Lepidopleuridae

Ribbed chiton Lepidopleurus cajetanus, copyright Wilfried Bay-Nouailhat.

Belongs within: Neoloricata.

The Lepidopleuridae are a group of small chitons in which the articulamentum is poorly developed (Smith 1960).

Characters (from Smith 1960): Relatively small (less than 5 cm long); valves generally sculptured with granules closely set in lines or quincunx; valve coverage complete; girdle narrow, minutely spiculose or scaly; insertion plates lacking, or if present, weak and unslit; articulamentum layer usually consisting only of weakly developed sutural laminae in valves ii to viii. Gills short, posterior.

<==Lepidopleuridae [Protochitonidae]
    |--Oldroydia Dall 1894 S60
    |    `--*O. percrassa Dall 1894 S60 [=Lepidopleurus (Oldroydia) percrassus O27]
    |--Protochiton Ashby 1925 S60
    |    `--*P. granulosus (Ashby & Torr 1901) [=Acanthochites (Notoplax) granulosus] S60
    |--Pterochiton Carpenter in Dall 1882 [incl. Anthracochiton Rochebrune 1883, Loricites Carpenter in Dall 1882] S60
    |    `--*P. eburonicus (Ryckholt 1845) [=Chiton eburonicus] S60
    |--Priscochiton Dall 1882 S60
    |    `--*P. canadensis (Billings 1865) [=Chiton canadensis] S60
    |--Parachiton Thiele 1909 P61
    |    |--*P. acuminatus [=Lepidopleurus acuminatus] P61
    |    |--P. subantarcticus Iredale & Hull 1930 P61
    |    `--P. textilis Powell 1937 P61
    |--Terenochiton Iredale 1914 P61
    |    |--*T. subtropicalis [=Lepidopleurus subtropicalis] P61
    |    |--T. fairchildi Iredale & Hull 1929 P61
    |    |--T. finlayi (Ashby 1929] P61
    |    |--T. inquinatus (Reeve 1847) P61 [=Lepidopleurus inquinatus F27; incl. L. iredalei Ashby 1920 F27]
    |    `--T. otagoensis Iredale & Hull 1929 P61
    `--Lepidopleurus Risso 1826 S-P86 (see below for synonymy)
         |--*L. cajetanus (Poli 1791) [=Chiton cajetanus] S60
         |--L. asellus MR96
         |--L. cancellatus Sowerby 1839 O27
         |--L. heathi Berry 1919 [=L. (Xiphiozona) heathi] O27
         |--L. intermedius Salvini-Plawen 1968 S-P86
         |--L. kerguelenensis F27
         |--L. laterodepressus S60
         |--L. pectinatus Carpenter 1864 C64
         |--L. regularis Cpr. 1855 C64
         `--L. tenuisculptus C64

Lepidopleurus Risso 1826 S-P86 [incl. Belchiton Ashby & Cotton 1939 S60, Deshayesiella Carpenter in Dall 1879 S60, Pilsbryella Nierstrasz 1905 S60, Xiphizona Berry 1919 S60]

*Type species of generic name indicated

REFERENCES

[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. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[MR96] Morse, M. P., & P. D. Reynolds. 1996. Ultrastructure of the heart-kidney complex in smaller classes supports symplesiomorphy of molluscan coelomic characters. In: Taylor, J. D. (ed.) Origin and Evolutionary Radiation of the Mollusca pp. 89–97. Oxford University Press: Oxford.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

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

[S-P86] Salvini-Plawen, L. v. 1986. Lower Mollusca. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 148–152. E. J. Brill/Dr W. Backhuys: Leiden.

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

Lepidochitonidae

Lepidochitona cinerea, copyright S. Rae.

Belongs within: Ischnochitonina.

The Lepidochitonidae are a group of chitons in which the valve margins bear distinctly separated apophyses.

<==Lepidochitonidae [Trachyderminae] P61
    |--Eudoxochiton Shuttleworth 1853 P61
    |    |--*E. nobilis (Gray 1843) [=Acanthopleura nobilis] S60
    |    `--E. huttoni Pilsbry 1893 P61
    |--Paricoplax Iredale & Hull 1929 P61
    |    |--*P. crocina (Reeve 1847) [=Chiton crocinus] P61
    |    |--P. mortenseni (Odhner 1924) P61
    |    |--P. perscrutanda Iredale & Hull 1929 P61
    |    `--P. profundior Dell 1956 P61
    |--Icoplax Thiele 1893 P61
    |    |--*I. punicea (Gould 1846) [=Chiton puniceus] P61
    |    |--I. chathamensis Dell 1960 P61
    |    |--I. empleura (Hutton 1872) P61 [=Callochiton empleurus F27]
    |    |--I. kapitiensis Mestayer 1926 P61 [=Callochiton kapitiensis F27]
    |    |--I. subeudoxa Iredale & Hull 1930 P61
    |    `--I. sulculata (Suter 1907) P61 [=Callochiton sulculatus F27]
    `--Lepidochitona Gray 1821 (see below for synonymy) S60
         |--L. cinerea (Linné 1767) (see below for synonymy) S60
         |--L. alba Linnaeus 1767 O27
         |--L. aleutica (Dall 1878) [=Trachyradsia aleutica, L. (Spongioradsia) aleutica] O27
         |--L. cavernae GC99
         |--L. corrugata GR98
         |--L. dentiens (Gould 1846) O27 [=Chiton (Onithochiton) dentiens O27, C64, Trachydermon dentiens C64]
         |--L. flectens (Carpenter 1864) O27 [=Ischnochiton (Trachydermon) flectens C64, O27]
         |--‘Trachydermon’ hartwegii Cpr. 1855 C64
         |--‘Trachydermon’ interstinctus C64
         |--‘Trachydermon’ nuttallii Cpr. 1855 C64
         |--‘Trachydermon’ pseudodentiens Carpenter 1864 C64
         `--L. sharpei (Pilsbry 1896) [=Trachydermon sharpii] O27

Lepidochitona Gray 1821 [=Lepidochiton Thiele 1928 non Carpenter 1857, Leptochitona (l. c.); incl. Adriella Thiele 1893, Craspedochilus Sars 1878, Spongioradsia Pilsbry 1894, Trachydermon Carpenter 1864] S60

Lepidochitona (Lepidochitona) cinerea (Linné 1767) [=Chiton cinereus; incl. C. marginatus Pennant 1777, *Lepidochitona marginata] S60

*Type species of generic name indicated

REFERENCES

[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. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[GC99] Giribet, G., & C. S. Carranza. 1999. Point Counter Point. What can 18S rDNA do for bivalve phylogeny? Journal of Molecular Evolution 48: 256–258.

[GR98] Giribet, G., & C. Ribera. 1998. The position of arthropods in the animal kingdom: a search for a reliable outgroup for internal arthropod phylogeny. Molecular Phylogenetics and Evolution 9: 481–488.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

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

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

Ischnochitonina

Lorica volvox, copyright Des Beechey.

Belongs within: Neoloricata.
Contains: Lepidochitonidae, Plaxiphoridae, Acanthochitona, Notoplax, Mopaliidae, Ischnochitonidae, Chitonidae, Chaetopleuridae, Callistochiton.

The Ischnochitonina are a group of chitons, including the greater number of living species, in which the ventral layer of the valves forms a well developed articulamentum (Smith 1960). The Cryptoconchidae, in which the dorsal tegmentum is reduced relative to the articulamentum, have been treated as their own separate suborder but appear to be phylogenetically nested within the Ischnochitonina. Members of the genus Callochiton have a fairly wide, leathery girdle covered with longitudinally packed minute narrow scales or spicules with only the tips showing (Smith 1960).

<==Ischnochitonina S60
    |--Callochiton Gray 1847 GO06, S60 (see below for synonymy)
    |    |--*C. laevis (Montagu 1803) [=Chiton laevis] S60
    |    |--C. castaneus (Wood 1815) [=C. (Stereochiton) castaneus, C. (Trachyradsia) castaneus, Eudoxochiton castaneus] L70
    |    |--C. chattonensis Ashby 1929 BE93
    |    |--C. inornatus Ten.-Woods 1881 L70 (see below for synonymy)
    |    `--C. septemvalvis GO06
    `--+--Cryptoconchidae (see below for synonymy) P61
       |    |--Acanthochitona GO06
       |    |--Notoplax P61
       |    |--Cryptoconchus Burrow 1815 P61 [=Cryptochoncus Clessin 1903 S60, Cryptoconctus Sowerby 1852 S60]
       |    |    `--*C. porosus (Burrow 1815) [=Chiton porosus] P61
       |    |--Choneplax Carpenter in Dall 1882 (see below for synonymy) S60
       |    |    `--C. latus (Guilding 1829) (see below for synonymy) S60
       |    |--Austrochiton Bergenhayn 1945 S60
       |    |    `--*A. rostratus (Ashby & Torr 1901) [=Acanthochites rostratus] S60
       |    |--Cryptoplax deBlainville 1818 (see below for synonymy) S60
       |    |    |--*C. larvaeformis (Burrow 1815) [=Chiton larvaeformis] S60
       |    |    |--C. burrowi (Smith 1884) [=Chiton burrowi] H09
       |    |    |--C. japonica GO06
       |    |    |--C. numicus S60
       |    |    |--C. oculatus (Burrow 1815) BW09 [=Chiton oculatus H09]
       |    |    `--C. pritchardi S60
       |    `--Craspedochiton Shuttleworth 1853 P61 (see below for synonymy)
       |         |--*C. laqueatus (Sowerby 1841) S60 [=Chiton laqueatus P61]
       |         |--‘Pseudotonicia’ cuneata HS01
       |         |--‘*Lophoplax’ finlayi Ashby 1926 F27
       |         |--‘Notoplax (Amblyplax)’ foveauxensis Ashby 1926 (see below for synonymy) F27
       |         |--‘Notoplax (*Amblyplax)’ oliveri Ashby 1926 [=Macandrellus oliveri Mestayer 1926] F27
       |         `--C. rubiginosus (Hutton 1872) P61
       `--+--+--+--Mopaliidae GO06
          |  |  `--Cryptochiton Middendorff 1847 GO06, S60 [Cryptochitonidae]
          |  |       `--*C. stelleri (Middendorff 1847) S60 (see below for synonymy)
          |  `--Tonicella Carpenter 1873 GO06, S60 (see below for synonymy)
          |       |--*T. marmorea (Fabricius 1780) S60 (see below for synonymy)
          |       |--T. lineata (Wood 1815) [=Chiton lineatus, Lepidochitona (Tonicella) lineata] O27
          |       |--T. rubra (Linnaeus 1767) [=Chiton ruber, Lepidochitona (Tonicella) ruber] O27
          |       |--T. saccharina Dall 1878 [=Lepidochitona (Tonicella) saccharina] O27
          |       |--T. sitkensis (Middendorff 1846) [=Chiton sitkensis, Lepidochitona (Tonicella) sitkensis] O27
          |       `--T. submarmorea (Middendorff 1848) [=Lepidochitona (Tonicella) submarmorea] O27
          `--+--+--Ischnochitonidae GO06
             |  `--+--Chitonidae GO06
             |     `--Lorica Adams & Adams 1852 GO06, P61 (see below for synonymy)
             |          |--*L. cimolia (Reeve 1847) [=Chiton cimolius] S60
             |          |--L. atkinsoni S60
             |          |--L. cudmorei S60
             |          |--L. duniana Hull 1910 F71
             |          |--L. haurakiensis (Mestayer 1921) F27 [=Aulacochiton haurakiensis P61, *Zelorica haurakiensis F27]
             |          |--‘Aulacochiton’ praecursor Smith et al. 1968 BE93
             |          `--L. volvox (Reeve 1847) GO06, S60 [=Chiton volvox P61, *Aulacochiton volvox P61]
             `--+--Chaetopleuridae GO06
                `--Callistoplacidae [Callistochitonidae, Callistochitoninae, Callistoplacinae] SF77
                     |--Callistochiton GO06
                     |--Callistoplax Carpenter in Dall 1882 S60
                     |    `--*C. retusa (Sowerby 1832) [=Chiton retusus] S60
                     |--Ceratozona Dall 1882 (see below for synonymy) S60
                     |    `--C. rugosa (Sowerby 1841) S60 (see below for synonymy)
                     |--Middendorffia Carpenter in Dall 1882 (see below for synonymy) S60
                     |    |--*M. polii (Philippi 1836) [=Chiton polii; incl. C. cinereus Poli 1791 non Linné 1767] S60
                     |    `--M. caprearum PP64
                     `--Nuttallina Dall 1871 [=Nutallina (l. c.), Nuttalina (l. c.)] S60
                          |--N. fluxa (Carpenter 1864) (see below for synonymy) S60
                          |--N. californica (Reeve 1847) [=Chiton californicus] O27
                          `--N. thomasi Pilsbry 1898 O27

Ischnochitonina incertae sedis:
  Subterenochiton Iredale & Hull 1924 [Subterenochitonidae] S60
    `--*S. gabrieli (Hull 1912) [=Ischnochiton gabrieli] S60
  Schizoplax Dall 1878 [Schizoplacidae] S60
    |--*S. brandtii (Middendorff 1846) [=Chiton brandtii] S60
    `--S. multicolor Dall 1920 O27
  Lepidochitonidae S60
  Schizochitonidae [Prochitonidae] S60
    |--Schizochiton Gray 1847 S60
    |    `--*S. incisus (Sowerby 1841) [=Chiton incisus] S60
    `--Loricella Pilsbry 1893 [incl. Pseudoloricella Ashby 1925, Squamophora Nierstrasz 1905] S60
         |--*L. angasi (Adams & Angas 1864) [=Lorica angasi] S60
         `--L. sculpta S60
  Plaxiphoridae S60

Cryptoconchidae [Acanthochitae, Acanthochitidae, Acanthochitinae, Acanthochitonidae, Acanthochitonina, Acanthochitoninae, Cryptoplacidae, Cryptoplacinae, Pseudotonicinae] P61

Callochiton Gray 1847 GO06, S60 [=Collochiton (l. c.) S60; incl. Acutoplax Cotton & Weeding 1939 S60, Eudoxoplax Iredale & May 1916 S60, Ocellochiton Ashby 1939 S60, Quaestiplax Iredale & Hull 1929 S60, Stereochiton Carpenter in Dall 1882 S60, Trachyradsia Carpenter in Dall 1878 S60; Callochitonidae, Callochitoninae]

Callochiton inornatus Ten.-Woods 1881 L70 [=Ca. (Trachyradsia) inornatus L70, Chiton inornatus F27, Eudoplax inornatus L70, Eudoxochiton (*Eudoxoplax) inornatus L70, F27; incl. Callochiton (Trachyradsia) lobatus L70]

Ceratozona Dall 1882 [incl. Ceratophorus Carpenter in Dall 1882 nec Schuckard 1837 nec Diesing 1850, Newcombia Carpenter in Dall 1882] S60

Ceratozona rugosa (Sowerby 1841) S60 [=Chiton rugosus S60; incl. Ch. guildingi Reeve 1847 S60, *Ceratozona guildingi S60, Ch. squalidus G79]

Choneplax Carpenter in Dall 1882 [=Chnoeplax (l. c.); incl. Chitoniscus Carpenter in Dall 1882 nec Herrmannsen 1852 nec Stål 1875] S60

Choneplax latus (Guilding 1829) [=Chitonellus latus; incl. Chiton strigatus Sowerby 1840, *Choneplax strigatus] S60

Craspedochiton Shuttleworth 1853 P61 [incl. Amblyplax Ashby 1926 S60, Angasia Carpenter in Dall 1882 S60, Augasia (l. c.) S60, Aristochiton Thiele 1909 S60, Bassethullia Pilsbry 1928 S60, Craspedoplax Iredale & Hull 1925 S60, Glyptelasma Iredale & Hull 1925 non Pilsbry 1907 S60, Ikedaella Taki & Taki 1929 S60, Leptoplax Carpenter in Dall 1882 S60, Loboplax Pilsbry 1893 S60, Lophoplax Ashby 1926 S60, Macandrellus Carpenter in Dall 1879 S60, Mecynoplax Thiele 1893 S60, Phacellopleura Carpenter in Dall 1879 non Agassiz 1846 S60, Phacellozona Pilsbry 1894 S60, Pseudotonicia Ashby 1928 S60, Spongiochiton Carpenter in Dall 1882 S60, Thaumastochiton Thiele 1909 S60]

*Cryptochiton stelleri (Middendorff 1847) S60 [=Chiton (*Cryptochiton) stelleri S60; incl. Ch. amiculatus Newcomb in Carpenter 1864 (n. n.) C64]

Cryptoplax deBlainville 1818 [=Crytoplax Gray 1821; incl. Ametrogephyrus Middendorff 1848, Chitonellus Lamarck 1819, Chitonella (l. c.), Chitoniscus Herrmannsen 1846, Dichachiton Middendorff 1848, Dibachiton (l. c.), Oscabrella Broderip 1835, Phaenochiton Middendorff 1848; Cryptoplacidae] S60

Lorica Adams & Adams 1852 GO06, P61 [incl. Aulacochiton Shuttleworth 1852 P61, Protolorica Ashby 1925 S60, Zelorica Finlay 1927 S60; Aulacochitonidae P61, Loricidae]

Middendorffia Carpenter in Dall 1882 [=Middendorfia; incl. Beanella Dall 1882, Beania Carpenter in Dall 1879 non Johnston 1840, Dawsonia Carpenter 1873 nec Brown 1811 (ICBN) nec Hartt in Dawson 1868 nec Nicholson 1873] S60

‘Notoplax (Amblyplax)’ foveauxensis Ashby 1926 [=Acanthochiton foveauxensis Mestayer 1926; incl. A. foveauxensis var. kirki Mestayer 1926] F27

Nuttallina fluxa (Carpenter 1864) [=Acanthopleura fluxa; incl. Chiton scaber Reeve 1847 non de Blainville 1825, *Nuttallina scabra] S60

Tonicella Carpenter 1873 GO06, S60 [=Toniciella (l. c.) S60; incl. Boreochiton Sars 1878 S60, Clathropleura Tiberi 1877 S60, Platysemus Middendorff 1847 S60]

*Tonicella marmorea (Fabricius 1780) S60 [=Chiton marmoreus S60, Boreochiton marmaoreus CS77, Lepidochitona (Tonicella) marmorea O27]

*Type species of generic name indicated

REFERENCES

[BE93] Benton, M. J., & D. H. Erwin. 1993. Mollusca: Amphineura and ‘Monoplacophora’. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 125–130. Chapman & Hall: London.

[BW09] Bryce, C., & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177–208.

[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.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

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

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[G79] Guppy, R. J. L. 1879. First sketch of a marine invertebrate fauna of the Gulf of Paria and its neighbourhood. Part I.—Mollusca. Journal of Conchology 2: 151–172.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

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

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

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

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SF77] Smith, A. G., & A. J. Ferreira. 1977. Chiton fauna of the Galápagos Islands. Veliger 20 (2): 82–97.

Last updated: 18 December 2021.

Ischnochitonidae

Ischnochiton australis, copyright Nuytsia@Tas.

Belongs within: Ischnochitonina.
Contains: Stenoplax (Stenoradsia).

The Ischnochitonidae are a group of chitons in which the dorsum of the intermediate valves is divided into distinct lateral and central areas, often by a diagonal rib (Smith 1960).

Characters (from Smith 1960): Ovate to elongate chitons of variable size. Tegmentum of intermediate valves divided into lateral and central areas by diagonal rib (commonly indistinct) extending from apex to anterior outer angle of valves on each side. Articulamentum of head and tail valves multiple-slitted, that of intermediate valves with single slit or in some groups with two or more slits on each side. Teeth sharp-edged, not pectinated, grooved, or buttressed on outside. Eaves not porous. Sutural laminae sharp and well developed.

<==Ischnochitonidae
    |--Oligochiton Berry 1922 S60
    |    `--*O. lioplax Berry 1922 S60
    |--Tonicina Thiele 1906 S60
    |    `--*T. zschaui (Pfeffer 1886) [=Chiton zschaui] S60
    |--Basiliochiton Berry 1918 [incl. Lophochiton Berry 1925 non Ashby 1923, Ploiochiton Berry 1926] S60
    |    |--*B. heathii (Pilsbry 1898) [=Mopalia heathii] S60
    |    `--B. lobium Berry 1925 O27
    |--Gurjanovillia Jakoleva 1952 F77
    |    |--*G. albrechti (Schrenck 1867) [=Chiton albrechti] F77
    |    `--G. lindberghi Jakovleva 1952 F77
    |--Cyanoplax Pilsbry 1892 [incl. Mopaliella Thiele 1909, Mopaliopsis Thiele 1893] S60
    |    |--*C. hartwegii (Carpenter 1855) S60 (see below for synonymy)
    |    |    |--C. h. hartwegii O27
    |    |    `--C. h. nuttalli (Carpenter 1855) [=Chiton nuttalli, Lepidochitona (Cyanoplax) hartwegii nuttalli] O27
    |    |--C. fackenthallae Berry 1919 [=Lepidochitona (Cyanoplax) fackenthallae] O27
    |    |--C. lowei (Pilsbry 1918) [=Trachydermon lowei, Lepidochitona (Cyanoplax) lowei] O27
    |    `--C. raymondi (Pilsbry 1894) [=Trachydermon (Cyanoplax) raymondi, Lepidochitona (C.) raymondi] O27
    |--Stenoplax Carpenter in Dall 1879 S60
    |    |  i. s.: S. circumsenta Berry 1956 PP78
    |    |         S. heathiana GW02
    |    |--S. (Stenoplax) S60
    |    |    |--*S. (S.) limaciformis (Sowerby 1832) [=Chiton limaciformis] S60
    |    |    |--S. (S.) biarcuata (Dall 1903) [=Ischnochiton (Stenoplax) biarcuata] O27
    |    |    |--S. (S.) conspicua (Carpenter in Pilsbry 1892) S60, O27 [=Ischnochiton (Stenoradsia) conspicuus O27]
    |    |    `--S. (S.) fallax (Carpenter 1892) [=Ischnochiton (Stenoplax) fallax] O27
    |    |--S. (Stenochiton Adams & Angas 1864) [incl. Zostericola Ashby 1919] S60
    |    |    `--S. (*S.) juloides (Adams & Angas 1864) [=*Stenochiton juloides] S60
    |    `--S. (Stenoradsia) S60
    |--Lepidozona Pilsbry 1892 F77 (see below for synonymy)
    |    |--*L. mertensii (Millendorff 1847) F77 [=Chiton mertensii F77, Ischnochiton (Lepidopleurus) mertensii O27, C64]
    |    |--L. asthenes (Berry 1919) [=Ischnochiton (Lepidozona) asthenes] O27
    |    |--L. californiensis S60
    |    |--L. clathrata (Reeve 1847) SF77 [=Chiton clathratus O27, Ischnochiton (Lepidozona) clathratus O27]
    |    |--L. cooperi (Carpenter in Pilsbry 1892) [=Ischnochiton (Lepidozona) cooperi] O27
    |    |    |--L. c. cooperi O27
    |    |    `--L. c. acutior (Carpenter in Dall 1919) [=Ischnochiton (Lepidozona) cooperi acutior] O27
    |    |--L. crockeri (Willett in Hertlein & Strong 1951) PP78
    |    |--L. decipiens (Carpenter 1892) [=Ischnochiton (Lepidozona) decipiens] O27
    |    |--L. formosa Ferreira 1974 PP78
    |    |--L. golischi (Berry 1919) [=Ischnochiton (Lepidozona) golischi] O27
    |    |--L. ima Sirenko 1975 F77
    |    |--L. multigranosa Sirenko 1975 F77
    |    |--L. serrata (Carpenter 1864) PP78
    |    |--L. sinudentata (Carpenter 1892) [=Ischnochiton (Lepidozona) sinudentatus] O27
    |    |--L. stearnsii (Dall 1902) [=Ischnochiton (Lepidozona) stearnsii] O27
    |    |--L. thielei Sirenko 1975 F77
    |    `--L. willetti (Berry 1917) [=Ischnochiton (Lepidozona) willetti] O27
    `--Ischnochiton Gray 1847 SF77 (see below for synonymy)
         |  i. s.: I. adelaidensis (Reeve 1847) [=Chiton adelaidensis] H09
         |         I. allyni Ferreira 1977 F77
         |         I. beanii C64
         |         I. circumvallatus (Reeve 1847) P61
         |         I. comptus GO06
         |         I. elenensis C64
         |         I. granulifer Thiele 1909 P61
         |         I. limaciformis C64
         |         I. luteoroseus Suter 1907 P61
         |         I. magdalensis [incl. Chiton giganteus Tilesius 1824 (nom. inv.)] C64
         |         I. maorianus Iredale 1914 P61
         |         I. marloffsteinensis Fieldel & Keupp 1988 BE93
         |         I. mawsoni Cotton 1937 P61
         |         ‘Callochiton’ mortenseni Odhner 1924 F27
         |         I. parallelus C64
         |         ‘Chiton’ platessa [=*Levicoplax platessa] F27
         |         I. prasinatus C64
         |         I. punctulatissimus VN05
         |--I. (Ischnochiton) F77
         |    |--*I. (I.) crispus (Reeve 1847) [=Chiton crispus] F77
         |    |--I. (I.) longicymba O27
         |    |--I. (I.) numantius S60
         |    |--I. (I.) spiculosus S60
         |    `--I. (I.) textilis (Gray 1828) S60 [=Chiton textilis P61]
         |--I. (Anisoradsia Iredale & May 1916) S60
         |    `--I. (*A.) mawlei Iredale & May 1916 S60
         |--I. (Autochiton Iredale & Hull 1924) S60
         |    `--I. (*A.) torri Iredale & May 1916 S60
         |--I. (Chondropleura Thiele 1906) S60
         |    `--I. (*C.) exaratus (Sars 1878) [=Lophyrus exaratus] S60
         |--I. (Euporoplax Iredale & Hull 1924) S60
         |    `--I. (*E.) virgatus (Reeve 1847) [=Chiton virgatus] S60
         |--I. (Euretoplax Iredale & Hull 1924) S60
         |    `--I. (*E.) wilsoni Sykes 1896 S60
         |--I. (Haploplax Pilsbry 1894) (see below for synonymy) S60
         |    `--I. (*H.) smaragdinus (Angas 1867) [=Lophyrus smaragdinus] S60
         |--I. (Heterozona Carpenter in Dall 1879) S60
         |    `--I. (*H.) cariosus Carpenter in Pilsbry 1892 S60
         |--I. (Ischnoplax Carpenter in Dall 1879) S60
         |    `--I. (*I.) pectinatus (Sowerby 1840) S60 [=Chiton pectinatus S60]
         |--I. (Ischnoradsia Shuttleworth 1853) [incl. Lepidoradsia Carpenter in Dall 1879] S60
         |    `--I. (*I.) australis (Sowerby 1840) [=Chiton australis] S60
         |--I. (Isochiton Ashby & Cotton 1934) S60
         |    `--I. (*I.) bardwelli Ashby & Cotton 1934 S60
         |--I. (Lophyropsis Thiele 1893) S60
         |    `--I. (*L.) imitator (Smith 1881) [=Chiton (Ischnochiton) imitatrix] S60
         |--I. (Ovatoplax Cotton & Weeding 1939) S60
         |    `--I. (*O.) mayi Pilsbry 1895 [=I. (Haploplax) mayi] S60
         |--I. (Radsiella Pilsbry 1892) S60
         |    `--I. (*R.) tridentatus Pilsbry 1893 SF77
         |--I. (Rhodoplax Thiele 1893) SF77
         |    |--I. (R.) striolatus (Gray 1828) [incl. Chiton squamulosus Adams 1845, *Rhodoplax squamulosus] SF77
         |    |--I. (R.) eucosmius Dall 1919 SF77 [=Radsiella eucosmius PP78]
         |    `--I. (R.) petaloides (Gould 1846) (see below for synonymy) SF77
         |--I. (Rhombochiton Berry 1919) S60
         |    `--I. (*R.) regularis (Carpenter 1855) [=Chiton regularis] S60
         |--I. (Stenosemus Middendorff 1847) [incl. Lepidopleuroides Thiele 1893] S60
         |    `--I. (*S.) albus (Linné 1767) [=Chiton albus] S60
         |--I. (Strigichiton Hull 1923) S60
         |    `--I. (*S.) verconis Torr 1911 S60
         `--I. (Tripoplax Berry 1919) F77 [incl. Ischnoradsia Carpenter in Dall 1879 non Shuttleworth 1853 S60]
              `--I. (*T.) trifidus (Carpenter 1864) F77 [=Trachydermon trifidus S60]

*Cyanoplax hartwegii (Carpenter 1855) S60 [=Chiton hartwegii S60, Ischnochiton hartwegii C64, Lepidochitona (Cyanoplax) hartwegii O27]

Ischnochiton Gray 1847 SF77 [incl. Beanella Thiele 1893 non Dall 1882 S60, Diktuonus Ashby 1931 S60, Lepidochiton Carpenter 1857 S60, Leptopleura Thiele 1893 S60, Levicoplax Iredale & Hull 1925 S60, Lophyriscus Thiele 1893 S60, Stereoplax Thiele 1893 S60]

Ischnochiton (Haploplax Pilsbry 1894) [incl. Chartoplax Iredale & Hull 1924, Radsiella Thiele 1893 non Pilsbry 1892] S60

Ischnochiton (Rhodoplax) petaloides (Gould 1846) [=Chiton petaloides, Radsiella petaloides; incl. Ischnochiton mariposa Dall 1919] SF77

Lepidozona Pilsbry 1892 F77 [incl. Lepidopleurus Carpenter in Dall 1879 nec Risso 1826 nec Claparède 1868 S60, Solivaga Iredale & Hull 1925 S60]

*Type species of generic name indicated

REFERENCES

[BE93] Benton, M. J., & D. H. Erwin. 1993. Mollusca: Amphineura and ‘Monoplacophora’. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 125–130. Chapman & Hall: London.

[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.

[F77] Ferreira, A. J. 1977. A new species of chiton from the Aleutian Islands (Mollusca: Polyplacophora). Veliger 20 (1): 27–29.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[GW02] Giribet, G., & W. Wheeler. 2002. On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data. Invertebrate Biology 121 (4): 271–324.

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

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. 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 4^th ed. Whitcombe and Tombs Limited: Christchurch.

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SF77] Smith, A. G., & A. J. Ferreira. 1977. Chiton fauna of the Galápagos Islands. Veliger 20 (2): 82–97.

[VN05] Vinther, J., & C. Nielsen. 2005. The Early Cambrian Halkieria is a mollusc. Zoologica Scripta 34: 81–89.

Mopaliidae

Mopalia hindsii, copyright Al Kordesch.

Belongs within: Ischnochitonina.

The Mopaliidae are a group of chitons found in the Pacific Ocean and polar regions characterised by a more or less bristly girdle (Smith 1960).

Characters (from Smith 1960): Small to large chitons with tegmentum divided into usual lateral and central areas, sculpture combining radial and longitudinal ribs or lirae that vary in strength from low to high relief. Insertion-plate slits in all valves generally corresponding in position with ends of external ribs, teeth not pectinated. Head valve normally with eight slits but may be smaller or larger by fusion or splitting of one or more teeth, whereas in some forms slits are abnormally multiplied; intermediate and tail valves with single slit on each side, although in one group lacking in tail valve; tail valve with posteromedian sinus; sutural laminae well developed to very large. Girdle more or less hairy or bristly, never with scales, bristles simple or dendritic, in some species growing out of sutural pores, each of which bears one or more hairy bristles.

<==Mopaliidae [Amiculidae, Katharinidae]
    |--Katharina Gray 1847 [=Catharina Dunker 1882, Katherina (l. c.)] S60
    |    `--*K. tunicata (Wood 1815) O27 [=Chiton tunicatus O27; incl. Katherina douglasiae C64]
    |--Amicula Gray 1847 (see below for synonymy) S60
    |    |--*A. vestita (Broderip & Sowerby 1829) [=Chiton vestitus] S60
    |    |--A. amiculata Pallus 1786 [=Chlamydochiton amiculatus] O27
    |    `--A. pallasii (Middendorff 1846) [=Chiton pallasii] O27
    |--Placiphorella Carpenter in Dall 1879 SF77 (see below for synonymy)
    |    |--*P. velata Dall 1879 S60
    |    |--P. atlantica S60
    |    |--P. blainvillii (Broderip 1832) [=Chiton blainvillii] SF77
    |    |--P. borealis Pilsbry 1892 O27
    |    |--P. pacifica Berry 1919 O27
    |    |--P. rufa Berry 1917 O27
    |    `--P. stimpsoni Gould 1859 O27
    `--Mopalia Gray 1847 [=Molpalia Gray 1857; incl. Osteochiton Dall 1886] S60
         |  i. s.: M. acuta Cpr. 1855 [=Chiton acutus] C64
         |         M. blainvillei O27
         |         M. celetoides Dall 1919 O27
         |         M. chacei Berry 1919 O27
         |         M. chloris Dall 1919 O27
         |         M. ciliata (Sowerby 1840) [=Chiton ciliatus] O27
         |           |--M. c. ciliata O27
         |           |--M. c. elevata Pilsbry 1892 [=M. lignosa f. elevata] O27
         |           `--M. c. wosnessenskii Middendorff 1847 O27 [=Chiton wosnessenskii C64]
         |         M. cirrata Berry 1919 O27
         |         M. egretta Berry 1919 O27
         |         M. goniura Dall 1919 O27
         |         M. imporcata Carpenter 1864 O27
         |           |--M. i. imporcata O27
         |           `--M. i. lionotus Pilsbry 1918 O27
         |         M. lowei Pilsbry 1918 O27
         |         M. muscosa Gould 1846 O27 (see below for synonymy)
         |           |--M. m. muscosa O27
         |           |--M. m. acuta Carpenter 1855 O27
         |           |--M. m. kennerlyi Carpenter 1864 O27 [=M. grayii (preoc.) C64]
         |           |--M. m. laevior Pilsbry 1918 O27
         |           |--M. m. lignosa Gould 1846 O27 (see below for synonymy)
         |           |--M. m. porifera Pilsbry 1892 O27
         |           `--M. m. swanii Carpenter 1864 [=M. kennerleyi var. swanii] O27
         |         M. phorminx Berry 1919 O27
         |         M. simpsonii C64
         |         M. sinuata Carpenter 1864 O27
         |         M. stimpsoni (Gould 1859) [=Chiton (Molpalia) stimpsoni] C64
         |--*M. (Mopalia) hindsii (Sowerby in Reeve 1847) S60 [=Chiton hindsii S60, M. muscosa hindsii O27]
         |--M. (Dendrochiton Berry 1911) S60
         |    |--M. (*D.) thamnopora Berry 1911 S60 [=Chaetopleura (D.) thamnopora O27]
         |    |--‘Chaetopleura’ (D.) gothica (Carpenter 1864) O27 [=Ischnochiton (Trachydermon) gothicus C64, O27]
         |    `--‘Dendrochiton’ lirulatus Berry 1963 PP78
         |--M. (Hachijomopalia Taki 1954) S60
         |    `--M. (*H.) integra Taki 1954 S60
         `--M. (Semimopalia Dall 1919) S60
              `--H. (*S.) grisea Dall 1919 S60

Amicula Gray 1847 [incl. Chlamydochiton Dall 1878, Chlamydoconcha Pilsbry 1893 non Dall 1884, Cryptochiton Gray 1847 non Middendorff 1847, Stimpsoniella Carpenter 1873, Symmetrogephyrus Middendorff 1847] S60

Mopalia muscosa Gould 1846 O27 [=Acanthopleura muscosa C64, Chiton (Chaetopleura) muscosus C64; incl. C. armatus C64, C. consimilis Nutt. ms C64, Mopalia consimilis C64, C. ornatus Nutt. 1855 C64]

Mopalia muscosa lignosa Gould 1846 O27 [=Chiton lignosus C64; incl. C. merckii Middendorff 1847 C64, Mopalia merckii C64, C. montereyensis Cpr. 1855 C64, M. montereyensis C64, C. (Chaetopleura) vespertinus C64]

Placiphorella Carpenter in Dall 1879 SF77 [=Placophorella Fischer 1885 S60; incl. Euplacophora Verrill & Smith 1882 S60, Langfordiella Dall 1925 S60, Placophora Dall 1889 nec Agardh 1863 (ICBN) nec Fischer 1885 S60, Placophoropsis Pilsbry 1893 S60]

*Type species of generic name indicated

REFERENCES

[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.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. 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.

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SF77] Smith, A. G., & A. J. Ferreira. 1977. Chiton fauna of the Galápagos Islands. Veliger 20 (2): 82–97.

Chitonidae

West Indian green chiton Chiton tuberculatus, copyright James St. John.

Belongs within: Ischnochitonina.

The Chitonidae are a group of chitons characterised by the presence of well-developed pectination on the outside of the insertion plates (Smith 1960).

Characters (from Smith 1960): Small to large chitons with sculpture of tegmentum varying considerably in strength and character; articulamentum consisting of strong insertion plates and sutural laminae. Outside of insertion plates with well-developed pectination which ranges from coarsely grooved to fine and comb-like. Girdle may be scaly, with short bristles, spines, or spicules, or nude and leathery.

<==Chitonidae (see below for synonymy)
    |--Enoplochiton Gray 1847 SF77
    |    `--*E. niger (Barnes 1824) S60 [=Chiton niger SF77]
    |--Lucilina Dall 1882 (see below for synonymy) S60
    |    |--*L. confossa (Gould 1846) S60 [=Chiton confossus S60, Tonicia confossa H09]
    |    `--L. lamellosa (Quoy & Gaimard 1835) BW09
    |--Amaurochiton Thiele 1893 P61 [incl. Poeciloplax Thiele 1893 S60]
    |    |--A. magnificus (Deshayes 1827) S60, SF77 (see below for synonymy)
    |    `--A. glaucus (Gray 1828) P61
    |--Sypharochiton Thiele 1893 P61 [=Sympharochiton (l. c.) S60; incl. Triboplax Thiele 1893 S60]
    |    |--*S. pelliserpentis (Quoy & Gaimard 1835) [=Chiton pelliserpentis] P61
    |    |--S. sinclairi (Gray 1843) P61
    |    `--S. torri (Suter 1907) P61
    |--Liolophura Pilsbry 1893 [=Liolopleura (l. c.); incl. Clavarizona Hull 1923] S60
    |    |--*L. japonica (Lischke 1873) [=Chiton japonicus] S60
    |    |--L. curtisiana (Smith 1884) [=Chiton curtisianus] H09
    |    |--‘Clavarizona’ hirtosa M54
    |    `--L. gaimardi [incl. L. gaimardi var. queenslandica Pilsbry 1894] H09
    |--Onithochiton Gray 1847 P61 (see below for synonymy)
    |    |--O. neglectus Rochebrune 1881 P61 (see below for synonymy)
    |    |--‘Mopalia’ australis Suter 1907 P61
    |    |--O. lyellii (Sowerby 1832) [=Chiton lyellii] H09
    |    |--O. marmoratus Wissel 1904 P61
    |    |--O. opiniosus Iredale & Hull 1932 P61
    |    `--O. subantarcticus Suter 1907 P61 [=O. neglectus var. subantarcticus F27]
    |--Tonicia Gray 1847 SF77 (see below for synonymy)
    |    |--*T. elegans (Frembly 1827) [=Chiton elegans] S60
    |    |--T. forbesii Carpenter 1857 [incl. T. crenulata (Broderip 1832) non Chiton crenulatus Risso 1826] SF77
    |    |    |--T. f. forbesii SF77
    |    |    `--T. f. arnheimi Dall 1903 SF77
    |    |--T. fortilirata (Reeve 1847) [=Chiton fortiliratus] H09
    |    |--T. lineata C64
    |    |--T. lineolata C64
    |    |--T. picta (Reeve 1847) [=Chiton pictus] H09
    |    |--T. pustulifera Dall 1919 O27
    |    `--T. submarmorea [=Chiton submarmoreus] C64
    `--Chiton Linnaeus 1758 SF77 (see below for synonymy)
         |  i. s.: C. alveolus Sars in Lovén 1846 N79
         |         C. cancellatus N79
         |         C. caribeorum G79
         |         C. dispar [incl. Lepidopleurus adamsii] C64
         |         C. eschscholtzii Middendorff 1847 C64
         |         C. glaucus HS01
         |         C. granosus VN05
         |         C. hispidus Linnaeus 1758 L58
         |         C. latus Sowerby 1825 SF77
         |         C. lividus Middendorff 1847 C64
         |         C. marmoratus G79
         |         C. mendicarius Mighels & Adams 1842 J49
         |         C. olivaceus Spengler 1797 RP07
         |         C. platei Thiele 1910 BD86
         |         C. pulcherrimus Sowerby 1841 H09
         |         C. punctatus Linnaeus 1758 L58
         |         C. sanguineus C64
         |         C. scrobiculatus Middendorff 1847 C64
         |         C. segmentatus [incl. Chaetopleura apparata, Cha. rufocostata] G79
         |         C. sitchensis Middendorff 1847 C64
         |         C. squamosus G20
         |--C. (Chiton) S60
         |    |--*C. (C.) tuberculatus Linnaeus 1758 S60
         |    |--C. (C.) albolineatus S60
         |    |--C. (C.) berryi Smith et al. 1968 BE93
         |    |--C. (C.) cretaceus Berry 1940 BE93
         |    `--C. (C.) stokesii Broderip 1832 EO64, SF77 [=Lophyrus stokesii C64]
         |--C. (Anthochiton Thiele 1893) S60, P61 (see below for synonymy)
         |    |--C. (*A.) tulipa VWR04, P61
         |    |--‘Anthochiton’ aereus (Reeve 1847) P61 [=Rhyssoplax aerea F27; incl. R. oliveri Mestayer 1921 F27]
         |    |--‘Anthochiton’ canaliculatus (Quoy & Gaimard 1835) P61 [=Rhyssoplax canaliculata F27]
         |    |--‘Anthochiton’ clavatus (Suter 1907) P61
         |    |--‘Anthochiton’ huttoni (Suter 1906) P61 [=Rhyssoplax huttoni F27]
         |    |--‘C. (*Rhyssoplax)’ janierensis Gray 1821 S60
         |    |--‘Rhyssoplax’ limans F27
         |    |--C. (A.) macdonaldensis [=C. (Rhyssoplax) macdonaldensis] S60
         |    |--‘C. (Rhyssoplax)’ relatus S60
         |    |--‘Anthochiton’ stangeri (Reeve 1847) P61 [=Rhyssoplax stangeri HS01]
         |    `--‘Anthochiton’ suteri (Iredale 1910) P61
         |--C. (Delicatoplax Iredale & Hull 1926) S60
         |    `--C. (*D.) translucens Hedley & Hull 1909 S60 [=Rhyssoplax translucens F27]
         |--C. (Mucrosquama Iredale & Hull 1926) S60
         |    `--C. (*M.) carnosus Angas 1867 S60
         |--C. (Radsia Gray 1847) SF77, S60
         |    |--C. (R.) goodalli Broderip in Broderip & Sowerby 1832 (see below for synonymy) SF77
         |    `--C. (R.) sulcatus Wood 1815 [=Lophyrus sulcatus] SF77
         |--C. (Squamopleura Nierstrasz 1905) (see below for synonymy) S60
         |    `--C. (*S.) miles Carpenter in Pilsbry 1893 S60
         `--C. (Tegulaplax Iredale & Hull 1926) S60
              `--C. (*T.) howensis Hedley & Hull 1912 S60

Amaurochiton magnificus (Deshayes 1827) S60, SF77 [=Chiton magnificus S60; incl. C. olivaceus Frembly 1827 non Spengler 1797 S60, *Amaurochiton olivaceus P61]

Chiton Linnaeus 1758 SF77 [incl. Chondroplax Thiele 1893 S60, Diochiton Thiele 1893 S60, Georgus Thiele 1893 S60, Gymnoplax Gray 1821 S60, Lophyrus Poli 1791 S60, Scuterigulus Meuschen 1787 S60, Trachyodon Dall 1892 S60, Typhlochiton Dall 1921 S60]

Chiton (Anthochiton Thiele 1893) S60, P61 [incl. Rhyssoplax Thiele 1893 P61, Clathropleura Thiele 1893 non Tiberi 1877 S60]

Chiton (Radsia) goodalli Broderip in Broderip & Sowerby 1832 [=Lophyrus goodalli; incl. C. (R.) chierchiae Nierstrasz 1908] SF77

Chiton (Squamopleura Nierstrasz 1905) [incl. Sclerochiton Carpenter in Dall 1882 non Kraatz 1859, Sklerochiton (l. c.), Slerochiton (l. c.)] S60

Chitonidae [Amaurochitoninae, Chitonacea, Chitones, Chitonia, Chitonina, Chitoninae, Gymnoplacidae, Liolophurinae, Lophyridae, Ornithochitoninae, Rhyssoplacinae, Sypharochitoninae, Toniciinae, Tonicinae]

Lucilina Dall 1882 [incl. Lucia Gould 1862 non Swainson 1833, Onithoplax Thiele 1910, Toniciopsis Thiele 1893] S60

Onithochiton Gray 1847 P61 [=Onitochiton (l. c.) S60, Onythochiton (l. c.) S60, Ornithochiton (l. c.) S60; incl. Onithella Mackay 1933 S60, Pristochiton Clessin 1903 S60]

Onithochiton neglectus Rochebrune 1881 P61 [incl. Chiton undulatus Quoy & Gaimard 1835 non Wood 1828 S60, *O. undulatus P61]

Tonicia Gray 1847 SF77 [=Tonichia Gray 1840 (n. n.) S60, Conicia (l. c.) S60, Tonica (l. c.) S60, Tonicea (l. c.) S60; incl. Fannettia Dall 1882 S60, Fannia Dall 1882 non Robineau-Desvoidy 1830 S60, Fannyia Gray 1857 S60]

*Type species of generic name indicated

REFERENCES

[BD86] Barash, A., & Z. Danin. 1986. Further additions to the knowledge of Indo-Pacific Mollusca in the Mediterranean Sea (Lessepsian migrants). Spixiana 9 (2): 117–141.

[BE93] Benton, M. J., & D. H. Erwin. 1993. Mollusca: Amphineura and ‘Monoplacophora’. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 125–130. Chapman & Hall: London.

[BW09] Bryce, C., & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177–208.

[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.

[EO64] Emerson, W. K., & W. E. Old Jr. 1964. Additional records from Cocos Island. Nautilus 77 (3): 90–92.

[F27] Finlay, H. J. 1927. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[G79] Guppy, R. J. L. 1879. First sketch of a marine invertebrate fauna of the Gulf of Paria and its neighbourhood. Part I.—Mollusca. Journal of Conchology 2: 151–172.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

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

[J49] Johnson, R. I. 1949. Jesse Wedgwood Mighels with a bibliography and a catalogue of his species. Occasional Papers on Mollusks 1 (14): 213–231.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M54] Macpherson, J. H. 1954. Molluscs (sea shells and snails). In: Willis, J. M. (ed.) The Archipelago of the Recherche pp. 55–63. Australian Geographical Society: Melbourne.

[N79] Norman, A. M. 1879. The Mollusca of the fiords near Bergen, Norway. Journal of Conchology 2: 8–77.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

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

[RP07] Rousset, V., F. Pleijel, G. W. Rouse, C. Erséus & M. E. Siddall. 2007. A molecular phylogeny of annelids. Cladistics 23: 41–63.

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SF77] Smith, A. G., & A. J. Ferreira. 1977. Chiton fauna of the Galápagos Islands. Veliger 20 (2): 82–97.

[VWR04] Vendrasco, M. J., T. E. Wood & B. N. Runnegar. 2004. Articulated Palaeozoic fossil with 17 plates greatly expands disparity of early chitons. Nature 429: 288–291.

[VN05] Vinther, J., & C. Nielsen. 2005. The Early Cambrian Halkieria is a mollusc. Zoologica Scripta 34: 81–89.

Acanthochitona

Acanthochitona crinita, copyright Serge Gofas.

Belongs within: Ischnochitonina.

Acanthochitona is a cosmopolitan genus of chitons in which the dorsum of valves is ornamented with a sculpture of rounded nodules (Smith 1960).

Characters (from Smith 1960): Small to medium-sized chitons, rather narrow and elongated, tegmentum sculpture consisting of separated nodules or pustules of round to tear-drop shape, generally arranged in radial lines or rows. Valve coverage complete. Tail valves with submedian mucro and no dorsal ribs. Insertion plates generally with five slits in head valve, one on each side of intermediate valves, and in tail valve one on each side with a wide shallow sinus between; all teeth sharp and nearly smooth; sutural laminae large. Girdle varying from nude to densely hairy, but invariably with series of bristle-bearing pores (normally 18), situated around head valve with single row on each side at valve sutures.

<==Acanthochitona Gray 1821 SF77 (see below for synonymy)
    |--*A. fascicularis (Linné 1766) S60 [=Chiton fascicularis S60, *Acanthochiton fascicularis P61]
    |--A. achates [=Chiton (Acanthochaetes) achates] C64
    |--A. adelaidae S60
    |--A. arragonites (Carpenter 1857) SF77
    |--‘Acanthochites’ asbestoides (Smith 1884) [=Chiton asbestoides] H09
    |--A. avicula (Carpenter 1866) [=Acanthochites avicula] SF77
    |--‘Acanthochiton’ brookesi Ashby 1926 P61
    |--‘Acanthochiton’ costatus A23
    |--A. crinita GO06
    |--A. crocodilus S60
    |    |--A. c. crocodilus S60
    |    `--A. c. debilior S60
    |--A. diegoensis Pilsbry 1893 [=A. arragonites var. diegoensis] O27
    |--A. hirudiniformis (Sowerby 1832) (see below for synonymy) SF77
    |--A. jacquelinae Smith & Ferreira 1977 SF77
    |--A. mariae HS01
    |--A. pilsbryoides S60
    |--A. retrojecta H15
    |--‘Acanthochiton’ speciosus A23
    |--A. spiculosa S60
    |--A. stygma (Rochebrune 1889) SF77
    |--‘Acanthochiton’ subviridis A23
    |--‘Acanthochiton’ thileniusi Thiele 1909 P61
    |--A. trianguloides S60
    |--A. violacea HS01
    `--A. zelandica (Quoy & Gaimard 1835) HS01, P61 [=Acanthochiton zelandicus P61]
         |--A. z. zelandica P61
         |--‘Acanthochiton zelandicus’ amplificatus Iredale & Hull 1930 P61
         `--‘Acanthochiton zelandicus’ hookeri Gray 1843 P61

Acanthochitona Gray 1821 SF77 [=Acanthochaetes (l. c.) S60, Acanthochetes (l. c.) S60, Acanthochistes (l. c.) S60, Acanthochites Leach in Risso 1826 S60, Acanthochiton Iredale 1915 S60, Acanthochitus (l. c.) S60, Acanthochoetes (l. c.) S60, Acanthocites (l. c.) S60, Acantochiton (l. c.) S60, Acantoctites (l. c.) S60; incl. Amycula Adams in Tapparone-Canefri 1874 S60, Chitonellus deBlainville 1825 non Lamarck 1819 S60, Crocochiton Cotton & Weeding 1939 S60, Eoplax Ashby & Cotton 1936 S60, Hamachiton Middendorff 1848 S60, Meturoplax Pilsbry 1894 S60, Phacellopleura Agassiz 1846 S60, Phacelopleura (l. c.) S60, Phakellopleura Guilding 1830 S60, Pseudoacanthochiton Šulc 1934 S60, Stectoplax Carpenter in Dall 1882 S60, Strecochiton Adams in Tapparone-Canefri 1874 S60, Stretochiton (l. c.) S60]

Acanthochitona hirudiniformis (Sowerby 1832) [=Chiton hirudiniformis, Acanthochites hirudiniformis, Tonicia hirudiniformis; incl. Acanthochitona coquimbensis Leloup 1941, Acanthochitona peruvianus Leloup 1941, Acanthochitona tabogensis Smith 1961] SF77

*Type species of generic name indicated

REFERENCES

[A23] Ashby, E. 1923. A new species and sub-species of Acanthochiton. Journal of the Royal Society of Western Australia 10 (4): 13–16.

[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.

[GO06] Giribet, G., A. Okusu, A. R. Lindgren, S. W. Huff, M. Schrödl & M. K. Nishiguchi. 2006. Evidence for a clade composed of molluscs with serially repeated structures: monoplacophorans are related to chitons. Proceedings of the National Academy of Sciences of the USA 103 (20): 7723–7728.

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

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

[H15] Hedley, C. 1915. Presidential address. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 1–77, pls 1–7.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

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

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SF77] Smith, A. G., & A. J. Ferreira. 1977. Chiton fauna of the Galápagos Islands. Veliger 20 (2): 82–97.

Neoloricata

Hanleya hanleyi, from Ávila & Sigwart (2013).

Belongs within: Aculifera.
Contains: Leptochiton, Lepidopleuridae, Ischnochitonina.

The Neoloricata are the clade of chitons, including all living species, in which the valves of the shell have a more or less hard inner, semiporcelanous layer (the articulamentum) that usually projects to overlap with adjoining valves (Smith 1960).

Characters (from Smith 1960): Valve structure consisting of periostracum, tegmentum, articulamentum, and hypostracum layers, sometimes also with mesostracum layer; articulamentum usually extending out from under tegmentum in form of insertion plates and sutural laminae.

Neoloricata [Lepidopleurida, Lepidopleurina] SB12
    |--Pedanochiton SB12
    `--+--Permochitonidae BE93
       |    |--Euleptochiton lebescontei (Barrois 1889) BE93
       |    `--Permochiton Iredale & Hull 1926 SB12, S60
       |         `--*P. australianus Iredale & Hull 1926 S60
       `--+--+--Glaphurochiton Raymond 1910 SB12, S60
          |  |    `--G. carbonarius VP17
          |  `--Gryphochiton Gray 1847 SB12, S60 [=Gryptochiton (l. c.) S60; Gryphochitonidae]
          `--+--Leptochiton SB12
             `--+--Chitonida VP17
                |    |--Lepidopleuridae SB12
                |    `--Ischnochitonina SB12
                |--Heterochitonidae [Ochmazochitonidae] BE93
                |    |--Lobarochiton anomalus (Rowley 1908) BE93
                |    |--Ochmazochiton comptus Hoare & Smith 1984 VP17, BE93
                |    |--Allochiton Fucini 1912 BE93, S60
                |    |    `--*A. gemmellaroi Fucini 1912 S60
                |    `--Heterochiton Fucini 1912 S60
                |         |--*H. giganteus Fucini 1912 S60
                |         `--H. vinassai S60
                `--Hanleya Gray 1857 SB12, S60 [=Hanleia (l. c.) S60, Hanleyia (l. c.) S60; Hanleyidae]
                     |--H. hanleyi (Bean in Thorpe 1844) S60 (see below for synonymy)
                     |    |--H. h. hanleyi O27
                     |    `--H. h. spicata Berry 1919 O27
                     |--H. brachyplax Simone & Jardim 2009 S11
                     |--H. multigranosa (Reuss 1860) BE93
                     `--H. nagelfar KC11

Neoloricata incertae sedis:
  Choriplax Pilsbry 1894 (see below for synonymy) S60
    `--*C. grayi (Adams & Angas 1864) [=Microplax grayi] S60
  Afossochiton Ashby 1925 [incl. Telochiton Ashby & Cotton 1939; Afossochitonidae, Afossochitonina] S60
    `--*A. cudmorei Ashby 1925 S60
  Llandeilochiton Bergenhayn 1955 [Llandeilochitonidae] S60
    `--*L. ashbyi Bergenhayn 1955 S60
  Beloplaxus Oehlert 1881 S60
    `--*B. sagittalis (Sandberger 1853) [=Chiton sagittalis] S60
  Glyptochiton deKoninck 1883 [Glyptochitonidae] S60
    |--*G. cordifer (de Koninck 1844) [=Chiton cordifer] S60
    |--G. quadratus (Etheridge 1882) BE93
    |--G. subquadratus (Kirkby & Young 1867) BE93
    `--G. youngianus (Kirkby in Young 1865) BE93
  Lavenachiton Cotton & Godfrey 1940 S60
    `--*L. cliftonensis (Ashby & Cotton 1939) [=Ischnochiton (Radsiella) cliftonensis] S60
  Lirachiton Ashby & Cotton 1939 [incl. Molachiton Ashby & Cotton 1939] S60
    `--*L. inexpectus (Ashby & Cotton 1931) [=Acanthochiton (*Lirachiton) inexpectus] S60
  Oochiton Ashby 1929 S60
    `--*O. halli Ashby 1929 S60
  Protalochiton Rochebrune 1883 S60
    `--*P. setlensis Rochebrune 1883 S60
  Pseudoischnochiton Ashby 1930 S60
    `--*P. wynyardensis Ashby 1930 S60
  Pterygochiton Rochebrune 1883 S60
    |--*P. terquemi (Deslongchamps 1859) [=Chiton terquemi] S60
    `--P. busambrensis S60
  Sulcochiton Ryckholt 1862 S60
    `--*S. grayi Ryckholt 1862 S60
  Trachypleura Jaeckel 1900 S60
    `--*T. triadomarchica Jaeckel 1900 S60
  Cymatochiton Dall 1882 S60 [Cymatochitonidae BE93]
    |--*C. loftusianus (King 1850) S60, BE93 [=Chiton loftusianus S60]
    |--C. howseanus (Kirkby 1857) BE93
    `--C. kirkbyi (de Koninck 1883) BE93
  Lekiskochiton [Lekiskochitonidae] BE93
    |--L. cordatus (Kirkby 1859) BE93
    `--L. fornicis Hoare & Smith 1984 BE93

Choriplax Pilsbry 1894 [incl. Microplax Adams & Angas 1864 non Fieber 1861; Choriplacidae, Choriplaxidae] S60

Hanleya hanleyi (Bean in Thorpe 1844) S60 [=Chiton hanleyi S60; incl. *H. debilis Gray 1857 S60, Chiton debilis O27]

*Type species of generic name indicated

REFERENCES

[BE93] Benton, M. J., & D. H. Erwin. 1993. Mollusca: Amphineura and ‘Monoplacophora’. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 125–130. Chapman & Hall: London.

[KC11] Kocot, K. M., J. T. Cannon, C. Todt, M. R. Citarella, A. B. Kohn, A. Meyer, S. R. Santos, C. Schander, L. L. Moroz, B. Lieb & K. M. Halanych. 2011. Phylogenomics reveals deep molluscan relationships. Nature 477: 452–456.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt. 3. Stanford University Press: Stanford University (California).

[S11] Simone, L. R. L. 2011. Phylogeny of the Caenogastropoda (Mollusca), based on comparative morphology. Arquivos de Zoologia 42 (4): 161–323.

[S60] Smith, A. G. 1960. Amphineura. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt I. Mollusca 1: Mollusca—General Features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—General Features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia pp. I41–I76. Geological Society of America, and University of Kansas Press.

[SB12] Sutton, M. D., D. E. G. Briggs, D. J. Siveter, D. J. Siveter & J. D. Sigwart. 2012. A Silurian armoured aplacophoran and implications for molluscan phylogeny. Nature 490: 94–97.

[VP17] Vinther, J., L. Parry, D. E. G. Briggs & P. Van Roy. 2017. Ancestral morphology of crown-group molluscs revealed by a new Ordovician stem aculiferan. Nature 542: 471–474.