Belongs within: Conoidea.
Contains: Coronia, Cryptogemma, Pleuroliria, Eopleurotoma (Eopleurotoma), Amuletum, Polystira, Turridrupa, Turris, Lophiotoma, Gemmula (Gemmula).
The Turridae are a group of marine gastropods previously recognised as including all non-conid and non-terebrid conoids but now restricted to a monophyletic group with a usually narrowly fusiform shell bearing obsolete axial sculpture and a peripheral anal sinus (Bouchet et al. 2011). These include the warm-water genus Gemmula in which the fusiform shell bears a peripheral keel or keels strongly studded with gemmules. The Indo-West Pacific genus Unedogemmula resembles Gemmula in the presence of a peripheral keel but largely lacks gemmules beyond distinct granulations on the early whorls (Powell 1966).
Fifteen seconds of mediocrity
Published 14 June 2011
This is going to be another one of those posts where I try to actually build something on the back of some obscure taxon about which I’ve only managed to find enough to fill about two sentences. You have been warned.
Kuroshioturris asukana was described as Drillia asukana by Yokoyama in 1926 in the Journal of the Faculty of Science, Imperial University of Tokyo. That might in itself explain why I haven’t been able to access the original description (for my American readers, the appropriate issue is on Google Books here, in case this is another one that Google is only making available within the United States). The species is a Pliocene member from Japan of the gastropod superfamily Conoidea, the ‘poison-tongued’ gastropods, in the family Turridae. The exact position of asukana in the turrids has shifted around a bit: it has appeared in Clavatula, in Gemmula, in (the most horrifying turrid genus of all) Pleurotoma, before Powell (1966) placed it into Kuroshioturris (treated by Powell as a subgenus of Ptychosyrinx, but as a separate genus by later authors such as Beu 2011). A second specimen other than the holotype assigned by Yokoyama to Pleurotoma asukana in 1928 may not in fact be the same species, and MacNeil (1960) noted that the holotype appeared to be a juvenile, further complicating identification.
Powell (1966) stated that Kuroshioturris was restricted to the Miocene and Pliocene of Japan, but this was an error. Two of the species listed by Powell (including the type species of the genus) are in fact members of the Recent fauna. Beu (2011) also assigned two species from the Pliocene to Recent of New Zealand to Kuroshioturris. Powell distinguished Kuroshioturris from Ptychosyrinx sensu stricto on the basis of the protoconch morphology, noting that the adult shell was otherwise nearly identical. However, other cases of conoid genera being separated solely on the basis of protoconch form have since been regarded as invalid (Beu 2011). Protoconch morphology often reflects to mode of larval lifestyle for the animal: a tall, narrow protoconch (such as in Ptychosyrinx) indicates an actively feeding, planktotrophic larva, while a blunt, short protoconch indicates a lecithotrophic larva nourished by yolk reserves. Changes from one nutritional mode to another seem to have happened repeatedly, leading to cases such as the lecithotrophic ‘genus’ Maoritomella turning out to be a polyphyletic assemblage of Tomopleura species that had independently abandoned planktotrophy. Many such genera have now been synonymised, but many (such as Kuroshioturris?) still require examination.
The turrid core
Published 15 February 2018
At this point, I’ve made numerous references on this site to the gastropod family Turridae, discussing its members and non-members and alluding to its sordid history. So maybe I should set out the basics of the story properly.
The Conoidea are a diverse group of marine predatory gastropods with over 4000 known living species. They are best known for the production by many species of venom used to paralyse their prey, in some species being potent enought to threaten humans. In the majority of conoideans, this venom is delivered via a tooth that becomes detached from the radula and is held at the end of the retractable proboscis. Until relatively recently, Conoidea were commonly divided between three families. Two of these families, the Conidae (cone shells) and Terebridae (awl shells) were well defined and constrained. The third family was the Turridae, including by far the greater number of species but not really defined within Conoidea beyond ‘the rest’. Many of ‘the rest’ were small, many were restricted to deep water, many were poorly known. Different systems were proposed over the years in an attempt to break the turrid mass into more manageable units but each system differed significantly from the next and no one system became universally accepted. Some authors would focus on the protoconch as their guide to classification, others would focus on the radula, others might call out features of the operculum. One author commented in 1922 that turrids were “considered by those who meddle with them to be more perplexing than any other molluscan family“, and this complaint was still being upheld by Kilburn (1983) over sixty years later.
Though it had long been accepted that the ‘turrids’ probably did not represent an evolutionarily coherent group, it wasn’t really until the advent of molecular phylogenies that things started falling into place. Puillandre et al. (2011) identified two main lineages within the Conoidea, leading to the dissolution of the original Turridae into no less than 13 families in order to maintain the already-established Conidae and Terebridae. Turridae in the strict sense was restricted to a much smaller clade of a bit over a dozen genera, sister to the Terebridae (Bouchet et al. 2011).
In contrast to the bewilderment of the original turrid array, Turridae sensu Bouchet et al. is a morphologically quite coherent group. They are more or less fusiform (spindle-shaped) shells, often with a narrow, high spire and relatively weak sculpture. Indeed, but for the fact that most tend to have a long siphonal canal at the base of the shell, they often bear a distinct resemblance to their terebrid sisters. The majority of turrids have a multispiral protoconch, indicating an extended, planktonic-feeding larval stage in development, but there are some species with a paucispiral protoconch indicative of direct development.
The radula of turrids usually comprises three apparent teeth in each row. The central tooth is actually formed from three teeth (the original pointed central tooth and two plate-like lateral teeth) fused together; in some species the division between these teeth remains visible whereas in others the central tooth disappears entirely. The main business part of the radula is the single pair of marginal teeth which, as in other conoideans, are enlarged and modified for venom delivery. They have a distinctive ‘duplex’ form; in older publications, this was referred to as a ‘wishbone’ form because the tooth appears under light microscopy to be divided between two branches. After the advent of electron microscopy, it was discovered that these two ‘branches’ in fact represent the thickened margins of an undivided tooth. The larger of the two margins is mostly attached to the radular membrane with only the tip of the tooth being free; the smaller margin is held free of the radula. The thinner part of the tooth between the two margins forms a gutter along which venom can flow. However, the radula is placed in such a position that it cannot be protruded through the mouth in the manner of grazing gastropods. As with other conoideans, prey (in this case probably worms) is despatched through the use of a detached marginal tooth transferred to the end of the proboscis. However, whereas other conoideans such as cone shells may have the tooth functioning like a hypodermic syringe for delivering prey, turrids use their tooth to slash at the prey like a switchblade, with venom passively entering through the resulting cuts. The proboscis is then used to draw the prey back into the mouth, where the radula is used to grasp and swallow it, sucking the unlucky worm down the gullet like spaghetti.
Systematics of Turridae
Characters (from Bouchet et al. 2011): Shell of medium to large size (usually 20–30 mm, up to 110 mm high), short- to high-fusiform, usually with high spire and long (rarely short and truncated) siphonal canal. Axial sculpture weak or absent. Anal sinus on whorl periphery. Protoconch typically multispiral, up to six whorls, protoconch I smooth, protoconch II with arcuate axial riblets; reduced paucispiral protoconch usually smooth, may have arcuate axial riblets. Operculum fully developed, with terminal nucleus. Radular formula typically 1-(1:R:1)-1. Small and narrow rachidian and plate-like laterals fused together, together constituting central formation of different development, varying from well-defined broad central tooth to tooth clearly formed of three elements through gradual reduction of rachidian and/or laterals to complete absence. Marginals duplex, of variable morphology, from broadly oval and flattened with nearly equally developed limbs to awl-shaped and divided only in basal part; major limb most commonly large and knife-shaped, accessory limb dorsal and more weakly developed.
<==Turridae [Pleurotomaceae, Pleurotomidae, Pleurotomina, Pleurotominae, Turritidae, Turrini, Turritinae]PK11 |--AmuletinaeBR17 | |--AmuletumBD12 | `--Graphidula Stephenson 1941 non Norman 1853 (ICBN)BR17, BD12 [Graphidulidae] | `--*G. terebriformis Stephenson 1941BD12 `--TurrinaeP66 |--PolystiraPK11 `--+--Ptychosyrinx Thiele 1925PK11, BK11 [incl. Bathybermudia Haas 1949BK11] | |--*P. bisinuata (Martens 1901) [=Pleurotoma (Subulata) bisinuata]P66 | |--P. carynae (Haas 1949)PK11 [=*Bathybermudia carynaeP66] | |--P. chilensis Berry 1968BK11 | |--P. lobata (Sowerby 1903) [=Pleurotoma (Surcula) lobata]P66 | |--P. timorensis (Tesch 1915) [=Pleurotoma timorensis]P66 | | |--P. t. timorensisP66 | | `--P. t. teschi Powell 1964P66 | `--P. truncata (Schepman 1913) [=Pleurotoma truncata]P66 `--+--TurridrupaPK11 `--+--+--TurrisPK11 | `--+--Iotyrris Medinskaya & Sysoev 2001PK11, BK11 | | `--I. cingulifera (Lamarck 1822)PK11 | `--Xenuroturris Iredale 1929PK11, BK11 [incl. Clamturris Iredale 1931BK11] | |--X. cingulifera (Lamarck 1822) [=Pleurotoma cingulifera; incl. *X. legitima Iredale 1929]P66 | | |--X. c. cinguliferaP66 | | `--X. c. erythraea (Weinkauff 1875) [=Pleurotoma erythraea]P66 | |--X. castanella Powell 1964P66 | |--X. cerithiformis Powell 1964P66 | |--X. incredula (Iredale 1931) [=*Clamturris incredula]P66 | |--X. kingae Powell 1964P66 | |--X. legitima Iredale 1929PK11 | `--X. millepunctata (Sowerby 1908) [=Pleurotoma millepunctata]P66 `--+--LophiotomaPK11 `--Gemmula Weinkauff 1875PK11, BK11 [incl. Eugemmula Iredale 1931BK11] | i. s.: G. complicataA27 | G. cretacea Sohl 1964BD12 | G. lululimi Olivera 2000 (see below for synonymy)KFO12 | G. mystica Simone 2005S11 | G. rarimaculata (Kuroda & Oyama 1971)PK11 | G. westaustralis Kosuge 1990MG-H11 |--G. (Gemmula)BK11 `--G. (Pinguigemmula MacNeil 1960)BK11 |--‘*Pinguigemmula’ okinavensis MacNeil 1960P66 |--‘Pinguigemmula’ luzonica Powell 1964P66 |--‘Pinguigemmula’ philippinensis Powell 1964MG-H11 `--G. (P.) thielei Finlay 1930 (see below for synonymy)P66 Turridae incertae sedis: Unedogemmula MacNeil 1961BK11 |--*U. unedo (Kiener 1839)P66, MG-H11 (see below for synonymy) |--U. bemmeleni (Oostingh 1941) [=Turris (Gemmula) bemmeleni]P66 |--U. deshayesii (Doumet 1839) [=Pleurotoma deshayesii, Gemmula (Unedogemmula) deshayesii]P66 |--U. hastula (Reeve 1843) (see below for synonymy)P66 |--U. haydeni (Vredenburg 1925) [=Pleurotoma haydeni, Gemmula (Unedogemmula) haydeni]P66 |--U. ickei (Martin 1906) [=Pleurotoma ickei, Gemmula (Unedogemmula) ickei]P66 |--U. ina MacNeil 1960 [=Gemmula (Unedogemmula) ina]P66 |--U. koolhoveni (Oostingh 1938) [=Turris (Gemmula) koolhoveni]P66 `--U. sondeiana (Martin 1895) [=Pleurotoma sondeina, Gemmula (Unedogemmula) sondeiana]P66 CoroniaP66 CryptogemmaBK11 PleuroliriaP66 Eothesbia Finlay & Marwick 1937P66 `--*E. microtomoides Finlay & Mariwck 1937P66 Fusitoma Casey 1904P66 `--*F. sipho Casey 1904P66 Pyramitoma Martin 1914P66 `--*P. puruensis (Martin 1914) [=Pleurotoma (Pyramitoma) puruensis]P66 Tritonimangilia Martin 1914P66 `--*T. varicifera (Martin 1914) [=Mangilia (Tritonimangilia) varicifera]P66 Varicobela Casey 1904P66 `--*V. smithi (Aldrich 1885) [=Strombus smithi]P66 Parasyngenochilus Long 1981DK08 `--P. angustior Long 1981DK08 Trypanotopsis Gardner 1945P66 `--*T. texana (Gabb 1860) [=Turris texana]P66 Sinistrella Meyer 1887P66 `--*S. americana (Aldrich 1885) [=Triforis americanus, Trypanotoma americana]P66 Echinoturris Powell 1942P66 `--E. finlayi (Powell 1935) [=Turris finlayi]P66 Decollidrillia Habe & Ito 1965BK11 `--*D. nigra Habe & Ito 1965P66 Nekewis Stewart 1926P66 |--*N. washingtoniana (Weaver 1912) [=Fasciolaria washingtoniana]P66 `--N. io (Gabb 1864) (see below for synonymy)P66 Campylacrum Finlay & Marwick 1937P66 |--*C. sanum Finlay & Marwick 1937P66 `--C. debile Finlay & Marwick 1937P66 Hemipleurotoma Cossmann 1889P66 |--*H. archimedis (Bellardi 1878) [=Pleurotoma archimedis]P66 |--H. communis (Conrad 1830)P12 `--H. denticula (Basterot 1825) [=Pleurotoma denticula]P66 Epidirella Iredale 1913 [=Austrogemmula Laseron 1954]BK11 |--E. xanthophaes (Watson 1886)P66 (see below for synonymy) `--E. sayceana (Chapman 1912) [=Pleurotoma sayceana]P66 Heteroterma Gabb 1868P66 |--*H. trochoidea Gabb 1868P66 |--‘Cominella’ praecursor Wilckens 1907P66 `--H. zelandica Marshall 1917P66 Hesperiturris Gardner 1945P66 |--*H. nodocarinata (Gabb 1860) [=Turris nodocarinata]P66 |--H. amichel Gardner 1945P66 `--H. zacatensis Gardner 1945P66 Zexilia Finlay 1927P66 |--*Z. waihaoensis (Suter 1917) [=Exilia waihaoensis]P66 |--Z. cingulata [=Latirofusus cingulatus]F27 |--Z. crassicostata [=Exilia crassicostata]F27 `--Z. dalli (Suter 1915) [=Exilia dalli]F27 Eoturris Finlay & Marwick 1937P66 |--*E. complicatus (Suter 1917) [=Turris complicatus]P66 |--E. multicinctus (Marshall 1917) [=Turris multicinctus]P66 |--E. neglectus (Suter 1917)P66 (see below for synonymy) `--E. uttleyi (Suter 1917) [=Turris uttleyi]P66 Optoturris Powell 1944P66 |--*O. optata (Harris 1897) [=Pleurotoma optata]P66 |--O. edita Powell 1944P66 |--O. kyushuensis Shuto 1961P66 `--O. paracantha (Tenison-Woods 1877) [=Pleurotoma paracantha]P66 Kuroshioturris Shuto 1961BK11 |--*K. hyugaensis Shuto 1961 [=Gemmula (*Kuroshioturris) hyugaensis, Ptychosyrinx (*K.) hyugaensis]P66 |--K. asukana (Yokoyama 1926) [=Drillia asukana, Ptychosyrinx (Kuroshioturris) asukana]P66 |--K. nipponica (Shuto 1961) [=Gemmula (Ptychosyrinx) nipponicus]P66 `--K. totomiensis (Makiyama 1931) (see below for synonymy)P66 Epalxis Cossmann 1889P66 |--*E. crenulata (Lamarck 1804) [=Pleurotoma crenulata]P66 |--E. bilirata (Boury 1899) [=Pleurotoma bilirata]P66 |--E. lemoinei Boury 1899P66 |--E. multigyrata (Deshayes 1865) [=Pleurotoma multigyrata]P66 `--E. ventricosa (Lamarck 1804)P66 [=Pleurotoma ventricosaP66, Drillia ventricosaH09] Eopleurotoma Cossmann 1889 [incl. Eodrillia Casey 1904]P66 |--E. (Eopleurotoma)P66 `--E. (Oxyacrum Cossmann 1889)P66 |--E. (*O.) obliterata (Deshayes 1834) [=Pleurotoma obliterata]P66 |--E. (O.) constricta (Edwards 1861) [=Pleurotoma constricta]P66 |--E. (O.) inflexa (Lamarck 1804) [=Pleurotoma inflexa]P66 `--E. (O.) lepta (Edwards 1861)P66 Veruturris Powell 1944P66 |--V. (Veruturris)P66 | |--*V. (V.) quadricarinata (Powell 1944) [=Xenuroturris (V.) quadricarinatus]P66 | |--V. (V.) bisculpta (Powell 1944) [=Xenuroturris (V.) bisculptus]P66 | |--V. (V.) cochleata (Powell 1944) [=Xenuroturris (V.) cochleatus]P66 | |--V. (V.) subconcava (Harris 1897) [=Pleurotoma subconcava]P66 | `--V. (V.) tomopleuroides (Powell 1944) [=Xenuroturris (V.) tomopleuroides]P66 `--V. (Cinguliturris Powell 1964)P66 `--V. (*C.) tatei (Cossmann 1896) [=Asthenotoma tatei]P66 Trypanotoma Cossmann 1896P66 |--*T. terebriformis (Meyer 1886) [=Pleurotoma terebriformis]P66 | |--T. t. terebriformisP66 | |--T. t. cooperi Harris 1937P66 | `--T. t. curta Harris 1937P66 |--T. carlottae (Harris 1899) [=Pleurotoma carlottae]P66 |--T. longispira Casey 1904P66 |--T. melanella Harris 1937P66 |--T. obtusa Casey 1904P66 `--T. stocki (Dickerson 1916) [=Turris stocki]P66 Mariadrillia Petuch 1988P12 `--M. parvoidea (Martin 1904)P12 Nodisurculina Petuch 1988P12 `--N. engonata (Conrad 1862)P12 Chesaclava Petuch 1988P12 `--C. dissimilis (Conrad 1830)P12 Chesasyrinx Petuch 1988P12 `--C. rotifera (Conrad 1830)P12 Mariaturricula Petuch 1988P12 `--M. biscatenaria (Conrad 1830)P12 Calverturris Petuch 1992P12 Transmariaturris Petuch 1992P12
Eoturris neglectus (Suter 1917)P66 [=Turris neglectus nec Lesson 1837 (not preoc. as in homonymous genera) nec Pleurotoma neglecta Reeve 1842 (not preoc. if in dif. gen.)P66, T. insensus Finlay 1924F27, Gemmula insensaF27]
Epidirella xanthophaes (Watson 1886)P66 [=Pleurotoma xanthophaesP66, Hemipleurotoma xenthophaesH22; incl. Hemipleurotoma tasmanica May 1911P66, *Austrogemmula tasmanicaP66, *Epidirella tasmanicaP66]
Gemmula lululimi Olivera 2000 [incl. Turris (Annulaturris) munizi Vera-Peláez, Vega-Luz & Lozano-Francisco 2000]KFO12
Gemmula (Pinguigemmula) thielei Finlay 1930 [incl. Pleurotoma (Gemmula) fusiformis Thiele 1925 nec Pl. fusiforme Requien 1848 nec Pl. nebula fusiforme Marshall 1893]P66
Kuroshioturris totomiensis (Makiyama 1931) [=Turris (Gemmula) totomiensis, Ptychosyrinx (Kuroshioturris) totomiensis]P66
Nekewis io (Gabb 1864) [=Fasciolaria io; incl. Surcula alizensis Anderson & Hanna 1925, S. ioformis Anderson & Hanna 1925]P66
Unedogemmula hastula (Reeve 1843) [=Pleurotoma hastula, Gemmula (Unedogemmula) hastula; incl. P. trypanodes Melvill 1904]P66
*Unedogemmula unedo (Kiener 1839)P66, MG-H11 [=Pleurotoma unedoP66, Gemmula (*Unedogemmula) unedoP66; incl. Turris binda Garrard 1961P66]
*Type species of generic name indicated
References
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[P12] Petuch, E. J. 2012. Miocene asteroid impacts: proposed effects on the biogeography and extinction patterns of eastern North American gastropods. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 967–981. Springer.
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