Famennian

Famennian
	372.2 ± 1.6 – 358.9 ± 0.4 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Chronology
	Events of the Devonian Period -420 — – -415 — – -410 — – -405 — – -400 — – -395 — – -390 — – -385 — – -380 — – -375 — – -370 — – -365 — – -360 — – P; a; l; e; o; z; o; i; c S D; e; v; o; n; i; a; n C M Přídolí E; a; r; l; y M; i; d L; a; t; e E; M Lochkovian Pragian Emsian Eifelian Givetian Frasnian Famennian Tournai. ← Rhynie chert[1] ← Hangenberg event ← Kellwasser event(s)[2] ← Widespread[3]; shrubs & trees ← S. America; glaciation begins ← Hunsrück fauna; Key events of the Devonian Period. ; Axis scale: millions of years ago.
Etymology
Name formality	Formal
Usage information
Celestial body	Earth
Regional usage	Global (ICS)
Time scale(s) used	ICS Time Scale
Definition
Chronological unit	Age
Stratigraphic unit	Stage
Time span formality	Formal
Lower boundary definition	FAD of the Conodont Palmatolepis triangularis; LAD of the Conodonts Ancyrodella and Ozarkodina and the Goniatites Gephuroceratidae and Beloceratidae
Lower boundary GSSP	Coumiac quarry, Montagne Noire, France; 43.4613°N 3.0403°E
GSSP ratified	1993[4]
Upper boundary definition	FAD of the Conodont Siphonodella sulcata (discovered to have biostratigraphic issues as of 2006).[5]
Upper boundary GSSP	La Serre, Montagne Noire, France; 43.5555°N 3.3573°E
GSSP ratified	1990[6]

The Famennian is the latter of two faunal stages in the Late Devonian epoch. It lasted from 372.2 million years ago to 358.9 million years ago. It was preceded by the Frasnian stage and followed by the Tournaisian stage.

It was during this age that tetrapods first appeared. In the seas, a novel major group of ammonoid cephalopods called clymeniids appeared, underwent tremendous diversification and spread worldwide, then just as suddenly went extinct.

The beginning of the Famennian is marked by a major extinction event, the Kellwasser Event, and the end with a smaller but still quite severe extinction event, the Hangenberg Event.

North American subdivisions of the Famennian include the Chautauquan, Canadaway, Conneaut, Conneautan, Conewango and Conewangan.

Name and definition

The Famennian stage was proposed in 1855 by Belgian geologist André Hubert Dumont and was accepted for the upper stage of the Upper Devonian by the Subcommission on Devonian Stratigraphy in 1981.[7] It is named after Famenne, a natural region in southern Belgium.

Famennian life

Placoderms

Placoderms of the Fammenian
Taxa	Presence	Location	Description	Images
Dinichthys		Ohio Shale, Ohio & Louisiana Limestone, Missouri	A giant predatory arthrodire placoderm. It was previously used as a wastebasket taxon for many unrelated large arthrodires, including species now assigned to Eastmanosteus, Titanichthys and the similar but not closely related Dunkleosteus.	Dinichthys herzi Diplognathus Dunkleosteus terrelli Dunkleosteus marsaisi, this species might belong to Eastmanosteus instead. Gorgonichthys clarki Groenlandaspis pennsylvanica Gymnotrachelus hydei Selenosteus brevis Stenosteus angustopectis Titanichthys agassizi
Diplognathus	Upper Famennian	Cleveland Shale, Ohio	An arthrodire placoderm of the superfamily Dinichthyloidea.
Dunkleosteus		Lost Burro Formation & Ohio Shale, North America Belgium[8] Morocco[9]	A dunkleosteid arthrodire with many species, some of them being among the largest placoderms to have existed.
Eastmanosteus licharevi		Lipetsk, Russia	A poorly known species of dunkleosteid arthrodire in the genus Eastmanosteus.
Gorgonichthys	Upper Famennian	Cleveland Shale, Ohio	A giant yet relatively lightly build arthrodire placoderm of the superfamily Dinichthyloidea. Gorgonichthys clarki is estimated to have reached 6 meters in length.
Groenlandaspis		Cosmopolitan, except eastern Asia	An arthrodire placoderm
Gymnotrachelus		Cleveland Shale, Ohio	A superficially barracuda-like selenosteid arthrodire placoderm.
Heintzichthys		Europe North America	An arthrodire placoderm of the superfamily Dinichthyloidea.
Selenosteus		Cleveland Shale, Ohio	A selenosteid arthrodire placoderm.
Stenosteus		Cleveland Shale, Ohio	A selenosteid arthrodire placoderm.
Titanichthys	Late Devonian	Morocco; North America	A giant, possibly filter-feeding, arthrodire placoderm of the superfamily Dinichthyloidea.

Cartilaginous fish

Cartilaginous fish of the Fammenian
Taxa	Presence	Location	Description	Images
Stethacanthus		Shar'yu River, Chernyshev Ridge, Russia Cleveland Shale, Ohio

Ray-finned fish

Actinopterygii of the Fammenian
Taxa	Presence	Location	Description	Images
Tegeolepis	Late Devonian	Ohio, United States

Coelacanths

Actinistia of the Fammenian
Taxa	Presence	Location	Description	Images
Serenichthys		Witpoort Formation, Eastern Cape, South Africa	A basal actinistian of the family Diplocercidae

Tetrapodomorphs (non-ichthyostegalian)

Tetrapodomorph fish of the Fammenian
Taxa	Presence	Location	Description	Images
Cabonnichthys		Canowindra, Australia	A genus of tristichopterid fish	Gooloogongia Hyneria
Gooloogongia		Canowindra, Australia	A genus of rhizodontid fish that reached about 90 cm in length.
Hyneria	Upper Famennian	Catskill Formation & Red Hill Shale, Pennsylvania	Hyneria was a large tristichopterid fish, estimated at 2.5–3.7 m in total length
Mandageria		Mandagery Sandstone, Canowindra, Australia	A genus of tristichopterid fish

Ichthyostegalia

Ichthyostegalians of the Fammenian
Taxa	Presence	Location	Description	Images
Acanthostega		Britta Dal Formation, East Greenland		Acanthostega Hynerpeton Ichthyostega Tulerpeton Ventastega
Densignathus	Upper Famennian	Catskill Formation & Red Hill Shale, Pennsylvania
Hynerpeton	Upper Famennian	Catskill Formation & Red Hill Shale, Pennsylvania
Ichthyostega		Evieux Formation, Belgium Britta Dal Formation, East Greenland
Jakubsonia	Lower Famennian	Russia
Metaxygnathus		Cloghnan Shale, New South Wales, Australia	Previously thought to be a lobe-finned fish.
Sinostega		Zhongning Formation, Ningxia Hui Autonomous Region, China	The first Devonian tetrapod to be found in Asia
Tulerpeton		Andreyevka, Tula Oblast, Russia
Ventastega		Ketleri Formation, Courland Peninsula, Latvia
Ymeria		Celsius Bjerg Group, Greenland	A smaller close relative of Ichtyostega.

References

Parry, S. F.; Noble, S. R.; Crowley, Q. G.; Wellman, C. H. (2011). "A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications". Journal of the Geological Society. London: Geological Society. 168 (4): 863–872. doi:10.1144/0016-76492010-043.
Kaufmann, B.; Trapp, E.; Mezger, K. (2004). "The numerical age of the Upper Frasnian (Upper Devonian) Kellwasser horizons: A new U-Pb zircon date from Steinbruch Schmidt(Kellerwald, Germany)". The Journal of Geology. 112 (4): 495–501. Bibcode:2004JG....112..495K. doi:10.1086/421077.
Algeo, T. J. (1998). "Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants, weathering processes, and marine anoxic events". Philosophical Transactions of the Royal Society B: Biological Sciences. 353 (1365): 113–130. doi:10.1098/rstb.1998.0195.
Klapper, G.; Feist, R.; Becker, R.; House, M. (December 1993). "Definition of the Frasnian/Famennian Stage boundary". 16 (4): 433–441. Retrieved 19 December 2020. Cite journal requires |journal= (help)
Kaiser, Sandra (1 April 2009). "The Devonian/Carboniferous boundary stratotype section (La Serre, France) revisited". Newsletters on Stratigraphy. 43: 195–205. doi:10.1127/0078-0421/2009/0043-0195. Retrieved 7 December 2020.
Paproth, Eva; Feist, Raimund; Flajs, Gerd (December 1991). "Decision on the Devonian-Carboniferous boundary stratotype" (PDF). Episodes. 14: 331–336.
Thorez, Jacques; Dreesen, Roland; Streel, Maurice (2006). "Frasnian". Geologica Belgica. 9 (1–2): 27–45. Retrieved 16 March 2013.
Denison, Robert (1978). "Placodermi". Handbook of Paleoichthyology. 2. Stuttgart New York: Gustav Fischer Verlag. p. 128. ISBN 978-0-89574-027-4.
Murray, A.M. (2000). "The Palaeozoic, Mesozoic and Early Cenozoic fishes of Africa". Fish and Fisheries. 1 (2): 111–145. doi:10.1046/j.1467-2979.2000.00015.x.