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Hyoliths were a group of small shelled animals that mostly lived during the Cambrian and Ordovician (though probable hyoliths remained alive as late as the Permian—Peel & Yochelson 1984). The relationship of hyoliths to other animals is uncertain—they have most often been treated as molluscs, but seemingly mostly by default. Hyoliths can be divided into two main groups (Kruse 2002): Orthothecimorpha (containing Circothecida and Orthothecida), in which the aperture and conch growth lines are planar, and Hyolithomorpha (containing Sulcavitida and Hyolithida), in which the aperture has a prominent ventral ligula. Hyolithomorphs (at least) also possessed a pair of projecting struts called helens on either side of the operculum; being delicate, these are rarely preserved, so the absence to date of any records of helens in orthothecimorphs does not necessarily indicate their absence.

Characteristics (from Kruse 2002): Bilaterally symmetrical conical conch with aperture at one end only, closed by an operculum.

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Hyolitha [Calyptoptomatida, Hyolithoidea, Hyolithozoes]
   |--Orthothecimorpha [Orthothecomorpha] K02
   |    |-- K02
   |    `-- K02
   `--Hyolithomorpha K02
        |-- K02
        `--Hyolithida [Hyolithina] K02
             |-- K02
             `-- M02

Hyolitha incertae sedis:

 Cupittheca K02
   |--C. clathrata (Bengtson in Bengtson et al. 1990) K02
   |--C. hemicyclata (Bengtson in Bengtson et al. 1990) K02
   `--C. holocyclata (Bengtson in Bengtson et al. 1990) K02
 ‘Salterella’ planoconvexa Tate 1892 K02
 Conotheca Missarzhevksy in Rozanov et al. 1969 KB09
   |--C. australiensis Bengtson in Bengtson et al. 1990 K02
   `--C. circumflexa Missarzhevksy in Rozanov et al. 1969 KB09
 Parkula bounites Bengtson in Bengtson et al. 1990 K02
 Hyptiotheca karraculum Bengtson in Bengtson et al. 1990 K02
 Triplicatella disdoma Conway Morris in Bengtson et al. 1990 K02
 Nganki wumirri Kruse 1990 K02
 Globoritubulus Qian & Zhang 1985 K02
 Eumorpholites bouceki Marek 1967 K02
 Elegantilites tigris Marek 1967 K02
 Insignicornus Sysoyev 1973 K02
 Nevadotheca Malinky 1988 M02
   |--N. arenophilus M02
   |--‘Hyolithus’ socialis M02
   `--‘Hyolithus’ terranovicus Walcott 1891 M02
 Emarites Marek, Malinky & Geyer 1997 M02
 Ambrolinevitus Syssoiev 1958 M02
   `--*A. straitellus (Syssoiev 1958) [=Hyolithus striatellus] M02
 Recilites solitarius (Barrande 1867) M02
 Polylopia Clark 1925 M02
 Tetratheca M02
   `--‘Quadrotheca’ clinisepta Syssoiev 1960 [=Costatheca clinisepta] M02
 Bactrotheca Novák 1891 M02
 Ensitheca Duan 1984 M02
 ‘Hyolithes’ globiger Saito 1936 [Globorilida] M90
 Hyolithellida M90
 Toxeumorphoridae [Toxeumorphorida, Xenoconchia] PY84
   |--Pseudotoxeuma magnifica Shimanskiy 1963 PY84
   |--Toxeumorphora PY84
   |    |--T. dombarense Shimanskiy 1963 PY84
   |    `--T. langi Shimanskiy 1963 PY84
   `--Macrotheca Waagen 1880 PY84
        |--*M. wynnei Waagen 1880 PY84
        |--M. almgreeni Peel & Yochelson 1984 PY84
        `--M. timorensis Wanner 1941 PY84
 Helenia Walcott 1889 [incl. Rhombocorniculum Walliser 1958] DC71
   `--H. cancellata Cobbold 1921 [incl. *Rhombocorniculum comleyense Walliser 1958] DC71
 Gompholites striatulus (Barrande 1847) MB05
 Maxilites Marek 1972 MB05
   |--M. robustus (Barrande 1867) MB05
   `--M. snajdri Marek 1972 MB05
 Slapylites signatulus MB05
 Tizilites creber MB05
 Dilytes occultus MB05
 Leolites MB05
   |--L. cognatus MB05
   `--L. paucicostatus MB05
 Oboedalites MB05
   |--O. oboediens MB05
   `--O. peregrinus MB05
 Chimerolites temperatus MB05
 Paolites obvius MB05

* Type species of generic name indicated

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[DC71] Druce, E. C., & P. J. Jones. 1971. Cambro-Ordovician conodonts from the Burke River Structural Belt, Queensland. Commonwealth of Australia, Bureau of National Development, Bureau of Mineral Resources, Geology and Geophysics, Bulletin 110: 1-159.

[KB09] Kouchinsky, A., S. Bengtson, W. Feng, R. Kutygin & A. Val'kov. 2009. The Lower Cambrian fossil anabaritids: affinities, occurrences and systematics. Journal of Systematic Palaeontology 7 (3): 241-298.

[K02] Kruse, P. D. 2002. Biostratigraphic potential of Middle Cambrian hyoliths from the eastern Georgina Basin. Alcheringa 26: 353-398.

[M90] Malinky, J. M. 1990. Cambrian Hyolitha from northeast Canada: Reappraisal of the hyolith orders Camerothecida and Diplothecida. Journal of Paleontology 64 (4): 587-595.

[M02] Malinky, J. M. 2002. A revision of Early to Mid Ordovician hyoliths from Sweden. Palaeontology 45: 511-555.

[MB05] Mus, M. M. & J. Bergström. 2005. The morphology of hyolithids and its functional implications. Palaeontology 48 (6): 1139-1167.

[PY84] Peel, J. S., & E. L. Yochelson. 1984. Permian Toxeumorphorida from Greenland: an appraisal of the molluscan class Xenoconchia. Lethaia 17: 211-221.


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