Paradrillia lithoria, from Melvill & Standen (1903).

Belongs within: Conoidea.
Contains: Marshallena, Aoteadrillia, Austrodrillia, Inodrillia, Mauidrillia, Anacithara (Anacithara).

The Horaiclavidae are a group of conoid gastropods generally bearing small, stout shells with a short siphonal canal (Bouchet et al. 2011).

Published 26 August 2013
Paradrillia patruelis, from Joop Trausel and Frans Slieker.

Paradrillia is a genus of conoid gastropods found in the Indo-Pacific region, with a fossil record going back to the Miocene (Powell 1966). Species of Paradrillia are small shells, about one to three centimetres in length, with a relatively tall spire and short siphonal canal. The sculpture on the outside of the shell usually consists of nodular spirals; the operculum is leaf-like, with a terminal nucleus. The long, awl-shaped radular teeth are trough-shaped in cross-section (Kilburn 1988).

Paradrillia regia, from G. & Ph. Poppe.

The classification of Paradrillia has shifted around over the years, like most of the less differentiated conoids that were lumped by Powell (1966) under the heading of ‘Turridae’. Powell (1966) classed them as Turriculinae, Kilburn (1988) transferred them to the Stictispirinae. Kilburn also synonymised Powell’s separate genera Paradrillia and Vexitomina. Powell had distinguished them on the basis that Paradrillia supposedly had an operculum with a mediolateral nucleus instead of the terminal nucleus of Vexitomina. However, this was based on a single specimen that Kilburn regarded as teratological after he found terminal nuclei in Paradrillia melvilli. Most recently, Paradrillia was placed by by Bouchet et al. (2011) on the basis of molecular data in their new family Horaiclavidae.

Systematics of Horaiclavidae

Characters (from Bouchet et al. 2011): Shell generally small, 5–25 mm, usually 7–15 mm high, shortly claviform, with relatively low spire and a short, truncated, poorly differentiated siphonal canal. Subsutural ramp usually poorly differentiated. Axial sculpture almost always present, usually as strong sinuate ribs. Spiral sculpture normally weak or obsolete, often with glossy shell surface. Anal sinus on subsutural slope, weak to moderately deep, often constrained by callus. Protoconch of up to 3.5 medially carinate but otherwise smooth whorls when multispiral, but usually paucispiral and smooth. Operculum with terminal nucleus. Radular formula 1-0-0-0-1. Marginal teeth duplex, with lanceolate major limb and usually narrow accessory limb, which is inserted in a shallow socket. Major limb often with angulation lateral to the place of accessory limb insertion. Rarely the additional limb is of similar size to the major limb and the teeth become trough-shaped in transverse section, with a collar near the base. Radular apparatus occasionally absent.

    |--+--+--Carinapex Dall 1924PK11, BK11
    |  |  |    `--*C. minutissima (Garrett 1873)P66 [=Drillia minutissimaP66, Iredalea minutissimaH22]
    |  |  `--Ceritoturris Dall 1924PK11, BK11
    |  |       |--*C. bittium Dall 1924P66
    |  |       |--C. nataliae Kilburn 1988BD12
    |  |       `--C. pupiformis (Smith 1884)PK11
    |  `--+--Anguloclavus Shuto 1983PK11, BK11
    |     `--+--Horaiclavus Oyama 1954PK11, BK11
    |        |    |--*H. splendidus (Adams 1867) [=Mangelia splendida]P66
    |        |    `--H. madurensis (Schepman 1913) [=Drillia madurensis]P66
    |        `--Anacithara Hedley 1922PK11, BK11
    |             |  i. s.: A. dulcinea (Melvill & Standen 1895) [=Mangilia dulcinea]H22
    |             |         A. ione (Melvill & Standen 1896) [=Drillia ione]H22
    |             |         A. leptalea Hedley 1922H22
    |             |         A. nanisca (Hervier 1897) [=Mangilia nanisca]H22
    |             |         A. osumiensis (Sowerby 1913) [=Mangilia osumiensis]H22
    |             |         A. themeropsis (Melvill & Standen 1896)BK11
    |             |         A. undaticosta (Reeve 1845) [=Pleurotoma undaticosta, Mangelia undaticosta]H22
    |             |--A. (Anacithara)P66
    |             `--A. (Anacitharoida Shuto 1965)P66
    |                  `--A. (*A.) kurodae Shuto 1965P66
    `--Paradrillia Makiyama 1940PK11, BK11 [incl. Alticlavatula MacNeil 1960BK11, Vexitomina Powell 1942BK11]
         |--*P. dainichiensis (Yokoyama 1923) [=Drillia dainichiensis]P66
         |--P. agalma (Smith 1906) [=Pleurotoma (Surcula) agalma]P66
         |--P. alluaudi (Dautzenberg 1932) [=Drillia alluaudi]P66
         |--P. astuta (Yokoyama 1928) [=Drillia astuta]P66
         |--P. astutoida (Shuto 1961) [=Clavatula (Paradrillia) astutoida]P66
         |--P. celebensis (Schepman 1913) [=Surcula celebensis]P66
         |--P. convexiuscula (Shuto 1961) [=Clavatula (Paradrillia) convexiuscula]P66
         |--‘Inquisitor’ coriorudis Hedley 1922 [=Vexitomina coriorudis]P66
         |--‘Drillia’ coxi Angas 1867P66 [=Vexitomina coxiP66, Inquisitor coxiH22; incl. D. agnewi Tenison-Woods 1879H22]
         |--P. djocdjocartae (Martin 1884) [=Drillia (Pleurotoma) djocdjocartae]P66
         |--P. ermelingi (Martin 1884) [=Pleurotoma (Drillia) ermelingi]P66
         |--P. fulva (Hinds 1843) (n. d.) [=Clavatula fulva, Clavus fulvus]KD08
         |--‘Vexitomina’ garrardi Laseron 1954P66
         |--P. himea (Makiyama 1927) [=Clavatula himea]P66
         |--P. inconstans (Smith 1875) [=Pleurotoma inconstans; incl. Clavatula gaylordae Preston 1905]P66
         |    |--P. i. inconstansP66
         |    `--P. i. prunulum (Melvill & Standen 1901) [=Drillia prunulum]P66
         |--P. kakegawensis (Makiyama 1927) [=Clavatula kakegawaensis]P66
         |--P. lithoria (Melvill & Standen 1903) [=Drillia lithoria]P66
         |--‘Drillia’ metcalfei Angas 1867P66 [=Inquisitor metcalfeiH22, Pleurotoma metcalfeiH22, *Vexitomina metcalfeiP66]
         |--P. minoensis (Shuto 1961) [=Clavatula (Paradrillia) minoensis]P66
         |--P. nivalioides (Yokoyama 1920) [=Pleurotoma (Drillia) nivalioides]P66
         |--‘Drillia’ optabilis Murdoch & Suter 1906 [=Vexitomina optabilis]P66
         |--P. patruelis (Smith 1875) [=Pleurotoma patruelis, *Alticlavatula patruelis; incl. Pl. consimilis Smith 1879]P66
         |--‘Vexitomina’ pilazona Laseron 1954P66
         |--‘Pleurotoma’ regia Reeve 1842P66 (see below for synonymy)
         |--‘Pleurotoma (Drillia) suavis Smith 1888P66 (see below for synonymy)
         |--P. taiwanensis (Nomura 1935) [=Clavatula taiwanensis]P66
         `--‘Vexitomina’ torquata Laseron 1954P66
Horaiclavidae incertae sedis:
  Inkinga Kilburn 1988BK11
  Coronacomitas Shuto 1983BK11
  Cytharoclavus Kuroda & Oyama in Kuroda, Habe & Oyama 1971BK11
  Darrylia García 2008BK11
  Graciliclava Shuto 1983BK11
  Naskia Sysoev & Ivanov 1985BK11
  Nquma Kilburn 1988BK11
  Psittacodrillia Kilburn 1988BK11
  Striatoguraleus Kilburn 1994BK11
  Iwaoa Kuroda 1940BK11
    `--*I. reticulata Kuroda 1953P66
  Austrocarina Laseron 1954BK11
    `--*A. recta (Hedley 1903)P66 [=Leucosyrinx rectaP66, Filodrillia rectaL54]
  Micropleurotoma Thiele 1929BK11
    |--*M. spirotropoides (Thiele 1925) [=Pleurotoma spirotropoides]BW80
    |--M. ashiyaensis Shuto & Ueda 1963P66
    |--M. melvilli (Sykes 1906) [=Spirotropis melvilli; incl. Pleurotoma nana Thiel 1925]BW80
    `--M. travailleuri Bouchet & Waren 1980 [=Pleurotoma obtusa Locard 1897 non Reeve 1846]BW80
  Pseudexomilus Powell 1944BK11
    |--*P. caelatus Powell 1944P66
    `--P. costicapitata (Verco 1909)P66 [=Drillia costicapitataP66, Filodrilla costicapitataH22]
  Thelecythara Woodring 1928BK11
    |--*T. mucronata (Guppy 1896) [=Cythara mucronata]P66
    `--T. floridana Fargo 1953P66 [incl. T. cruzensis Usticke 1969BC01]
  Haedropleura Monterosato 1883BR17
    |--*H. septangularis (Montagu 1803)BR05 (see below for synonymy)
    |    |--H. s. septangularisP66
    |    `--H. s. rigida (Reeve 1846) [=Mangelia rigida]P66
    |--H. contii (Bellardi 1878) [=Bela contii]P66
    |--H. dora Thiele 1925P66
    |--H. proxima (Peyrot 1938) [=Bela (Haedropleura) proxima]P66
    `--H. secalina (Philippi 1844) [=Pleurotoma secalinum]P66
  Buchema Corea 1934BK11
    |--*B. tainoa (Corea 1934) [=Carinodrillia (*Buchema) tainoa]P66
    |--B. apitoa (Corea 1934) [=Carinodrillia (Buchema) apitoa]P66
    |--B. interpleura (Dall & Simpson 1901)BK11
    |--B. interstrigata (Smith 1882) [incl. Monilispira circumcincta Usticke 1969]BC01
    |--B. liella (Corea 1934) [=Carinodrillia (Buchema) liella]P66
    |--B. mamona (Corea 1934) [=Carinodrillia (Buchema) mamona]P66
    |--B. pylonia Fargo 1953 [=Carinodrillia (Buchema) pylonia]P66
    `--B. suimaca (Corea 1934) [=Carinodrillia (Buchema) suimaca]P66

*Haedropleura septangularis (Montagu 1803)BR05 [=Murex septangularisBR05, Bela septangularisBR05, Pleurotoma septangularisP66; incl. Mangilia goreensis Maltzan 1883P66, P. heptagona Scacchi 1835P66, Cythara maltzani Knudsen 1952P66, Mangelia plicatilis Risso 1826P66]

‘Pleurotoma’ regia Reeve 1842P66 [=Vexitomina regiaP66; incl. P. rougeyroni Souverbie 1894P66, Turridrupa rougeyroniH22]

‘Pleurotoma (Drillia) suavis Smith 1888P66 [=Inquisitor suavisH22, Vexitomina suavisP66; incl. Drillia prosuavis Hedley 1903P66]

*Type species of generic name indicated


[BD12] Bandel, K., & D. T. Dockery III. 2012. Protoconch characters of Late Cretaceous Latrogastropoda (Neogastropoda and Neomesogastropoda) as an aid in the reconstruction of the phylogeny of the Neogastropoda. Freiberger Forschungshefte C542 (psf 20): 93–128.

[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.

[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.

[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.

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

Kilburn, R. N. 1988. Turridae (Mollusca: Gastropoda) of southern Africa and Mozambique. Part 4. Subfamilies Drilliinae, Crassispirinae and Strictispirinae. Annals of the Natal Museum 29 (1): 167–320.

[KD08] Kilburn, R. N., & H. Dekker. 2008. New species of turrid conoideans (Gastropoda: Conoidea) from the Red Sea and Arabia. Basteria 72 (1–3): 1–19.

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

[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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