The phylogeny and classification of the phylum Cephalorhyncha

A.V. Adrianov & V.V. Malakhov

Abstract. The phylum Cephalorhyncha is a taxon including headproboscis worms: Priapulida, Loricifera, Kinorhyncha, Nematomorpha. The body of all Cephalorchyncha is divided into proboscis (= introvert) and trunk. The introvert is capable of invaginating into the trunk and is subdivided into the mouth cone, central part with sensory-locomotory scalids, and neck region. The central nervous system consists of nerve ring located in the introvert at the base of mouth cone, and ventral cord. The cephalorhynch worms of Middle Cambrian Burdgess Shale fauna are reviewed. Ancalagon and Fieldia are the most primitive cephalorhynchs. Their proboscis is not able to invaginate completely. Ancalagon and Fieldia could be included into a separate subphylum Protocephalorhyncha. Another subphylum, Eucephalorhyncha, is characterized by the introvert being able to invaginate into the trunk. Subphylum Eucephalorhyncha includes the classes Louisellida (MiddleCambrian only), Priapulida (with Middle Cambrian Ottoia and Selkirkia, Carboniferous Priapulites and several recent genera), Loricifera (considered to be neotenic group), Kinorhyncha, and Nematomorpha, the most advanced group, where only larvae retain the structure of headproboscis worm. Diagnoses of the phylum Cephalorhyncha, new subphyla Protocephalorhyncha and Eucephalorhyncha, and all classes and orders of extinct and extant cephalorhynchs are presented.

Zoosystematica Rossica, 1994, 3(2): 181–201  ▪  Published in print 1 August 1995


  

References

Adrianov, A.V. & Malakhov, V.V. 1990. Sense organs of Kinorhyncha by example of the order Homalorhagida (Cephalorhyncha, Kinorhyncha). Zool. Zhurn., 69(4): 5-14. (In Russian).

Adrianov, A.V. & Malakhov, V.V. 1991a. The nervous system of kinorhynchs (Cephalorhyncha, Kinorhyncha). Zool. Zhurn., 70(2): 5-12. (In Russian).

Adrianov, A.V. & Malakhov, V.V. 1991b. Body cavity of Kinorhyncha. Dokl. Akad. Nauk SSSR, 317(2): 506-509. (In Russian).

Adrianov, A.V. & Malakhov, V.V. 1991c. The fine structure of spermatozoids and peculiar features of spermatogenesis in Pycnophyes kielensis (Homalorhagida, Pycnophyidae). Zool. Zhurn., 70(4): 28-36. (In Russian).

Adrianov, A.V. & Malakhov, V.V. 1991d. Fine structure of the reproductive system in Cephalorhyncha, Kinorhyncha. Zool. Zhurn., 70(5): 28-38. (In Russian).

Adrianov, A.V. & Malakhov, V.V. 1994. Kinorinkhi: stroenie, razvitie, filogeniya i sistematika [Kinorhyncha: structure, development, phylogeny and taxonomy]. 276 p. Moscow: Nauka. (In Russian).

Adrianov, A.V., Malakhov, V.V. & Yushin, V.V. 1989a. Loricifera - a new taxon of marine invertebrates. Biologiya Morya, 2: 70-72. (In Russian).

Adrianov, A.V., Malakhov, V.V., Tchesunov, A.V. & Tzetlin, A.B. 1989b. A dwarf priapulida Tubiluchus arcticus sp. n. from the White Sea. Zool. Zhurn., 68(3): 126-131. (In Russian).

Adrianov, A.V., Malakhov, V.V. & Yushin, V.V. 1990a. Fine structure of the cuticle of Pycnophyes kielensis (Kinorhyncha, Homalorhagida). Zool. Zhurn., 69(1): 5-11. (In Russian).

Adrianov, A.V., Chesunov A.V. & Spiridonov, S.E. 1990. Post-embryonic development of Pycnophyes kielensis (Kinorhyncha, Homalorhagida). Zool. Zhurn., 69(6): 132-137. (In Russian).

Andrassy, I. A. 1976. Evolution as a basis for the systematization of nematodes. 228 p. Budapest: Akade'miai Kiado'.

Beklemishev, V.N. 1964. Osnovy sravnitelnoy anatomii bespozvonochnykh [Principles of comparative anatomy of invertebrates]. Vol.1. 432 p. Moscow: Nauka.

Bresciani, J. 1991. Nematomorpha. In: E. Harrison (ed.). Microscopic anatomy of invertebrates, 4: 197-218. London, New York: Wiley-Liss Inc.

Brien, P. 1961. Elements de zoologie. Notions dy anatomie comparee. 2. Les Coelomates Hyponeriens. 715 p. Liege.

Chitwood, B. & Chitwood, M. 1950. Introduction to nematology. 213 p. Baltimore (Md): Monum. Print. Co.

Conway Morris, S. 1977. Fossil priapulid worms. Spec. Pap. Paleontol., 20: 1-101.

Eakin, R.M. & Brandenburger, J.L. 1974. Ultrastructural features of a Gordian worm (Nematomorpha). J. Ultrastr. Res., 46: 351-374. https://doi.org/10.1016/S0022-5320(74)90062-8

Grasse, O. Poisson, R. & Tuzet, O. 1961. Zoologie. 1. Invertebres. 919 p. Paris.

Grobben, K. 1908. Die systematische Einteilung des Tierreiches. Verh. zool.-bot. Ges. Wien, 58(10): 491-511.

Hammarsten, O. 1913. Beitrage zur Entwicklung von Halicryptus spinulosus Sieb. Zool. Anz., 41: 501-505.

Hammarsten, O. 1915. Zur Entwicklungsgeschichte von Halicryptus spinulosus Sieb. Zeitschr. wiss. Zool., 112: 525-571.

Hammond, R. 1967. The fine structure of the trunk and praesoma wall of Acanthocephalus ranae (Schrank 1788) Lune, 1911. Parasitology, 57: 475-486. https://doi.org/10.1017/S003118200007236X

Higgins, R.P. & Kristensen, R.M. 1986. New Loricifera from Southeastern United States coastal waters. Smithson. Contr. Zool., 438: 1-70. https://doi.org/10.5479/si.00810282.438

Higgins, R.P. & Storch, V. 1989. Ultrastructural observations of the larva of Tubiluchus corallicola (Priapulida). Helgol. Meersuntersuch., 43: 1-11. https://doi.org/10.1007/BF02365545

Higgins, R.P. & Storch, V. 1991. Evidence for direct development of Meiopriapulus fijiensis (Priapulida). Trans, amer. micr. Soc., 110: 37-46. https://doi.org/10.2307/3226738

Hyman, L.H. 1951. The Invertebrates: Acanthocephala, Aschelminthes and Entoprocta. The pseudocoelomate Bilateria. Vol. 3. 572 p. New York, Toronto, London: McGraw-Hill.

Ioffe, B.I. & Kotikova, E.A. 1988. Nervous system of Priapulus caudatus and Halicryptus spinulosus (Priapulida). Trudy zool. Inst. Akad. Nauk SSSR, 183: 52-77. (In Russian).

Kaestner, A. 1965. Lehrbuch der speziellen Zoologie. I. Bd. 1. Wirbellose. 845 p. Stuttgart: G. Fischer.

Kristensen, R.M. 1983. Loricifera, a new phylum with Aschelminthes characters from the meiobenthos. Z. Zool. Syst. Evol., 21: 163-180. https://doi.org/10.1111/j.1439-0469.1983.tb00285.x

Kristensen, R.M. 1991a. Loricifera. In: E. Harrison (ed.). Microscopic anatomy of invertebrates, 4: 351 -375. London, New York: Wiley-Liss Inc.

Kristensen, R.M. 1991b. Loricifera - a general biological and phylogenetic overview. Zoological systematics Symposium: New higher taxa presented by their discoverers. Verh. dtsch. zool Ges., 84: 231-246.

Kristensen, R.M. & Higgins, R.P. 1991. Kinorhyncha. In: E. Harrison (ed.). Microscopic anatomy of invertebrates, 4: 377-404. London, New York: Wiley-Liss Inc.

Kristensen, R.M. & Shirayama, Y. 1988. Pliciloricus hadalis (Pliciloricidae), a new loriciferan species collected from the Izu-Ogasawara trench, Western Pacific. Zool. Science, 5: 875-881.

Land, J. van der. 1968. A new Aschelminth, probably related to the Priapulida. Zool. Meded. Leiden, 42: 237-250.

Land, J. van der. 1970. Priapulida. Zool. Verhandl, 112: 1-118.

Land, J. van der. 1982. A new species of Tubiluchus (Priapulida) from the Red Sea. Netherl. J. Zool., 32: 324-335. https://doi.org/10.1163/002829681X00365

Land, J. van der. 1985. A new species of Tubiluchus (Priapulida) from the Pacific Ocean. Proc. ned. Akad. Wet. (C), 88: 371-377.

Lang, K. 1953. Die Entwicklung des Eies von Priapulus caudatus Lam. und die systematische Stellung der Priapuliden. Arkiv Zool. (2), 4: 321-348.

Malakhov, V.V. 1980. Cephalorhyncha-a new phylum of the animal kingdom uniting Priapulida, Kinorhyncha, and Gordiacea, and a system of Aschelminthes worms. Zool. Zhurn., 59(4): 485-499. (In Russian).

Malakhov, V.V. 1986. Nematody: stroenie, razvitie, sistemaifilogeniya [Nematodes: structure, development, classification, and phylogeny]. 215 p. Moscow: Nauka. (In Russian).

Malakhov, V.V. 1994. Nematodes: structure, development, classification, and phylogeny. W. Duane Hope (ed). 286 p. Washington & London: Smithsonian Institution Press.

May, H.G. 1919. Contributions to the life histories of Gordius robustus Leidy and Paragordius varius (Leidy). Biol. Monogr., 5: 1-119. https://doi.org/10.5962/bhl.title.16768

Morse, M.P. 1981. Meiopriapulus fijiensis n. gen., n. sp.: an interstitial priapulid from coarse sand in Fiji. Trans. amer. Micr. Soc., 100(3): 239-252. https://doi.org/10.2307/3225549

Neuhaus, B. 1988. Ultrastrueture of theprotonephridia in Pycnophyes kielensis (Kinorhyncha, Homalorhagida). Zoomorphology, 108: 245-253.

Neuhaus, B. 1991. Zur Ultrastruktur, Postembryonalentwicklung und phylogenetischen Verwandtschaft der Kinorhyncha, Dissertation zur Erlangung des Doktorgrades. Universitat Gottingen. 190 p.

Osipova, M.W. 1982. To the study of the hydraulic locomotion of Priapulus caudatus. Zool. Zhurn., 61(2): 170-181. (In Russian).

Pagenstecher, H.A. 1875. Allgemeine Zoologie oder Grundgesetze des tierischen Baus und Lebens. 275 p. Berlin: Wiegandt & al.

Por, F.D. & Bromley, H.J. 1974. Morphology and anatomy of Maccabeus tenteculatus (Priapulida, Seticoronaria). J. zool. Soc. London, 173: 173-197. https://doi.org/10.1111/j.1469-7998.1974.tb03125.x

Salvini-Plawen, L. von. 1974. Zur Morphologie und Systematik der Priapulidae: Chaetostephanus praeposteriens, der Vertreter einer neuen Ordnung Seticoronaria. Zeitschr. zool. Syst. Evolutionsforsch., 12: 31-54. https://doi.org/10.1111/j.1439-0469.1974.tb00156.x

Schepotieff, A. 1907. Die Echinoderiden. Zeitschr. wiss. Zool., 88: 291-326.

Schram, F.R. 1973. Pseudocoelomates and a nemertine from the Illinois pennsylvanian. J. Paleont., 47: 985-989.

Skryabin, K.I. & Schulz, R.S. 1931. Gelmintozy cheloveka [Helminthoses of man]. In: Osnovy meditsinskoy gelmintologii [Fundamentals of medical helminthology ]. 378 p. Moscow: Gosmedizdat.

Storch, V., Higgins, R.P. & Morse, P. 1989. Internal anatomy of Meiopriapulus fijiensis (Priapulida). Trans. amer. micr. Soc., 108: 245-261. https://doi.org/10.2307/3226343

Storch, V.M., Higgins, R.P. & Rumohr, H. 1990. Ultrastructure of introvert and pharynx of Halicryptus spinulosus (Priapulida). J. Morphol., 206: 163-171. https://doi.org/10.1002/jmor.1052060203

Storch, V., Higgins, R.P., Malakhov, V.V. & Adrianov, A.V. 1994. Microscopic anatomy and ultrastructure of the introvert of Priapulus caudatus and P. tuberculatospinosus (Priapulida). J. Morphol, 220: 281-293. https://doi.org/10.1002/jmor.1052200307

Stormer, L. 1944. On the relationships and phylogeny of fossil and recent Arachnomorpha, a comparative study of Arachnida, Xiphosura, Eurypterida, Trilobita and other fossil Arthropoda. Skr. norske Vidensk. Akad., 5: 1-158.

Walcott, C. 1911a. Middle Cambrian annelids. Cambrian geology and paleontology. II. Smithson. misc. Collns., 57: 109-144.

Walcott, C. 1911b. Middle Cambrian holothurians and medusae. Cambrian geology and paleontology. II. Smithson. misc. Collns., 57: 41-68.

Walcott, C. 1911c. Middle Cambrian annelids. Cambrian geology and paleontology. II. Smithson. misc. Collns., 57: 109-144.

Walcott, C. 1912. Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merosomata. Cambrian geology and paleontology. II. Smithson. misc. Collns., 57: 145-228.

Walcott, C. 1931. Addenda to description of Burgess Shale fossils (with explanatory notes by C.E. Resser). Smithson. misc. Collns., 85: 1-46.

Zapotosky, J.E. 1971. The cuticular ultrastrueture of Paragordius varius (Leidy 1851). Proc. helminthol. Soc. Wash., 38: 228-236.

Zapotosky, J.E. 1974. Fine structure of the larval stage of Paragordius varius. I. Preseptum. Proc. helminthol. Soc. Wash., 41: 209-221.

Zapotosky, J.E. 1975. Fine structure of the larval stage of Paragordius varius (Leidy 1851). II. The Postseptum. Proc. helminthol Soc. Wash., 42: 103-111.

Zelinka, C. 1928. Monographie der Echinodera. 396 p. Leipzig: W. Engelmann.

Zenkevich, L.A. 1944. Essay on the evolution of locomotory apparates of animals. Zhurn. obshchey Biol., 5(3): 129-148. (In Russian).

Volume, Year:
Vol. 33. 2024
Vol. 32. 2023
Vol. 31. 2022
Vol. 30. 2021
Vol. 29. 2020
Vol. 28. 2019
Vol. 27. 2018
Vol. 26. 2017
Vol. 25. 2016
Vol. 24. 2015
Vol. 23. 2014
Vol. 22. 2013
Vol. 21. 2012
Vol. 20. 2011
Vol. 19. 2010
Vol. 18. 2009
Vol. 17. 2008
Vol. 16. 2007
Vol. 15. 2006
Vol. 14. 2005
Vol. 13. 2004
Vol. 12. 2003
Vol. 11. 2002
Vol. 10. 2001
Vol. 9. 2000
Vol. 8. 1999
Vol. 7. 1998
Vol. 6. 1997
Vol. 5. 1996
Vol. 4. 1995
Vol. 3. 1994
Vol. 2. 1993
Vol. 1. 1992
Supplementa:
Suppl. 3. 2020
Suppl. 2. 2018