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Agnathans
Lampetra fluviatilis
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Superclass: Agnatha
Cope, 1889
Excluded groups

Gnathostomata[1]

Agnatha (Latin, "no jaws") is a superclass of jawless fish in the phylum Chordata, subphylum Vertebrata. The group excludes all vertebrates with jaws, known as gnathostomes.

The agnathans as a whole are paraphyletic,[2] because most extinct agnathans belong to the stem group of gnathostomes.[3][4] Recent molecular data, both from rRNA[5] and from mtDNA[6] strongly supports the theory that living agnathans, known as cyclostomes, are monophyletic.[7]

The oldest fossil agnathans appeared in the Cambrian, and two groups still survive today: the lampreys and the hagfish, with about 100 species in total. Hagfish are considered members of the subphylum Vertebrata, because they secondarily lost vertebrae; before this event was inferred from molecular [5][6][8]and developmental [9] data, the group Craniata was created by Linnaeus (and is still sometimes used as a strictly morphological descriptor) to reference hagfish plus vertebrates. In addition to the absence of jaws, modern agnathans are characterised by absence of paired fins; the presence of a notochord both in larvae and adults; and seven or more paired gill pouches. There is a light sensitive pineal eye (homologous to the pineal gland in mammals). All living and most extinct Agnatha do not have an identifiable stomach or any appendages. Fertilization and development are both external. There is no parental care in the Agnatha class. The Agnatha are ectothermic or cold blooded, with a cartilaginous skeleton, and the heart contains 2 chambers.

While a few scientists still regard the living agnaths as only superficially similar, and argue that many of these similarities are probably shared basal characteristics of ancient vertebrates, recent classifications clearly place hagfish (the Myxini or Hyperotreti), with the lampreys (Hyperoartii) as being more closely related to each other than either is to the jawed fishes.

Metabolism

Agnathans are ectothermic, meaning they do not regulate their own body temperature. Agnathan metabolism is slow in cold water, and therefore they do not have to eat very much. They have no distinct stomach, but rather a long gut, more or less homogenous throughout its length. Lampreys feed off other fish and mammals. They rely on a row of sharp teeth to shred their host. Anticoagulant fluids preventing clotting are injected into the host, causing the host to yield more blood. Hagfish are decomposers, eating mostly dead animals. They also use a sharp set of teeth to break down the animal. The fact that all Agnathan teeth are not able to move up and down limit their possible food types.

Body covering

The only modern Agnathan body covering is skin, with neither dermal or epidermal scales. The skin of hagfish has copious slime glands, the slime constituting their defense mechanism, the slime can sometimes clog up enemy fish's gills, causing them to die. Many extinct agnathans sported heavy dermal armour or small mineralized scales (see below).

Appendages

Most agnathans, including all those living today have no paired appendages, although they do have a dorsal and a caudal fin. Some fossil agnathans, such as osteostracans, did have paired fins, a trait inherited in their jawed descendants.[10]

Reproduction

Fertilization in lampreys is external. Mode of fertilization in hagfishes is not known. Development in both groups probably is external. There is no known parental care. Not much is known about the hagfish reproductive process. It is believed that hagfish only have 30 eggs over a lifetime.[11] Most species are hermaphrodites. There is very little of the larval stage that characterizes the lamprey. Lamprey are only able to reproduce once. After external fertilization, the lamprey's cloacas remain open, allowing a fungus to enter their intestines, killing them. Lampreys reproduce in freshwater riverbeds, working in pairs to build a nest and burying their eggs about an inch beneath the sediment. The resulting hatchlings go through four years of larval development before becoming adults. They also have a certain unusual form of reproduction.

Fossil agnathans

Although a minor element of modern marine fauna, Agnatha were prominent among the early fish in the early Paleozoic. Two types of Early Cambrian animal apparently having fins, vertebrate musculature, and gills are known from the early Cambrian Maotianshan shales of China: Haikouichthys and Myllokunmingia. They have been tentatively assigned to Agnatha by Janvier. A third possible agnathid from the same region is Haikouella. A possible agnathid that has not been formally described was reported by Simonetti from the Middle Cambrian Burgess Shale of British Columbia.

Many Ordovician, Silurian, and Devonian agnathans were armored with heavy bony-spiky plates. The first armored agnathans—the Ostracoderms, precursors to the bony fish and hence to the tetrapods (including humans)—are known from the middle Ordovician, and by the Late Silurian the agnathans had reached the high point of their evolution. Most of the ostracoderms, such as thelodonts, osteostracans, and galeaspids, were more closely related to the gnathostomes than to the surviving agnathans, known as cyclostomes. Cyclostomes apparently split from other agnathans before the evolution of dentine and bone, which are present in many fossil agnathans, including conodonts.[12] Agnathans declined in the Devonian and never recovered.

Groups

See also

References

  1. ^ Colbert, E.H. & Morales, M. (2001): Colbert's Evolution of the Vertebrates: A History of the Backboned Animals Through Time. 4th edition. John Wiley & Sons, Inc, New York - ISBN 978-0-471-38461-8.
  2. ^ Purnell, M. A. (2001). Derek E. G. Briggs and Peter R. Crowther, ed. Palaeobiology II. Oxford: Blackwell Publishing. p. 401. ISBN 0-632-05149-3. 
  3. ^ Zhao Wen-Jin, Zhu Min (2007). "Diversification and faunal shift of Siluro-Devonian vertebrates of China". Geological Journal. 42 ((3-4)): 351–369. doi:10.1002/gj.1072. 
  4. ^ Sansom, Robert S. (2009). "Phylogeny, classification, & character polarity of the Osteostraci (Vertebrata)". Journal of Systematic Palaeontology. 7: 95–115. doi:10.1017/S1477201908002551. 
  5. ^ a b Mallatt, J., and J. Sullivan. 1998. (1998). "28S and 18S ribosomal DNA sequences support the monophyly of lampreys and hagfishes.". Molecular Biology and Evolution. 15 (12): 1706–1718. PMID 9866205. 
  6. ^ a b DeLarbre Christiane ; Gallut Cyril ; Barriel Veronique ; Janvier Philippe ; Gachelin Gabriel (2002). "Complete mitochondrial DNA of the hagfish, Eptatretus burgeri: The comparative analysis of mitochondrial DNA sequences strongly supports the cyclostome monophyly.". Molecular Phylogenetics & Evolution. 22 (2): 184–192. PMID 11820840. doi:10.1006/mpev.2001.1045. 
  7. ^ Janvier, P. 2010. "MicroRNAs revive old views about jawless vertebrate divergence and evolution." Proceedings of the National Academy of Sciences (USA) 107:19137-19138. [1] "Although I was among the early supporters of vertebrate paraphyly, I am impressed by the evidence provided by Heimberg et al. and prepared to admit that cyclostomes are, in fact, monophyletic. The consequence is that they may tell us little, if anything, about the dawn of vertebrate evolution, except that the intuitions of 19th century zoologists were correct in assuming that these odd vertebrates (notably, hagfishes) are strongly degenerate and have lost many characters over time."
  8. ^ Stock, David; Whitt GS (7). "Evidence from 18S ribosomal RNA sequences that lampreys and hagfishes form a natural group". Science. 257 (5071). PMID 1496398. doi:10.1126/science.1496398. Retrieved 22 November 2011.  Cite uses deprecated parameter |coauthors= (help); Unknown parameter |month= ignored (help); Check date values in: |date=, |year= / |date= mismatch (help)
  9. ^ Ota, Kinya; et al (28). "Identification of vertebra-like elements and their possible differentiation from sclerotomes in the hagfish". nature communications. 2 (6). doi:10.1038/ncomms1355.  Cite uses deprecated parameter |coauthors= (help); Unknown parameter |month= ignored (help); Check date values in: |year=, |date=, |year= / |date= mismatch (help)
  10. ^ Romer, A.S. & Parsons, T.S. (1985): The Vertebrate Body. (6th ed.) Saunders, Philadelphia.
  11. ^ http://www.aquaticcommunity.com/mix/hagfish.php
  12. ^ Baker, Clare V.H. (2008). "The evolution and elaboration of vertebrate neural crest cells". Current Opinion in Genetics & Development. 18 (6): 536–543. PMID 19121930. doi:10.1016/j.gde.2008.11.006. 
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