Conodonts (Greek kōnos, "cone", + odont, "tooth") are extinct agnathan chordates resembling eels, classified in the class Conodonta. For many years, they were known only from tooth-like microfossils found in isolation and now called conodont elements. Knowledge about soft tissues remains limited. The animals are also called Conodontophora (conodont bearers) to avoid ambiguity.
Temporal range: Cambrian to Late Triassic (Rhaetian)
|Reconstruction of a conodont|
|Two conodont "teeth" and a reconstruction of a conodont|
Conodonts are considered index fossils, fossils used to define and identify geological periods.
The conodonts first appeared during the Cambrian Stage 2 (also referred as Tommotian). The still unnamed Cambrian Stage 10 can be defined as the first appearance of Eoconodontus notchpeakensis. The upper boundary is defined as the appearance of Iapetognathus fluctivagus which marks the beginning of the Tremadocian and is radiometrically dated as 485.4 ± 1.9 million years ago.
The Cambrian–Ordovician extinction event occurred approximately 488 million years ago. This early Paleozoic extinction event extirpated many conodonts.
The Lau event, about 420 million years ago, a relatively minor mass extinction during the Silurian period, had a major impact on conodont populations.
The entire class is postulated to have been wiped out in the Triassic–Jurassic extinction event, which occurred roughly 200 million years ago. Near the end of the Triassic deadly marine biocalcification began to occur, along with oceanic acidification, sea-level fluctuations and the Central Atlantic Magmatic Province (CAMP) releasing carbon dioxide, sulfur dioxide and aerosols. These environmental catastrophes caused the extinction of the conodonts, along with 34% of other marine genera.
The 11 known fossil imprints of conodont animals record an eel-like creature with 15 or, more rarely, 19 elements that form a bilaterally symmetrical array in the head.
The organisms range from a centimeter or so[verification needed] to 40 cm (Promissum) in length. It is now widely agreed that conodonts had large eyes, fins with fin rays, chevron-shaped muscles and a notochord.
Conodont teeth are the earliest found in the fossil record. The evolution of mineralized tissues has been puzzling for more than a century. It has been hypothesized that the first mechanism of mammalian tissue mineralization began either in the oral skeleton of conodont or the dermal skeleton of early agnathans.
The element array constituted a feeding apparatus that is radically different from the jaws of modern animals. They are now termed "conodont elements" to avoid confusion. The three forms of teeth, i.e., coniform cones, ramiform bars, and pectiniform platforms, probably performed different functions.
For many years, conodonts were known only from enigmatic tooth-like microfossils (200 micrometers to 5 millimeters in length), which occur commonly, but not always in isolation, and were not associated with any other fossil. Until the early 1980s, conodont teeth had not been found in association with fossils of the host organism, in a konservat lagerstätte. This is because most of the conodont animal was soft-bodied, thus everything but the teeth was unsuited for preservation under normal circumstances.
They are widely used in biostratigraphy. Conodont elements are also used as paleothermometers, a proxy for thermal alteration in the host rock, because under higher temperatures, the phosphate undergoes predictable and permanent color changes, measured with the conodont alteration index. This has made them useful for petroleum exploration where they are known, in rocks dating from the Cambrian to the Late Triassic.
The conodont apparatus may comprise a number of discrete elements, including the spathognathiform, ozarkodiniform, trichonodelliform, neoprioniodiform, and other forms.
Elements of ozarkodinidsEdit
The feeding apparatus of ozarkodinids is composed at the front of an axial Sa element, flanked by two groups of four close-set elongate Sb and Sc elements which were inclined obliquely inwards and forwards. Above these elements lay a pair of arched and inward pointing (makellate) M elements. Behind the S-M array lay transversely oriented and bilaterally opposed (pectiniform, i.e. comb-shaped) Pb and Pa elements.
The "teeth" of some conodonts have been interpreted as filter-feeding apparatuses, filtering plankton from the water and passing it down the throat. Others have been interpreted as a "grasping and crushing array". The lateral position of the eyes makes it unlikely that conodonts were active predators. The preserved musculature suggests that some conodonts (Promissum at least) were efficient cruisers, but incapable of bursts of speed.
Classification and phylogenyEdit
Milsom and Rigby envision them as vertebrates similar in appearance to modern hagfish and lampreys, and phylogenetic analysis suggests they are more derived than either of these groups. However, this analysis comes with one caveat: early forms of conodonts, the protoconodonts, appear to form a distinct clade from the later paraconodonts and euconodonts. Protoconodonts likely represent a stem group to the phylum that includes chaetognath worms; this conclusion suggests that chaetognaths are not close relatives of true conodonts. Moreover, some analyses do not regard conodonts as either vertebrates or craniates, because they lack the main characteristics of these groups.
Conodonta Pander 1856 non Eichenberg 1930 sensu Sweet & Donoghue 2001 [Conodontia; Conodontophorida Eichenberg 1930; Conodontochordata]
- Paraconodontida Müller 1962
- Conodontophora Eichenberg 1930
- Caviodonti Sweet 1988
- Conodonti Pander 1856 non Branson 1938
- ?Strachanognathidae Bergström 1981
- Oneotodontidae Miller 1981 [Teridontidae Miller 1981]
- Protopanderodontida Sweet 1988 [Distacodontida; Panderodontida]
- Pronodontidae Lindström 1970
- Cornuodontidae Faohraeus1966
- Protopanderodontidae Lindström 1970 [Juanognathidae Bergström 1981; Scolopodontidae Bergström 1981; Ulrichodinidae Bergström 1981]
- Clavohamulidae Lindström 1970
- Drepanoistodontidae Faohraeus 1978
- Acanthodontidae Lindström 1970
- Panderodontidae Lindström 1970
- Prioniodontida Dzik 1976 (paraphyletic)
- Oepikodontidae Bergström 1981
- Paracordylodontidae Bergström 1981
- Rossodus Repetski & Ethington 1983
- Multioistodontidae Harris 1964
- Oistodontidae Lindström 1970 [Tripodontidae]
- Periodontidae Lindström 1970
- Rhipidognathidae Lindström 1970 sensu Sweet 1988
- Prioniodontidae Bassler 1925
- Balognathidae Hass 1959
- Polyplacognathidae Bergström 1981
- Distomodontidae Klapper 1981
- Icriodellidae Sweet 1988
- Icriodontidae Müller & Müller 1957
- Cyrtoniodontidae Hass 1959 [Phragmodontidae Bergström 1981]
- Plectodinidae Sweet 1988
- Prioniodinida Sweet 1988
- Ozarkodinida Dzik 1976 [Polygnathida]
- Cryptotaxidae Klapper & Philip 1971
- Spathognathodontidae Hass 1959
- Pterospathodontidae Cooper 1977
- Kockelellidae Klapper 1981
- Polygnathidae Bassler 1925
- Palmatolepidae Sweet 1988
- Anchignathodontidae Clark 1972
- Elictognathidae Austin & Rhodes 1981
- Gnathodontidae Sweet 1988
- Idiognathodontidae Harris & Hollingsworth 1933
- Mestognathidae Austin & Rhodes 1981
- Cavusgnathidae Austin & Rhodes 1981
- Sweetognathidae Ritter 1986
- Here, the hagfish are treated as a separate clade, as in Sweet and Donoghue's 2001 tree produced without cladistic analysis. However, it has been recognised by some  that the hagfish and lampreys may be closer to one another in their own clade, the Cyclostomata.
- The clade Proconodontida is also known as Cavidonti.
- Euconodonta is referred to as "Conodonti" by Sweet and Donoghue, although this is not widely used[original research?].
- Conodonten aus dem Culm des Harzes. W Eichenberg, Paläontologische Zeitschrift, 1930, page 177-182, doi:10.1007/BF03044446
- J. J. Sepkoski. 2002. A compendium of fossil marine animal genera. Bulletins of American Paleontology 363:1-560
- Early Cambrian (Tommotian) conodonts and other shelly microfauna from the Upper Krol of Mussoorie Syncline. RJ Azmi, VP Pancholi, Lesser Himalaya, with remarks on the Precambrian, 1983
- DeSantis, M.K.; Brett C.E. (2011). "Late Eifelian (Middle Devonian) biocrises: Timing and signature of the pre-Kačák Bakoven and Stony Hollow Events in eastern North America". Palaeogeography, Palaeoclimatology, Palaeoecology. Elsevier. 304 (1–2): 113–135. doi:10.1016/j.palaeo.2010.10.013. Retrieved 4 March 2012.
- The extinction of conodonts —in terms of discrete elements— at the Triassic-Jurassic boundary
- Graham Ryder; David E. Fastovsky; Stefan Gartner (1996). The Cretaceous-Tertiary Event and Other Catastrophes in Earth History. Geological Society of America. p. 19. ISBN 9780813723075.
- Gabbott, S.E.; R. J. Aldridge; J. N. Theron (1995). "A giant conodont with preserved muscle tissue from the Upper Ordovician of South Africa". Nature. 374 (6525): 800–803. Bibcode:1995Natur.374..800G. doi:10.1038/374800a0.
- Shubin, Neil (2009). Your Inner Fish: A Journey into the 3.5 Billion Year History of the Human Body (reprint ed.). New York: Pantheon Books. pp. 85–86. ISBN 9780307277459.
- MIRACLE. "Conodonts". Retrieved 26 August 2014.
- Briggs, D. E. G.; Clarkson, E. N. K.; Aldridge, R. J. (1983). "The conodont animal". Lethaia. 16 (1): 1–14. doi:10.1111/j.1502-3931.1983.tb01993.x.
- Chemical systematics of conodont apatite determined by laser ablation ICPMS. Julie A. Trotter and Stephen M. Eggins, Chemical Geology, Volume 233, Issues 3–4, 15 October 2006, Pages 196–216, doi:10.1016/j.chemgeo.2006.03.004
- A Buffered Formic Acid Technique for Conodont Extraction. Lennart Jeppsson and Rikard Anehus, Journal of Paleontology, Vol. 69, No. 4 (Jul., 1995), pages 790-794 (Stable URL)
- Extraction Techniques for Phosphatic Fossils. Owen R. Green in A Manual of Practical Laboratory and Field Techniques in Palaeobiology. pages 318-330, doi:10.1007/978-94-017-0581-3_27
- Effects of extraction protocols on the oxygen isotope composition of conodont elements. Page C. Quinton, Stephen A. Leslie, Achim D. Herrmann and Kenneth G. MacLeod, Chemical Geology, Volume 431, 1 August 2016, Pages 36–43, doi:10.1016/j.chemgeo.2016.03.023
- Bergström, S. M.; Carnes, J. B.; Ethington, R. L.; Votaw, R. B.; Wigley, P. B. (1974). "Appalachignathus, a New Multielement Conodont Genus from the Middle Ordovician of North America". Journal of Paleontology. Paleontological Society. 48 (2): 227–235. doi:10.2307/1303249 (inactive 2017-01-16). JSTOR 1303249.
- Conodonten-Funde in ursprünglichem Zusammenhang. Hermann Schmidt, Palaeontologische Zeitschrift, 1934, volume 16, Issue 1-2 , pages 76-85, doi:10.1007/BF03041668
- The Zoological Relationships of the Conodonts. Harold W. Scott, Journal of Paleontology, Vol. 8, No. 4 (Dec., 1934), pages 448-455 (Stable URL)
- Conodont Assemblages from the Heath Formation, Montana. Harold W. Scott, Journal of Paleontology, Vol. 16, No. 3 (May, 1942), pages 293-300 (Stable URL)
- Late Mississippian conodonts from the Bird Spring Formation in Nevada. David L. Dunn, Journal of Paleontology, November 1965, volume 39, issue 6 (abstract)
- A Questionable Natural Conodont Assemblage from Middle Ordovician Limestone, Ottawa, Canada. Christopher R. Barnes, Journal of Paleontology, Vol. 41, No. 6 (Nov., 1967), pages 1557-1560 (Stable URL)
- Purnell, M. A.; Donoghue, P. C. (1997). "Architecture and functional morphology of the skeletal apparatus of ozarkodinid conodonts". Philosophical Transactions of the Royal Society B: Biological Sciences. 352 (1361): 1545–1564. doi:10.1098/rstb.1997.0141. PMC .
- Briggs, D. (May 1992). "Conodonts: a major extinct group added to the vertebrates". Science. 256 (5061): 1285–1286. Bibcode:1992Sci...256.1285B. doi:10.1126/science.1598571. PMID 1598571.
- Milsom, Clare; Rigby, Sue (2004). "Vertebrates". Fossils at a Glance. Victoria, Australia: Blackwell Publishing. p. 88. ISBN 0-632-06047-6.
- Donoghue, P.C.J.; Forey, P.L.; Aldridge, R.J. (2000). "Conodont affinity and chordate phylogeny". Biological Reviews. 75 (2): 191–251. doi:10.1111/j.1469-185X.1999.tb00045.x. PMID 10881388. Retrieved 2008-04-07.
- Szaniawski, H. (2002). "New evidence for the protoconodont origin of chaetognaths" (PDF). Acta Palaeontologica Polonica. 47 (3): 405.
- Turner, S., Burrow, C.J., Schultze, H.P., Blieck, A., Reif, W.E., Rexroad, C.B., Bultynck, P., Nowlan, G.S.; Burrow; Schultze; Blieck; Reif; Rexroad; Bultynck; Nowlan (2010). "False teeth: conodont-vertebrate phylogenetic relationships revisited" (PDF). Geodiversitas. 32 (4): 545–594. doi:10.5252/g2010n4a1.
- Sweet, W. C.; Donoghue, P. C. J. (2001). "CONODONTS: PAST, PRESENT, FUTURE". Journal of Paleontology. 75 (6): 1174–1184. doi:10.1666/0022-3360(2001)075<1174:CPPF>2.0.CO;2.
- Bourlat, S. J; T. Juliusdottir, C. J Lowe, R. Freeman, J. Aronowicz, M. Kirschner, E. S Lander, M. Thorndyke, H. Nakano, A. B Kohn, others (2 November 2006). "Deuterostome phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida". Nature. 444 (7115): 85–88. Bibcode:2006Natur.444...85B. doi:10.1038/nature05241. ISSN 0028-0836. PMID 17051155.
- Sweet, W. C. (1988). "The Conodonta: morphology, taxonomy, paleoecology and evolutionary history of a long-extinct animal phylum". Oxford monographs on geology and geophysics (10): 1–211.
- Mikko's Phylogeny Archive  Haaramo, Mikko (2007). "Conodonta - conodonts". Retrieved 2015-12-30.
- Aldridge, R. J.; Briggs, D. E. G.; Smith, M. P.; Clarkson, E. N. K.; Clark, N. D. L. (1993). "The anatomy of conodonts". Philosophical Transactions of the Royal Society of London, Series B. 340 (1294): 405–421. doi:10.1098/rstb.1993.0082.
- Aldridge, R. J.; Purnell, M. A. (1996). "The conodont controversies". Trends in Ecology and Evolution. 11 (11): 463–468. doi:10.1016/0169-5347(96)10048-3. PMID 21237922.
- Donoghue, P. C. J.; Forey, P. L.; Aldridge, R. J. (2000). "Conodont affinity and chordate phylogeny". Biological Reviews. 75 (2): 191–251. doi:10.1111/j.1469-185X.1999.tb00045.x. PMID 10881388.
- Gould, Stephen Jay (1985). "Reducing Riddles". In The Flamingo's Smile, 245-260. New York, W.W. Norton and Company. ISBN 0-393-30375-6.
- Janvier, P (1997). "Euconodonta". The tree of life web project. Retrieved 2007-09-05.
- Knell, Simon J. The Great Fossil Enigma: The Search for the Conodont Animal (Indiana University Press; 2012) 440 pages
- Sweet, Walter. The Conodonta.
- Sweet, W. C.; Donoghue, P. C. J. (2001). "Conodonts: past, present and future". Journal of Paleontology. 75 (6): 1174–1184. doi:10.1666/0022-3360(2001)075<1174:CPPF>2.0.CO;2. ISSN 0022-3360.
- A suprageneric taxonomy of the conodonts. Maurits Lindström, Lethaia, Volume 3, Issue 4, pages 427–445, October 1970, doi:10.1111/j.1502-3931.1970.tb00834.x
- Mark Purnell. "An oblique anterior view of a model of the apparatus of the Pennsylvanian conodont Idiognathodus".
- "'The Jaws That Catch': an Introduction to the Conodonta". Palæos. Retrieved 2013-07-01.
- Jim Davison (2002-10-15). "Ordovician conodonts". Retrieved 2009-07-07.