Drilliidae

Cruziturricula arcuata, copyright Smithsonian Institution.

Belongs within: Conoidea.
Contains: Tylotiella, Fenimorea, Austroclavus, Elaeocyma, Cymatosyrinx, Syntomodrillia, FusiturriculaIredalea, Clavus, Cerodrillia, Splendrillia (Splendrillia), Agladrillia, Clathrodrillia, Drillia (Drillia).

The Drilliidae are a group of slender-shelled predatory gastropods, with a fossil record extending to the Upper Cretaceous (Powell 1966). Most retain a radula with well-developed lateral teeth (Bouchet et al. 2011), probably for active processing of prey, in contrast to the specialised venom-injecting radulae of other conoids. The basal genus Cruziturricula, however, lacks laterals and has a radula with wishbone-type marginal teeth only. It and the related Fusiturricula (for which the radula is currently unknown) may be misplaced in the Drilliidae (Bouchet et al. 2011). Members of the type genus Drillia are characterised by a large, solid shell with a tall spire and truncated body whorl, and are most diverse in western Africa (Powell 1966).

Characters (from Bouchet et al. 2011): Shell small- to medium-sized, usually 15–25 mm, up to 50 mm high, with rather high spire and usually truncated base. Spiral sculpture often obsolete. Anal sinus on subsutural ramp, deep, (sub)symmetrical, sometimes tubular. Protoconch usually paucispiral (up to two whorls), smooth or abapically carinate. Operculum with terminal nucleus. Radular formula 1-1-R-1-1, rarely 1-1-0-1-1, central tooth small, from narrow unicuspid to subrectangular with additional cusps, rarely reduced to completely absent. Lateral teeth broad, pectinate, and arched, marginal teeth from simple flat and sharply pointed to duplex with slightly thickened edges and to loosely enrolled with small barb near tip.

<==Drilliidae [Clavinae, Clavini, Clavusinae, Drilliinae, Drillinae]
    |--+--FusiturriculaPK11
    |  `--Cruziturricula Marks 1951PK11, BK11
    |       |--*C. cruziana (Olsson 1932) [=Turricula (Pleurofusia) cruziana]P66
    |       |--C. andesita (Olsson 1942) [=Turricula (Knefastia) andesita]P66
    |       |--C. arcuata (Reeve 1843)PK11 [=Pleurotoma arcuataP66]
    |       |--C. eolavinia (Olsson 1930) [=Turricula (Pleurofusia) eolavinia]P66
    |       |--C. lavinia (Dall 1919) [=Turricula (Surcula) lavinia]P66
    |       `--C. piura (Olsson 1931) [=Turricula (Surcula) piura]P66
    `--+--+--Conopleura Hinds 1844PK11, BK11
       |  |    `--*C. striata Hinds 1844 [=Pleurotoma partita Reeve 1846]P66
       |  `--+--+--IredaleaPK11
       |     |  `--+--ClavusPK11
       |     |     `--CerodrilliaPK11
       |     `--Splendrillia Hedley 1922PK11, BK11
       |          |  i. s.: S. bratcherae McLean & Poorman 1971PP78
       |          |         S. petiti (Tippett 1995) [=Fenimorea petiti]BC01
       |          |--S. (Splendrillia)BK11
       |          `--S. (Hauturua Powell 1942)BK11
       |               |--S. (*H.) vivens (Powell 1942) [=Syntomodrillia (*H.) vivens]P66
       |               |--S. (H.) exiguescens (Marwick 1931) [=Austrodrillia exiguescens]P66
       |               |--S. (H.) exuta Powell 1944P66
       |               `--S. (H.) laevella (Marwick 1931) [=Austrodrillia laevella]P66
       `--+--+--AgladrilliaPK11
          |  `--+--Calliclava McLean 1971PK11, BK11
          |     |    |--C. albolaqueata (Carpenter 1865)SK81
          |     |    |--C. alcmene (Dall 1919)PP78
          |     |    `--C. jaliscoensis McLean & Poorman 1971BC01
          |     `--Imaclava Bartsch 1944PK11, BK11
          |          |--I. pembertoni (Lowe 1935) (see below for synonymy)P66
          |          |--I. pilsbryi Bartsch 1950BK11 [=Clathrodrillia (Imaclava) pilsbryi non C. pilsbryi Lowe 1935P66]
          |          `--I. unimaculata (Sowerby 1834)PK11 (see below for synonymy)
          `--+--ClathrodrilliaPK11
             `--Drillia Gray 1838PK11, BK11
                  |  i. s.: D. alabaster (Reeve 1843) [=Pleurotoma alabaster]H09
                  |         D. albomaculataM23
                  |         D. anthamilla Melvill 1923M23
                  |         D. appeliiM23
                  |         D. chaaci Espinosa & Rolán 1995BC01
                  |         D. cubana Melvill 1923M23
                  |         D. digitalis Reeve 1843M23
                  |         D. euchroes Melv. 1912M23
                  |         D. exigua (Hombron & Jacq. 1853) [=Pleurotoma exigua]H09
                  |         D. gracillimaC64
                  |         D. griffithiiM23
                  |         D. hexagonaC64
                  |         D. incrassataC64
                  |         D. inermisC64
                  |         D. infrafusca Sowerby 1893M23
                  |         D. innocens Melvill 1923M23
                  |         D. insignita Melvill 1923M23
                  |         D. kleinrosa Usticke 1969BC01
                  |         D. luctuosaC64
                  |         D. majorM23
                  |         D. mauraC64
                  |         D. nigerrimaC64
                  |         D. nitens (Hinds 1843) [=Clavatula nitens]H09
                  |         D. nodifera Pease 1860 [=Pleurotoma nodifera non Lamarck 1822 (not preoc. if in dif. gen.)]K65
                  |         D. nodulosa Pease 1863K65
                  |         D. ochroleuca Melv. & Sykes 1897M23
                  |         D. ornataM23
                  |         D. papillosa Garrett 1873M23
                  |         D. paria Reeve 1846M23
                  |         D. paroeca Melvill 1923M23
                  |         ‘Pleurotoma (Drillia) parva’ Smith 1888 non P. parva Conrad 1830F27
                  |         D. poncianaBC01
                  |           |--D. p. poncianaBC01
                  |           `--D. p. virgata Usticke 1969 [=D. bandata Usticke 1971]BC01
                  |         D. primula Melvill 1923M23
                  |         D. roseola (Hertlein & Strong 1955)BC01 (see below for synonymy)
                  |         D. rudisC64
                  |         D. stiza Tenison-Woods 1879F71
                  |         D. wanganuiensisH86
                  |         D. wolfei Tippett 1995BC01
                  |         D. zebraM23
                  |--D. (Drillia)BC01
                  `--D. (Pseudodrillia Lukovic 1924)P66
                       |--D. (P.) abnormalis Lukovic 1924P66
                       |--D. (P.) arabica Lukovic 1924P66
                       `--D. (P.) longa Lukovic 1924P66
Drilliidae incertae sedis:
  Clavatoma Powell 1942P66
    `--*C. pulchra Powell 1942P66
  Maoricrassus Vella 1954P66
    `--*M. carinatus Vella 1954P66
  Tahudrillia Powell 1942P66
    `--*T. simplex Powell 1942P66
  Integradrillia Powell 1942P66
    `--*I. integra (Tenison-Woods 1880) [=Drillia integra]P66
  Vixinquisitor Powell 1942P66
    `--*V. vixumbilicata (Harris 1897) [=Drillia vixumbilicata]P66
  Globidrillia Woodring 1928BK11
    `--*G. ula Woodring 1928P66
  Tripia Gregorio 1890 (n. d.)P66
    `--*T. anteatripla de Gregorio 1890 (n. d.) (see below for synonymy)P66
  Boreodrillia Sorgenfrei 1958P66
    `--*B. toftlundensis Sorgenfrei 1958P66
  Mitrelloturris Eames 1957P66
    `--*M. casteri (Chavan 1952) [=Asthenotoma (Endiatoma) casteri]P66
  Terebritoma Cossmann 1894P66
    `--*T. solitaria [=Mangelia solitaria]P66
  Neodrillia Bartsch 1943BK11
    |--*N. cydia Bartsch 1943 (see below for synonymy)P66
    |--N. blacki Petuch 2004PMB15
    `--N. euphanes (Melvill 1923) [=Drillia euphanes]P66
  Wairarapa Vella 1954P66
    |--*W. rebecca Vella 1954P66
    `--‘Melatoma’ duplaris Hedley 1922P66
  Douglassia Bartsch 1934BK11
    |--*D. enae Bartsch 1934P66
    `--D. bealiana Schwengel & McGinty 1942P66
  Crassopleura Monterosato 1884BK11 [=Crossopleura (l. c.)P66]
    |--C. costulatum (Cantraine 1835) (see below for synonymy)P66
    `--C. brevis (Yokoyama 1922) [=Drillia glabriuscula brevis]P66
  Leptadrillia Woodring 1928BK11
    |--*L. parkeri (Gabb 1873) [=Turris (Surcula) parkeri]P66
    |--L. loria Bartsch 1934P66
    `--L. splendida Bartsch 1934P66
  Kylix Dall 1919BK11
    |--*K. alcyone (Dall 1919) [=Clathrodrillia (Kylix) alcyone]P66
    |--K. alcmene (Dall 1919) [=Clathrodrillia (Kylix) alcmene]P66
    |--K. contracta McLean & Poorman 1971BC01
    |--K. paziana (Dall 1919)PP78
    |--K. turveri Hertlein & Strong 1951P66
    `--K. zacae Hertlein & Strong 1951P66
  TylotiellaBK11
  Plagiostropha Melvill 1927BK11
    |--*P. quintuplex Melvill 1927 [=Clavus (*Plagiostropha) quintuplex]P66
    |--P. caledoniensis Wells 1995BK11
    |--‘Pleurotoma’ ebur Reeve 1845P66
    `--‘Pleurotoma’ opalus Reeve 1845P66
  Paracuneus Laseron 1954BK11
    |--*P. immaculatus (Tenison-Woods 1876)P66 (see below for synonymy)
    |    |--P. i. immaculatusP66
    |    `--P. i. peronianus Laseron 1954P66
    |--P. kemblensis Laseron 1954P66
    `--P. spadix (Watson 1886)P66 [=Pleurotoma (Drillia) spadixP66, Inquisitor spadixH22]
         |--P. s. spadixP66
         `--P. s. tumulus Laseron 1954P66
  Spirotropis Sars 1878BK11
    |--S. monterosatoi (Locard 1897) (see below for synonymy)BW80
    |--S. azorica Bouchet & Waren 1980BW80
    |--S. centimata (Dall 1889) (see below for synonymy)BW80
    |--S. ephamilla Verrill 1884P66
    |--S. modiola (Jan 1832)BW80 (see below for synonymy)
    |--S. remota Powell 1958P66
    `--S. studeriana (Martens 1878) [=Pleurotoma studeriana]P66
  Sedilia Fargo 1953P66
    |--*S. sedilia (Dall 1890) [=Drillia sedilia]P66
    |--S. hoplophorus (Dall 1892) [=Drillia hoplophorus]P66
    |--S. ochoidia Fargo 1953P66
    |--S. ondulum Fargo 1953P66
    |--S. sapa Fargo 1953P66
    |    |--S. s. sapaP66
    |    `--S. s. sapala Fargo 1953P66
    |--S. schismatica (Dall 1892) [=Drillia schismatica]P66
    `--S. transa Fargo 1953P66
  FenimoreaBK11
  Eumetadrillia Woodring 1928BK11
    |--*E. serra (Woodring 1928) [=Agladrillia (*Eumetadrillia) serra]P66
    |--E. centrodes (Gardner 1937) [=Drillia centrodes, Agladrillia (Eumetadrillia) centrodes]P66
    |--E. dodona Gardner 1937 [=Agladrillia (Eumetadrillia) dodona]P66
    |--E. fuegiensis (Smith 1888) [=Pleurotoma (Surcula) fuegiensis, Agladrillia (Eumetadrillia) fuegiensis]P66
    |--E. isthmica (Brown & Pilsbry 1911) [=Drillia isthmica, Agladrillia (Eumetadrillia) isthmica]P66
    |--E. newmani (Dall 1890) [=Drillia newmani, Agladrillia (Eumetadrillia) newmani]P66
    |--E. rabdotacona Gardner 1937 [=Agladrillia (Eumetadrillia) rabdotacona]P66
    `--E. waltoniana (Gardner 1937) [=Drillia waltoniana, Agladrillia (Eumetadrillia) waltoniana]P66
  AustroclavusP66
  ElaeocymaBK11
  CymatosyrinxBK11
  Acinodrillia Kilburn 1988BK11
  Bellaspira Conrad 1868BK11
    |--*B. virginiana (Conrad 1862) [=Mangelia virginiana]P66
    |--B. acclivicosta McLean & Poorman 1970BC01
    `--B. pentapleura Schwengel 1940P66
  Orrmaesia Kilburn 1988BK11
  Stenodrillia Korobkov 1955BK11
    `--*S. allionii (Bellardi 1877) [=Drillia allionii]P66
  SyntomodrilliaBK11
Nomen nudum: Drillia albolaqueata Carpenter 1864C64

Crassopleura costulatum (Cantraine 1835) [=Pleurotoma costulatum; incl. P. elegans Scacchi 1835, P. incisa Reeve 1843, P. incrassata Dujardin 1837 non Sowerby 1834, P. maravignae Bivona 1838, *Crassopleura maravignae]P66

Drillia roseola (Hertlein & Strong 1955)BC01 [=Cymatosyrinx roseolaBC01; incl. Pleurotoma rosea Sowerby 1834 non Quoy & Gaimard 1833P66]

Imaclava pembertoni (Lowe 1935) [=Clavus pembertoni, Clathrodrillia (Imaclava) pembertoni; incl. *Imaclava ima Bartsch 1944]P66

Imaclava unimaculata (Sowerby 1834)PK11 [=Pleurotoma unimaculataP66, Clathrodrillia (Imaclava) unimaculatusP66]

*Neodrillia cydia Bartsch 1943 [incl. N. antiguensis Bartsch 1943, N. barbadensis Bartsch 1943, N. encia Bartsch 1943, N. jamaicensis Bartsch 1943]P66

*Paracuneus immaculatus (Tenison-Woods 1876)P66 [=Mangelia immaculataP66, Drillia immaculataH22, Inquisitor immaculatusL54; incl. Drillia gabrieli Pritchard & Gatliff 1899P66]

Spirotropis centimata (Dall 1889) [=Drillia (Cymatosyrinx) centimata; incl. Pleurotoma centimata var. major Locard 1897]BW80

Spirotropis modiola (Jan 1832)BW80 [=Fusus modiolusBW80, Pleurotoma modiolaBW80; incl. P. acuta Bellardi 1842 non Perry 1811P66, P. carinata Bivona 1838 non Gray in Griffith & Pidgeon 1834P66, P. scalaris Partsch 1837P66]

Spirotropis monterosatoi (Locard 1897) [=Pleurotoma monterosatoi; incl. P. carinata Sars 1878 nec Gray in Griffith & Pidgeon 1834 nec Bivona 1838, *Spirotropis carinata, Mangelia eburnea Sars 1859 non Bivona 1838, S. sarsi Warén 1975]BW80

*Tripia anteatripla de Gregorio 1890 (n. d.) [=Pleurotoma (Tripia) anteatripla, Crassispira (Tripia) anteatripla]P66

*Type species of generic name indicated

References

[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.

[BW80] Bouchet, P., & A. Waren. 1980. Revision of the north-east Atlantic bathyal and abyssal Turridae (Mollusca, Gastropoda). Journal of Molluscan Studies Supplement 8: 1–119.

[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. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488–533.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

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

[H22] Hedley, C. 1922. A revision of the Australian Turridae. Records of the Australian Museum 13 (6): 213–359, pls 42–56.

[H86] Hutton, F. W. 1886. The Mollusca of the Pareora and Oamaru systems of New Zealand. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 205–237.

[K65] Kay, E. A. 1965. Marine molluscs in the Cuming collection, British Museum (Natural History) described by William Harper Pease. Bulletin of the British Museum (Natural History): Zoology Supplement 1: 1–96, 14 pls.

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

[M23] Melvill, J. C. 1923. Descriptions of twenty-one species of Turridae (Pleurotomidae) from various localities in the collection of Mr. E. R. Sykes. Proceedings of the Malacological Society of London 15 (4): 162–171.

[PMB15] Petuch, E. J., R. F. Myers & D. P. Berschauer. 2015. The Living and Fossil Busycon Whelks: Iconic Mollusks of Eastern North America. San Diego Shell Club, Inc.

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

[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.

[SK81] Shimek, R. L., & A. J. Kohn. 1981. Functional morphology and evolution of the toxoglossan radula. Malacologia 20 (2): 432–438.

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