Benthomangelia macra, copyright Natural History Museum, London.

Belongs within: Conoidea.
Contains: Marita, Neoguraleus (Neoguraleus), Neoguraleus (Fusiguraleus), Brachycythara, Bela, Tenaturris, Citharomangelia, Leiocithara, Guraleus (Guraleus), Amblyacrum, Propebela, Cryoturris, Ithycythara, Agathotoma, Pseudorhaphitoma, Pyrgocythara, Kurtziella, Mangelia, Oenopota, Heterocithara, Eucithara.

The Mangeliidae are a variable group of conoid gastropods with strongly sculptured shells, many of which have a multispiral protoconch. Most species lack an operculum though one with a terminal nucleus is present in Oenopota and related genera (Bouchet et al. 2011). Benthomangelia is a widespread deep-water genus characterised by angulated whorls and a greenish, yellowish or transparent periostracum giving the shell a polished appearance (Bouchet & Waren 1980).

Characters (from Bouchet et al. 2011): Shell small to medium size, 3–30 mm, usually 6–12 mm, oval to low- or high-fusiform, usually with comparatively low spire, often with shoulder angulation. Spiral and axial sculpture both well developed. Shell surface often bearing microsculpture of spirally aligned granules. Subsutural ramp usually not separated sculpturally. Anal sinus on subsutural ramp, shallow to rather deep, rarely tubular. Aperture normally not constrained with strong outgrowths or callus pads, rarely denticulate. Siphon rather short to moderately long. Protoconch typically multispiral, of up to five whorls, with axially ribbed protoconch II; spiral cords on protoconch II present or absent. When paucispiral, protoconch usually spirally lirate. Operculum present, with terminal nucleus, or normally absent. Radula of marginal teeth of very variable morphology; tooth morphology from semi-enrolled to true hypodermic, side projections frequently present around base with large irregularly shaped ‘root’ projecting from base; barbs may be present, from small to very large, or absent. Tooth canal opening laterally.

<==Mangeliidae [Mangeliinae, Mangiliinae, Oenopotinae]BK11
    |--Benthomangelia Thiele 1925PK11, BK11 [incl. Metuonella Sorgenfrei 1958BW80]
    |    |--*B. trophonoidea (Schepman 1913) [=Surcula trophonoidea]BW80
    |    |--B. antonia (Dall 1881)BW80 (see below for synonymy)
    |    |--B. bandellaBW80
    |    |--B. decapitata Bouchet & Waren 1980BW80
    |    |--‘Daphnella’ grippi Kautsky 1925 [=*Metuonella grippi]BW80
    |    `--B. macra (Watson 1881)GK02 [=Pleurotoma (Mangelia) macraBW80, Pleurotomella macraP66]
       |  `--Toxicochlespira Sysoev & Kantor 1990PK11, BK11
       |       `--T. pagoda Sysoev & Kantor 1990PK11
             `--Anticlinura Thiele 1934PK11, BK11 [=Clinuromella Beets 1943BK11, Clinuropsis Thiele 1929 non Vincent 1913BW80]
                  |--*A. monochorda (Dall 1908)BW80 (see below for synonymy)
                  `--A. peruviana (Dall 1908) [=Clinura peruviana, Pleurotomella (Anticlinura) peruviana]P66
Mangeliidae incertae sedis:
  ‘Pleurotoma’ triticea Kiener 1840K92
  Granoturris Fargo 1953P66
    |--*G. padolina Fargo 1953P66
    `--G. antipyrgus (Pilsbry & Lowe 1932) [=Kurtziella (Granoturris) antipyrgus]PP78
  Stellatoma Bartsch & Rehder 1939BK11
    `--*S. stellata (Stearns 1872) [=Mangelia stellata]P66
  Vexiguraleus Powell 1942P66
    `--*V. clifdenensis Powell 1942P66
  Mangaoparia Vella 1954P66
    `--*M. powelli Vella 1954P66
  Apitua Laseron 1954BK11
    `--*A. delicatula Laseron 1954P66
  Adelocythara Woodring 1928P66
    `--*A. primolevis Woodring 1928P66
  Pachycythara Woodring 1928P66
    `--*P. cryptonata Woodring 1928P66
  Amekicythara Eames 1957P66
    `--*A. douvillei (Newton 1922) [=Cominella douvillei]P66
  Buchozia Bayan 1873 [=Etallonia Deshayes 1862 non Oppel 1861]P66
    |--*B. citharella (Lamarck 1803) [=Fusus citharellus, Auricula citharella, *Etallonia citharella]P66
    `--‘Etallonia’ prisca Deshayes 1862P66
  Glyphoturris Woodring 1928BK11
    |--*G. rugirima (Dall 1889) [=Mangilia quadrata rugirima]P66
    |--G. lampra Woodring 1928P66
    `--G. quadrata (Reeve 1846) [incl. Cryoturris cruzana Usticke 1969]BC01
  Bactrocythara Woodring 1928BK11
    |--*B. obtusa (Guppy 1896) [=Cythara obtusa]P66
    `--B. asarca (Dall & Simpson 1901) [=Mangilia (Cythara) asarca]P66
  Mappingia Ludbrook 1941P66
    |--*M. acutispira Ludbrook 1941P66
    `--M. matronalis Ludbrook 1958P66
  Papillocithara Kilburn 1992K92
    |--*P. hebes Kilburn 1992K92
    `--P. semiplicata Kilburn 1992K92
  Cytharopsis Adams 1865BK11
    |--*C. cancellata (Adams 1865) (n. d.) [=Mangilia cancellata]P66
    |--C. butonensis (Schepman 1913) [=Mangilia butonensis, Leiocithara butonensis]K92
    `--‘Pleurotoma (Glyphostoma)’ exquisita Smith 1882K92 [=Raphitoma exquisitaH22]
  Acmaturris Woodring 1928BK11
    |--*A. comparata Woodring 1928P66
    |--A. brisis Woodring 1928P66
    `--A. scalida Woodring 1928P66
  Saccharoturris Woodring 1928BK11
    |--*S. consentanea (Guppy 1896) [=Mangilia consentanea]P66
    |--S. centrodes Gardner 1937P66
    `--S. monocingulata (Dall 1889) [=Mangilia monocingulata]P66
  Etremella Makiyama 1927P66
    `--*E. totomiensis (Makiyama 1927) [=Cytharella totomiensis]P66
         |--E. t. totomiensisP66
         `--E. t. tachymorpha (Makiyama 1927) [=Cytharella tachymorpha]P66
  Apispiralia Laseron 1954BK11
    |--*A. albocincta (Angas 1871)P66 [=Clathurella albocinctaP66, Asperdaphne albocinctaL54]
    |--A. catena Laseron 1954P66
    `--A. maxima Laseron 1954P66
  Eoclathurella Casey 1904P66
    |--*E. jacksonica Casey 1904P66
    |--E. meridionalis (Meyer 1886) [=Mangilia meridionalis]P66
    `--E. obesula Casey 1904P66
  Enatoma Rovereto 1899 [=Atoma Bellardi 1875 non Latreille 1817]P66
    |--*E. hypothetica (Bellardi 1847) [=Raphitoma hypothetica, *Atoma hypothetica]P66
    |--E. costata (Peyrot 1932) [=Mangilia (Atoma) costata]P66
    |--E. gallica (Peyrot 1932) [=Mangilia (Atoma) gallica]P66
    `--E. inopinatus (Peyrot 1938) [=Mangelia (Atoma) inopinatus]P66
  Kurtzia Bartsch 1944BK11
    |--*K. arteaga (Dall & Bartsch 1910)P66 [=Mangilia arteagaP66, M. (Kurtzilla) arteagaO27]
    |    |--K. a. arteagaP66
    |    `--K. a. roperi (Dall 1919) [=Mangilia (Kurtziella) arteaga roperi]P66
    |--K. gordoni Bartsch 1944P66
    `--K. granulatissima (Mörch 1860)PP78
  Macteola Hedley 1918BK11
    |--*M. anomala (Angas 1877)P66 [=Purpura (Cronia) anomalaP66, Mangelia anomalaH22, Murex anomalaH22]
    |--M. interrupta (Reeve 1846) [=Mangelia interrupta]P66
    |--M. sandersonae (Bucknill 1927) [=Scrinium sandersonae]P66
    |--M. segesta (Duclos in Chenu 1850)BK11 (see below for synonymy)
    |--M. theskela (Melvill & Standen 1895)P66 [=Mangelia (Glyphostoma) theskelaH22]
    `--M. thiasotes [=Mangilia thiasotes]H22
  Notocytharella Hertlein & Strong 1955BK11
    |--*N. niobe (Dall 1919) [=Cytharella niobe]P66
    |--N. ephaedra (Dall 1919) [=Philbertia ephaedra]P66
    |--N. hastula (Pilsbry & Lowe 1932) [=Cytharella hastula]P66
    |--N. hippolita (Dall 1919) [=Cytharella hippolita]P66
    `--N. phaethusa (Dall 1919) [=Cytharella phaethusa]P66
  Kurtzina Bartsch 1944BK11
    |--*K. beta (Dall 1919) [=Mangilia (Kurtziella) beta, M. (K.) beata (l. c.)]P66
    |--K. antiochroa (Pilsbry & Lowe 1932) [=Mangelia antiochroa]P66
    |--K. antipyrgus (Pilsbry & Lowe 1932) [=Mangelia antipyrgus]P66
    |--K. cymatias (Pilsbry & Lowe 1932) [=Mangelia cymatias]P66
    `--K. cyrene (Dall 1919) [=Mangilia (Kurtziella) cyrene]P66
  Neoguraleus Powell 1939BK11
    |  i. s.: ‘Daphnella (Raphitoma)’ calais (Kautsky 1925)P66
    |         ‘Mangilia’ gurichi (Kautsky 1925)P66
    |         N. kutekinensis MacNeil 1960P66
    |         ‘Pleurotoma’ tenella Mayer 1858P66
    |--N. (Neoguraleus)P66
    `--N. (Fusiguraleus)P66
  Obesotoma Bartsch 1941BK11
    |--*O. japonica Bartsch 1941P66
    |--O. arctica (Adams 1855) [=Bela arctica]P66
    |--O. hanazakiensis Habe 1958P66
    |--O. lawrenciana (Dall 1919)P66 [=Lora lawrencianaP66, Bela tenuilirata Krause 1886 non Dall 1871D19]
    |--O. murdochiana (Dall 1885) [=Bela murdochiana]P66
    |--O. schantaricum (Middendorff 1849) [=Pleurotoma schantaricum]P66
    `--O. uchidai Habe 1958P66
  Gingicithara Kilburn 1992K92
    |--*G. lyrica (Reeve 1846)K92 (see below for synonymy)
    |--G. albivestis (Pilsbry 1934)K92 (see below for synonymy)
    |--G. cylindrica (Reeve 1846)K92 [=Mangelia cylindricaK92, Cythara cylindricaH22, Eucithara cylindricaK92]
    |--G. maraisi Kilburn 1992K92
    |--G. notabilis (Smith 1888) (see below for synonymy)K92
    |--G. pessulata (Reeve 1846)K92 [=Mangelia pessulataK92, Cythara pessulataH22, Rhaphitoma (l. c.) pessulataK92]
    `--G. ponderosa (Reeve 1846)K92 [=Mangelia ponderosaK92, Eucithara ponderosaP66]
  Liracraea Odhner 1924BK11
    |--*L. epentroma (Murdoch 1905) [=Clathurella epentroma]P66
    |--L. benthicola Dell 1956P66 [=L. epentroma benthicolaP66, L. odhneri benthicolaP61]
    |--L. dictyota (Hutton 1885)P66 [=Clathurella dictyotaP66, Asperdaphne dictyotaH22]
    |--L. odhneri Powell 1942P66
    |--L. otakauica Powell 1942P66
    |--L. subantarctica Powell 1942P66
    |--L. titirangiensis Marwick 1928P66
    `--L. whangaroaensis (Murdoch 1905) [=Clathurella whangaroaensis]P66
  Guraleus Hedley 1918BK11
    |  i. s.: G. amplexus [=Clathurella amplexa]H22
    |         G. lallemantianus (Crosse & Fischer 1865) (see below for synonymy)H22
    |         G. octangulata (Dunker 1860)P66
    |--G. (Guraleus)BK11
    `--G. (Euguraleus Cotton 1947)BK11
         |--G. (*E.) anisus (Cotton 1947) [=*Euguraleus anisus]P66
         |--G. (E.) jacksonensis (Angas 1877)P66 [=Mangelia jacksonensisP66, Daphnella jacksonensisH22]
         |--G. (E.) letourneuxianus (Crosse & Fischer 1865)H22, L54, H22 (see below for synonymy)
         |--G. (E.) tasmanicus (Tenison-Woods 1876)P66 (see below for synonymy)
         |    |--G. t. tasmanicusP66
         |    `--G. t. peronianus (Laseron 1954) [=Euguraleus tasmanicus peronianus]P66
         `--G. (E.) thornleyanus (Laseron 1954) [=Euguraleus thornleyanus]P66
  Bellacythara McLean 1971BK11
  Cacodaphnella Pilsbry & Lowe 1932BK11
    `--*C. delgada Pilsbry & Lowe 1932P66
  Curtitoma Bartsch 1941 [incl. Nematoma Bartsch 1941, Widalli Bogdanov 1986]BK11
    |--*C. hecuba Bartsch 1941BW80 [=Nodotoma hecubaP66]
    `--*Nematoma’ hokkaidoensis Bartsch 1941BW80 [=Nodotoma hokkaidoensisP66]
  Euryentmema Woodring 1928BK11
    `--*E. cigclis Woodring 1928P66
  Fehria van Aartsen 1988BK11
  Genotina Vera-Peláez 2004BK11
    `--G. adamii (Bozzetti 1994)BK11
  Granotoma Bartsch 1941BK11
    `--*G. krausei (Dall 1886)BW80 [=Bela krauseiBW80, Lora krauseiD19, Oenopota (Funitoma) krauseiP66]
  Hemicythara Kuroda & Oyama in Kuroda, Habe & Oyama 1971BK11
  Mitraguraleus Laseron 1954BK11
    |--*M. mitralis (Adams & Angas 1864)P66 (see below for synonymy)
    `--M. australis (Adams & Angas 1864)P66 (see below for synonymy)
  Oenopotella Sysoev 1988BK11
    `--O. ultraabyssalis Sysoev 1988BK11
  Paraspirotropis Sysoev & Kanto 1984BK11
    `--P. simplicissima (Dall 1907)BK11
  Perimangelia McLean 2000BK11
  Platycythara Woodring 1928BK11
    `--*P. eurystoma Woodring 1928 [=Pyrgocythara (*Platycythara) eurystoma]P66
  Rubellatoma Bartsch & Rehder 1939BK11
    |--*R. rubella [=Mangelia rubella]P66
    |--R. diomedea Bartsch & Rehder 1939P66
    `--R. powelli (Shasky 1971) [=Kurtziella (Rubellatoma) powelli]BC01
  Suturocythara García 2008BK11
  Venustoma Bartsch 1941BK11
    `--*V. harucoa Bartsch 1941BW80 [=Nodotoma harucoaP66]
  Vitjazinella Sysoev 1988BK11
    `--V. multicostata Sysoev 1988BK11
  Vitricythara Fargo 1953BK11
    |--*V. metria (Dall 1903) [=Cythara metria, Pyrgocythara (Vitricythara) metria]P66
    |    |--V. m. metriaP66
    |    `--V. m. micromeris (Dall 1903) (see below for synonymy)P66
    `--V. klasmidia (Shasky 1971) (see below for synonymy)BC01

*Anticlinura monochorda (Dall 1908)BW80 [=Clinura monochordaBW80, *Clinuromella monochordaBW80, *Clinuropsis monochordaBW80, Pleurotomella (*Anticlinura) monochordaP66]

Benthomangelia antonia (Dall 1881)BW80 [=Pleurotoma (Mangilia) antoniaBW80; incl. Pleurotomella diomedae Verrill 1884BW80, Pleurotoma (Mangelia) incincta Watson 1881BW80, Pleurotomella incinctaP66, Mangelia innocens Thiele 1925BW80, Pleurotoma mericiacum Locard 1897BW80]

Gingicithara albivestis (Pilsbry 1934)K92 [=Mangilia albivestisK92, Eucithara albivestisP66; incl. M. pura Pilsbry 1904 non Reeve 1846K92]

*Gingicithara lyrica (Reeve 1846)K92 [=Mangelia lyricaK92, Cithara lyricaH22, Cythara cylindrica var. lyricaH22, Eucithara lyricaP66, Mangilia lyricaK92]

Gingicithara notabilis (Smith 1888) [=Pleurotoma (Mangilia) notabilis; incl. Mangilia (Cythara) brunneolineata Preston 1905, M. eudeli Sowerby 1901, M. (C.) euselma Melvill & Standen 1896, M. euselma var. evanescens Bouge & Dautzenberg 1914, M. quadrasi Boettger 1895]K92

Guraleus lallemantianus (Crosse & Fischer 1865) [=Pleurotoma (Mangelia) lallemantiana, Clathurella lallemantiana]H22

Guraleus (Euguraleus) letourneuxianus (Crosse & Fischer 1865)H22, L54, H22 [=Pleurotoma (Clathurella) letourneuxianaH22, Daphnella letourneuxianaH22, Mangelia letourneuxianaH22]

Guraleus (Euguraleus) tasmanicus (Tenison-Woods 1876)P66 [=Cithara tasmanicaP66, Mangilia tasmanicaH22; incl. M. alternata Tenison-Woods 1879H22, Cythara maccoyi Petterd 1879H22]

Macteola segesta (Duclos in Chenu 1850)BK11 [=Columbella segestaP66; incl. Daphnella bella Pease 1860P66, Mangilia (Cythara) bellaK65, Pleurotoma gemmulata Deshayes 1863P66]

Mitraguraleus australis (Adams & Angas 1864)P66 [=Bela australisP66, Euguraleus australisL54, Guraleus mitralis var. australisH22, G. (Mitraguraleus) australisP66, Mangilia australisH22]

*Mitraguraleus mitralis (Adams & Angas 1864)P66 [=Bela mitralisP66, Daphnella mitralisH22, Euguraleus mitralisL54, Guraleus (Marita) mitralisL54, Mangilia mitralisH22]

Vitricythara metria micromeris (Dall 1903) [=Glyphostoma metria micromeris, Pyrgocythara (Vitricythara) metria micromeris]P66

Vitricythara klasmidia (Shasky 1971) [=Agathotoma (Vitricythara) klasmidia, Pyrgocythara (V.) klasmidia; incl. A. (V.) secalis Shasky 1971]BC01

*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.

[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.

[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.

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

[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.

[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.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 1. Stanford University Press: Stanford University (California).

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

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[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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