Belongs within: Conoidea.
Contains: Fenimorea, Austroclavus, Elaeocyma, Cymatosyrinx, Syntomodrillia, Fusiturricula, Clathrodrillia, Drillia (Drillia), Agladrillia, Splendrillia (Splendrillia), Clavus, Cerodrillia.
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).
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]
|--+--Cruziturricula Marks 1951 PK11, BK11
| | |--*C. cruziana (Olsson 1932) [=Turricula (Pleurofusia) cruziana] P66
| | |--C. andesita (Olsson 1942) [=Turricula (Knefastia) andesita] P66
| | |--C. arcuata (Reeve 1843) [=Pleurotoma arcuata] P66
| | |--C. eolavinia (Olsson 1930) [=Turricula (Pleurofusia) eolavinia] P66
| | |--C. lavinia (Dall 1919) [=Turricula (Surcula) lavinia] P66
| | `--C. piura (Olsson 1931) [=Turricula (Surcula) piura] P66
| `--Fusiturricula PK11
`--+--+--+--Clathrodrillia PK11
| | `--Drillia Gray 1838 PK11, BR05
| | | i. s.: D. alabaster (Reeve 1843) [=Pleurotoma alabaster] H09
| | | D. chaaci Espinosa & Rolán 1995 BC01
| | | D. essingtonensis (Smith 1888) [=Pleurotoma essingtonensis] H09
| | | D. exigua (Hombron & Jacq. 1853) [=Pleurotoma exigua] H09
| | | D. kleinrosa Usticke 1969 BC01
| | | D. laterculata (Sowerby 1870) [=Pleurotoma laterculata] H09
| | | D. nitens (Hinds 1843) [=Clavatula nitens] H09
| | | ‘Pleurotoma (Drillia) parva’ Smith 1888 non P. parva Conrad 1830 F27
| | | D. ponciana BC01
| | | |--D. p. ponciana BC01
| | | `--D. p. virgata Usticke 1969 [=D. bandata Usticke 1971] BC01
| | | D. ‘punctata’ (Reeve 1845) [=Pleurotoma punctata non Schubert & Wagner 1829] H09
| | | D. putilla (Reeve 1845) [=Pleurotoma putilla] H09
| | | D. radula (Hinds 1843) [=Pleurotoma radula] H09
| | | D. roseola (Hertlein & Strong 1955) BC01 (see below for synonymy)
| | | D. spaldingi Brazier 1876 H09
| | | D. stiza Tenison-Woods 1879 F71
| | | D. wolfei Tippett 1995 BC01
| | |--D. (Drillia) P66
| | `--D. (Pseudodrillia Lukovic 1924) P66
| | |--D. (P.) abnormalis Lukovic 1924 P66
| | |--D. (P.) arabica Lukovic 1924 P66
| | `--D. (P.) longa Lukovic 1924 P66
| `--+--Agladrillia PK11
| `--+--Imaclava Bartsch 1944 PK11, BK11
| | |--I. pembertoni (Lowe 1935) (see below for synonymy) P66
| | |--I. pilsbryi Bartsch 1950 [=Clathrodrillia (Imaclava) pilsbryi non C. pilsbryi Lowe 1935] P66
| | `--I. unimaculata (Sowerby 1834) BK11 (see below for synonymy)
| `--Calliclava McLean 1971 PK11, BK11
| |--C. alcmene (Dall 1919) PP78
| `--C. jaliscoensis McLean & Poorman 1971 BC01
`--+--Conopleura Hinds 1844 PK11, BK11
| `--*C. striata Hinds 1844 [=Pleurotoma partita Reeve 1846] P66
`--+--Splendrillia Hedley 1922 PK11, P66
| | i. s.: S. bratcherae McLean & Poorman 1971 PP78
| | S. petiti (Tippett 1995) [=Fenimorea petiti] BC01
| |--S. (Splendrillia) P66
| `--S. (Hauturua Powell 1942) P66
| |--S. (*H.) vivens (Powell 1942) [=Syntomodrillia (*H.) vivens] P66
| |--S. (H.) exiguescens (Marwick 1931) [=Austrodrillia exiguescens] P66
| |--S. (H.) exuta Powell 1944 P66
| `--S. (H.) laevella (Marwick 1931) [=Austrodrillia laevella] P66
`--+--Iredalea Oliver 1915 PK11, P66 [incl. Brephodrillia Pilsbry & Lowe 1932 P66]
| |--*I. subtropicalis Oliver 1915 P66
| |--I. acuminata (Mighels 1848) [=Pleurotoma acuminata] P66
| |--I. ella (Pilsbry & Lowe 1932) [=Brephodrillia ella] P66
| |--I. macleayi (Brazier 1876) [=Clathurella macleayi] P66
| |--I. perfectus (Pilsbry & Lowe 1932) [=*Brephodrillia perfectus] P66
| |--I. pupoidea (Adams 1872) [=Zafra pupoidea; incl. Mangilia victor Sowerby 1894] P66
| `--I. pusilla (Garrett 1873) [=Drillia pusilla] P66
`--+--Clavus PK11
`--Cerodrillia PK11
Drilliidae incertae sedis:
Clavatoma Powell 1942 P66
`--*C. pulchra Powell 1942 P66
Maoricrassus Vella 1954 P66
`--*M. carinatus Vella 1954 P66
Tahudrillia Powell 1942 P66
`--*T. simplex Powell 1942 P66
Integradrillia Powell 1942 P66
`--*I. integra (Tenison-Woods 1880) [=Drillia integra] P66
Vixinquisitor Powell 1942 P66
`--*V. vixumbilicata (Harris 1897) [=Drillia vixumbilicata] P66
Globidrillia Woodring 1928 P66
`--*G. ula Woodring 1928 P66
Tripia Gregorio 1890 (n. d.) P66
`--*T. anteatripla de Gregorio 1890 (n. d.) (see below for synonymy) P66
Boreodrillia Sorgenfrei 1958 P66
`--*B. toftlundensis Sorgenfrei 1958 P66
Mitrelloturris Eames 1957 P66
`--*M. casteri (Chavan 1952) [=Asthenotoma (Endiatoma) casteri] P66
Terebritoma Cossmann 1894 P66
`--*T. solitaria [=Mangelia solitaria] P66
Neodrillia Bartsch 1943 P66
|--*N. cydia Bartsch 1943 (see below for synonymy) P66
`--N. euphanes (Melvill 1923) [=Drillia euphanes] P66
Wairarapa Vella 1954 P66
|--*W. rebecca Vella 1954 P66
`--‘Melatoma’ duplaris Hedley 1922 P66
Douglassia Bartsch 1934 P66
|--*D. enae Bartsch 1934 P66
`--D. bealiana Schwengel & McGinty 1942 P66
Crassopleura Monterosato 1884 [=Crossopleura (l. c.)] P66
|--C. costulatum (Cantraine 1835) (see below for synonymy) P66
`--C. brevis (Yokoyama 1922) [=Drillia glabriuscula brevis] P66
Leptadrillia Woodring 1928 P66
|--*L. parkeri (Gabb 1873) [=Turris (Surcula) parkeri] P66
|--L. loria Bartsch 1934 P66
`--L. splendida Bartsch 1934 P66
Kylix Dall 1919 P66
|--*K. alcyone (Dall 1919) [=Clathrodrillia (Kylix) alcyone] P66
|--K. alcmene (Dall 1919) [=Clathrodrillia (Kylix) alcmene] P66
|--K. contracta McLean & Poorman 1971 BC01
|--K. paziana (Dall 1919) PP78
|--K. turveri Hertlein & Strong 1951 P66
`--K. zacae Hertlein & Strong 1951 P66
Tylotiella Habe 1964 BK11
|--*T. subobliquata (Smith 1879) [=Drillia subobliquata] P66
|--T. humilis (Smith 1879) [=Drillia humilis] P66
|--‘Drillia’ longispira Smith 1879 [=Leiocithara longispira, Rubellatoma longispira] K92
|--T. pica (Reeve 1843) [=Pleurotoma pica] P66
`--T. sacra (Reeve 1845) [=Pleurotoma sacra] P66
Plagiostropha Melvill 1927 BK11
|--*P. quintuplex Melvill 1927 [=Clavus (*Plagiostropha) quintuplex] P66
|--P. caledoniensis Wells 1995 BK11
|--‘Pleurotoma’ ebur Reeve 1845 P66
`--‘Pleurotoma’ opalus Reeve 1845 P66
Paracuneus Laseron 1954 P66
|--*P. immaculatus (Tenison-Woods 1876) P66 (see below for synonymy)
| |--P. i. immaculatus P66
| `--P. i. peronianus Laseron 1954 P66
|--P. kemblensis Laseron 1954 P66
`--P. spadix (Watson 1886) [=Pleurotoma (Drillia) spadix] P66
|--P. s. spadix P66
`--P. s. tumulus Laseron 1954 P66
Spirotropis Sars 1878 P66
|--*S. ‘carinata’ (Philippi 1844) (see below for synonymy) P66
|--S. clytotropis Sykes 1906 P66
|--S. ephamilla Verrill 1884 P66
|--S. melvilli Sykes 1906 P66
|--S. modiolus (Cristofori & Jan 1832) (see below for synonymy) P66
|--S. monterosatoi (Locard 1897) [=Pleurotoma monterosatoi] P66
|--S. remota Powell 1958 P66
`--S. studeriana (Martens 1878) [=Pleurotoma studeriana] P66
Sedilia Fargo 1953 P66
|--*S. sedilia (Dall 1890) [=Drillia sedilia] P66
|--S. hoplophorus (Dall 1892) [=Drillia hoplophorus] P66
|--S. ochoidia Fargo 1953 P66
|--S. ondulum Fargo 1953 P66
|--S. sapa Fargo 1953 P66
| |--S. s. sapa P66
| `--S. s. sapala Fargo 1953 P66
|--S. schismatica (Dall 1892) [=Drillia schismatica] P66
`--S. transa Fargo 1953 P66
Fenimorea P66
Eumetadrillia Woodring 1928 BK11
|--*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) (see below for synonymy) 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
Austroclavus P66
Elaeocyma P66
Cymatosyrinx P66
Acinodrillia Kilburn 1988 BK11
Bellaspira Conrad 1868 BK11
|--*B. virginiana (Conrad 1862) [=Mangelia virginiana] P66
|--B. acclivicosta McLean & Poorman 1970 BC01
`--B. pentapleura Schwengel 1940 P66
Orrmaesia Kilburn 1988 BK11
Stenodrillia Korobkov 1955 BK11
`--*S. allionii (Bellardi 1877) [=Drillia allionii] P66
Syntomodrillia BK11
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 roseola BC01; incl. Pleurotoma rosea Sowerby 1834 non Quoy & Gaimard 1833 P66]
Eumetadrillia fuegiensis (Smith 1888) [=Pleurotoma (Surcula) fuegiensis, Agladrillia (Eumetadrillia) fuegiensis] P66
Imaclava pembertoni (Lowe 1935) [=Clavus pembertoni, Clathrodrillia (Imaclava) pembertoni; incl. *Imaclava ima Bartsch 1944] P66
Imaclava unimaculata (Sowerby 1834) BK11 [=Pleurotoma unimaculata P66, Clathrodrillia (Imaclava) unimaculatus P66]
*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 immaculata P66, Inquisitor immaculatus L54; incl. Drillia gabrieli Pritchard & Gatliff 1899 P66]
*Spirotropis ‘carinata’ (Philippi 1844) [=Pleurotoma carinatum nec Gray in Griffith & Pidgeon 1834 nec Bivona 1838] P66
Spirotropis modiolus (Cristofori & Jan 1832) [=Fusus modiolus; incl. Pleurotoma acuta Bellardi 1842 non Perry 1811, P. carinata Bivona 1838 non Gray in Griffith & Pidgeon 1834, P. scalaris Partsch 1837] P66
*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.
[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.
[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.
[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.
[K92] Kilburn, R. N. 1992. Turridae (Mollusca: Gastropoda) of southern Africa and Mozambique. Part 6. Subfamily Mangeliinae, section 1. Annals of the Natal Museum 33 (2): 461–575.
[L54] Laseron, C. F. 1954. Revision of the New South Wales Turridae (Mollusca). Royal Zoological Society of New South Wales: Sydney.
[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.
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