) subula (Reeve 1845) [=Pleurotoma subula] P66
                                |--D. (H.) subuloides Schepman 1913 P66
                                `--D. (H.) supracancellata Schepman 1913 P66

Raphitomidae incertae sedis:
  Andonia Harris & Burrows 1891 [Andoniinae] BR05
  Austrodaphnella Laseron 1954 P66
    `--*A. clathrata Laseron 1954 P66
  Hokianga Laws 1947 P66
    `--*H. nodulata Laws 1947 P66
  Favriella Hornung 1920 P66
    `--*F. weberi (Hornung 1920) [=Daphnella (*Favriella) weberi] P66
  Metuonella Sorgenfrei 1958 P66
    `--*M. grippi (Kautsky 1925) [=Daphnella grippi] P66
  Microgenia Laseron 1954 P66
    `--*M. edwini (Brazier 1894) [=Clathurella edwini] P66
  Isodaphne Laseron 1954 P66
    `--*I. garrardi Laseron 1954 P66
  Fenestrodaphne Powell 1944 P66
    `--*F. pulchra Powell 1944 P66
  Magnella Dittmer 1960 P66
    `--*M. andersoni Dittmer 1960 P66
  Phandella Casey 1903 P66
    `--*P. nepionica Casey 1903 P66
  Syngenochilus Powell 1944 P66
    `--*S. radiapex Powell 1944 P66
  Daphnobela Cossmann 1896 P66
    `--*D. juncea (Sowerby 1822) [=Buccinum junceum] P66
  Thesbia Jeffreys 1867 P66
    |--*T. nana (Loven 1846) [=Tritonium nanum] P66
    `--T. microtoma Cossmann 1899 P66
  Leiosyrinx Bouchet & Sysoev 2001 BS01
    |--*L. immedicata Bouchet & Sysoev 2001 BS01
    |--L. apheles Bouchet & Sysoev 2001 BS01
    |--L. liphaima Bouchet & Sysoev 2001 BS01
    `--L. matsukumai Bouchet & Sysoev 2001 BS01
  Teretia Norman 1888 BK11 [incl. Teres Bucquoy, Dautzenberg & Dollfus 1883 non Boettger 1878 P66]
    |--*T. anceps (Eichwald 1830) (see below for synonymy) P66
    |--T. amoena (Sars 1878) [=Rhaphitoma amoena] P66
    |--T. megalembryon (Dautzenberg & Fischer 1896) [=Pleurotoma megalembryon] P66
    `--T. nana (Hornung 1920) [=Daphnella (Teres) nana] P66
  Asperdaphne Hedley 1922 [incl. Scabrella Hedley 1918 non Sacco 1890] P66
    |--A. (Asperdaphne) P66
    `--A. (Aspertilla Powell 1944) P66
         |--A. (*A.) legrandi (Beddome 1883) [=Drillia legrandi] P66
         `--A. (A.) exsculpta Powell 1944 P66
  Typhlosyrinx Thiele 1925 BS01
    |--*T. vepallida (Martens 1902) (see below for synonymy) BS01
    |--T. neocaledoniensis Bouchet & Sysoev 2001 BS01
    |--T. panamica Bouchet & Sysoev 2001 BS01
    |--T. praecipua (Smith 1899) [=Pleurotoma (Surcula) praecipua] BS01
    `--T. supracostata (Schepman 1913) [=Surcula supracostata] BS01
  Xanthodaphne Powell 1942 P66
    |--*X. membranacea (Watson 1886) [=Pleurotoma (Thesbia) membranacea] P66
    |--X. maoria (Dell 1956) P66
    |--X. pyrropelex (Barnard 1963) [=Typhlosyrinx pyrropelex] BS01
    |--X. subrosea (Barnard 1963) [=Typhlosyrinx subrosea] BS01
    `--X. xanthias (Watson 1886) [=Pleurotoma (Thesbia) xanthias] P66
  Stilla Finlay 1926 P66
    |--*S. flexicostata (Suter 1899) [=Mangilia flexicostata] P66
    |--S. anomala Powell 1955 P66
    |--S. delicatula Powell 1927 P66
    |--S. fiordlandica Fleming 1948 P66
    `--S. paucicostata Powell 1937 P66
  Cryptodaphne Powell 1942 P66
    |--*C. pseudodrillia Powell 1942 P66
    |--C. abbreviata (Schepman 1913) [=Pleurotomella abbreviata] P66
    |--C. affinis (Schepman 1913) [=Pleurotomella affinis] P66
    |--C. biconica (Schepman 1913) [=Pleurotomella biconica] P66
    `--C. gradata (Schepman 1913) [=Pleurotomella gradata] P66
  Puha Marwick 1931 P66
    |--*P. fulgida Marwick 1931 P66
    |--P. hebes (Hutton 1873) [=Pleurotoma hebes] P66
    |--P. japonica Shuto 1961 P66
    |--P. pusula Laws 1947 P66
    `--P. sinusigera (Powell 1942) P66
  Phymorhynchus Dall 1908 P66
    |--*P. castanea (Dall 1895) [=Pleurotomella castanea] P66
    |--P. argeta (Dall 1889) [=Pleurotomella argeta] P66
    |--P. cingulata (Dall 1889) [=Pleurotomella cingulata] P66
    |--P. clarinda (Dall 1908) [=Pleurotomella (Phymorhynchus) clarinda] P66
    |--P. major Warén & Bouchet 2001 BK11
    `--P. oceanica (Dall 1908) [=Pleurotomella (Phymorhynchus) oceanica] P66
  Pontiothauma Smith 1895 P66
    |--*P. mirabile Smith 1895 P66
    |--P. abyssicola Smith 1895 P66
    |--P. ergata Hedley 1916 P66
    |--P. minus Smith 1906 P66
    `--P. pacei Smith 1906 P66
  Zenepos Finlay 1928 P66
    |--*Z. totolirata (Suter 1908) [=Daphnella totolirata] P66
    |--Z. chariessa (Suter 1908) [=Daphnella chariessa] P66
    |--Z. lacunosa (Hutton 1885) [=Daphnella lacunosa] P66
    |--Z. mimica (Sowerby 1896) [=Daphnella mimica] P66
    |--Z. minuta (Tenison-Woods 1877) [=Drillia minuta] P66
    `--Z. miranda (Thiele 1925) [=Taranis (Nepotilla) miranda] P66
  Exomilus Hedley 1918 P66
    |--*E. lutrarius (Hedley 1907) [=Mangelia lutraria] P66
    |--E. anxius (Hedley 1909) [=Mangilia anxia] P66
    |--E. ‘cancellatus’ (Beddome 1882) [=Mangilia cancellata non Adams 1865] P66
    |--E. cylindricus Laseron 1954 P66
    |--E. pentagonalis (Verco 1896) [=Drillia pentagonalis] P66
    `--E. telescopialis (Verco 1896) [=Drillia telescopialis] P66
  Teleochilus Harris 1897 [incl. Litachilus Powell 1944] P66
    |--*T. gracillimus (Tenison-Woods 1877) [=Daphnella gracillima] P66
    |--T. balcombensis Powell 1944 P66
    |--T. comptus Powell 1944 P66
    |--T. denseliratus Powell 1944 P66
    |--T. duplicatus Powell 1944 P66
    `--T. royanus Iredale 1924 [=*Litachilus royanus] P66
  Maoridaphne Powell 1942 P66
    |--*M. clifdenica (Laws 1939) [=Daphnella clifdenica] P66
    |--M. haroldi Powell 1942 P66
    `--M. kaiparica (Laws 1939) [=Daphnella kaiparica] P66
  Spergo Dall 1895 P66
    |--S. (Spergo) P66
    |    |--*S. (S.) glandiniformis Dall 1895 P66
    |    |--S. (S.) daphnelloides Dall 1895 P66
    |    |--S. (S.) fusiformis (Kuroda & Habe in Habe 1962) P66, BK11
    |    `--S. (S.) fusus MacNeil 1960 P66
    `--S. (Speoides Kuroda & Habe in Habe 1962) BK11
         `--S. (*S.) yoshidai (Kuroda & Habe 1961) [=*Speoides yoshidai] P66
  Clinura Bellardi 1875 P66
    |--*C. calliope (Brocchi 1914) [=Murex (Pleurotoma) calliope] P66
    |--C. anassa Murphy & Rodda 1960 P66
    |--C. bituminata (Beets 1843) [=Surculites bituminata] P66
    |--C. controversa (Bellardi 1847) [=Pleurotoma controversa] P66
    |--C. generosa (Marwick 1931) [=Waitara generosa] P66
    |--C. sabatiorum Bellardi 1877 P66
    |--C. sopronensis (Wolf 1870) [=Pleurotoma sopronensis] P66
    |--C. subtrochlearis (Friedberg 1912) [=Surcula (Clinura) subtrochlearis] P66
    `--C. trochlearis (Hörnes 1854) [=Pleurotoma trochlearis] P66
  Pseudodaphnella P66
  Eubela P66
  Rugobela P66
  Buccinaria BK11
  Nepotilla P66
  Thatcheriasyrinx Powell 1969 BK11
    `--T. orientis (Melvill 1904) BK11
  Rocroithys Sysoiev & Bouchet 2001 BK11
    `--R. niveus Sysoev & Bouchet 2001 BK11
  Microdaphne McLean 1971 BK11
    `--M. morrisoni Rehder 1980 BK11
  Abyssobela Kantor & Sysoev 1986 BK11
  Abyssothauma Sysoev 1996 BK11
  Acamptodaphne Shuto 1971 BK11
  Acanthodaphne Bonfitto & Morassi 2006 BK11
  Aliceia Dautzenberg & Fischer 1897 BK11
    `--*A. aenigmatica Dautzenberg & Fischer 1897 [=Clavus aenigmatica] P66
  Austropusilla Laseron 1954 BK11
    |  i. s.: A. profundis Laseron 1954 P66
    |--*A. (Austropusilla) hilum (Hedley 1908) P66 [=Mangelia hilum P66, Filodrillia hilum L54]
    `--A. (Metaclathurella Shuto 1983) BK11
  Bathybela Kobelt 1905 [incl. Bathypota Nordsieck 1968] BK11
    |--*B. nudator (Locard 1897) [=Thesbia nudator] P66
    `--B. folini (Locard 1897) [=Thesbia folini] P66
  Cenodagreutes Smith 1967 BK11
  Clathurina Melvill 1917 BK11
    `--*C. foraminata (Reeve 1845) [=Pleurotoma foraminata, Kermia foraminata] P66
  Famelica Bouchet & Warén 1980 BK11
  Kuroshiodaphne Shuto 1965 BK11
    |--*K. fuscobalteata (Smith 1879) (see below for synonymy) P66
    `--K. subzonata (Smith 1879) [=Daphnella subzonata, Maoridaphne (Kuroshiodaphne) subzonata] P66
  Lusitanops Nordsieck 1968 BK11
  Mioawateria Vella 1954 BK11
    |--*M. personata (Powell 1942) [=Awateria (*Mioawateria) personata] P66
    |--M. expalliata (Laws 1947) [=Awateria (Mioawateria) expalliata] P66
    |--M. experta (Laws 1947) [=Awateria (Mioawateria) experta] P66
    |--M. karakaensis (Marwick 1931) [=Awateria (Mioawateria) karakaensis] P66
    `--M. pahaoaensis (Vella 1954) [=Awateria (Mioawateria) pahaoaensis] P66
  Neopleurotomoides Shuto 1971 BK11
  Pagodidaphne Shuto 1983 BK11
  Paramontana Laseron 1954 P66
    |--*P. modesta (Angas 1877) P66 [=Clathurella modesta P66, Pseudodaphnella modesta L54]
    |--P. fusca Laseron 1954 P66
    |--P. mayana (Hedley 1922) [=Pseudodaphnella mayana] P66
    |--P. oligoina (Hedley 1922) [=Pseudodaphnella oligoina] P66
    |--P. punicea (Hedley 1922) [=Pseudodaphnella punicea] P66
    `--P. rufozonata (Angas 1877) P66 [=Clathurella rufozonata P66, Pseudodaphnella rufozonata L54]
  Taranidaphne Morassi & Bonfitto 2001 BK11
  Thatcherina Verez-Peláez 1998 BK11
  Truncadaphne McLean 1971 BK11
  Tuskaroria Sysoev 1988 BK11
  Vepridaphne Shuto 1983 BK11
  Philbertia BK11

Daphnella Hinds 1844 PK11, BR05 [incl. Eudaphne Bartsch 1931 non Reuss 1922 P66, Eudaphnella Bartsch 1933 P66]

*Kuroshiodaphne fuscobalteata (Smith 1879) [=Daphnella fuscobalteata, Maoridaphne (*Kuroshiodaphne) fuscobalteata] P66

Pleurotomella Verrill 1873 PK11, BK11 [incl. Azorilla Nordsieck 1968 BK11, Azorita Nordsieck 1968 BK11, Fusidaphne Laseron 1954 BK11, Tasmadaphne Laseron 1954 BK11]

*Teretia anceps (Eichwald 1830) [=Pleurotoma anceps, *Teres anceps; incl. Rhaphitoma barbieri Brusina 1866, Pleurotoma boreale Loven 1846, P. fusiforme Requien 1848, Fusus laviae Calcara 1845, P. minutum Aradas 1847, Clathurella anceps soluta Marshall 1912, P. teres Forbes in Reeve 1844, P. trecchi Testa in Jeffreys 1867] P66

*Typhlosyrinx vepallida (Martens 1902) [=Pleurotoma (Leucosyrinx) vepallida, Pleurotoma (Surcula) vepallida, Pleurotomella (Typhlosyrinx) vepallida] BS01

*Type species of generic name indicated


[BK11] Bouchet, P., Y. I. Kantor, A. Sysoev & N. Puillandre. 2011. A new operational classification of the Conoidea (Gastropoda). Journal of Molluscan Studies 77: 273–308.

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

[BS01] Bouchet, P., & A. Sysoev. 2001. Typhlosyrinx-like tropical deep-water turriform gastropods (Mollusca, Gastropoda, Conoidea). Journal of Natural History 35: 1693–1715.

[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.

[D19] Dall, W. H. 1919. Descriptions of new species of mollusks of the family Turritidae from the west coast of America and adjacent regions. Proceedings of the United States National Museum 56 (2288): 1–86, pls 1–24.

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

[L54] Laseron, C. F. 1954. Revision of the New South Wales Turridae (Mollusca). Royal Zoological Society of New South Wales: Sydney.

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

[P66] Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae: an evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Auckland Institute and Museum 5: 1–184, pls 1–23.

[PK11] Puillandre, N., Y. I. Kantor, A. Sysoev, A. Couloux, C. Meyer, T. Rawlings, J. A. Todd & P. Bouchet. 2011. The dragon tamed? A molecular phylogeny of the Conoidea (Gastropoda). Journal of Molluscan Studies 77: 259–272.

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


  1. Is there any snail cooler than Thatcheria mirabilis? I think not.


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