ülemklass

Agnatha Cope, 1889

Lõuatud e agnaadid

Taksoni iseloomustus
main agnathan groups

Telodontide (Thelodonti) välisskelett koosnes soomuselaadsetest moodustistest, nn nahahammastest, suu oli pea eesääres ning silmad külgedel.

Erikilbiliste (Heterostraci) pead ja keha esimest osa kattis plaatidest rüü, keha tagumist osa ja saba katsid soomused.

Luukilbiliste (Osteostraci) pead kattis ühtlane kilp, silmad olid lähestikku seljapoolel. Silmade taga ja pea äärel olid tundeelundid, keha tagaosa oli kaetud soomustega.

Kilbitute (Anaspida) pead katsid väikesed plaadid, keha kõrged soomused. Suu oli pea eesääres kõhupoolel, silmad külgedel.

Lõuatud ehk agnaadid on primitiivsed kalalaadsed selgroogsed, kelle põhitunnuseks on lõualuude puudumine ja kõhrest sisetoes. Algelisematel lõuatutel olid rinnauimed ja sabauim. Väljasurnud lõuatud jagunevad: erikilbilisteks (heterostraagid), telodondid, luukilbilised (osteostraagid), kilbitud (anaspiidid) ja galeaspiidid. Esimesed lõuatud ilmusid Vara-Ordoviitsiumis, nende õitseaeg oli Siluri ja Devoni ajastul. Tänapäeva on jõudnud vaid vähesed lõuatud — sõõrsuused (pihkased ja silmud).

Maailma vanimate lõuatute ja ka selgroogsete hulka kuuluvad Arandaspis Austraaliast ja Sacabambaspis Boliiviast. Lõuatuid olid enamasti madalmerede asukad, nad toitusid vees olevast planktonist ja põhjamudast.

Siluri ajastu lõuatute suurus jäi enamasti vahemikku 7-40 cm. Mõned Devoni ajastu erikilbilised kasvasid aga enam kui 2 m pikkuseks. Eestist on leitud ka kõigi aegade suurima lõuatute perekonna Tartuosteus esindajate luuplaate.

Eesti Siluri ja Devoni settekivimitest võib leida kõikide väljasurnud lõuatute esindajaid v.a. galeaspiide. Tervete lõuatute leiud on Eestis väga haruldased, enamasti leitakse nende mikrojäänused ja üksikuid luuplaate või nende osi.

NB! Kõik lõuatute kivistised on Eestis looduskaitse all ning nende kogumiseks on vajalik Keskkonnaameti eriluba.

Phlebolepis elegans, Saaremaa Siluri merest Teledont Phlebolepis elegans Saaremaa Siluri ajastu meres. Rekonstruktsioon: Wilson & Märss, 2012.
Valik taksonit käsitlevast teaduskirjandusest
Märss, T., Wilson, M. V., Viljus, M. 2022. Endolymphatic structures in headshields of the osteostracan genus Tremataspis (Agnatha) from the Silurian of Estonia. Estonian Journal of Earth Sciences 71, 3, 135-156. DOI:10.3176/earth.2022.10
Bremer, O., Qu, Q., Sanchez, S., Märss, T., Fernandez, V., Blom, H. 2021. The emergence of a complex pore‐canal system in the dermal skeleton of Tremataspis (Osteostraci). Journal of Morphology 282, 8, 1141-1157. DOI:10.1002/jmor.21359
Bremer, O., Jarochowska, E., Märss, T. 2020. Vertebrate remains and conodonts in the upper Silurian Hamra and Sundre formations of Gotland, Sweden. GFF 142, 1, 52-80. DOI:10.1080/11035897.2019.1655790
Märss, T. 2019. Silurian cyathaspidid heterostracans of Northern Eurasia. Estonian Journal of Earth Sciences 68, 3, 113-146. DOI:10.3176/earth.2019.11
Tinn, O., Märss, T. 2018. The earliest osteostracan Kalanaspis delectabilis gen. et sp. nov. from the mid-Aeronian (mid-Llandovery, lower Silurian) of Estonia. Journal of Vertebrate Paleontology 38, 1, e1425212. DOI:10.1080/02724634.2017.1425212
Glinskiy, V. 2018. Phylogenetic relationships of psammosteid heterostracans (Pteraspidiformes), Devonian jawless vertebrates. Biological Communications 62, 4, 219-243. DOI:10.21638/11701/spbu03.2017.402
Glinskiy, V. N., Nilov, S. P. 2017. A new psammosteid (Agnatha, Heterostraci) from the Amata Regional Stage of the Main Devonian Field and morpho-histological types of discrete micromeric elements in the family Psammosteidae. Estonian Journal of Earth Sciences 66, 2, 59-76. DOI:10.3176/earth.2017.05
Glinskiy, V., Mark-Kurik, E. 2016. Revision of Psammosteus livonicus Obruchev (Agnatha, Heterostraci) from the Devonian Amata Regional Stage of the NW of the East European Platform. Estonian Journal of Earth Sciences 65, 1, 1-18. DOI:10.3176/earth.2016.02
Dec, M. 2015. A new Tolypelepidid (Agnatha, Heterostraci) from the Late Silurian of Poland. Journal of Paleontology 89, 4, 1-8. DOI:10.1017/jpa.2015.28
Tuuling, T. 2015. A specimen of Psammolepis in the Gauja Formation (Estonia). bakalaureusetöö, pp. 1-31.
Keating, J. N., Marquart, C. L., Donoghue, P. C. J. 2015. Histology of the heterostracan dermal skeleton: Insight into the origin of the vertebrate mineralised skeleton. Journal of Morphology 276, 6, 657-680. DOI:10.1002/jmor.20370
Fadel, A., Žigaite, Ž., Blom, H., Perez-Huerta, A., Jeffries, T., Märss, T., Ahlberg, P. E. 2015. Palaeoenvironmental signatures revealed from rare earth element (REE) compositions of vertebrate microremains of the Vesiku Bone Bed (Homerian, Wenlock), Saaremaa Island, Estonia. Estonian Journal of Earth Sciences 64, 1, 36-41. DOI:10.3176/earth.2015.07
Märss, T., Afanassieva, O., Blom, H. 2014. Biodiversity of the Silurian osteostracans of the East Baltic. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 105, 2, 73-148. DOI:10.1017/S1755691014000218
Afanassieva, O. B., Märss, T. 2014. New Data on the Exoskeleton of the Osteostracan Genus Aestiaspis (Agnatha) from the Silurian of Saaremaa Island (Estonia) and the Severnaya Zemlya Archipelago (Russia). Paleontologicheski Zhurnal 48, 1, 74-78. DOI:10.1134/S003103011401002X
Mark-Kurik, E., Blieck, A., Burrow, C. J., Turner, S. 2013. Early Devonian fishes from coastal De Long Strait, central Chukotka, Arctic Russia. Geodiversitas 35, 3, 545-578. DOI:10.5252/g2013n3a3
Märss, T. 2011. A unique Late Silurian Thelodus squamation from Saaremaa (Estonia) and its ontogenetic development. Estonian Journal of Earth Sciences 60, 3, 137-146. DOI:10.317/earth.2011.3.02
Märss, T., Karatajūtė-Talimaa, V. 2009. Late Silurian - Early Devonian tessellated heterostracan Oniscolepis Pander, 1856 from the East Baltic and North Timan. Estonian Journal of Earth Sciences 58, 1, 43-62. DOI:10.3176/earth.2009.1.05
Lebedev, O. A., Mark-Kurik, E., Karatajūtė-Talimaa, V. N., Lukševičs, E., Ivanov, A. 2009. Bite marks as evidence of predation in early vertebrates. Acta Zoologica 90, Suppl. 1, 344-356. DOI:10.1111/j.1463-6395.2008.00344.x
Lukševics, E., Lebedev, O., Mark-Kurik, E., Karatajūtė-Talimaa, V. 2009. The earliest evidence of host-parasite interactions in vertebrates. Acta Zoologica 90, Suppl. 1, 335-343. DOI:10.1111/j.1463-6395.2008.00362.x
Märss, T., Kleesment, A., Niit, M. 2008. Karksilepis parva gen. et sp. nov. (Chondrichthyes) from the Burtnieki Regional Stage, Middle Devonian of Estonia. Estonian Journal of Earth Sciences 57, 4, 219-230. DOI:10.3176/earth.2008.4.02
Märss, T., Turner, S., Karatajūtė-Talimaa, V. 2007. "Agnatha" II. Thelodonti. Handbook of Paleoichthyology, pp. 1-143. Verlag Dr. Friedrich Pfeil.
Märss, T. 2006. Thelodonts (Agnatha) from the basal beds of the Kuressaare Stage, Ludlow, Upper Silurian of Estonia. Proceedings of the Estonian Academy of Sciences. Geology 55, 1, 43-66. DOI:10.3176/geol.2006.1.03
Märss, T. 2006. Exoskeletal ultrasculpture of early vertebrates. Journal of Vertebrate Paleontology 26, 2, 235-252. DOI:10.1671/0272-4634(2006)26[235:EUOEV]2.0.CO;2
Märss, T. 2005. Thelodont Oeselia mosaica gen. et sp. nov. from the Wenlock and Ludlow of the East Baltic. Proceedings of the Estonian Academy of Sciences. Geology 54, 3, 181-190. DOI:10.3176/geol.2005.3.03
Elliott, D. K., Mark-Kurik, E. 2005. A review of the lateral line sensory system in psammosteid heterostracans. Revista Brasileira de Paleontologia 8, 2, 99-108. DOI:10.4072/rbp.2005.2.02
Novitskaya L. I. 2004. Subclass Heterostraci. Fossil vertebrates of Russia and adjacent countries. Agnathans and early fisches. The reference book for palaeontologists and geologists, pp. 69-207. GEOS.
Märss, T., Miller, C. G. 2004. Thelodonts and distribution of associated conodonts from the Llandovery-lowermost Lochkovian of the Welsh Borderland. Palaeontology 47, 5, 1211-1265. DOI:10.1111/j.0031-0239.2004.00409.x
Karatajūtė-Talimaa, V., Märss, T. 2004. Subclass Thelodonti. Fossil vertebrates of Russia and adjacent countries. Agnathans and early fisches. The reference book for palaeontologists and geologists, pp. 12-68. GEOS.
Afanassieva, O. B. 2004. Subclass Osteostraci. Agnathans and Early Fishes, pp. 210-267. GEOS.
Miller, C. G., Märss, T., Blom, H. 2004. New anaspid material from the Late Silurian of Britain and Estonia. Acta Universitatis Latviensis 679, 46-56. Latvijas Universitāte.
Märss, T. 2003. Paralogania from the Rootsiküla (Wenlock) and Paadla (Ludlow) stages of Estonia. Proceedings of the Estonian Academy of Sciences. Geology 52, 2, 98-112. DOI:10.3176/geol.2003.2.03
Blom, H., Märss, T., Miller, C. G. 2003. A new birkeniid anaspid from the Upper Silurian of Skåne, south Sweden. GFF 125, 2, 57-61. DOI:10.1080/11035890301252057
Märss, T., Wilson, M. V. H., Thorsteinsson, R. 2002. New thelodont (Agnatha) and possible chondrichthyan (Gnathostomata) taxa established in the Silurian and Lower Devonian of the Canadian Arctic Archipelago. Proceedings of the Estonian Academy of Sciences. Geology 51, 2, 88-120. DOI:10.3176/geol.2002.2.02
Karatajūtė-Talimaa, V., Märss, T. 2002. Upper Silurian thelodonts from Severnaya Zemlya Archipelago (Russia). Geodiversitas 24, 2, 405-443.
Märss, T., Karatajūtė-Talimaa, V. 2002. Ordovician and Lower Silurian thelodonts from Severnaya Zemlya Archipelago (Russia). Geodiversitas 24, 2, 381-404.
Karatajūtė-Talimaa, V. 2002. Lower Devonian (Lochkovian) thelodonts from the October Revolution Island (Severnaya Zemlya Archipelago). Geodiversitas 24, 4, 791-804.
Blom, H., Märss, T., Miller, C. G. 2002. Silurian and earliest Devonian birkeniid anaspids from the Northern Hemisphere. Transactions of the Royal Society of Edinburgh Earth Sciences 92, 3, 263-323. DOI:10.1017/S0263593300000250
Märss, T. 2002. Silurian and Lower Dewonian anaspids (Agnatha) from Severnaya Zemlya (Russia). Geodiversitas 24, 1, 123-137.
Upeniece, I. 2001. The unique fossil assemblage from the Lode Quarry (Upper Devonian, Latvia). Mitteilungen aus dem Museum für Naturkunde der Humboldt-Universität in Berlin 4, 101-119. DOI:10.5194/fr-4-101-2001
Soehn, K. L., Märss, T., Caldwell, M. W., Wilson, M. V. H. 2001. New and biostratigraphically useful thelodonts from the Silurian of the Mackenzie Mountains, Northwest Territories, Canada. Journal of Vertebrate Paleontology 21, 4, 651-659. DOI:10.1671/0272-4634(2001)021[0651:NABUTF]2.0.CO;2
Mark-Kurik, E. 2000. The Middle Devonian fishes of the Baltic States (Estonia, Latvia) and Belarus. Courier Forschungsinstitut Senckenberg 223, 309-324.
Märss, T. 1999. A new Late Silurian or Early Devonian thelodont from the Boothia Peninsula, Arcitc Canada. Palaeontology 42, 6, 1079-1099. DOI:10.1111/1475-4983.00111
Otto, M., Laurin, M. 1999. Osteostracan tesserae from the Baltic Middle Devonian: morphology and microanatomy. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie. Monatshefte: Abt. B. Geologie, Paläontologie 1999, 8, 464-476. DOI:10.1127/njgpm/1999/1999/464
Märss, T., Ritchie, A. 1998. Articulated thelodonts (Agnatha) of Scotland. Transactions of the Royal Society of Edinburgh Earth Sciences 88, 3, 143-195. DOI:10.1017/S026359330000691X
Mark-Kurik, E., Janvier, P. 1997. A new tremataspidid (Vertebrata, Osteostraci, Thyestiida) from the Devonian of northern Urals, with remarks on tannuaspidids. Neues Jahrbuch für Geologie und Paläontologie. Abhandlungen 206, 3, 405-421. DOI:10.1127/njgpa/206/1997/405
Märss, T. 1997. Vertebrates of the Pridoli and Silurian-Devonian boundary beds in Europe. Modern Geology 21, 1, 17-41.
Afanassieva, O. B., Märss, T. 1997. Exoskeleton Structure and the Distribution of Aestiaspis viitaensis (Agnatha) from the Silurian of Estonia. Paleontological Journal 31, 6, 641-647.
Märss, T. 1996. Loganellia (Thelodonti, Agnatha) from the Jaagarahu Stage, Wenlock, Estonia. Proceedings of the Estonian Academy of Sciences. Geology 45, 4, 189-202. DOI:10.3176/geol.1996.4.02
Mark-Kurik, E., Janvier, P. 1995. Early Devonian osteostracans from Severnaya Zemlya, Russia. Journal of Vertebrate Paleontology 15, 3, 449-462. DOI:10.1080/02724634.1995.10011241
Janvier, P., Lelievre, H. 1994. A new tremataspid osteostracan, Aestiaspis viitaensis n. g., n. sp., from the Silurian of Saaremaa, Estonia. Proceedings of the Estonian Academy of Sciences. Geology 43, 3, 122-128. DOI:10.3176/geol.1994.3.02
Mark-Kurik, E. 1993. Notes on the squamation in psammosteids. Modern Geology 18, 1, 107-114. ..
Märss, T. 1992. The structure of growth layers of Silurian fish scales as a potential evidence of the environmental changes. Fossil Fishes as Living Animals, pp. 41-48. Academy of Sciences of Estonia.
Fredholm, D. 1990. Agnathan vertebrates in the Lower Silurian of Gotland, Sweden. Geologiska Föreningens i Stockholm Förhandlingar 112, 1, 61-84. DOI:10.1080/11035899009453162
Afanassieva, O. B. 1990. Novyj Brenneviaspis (Agnatha) iz nižnedevonskih otloženij Podolii. Paleontologicheski Zhurnal 4, 128-131.
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Märss, T. 1989. Vertebrates. A Global Standard for The Silurian System. National Museum of Wales, Geological Series 9, pp. 284-289. National Museum of Wales.
Märss, T. 1986. Silurian Vertebrates of Estonia and West Latvia. Fossilia Baltica 1. Fossilia Baltica 1, 1-104. Valgus.
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Klassifikatsioon
organismirühmBiota
riikAnimalia
hõimkondChordata
alamhõimkondVertebrata
organismirühmGnathostomata
Pisces
ülemklassAgnatha
klassincertae sedis
Cephalaspidomorphi
Pteraspidomorphi