2021 in paleontology

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Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 2021.

Flora

Plants

Main article: 2021 in paleobotany

Fungi

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bleximothyrium	Gen. et sp. nov	Valid	Le Renard et al.	Early Cretaceous (Aptian)	Potomac Group	United States (Virginia)	A dothideomycete fly‐speck fungus. Type species B. ostiolatum.
Columnomyces electri	Sp. nov	Valid	Haelewaters & Perreau in Perreau, Haelewaters & Tafforeau	Miocene	Dominican amber	Dominican Republic	A laboulbeniale fungus, parasitic on the beetle Proptomaphaginus alleni.
Rhizophydites	Gen. et sp. nov	Valid	Krings, Serbet & Harper	Early Devonian	Rhynie chert	United Kingdom	A Chytridiomycotan fungus. Type species R. matryoshkae.

Research

• Exceptionally preserved specimens of Tawuia, providing new information on the anatomy of this organism, are described from the Tonian Liulaobei and Shiwangzhuang formations (China) by Tang et al. (2021), who interpret Tawuia as a coenocytic eukaryote, possibly a macroalga.
• Microfossils which may represent early terrestrial fungi are described from the Ediacaran Doushantuo Formation (China) by Gan et al. (2021).
• A Rhynie chert fossil Mycokidstonia sphaerialoides, originally interpreted as an ascomycete, is reclassified as a member of Glomeromycota belonging to the family Ambisporaceae by Walker et al. (2021).
• Carboniferous organism Oochytrium lepidodendri, originally classified as a fungus, is reinterpreted as an oomycete by Strullu-Derrien et al. (2021).
• Probable fossils of multicellular eukaryotic macroalgae (possibly with a green algal affinity) are described from the Tonian Dolores Creek Formation in the Wernecke Mountains (Canada) by Maloney et al. (2021), who interpret these fossils as likely to be some of the few green algae and some of the largest macroscopic eukaryotes yet recognized in the early Neoproterozoic, indicating that eukaryotic algae colonized marine environments by the early Neoproterozoic.

Cnidarians

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Columnocoenia falkenbergensis	Sp. nov	Valid	Baron-Szabo	Early Cretaceous (Aptian)	Schrattenkalk Formation	Germany Romania	A stony coral.
Confusaforma prima	Sp. nov	Valid	Löser in Löser et al.	Early Cretaceous (Valanginian)	Sierra del Pozo Formation	Spain	A coral belonging to the family Solenocoeniidae.
Decimoconularia	Gen. et sp. nov	Valid	Guo et al.	Cambrian Stage 2	Yanjiahe Formation	China	A hexangulaconulariid. Genus includes new species D. isofacialis.
Eopreverastrea	Gen. et sp. nov	Valid	Löser in Löser et al.	Early Cretaceous (Valanginian)	Sierra del Pozo Formation	Spain	A coral belonging to the family Aulastraeoporidae. The type species is E. llanoensis.
Floriastrea iberica	Sp. nov	Valid	Löser in Löser et al.	Early Cretaceous (Valanginian)	Sierra del Pozo Formation	Spain	A coral belonging to the family Actinastreidae.
Monopachyphyllia	Gen. et sp. nov	Valid	Kołodziej & Marian	Early Cretaceous (Aptian)		Romania	A colonial coral belonging to the group Pachythecaliina, possibly belonging to the superfamily Heterocoenioidea and the family Carolastraeidae. Genus includes new species M. roniewiczae.
Palaeodiphasia	Gen et comb. nov	Valid	Song et al.	Late Cambrian	Fengshan Formation	China	A member of Leptothecata belonging to the group Macrocolonia; a new genus for "Siberiograptus" simplex Lin (1985).
Siderohelia	Gen. et sp. nov	Valid	Löser in Löser et al.	Cretaceous (Hauterivian to Santonian)		Spain	A stony coral belonging to the family Rhizangiidae. The type species is S. aquilai.

Research

• A study on the morphology, embryonic development and phylogenetic relationships of Quadrapyrgites is published by Zhao et al. (2021), who interpret this taxon and its probable relative Olivooides as more likely to be diploblastic cnidarians than triploblastic cycloneuralians.
• An exceptionally preserved conulariid specimen, keeping its aperture semi-closed and making it possible to see most of the internal part of the closure with rib continuation inwards, is described from the Ordovician of southeastern Brandenburg (Germany) by Sendino & Bochmann (2021).

Arthropods

Main articles: 2021 in arthropod paleontology and 2021 in insect paleontology

Brachiopods

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bellistrophia askarensis	Sp. nov	Valid	Popov & Nikitina in Popov et al.	Cambrian (Wuliuan)	Athei Formation	Kazakhstan	A kutorginide brachiopod.
Carinagypa robecki	Sp. nov	Valid	Blodgett et al.	Devonian (Emsian)		United States (Alaska)	A member of Pentamerida belonging to the family Gypidulidae.
Crinisarina pseudoglobularis	Sp. nov	Valid	Serobyan et al.	Devonian (Famennian)		Armenia	An athyride brachiopod.
Eoconulus tucunucoensis	Sp. nov	Valid	Lavié, Mestre & Carrera	Ordovician	San Juan Formation	Argentina	An acrotretid brachiopod.
Kermanirhyncha	Gen. et sp. nov	Valid	Popov et al.	Silurian (Aeronian)	Shabdjereh Formation	Iran	A rhynchonellide brachiopod. Genus includes new species K. granulata.
Levanispirifer	Gen. et sp. nov	Valid	Popov et al.	Silurian (Aeronian)	Shabdjereh Formation	Iran	A spiriferide brachiopod. Genus includes new species L. alatus.
Luthieria	Gen. et sp. nov	Valid	Lavié, Mestre & Carrera	Ordovician	San Juan Formation	Argentina	An obolid brachiopod. Genus includes new species L. diminuta.
Mictospirifer obtusus	Sp. nov	Valid	Popov et al.	Silurian (Aeronian)	Shabdjereh Formation	Iran	A spiriferide brachiopod.
Psiloria karasuensis	Sp. nov	Valid	Popov & Nikitina in Popov et al.	Cambrian (Wuliuan)	Athei Formation	Kazakhstan	A protorthide brachiopod.
Schellwienella clarkei	Sp. nov	Valid	Rezende & Isaacson	Devonian	Ponta Grossa Formation	Brazil	A member of Orthotetida.
Xanastur	Nom. nov	Valid	García-Alcalde	Early Devonian		Spain	A terebratulid brachiopod; a replacement name for Xana García-Alcalde (1972).
Xinjiangiproductus? junggarensis	Sp. nov	In press	Guo, Chen & Liao	Early Carboniferous	Hongshanzui Formation	China

Molluscs

Main article: 2021 in paleomalacology

Echinoderms

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Archiacia ramitaensis	Sp. nov	Valid	Néraudeau & Mouty	Late Cretaceous (Cenomanian)		Syria	A sea urchin belonging to the family Archiaciidae.
Barbaraster	Gen. et 2 sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophiurida. The type species is B. colbachi; genus also includes B. muenzbergerae.
Calvaticrinus	Gen. et comb. nov	In press	Gale & Matrion	Early Cretaceous (Albian)		France United Kingdom	A microcrinoid belonging to the family Roveacrinidae. The type species is "Plotocrinus" monocarinatus Destombes (1984); genus also includes C. subplanatus (Destombes, 1984) and "Discocrinus integer Hess (2010).
Cantabrigiaster	Gen. et sp. nov		Hunter & Ortega-Hernández	Early Ordovician	Fezouata Formation	Morocco	A somasteroid asterozoan. The type species is C. fezouataensis.
Cherbonniericrinus requiensis	Sp. nov	Valid	Roux, Martinez & Vizcaïno	Eocene (Ypresian)		France	A crinoid belonging to the family Rhizocrinidae.
Costatocrinus fragilis	Sp. nov	In press	Gale	Late Cretaceous (Campanian)		United Kingdom	A crinoid.
Dermacantha reolidi	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the family Ophionereididae.
Douglasicrinus	Gen. et sp. nov	In press	Gale	Late Cretaceous (Campanian)		United Kingdom	A crinoid. Genus includes new species D. alumensis.
Globator aegyptiaca	Sp. nov	In press	El Qot	Early Cretaceous (Albian)		Egypt	A sea urchin.
Globulocrinus	Gen. et sp. nov	Valid	Roux, Martinez & Vizcaïno	Eocene (Ypresian)		France	A crinoid belonging to the family Rhizocrinidae. Genus includes new species G. amphoraformis.
Hessicrinus vectensis	Sp. nov	In press	Gale	Late Cretaceous (Campanian)		United Kingdom	A crinoid.
Holopus plaziati	Sp. nov	Valid	Roux, Martinez & Vizcaïno	Eocene (Ypresian)		France	A crinoid belonging to the family Holopodidae.
Hrabalicrinus	Gen. et sp. nov	Valid	Salamon et al.	Late Jurassic (Oxfordian)		Czech Republic	A comatulid crinoid. Genus includes new species H. zitti.
Ikerus	Gen. et sp. nov	In press	Jell & Sprinkle	Cambrian	Thorntonia Limestone	Australia	An edrioblastoid. Genus includes new species I. edgari
Inexpectacantha ullmanni	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Euryophiurida.
Lapidaster hougardae	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophioscolecida and the family Ophioscolecidae.
Ophiomisidium pratchettae	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophiurida and the family Astrophiuridae.
Ophiomusa perezi	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophiurida and the family Ophiomusaidae.
Ophiotardis	Gen. et sp. et comb. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Sinemurian-Toarcian)	Posidonia Shale Jurensismergel Formation? Luxembourg Sandstone Formation Middle Lias Formation	Luxembourg United Kingdom France? Germany?	A brittle star belonging to the group Ophiurida and the family Ophiopyrgidae. The type species is O. tennanti; genus also includes "Ophiura" astonensis Hess (1964).
Palaeocoma kortei	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophiurida and the family Ophiopyrgidae.
Pseudoconocrinus lavadensis	Sp. nov	Valid	Roux, Martinez & Vizcaïno	Eocene (Ypresian)		France	A crinoid belonging to the family Rhizocrinidae.
Sagittacrinus rotundacutus	Sp. nov	In press	Gale	Late Cretaceous (Campanian)		United Kingdom	A crinoid.
Sidericrinus (col.) plymouthensis	Sp. nov	Valid	Donovan & Fearnhead	Early Devonian		United Kingdom	A crinoid.
Sinaiosalenia	Gen. et sp. nov	In press	El Qot	Late Cretaceous (Cenomanian)		Egypt	A sea urchin. Genus includes new species S. rhombohedralis.
Sinosura dieschbourgae	Sp. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Toarcian)	Posidonia Shale	Luxembourg	A brittle star belonging to the group Ophioscolecida and the family Ophioleucidae.
Thanataster	Gen. et sp. et comb. nov	In press	Thuy & Numberger-Thuy	Early Jurassic (Sinemurian to Toarcian)	Posidonia Shale Luxembourg Sandstone Formation	Luxembourg	A brittle star belonging to the group Ophiurida. The type species is T. desdemonia; genus also includes "Ophiomusium" sinemurensis Kutscher & Hary (1991).
Thorntonites	Gen. et sp. nov	In press	Jell & Sprinkle	Cambrian	Thorntonia Limestone	Australia	A stalked eocrinoid. Genus includes new species T. dowlingi
Trecrinus	Gen. et sp. nov	Valid	Semenov et al.	Ordovician (Darriwilian)		Russia	A hybocrinid crinoid. Genus includes new species T. schmidti.
Zoroaster marambioensis	Sp. nov	Valid	Palópolo et al.	Eocene	La Meseta Formation	Antarctica	A starfish belonging to the family Zoroasteridae.

Research

• A study on the functional efficiency of hydrospires of blastoids, evaluating their potential significance for longer survival of blastoids than other blastozoan echinoderms, is published by Paul (2021).
• A study on extinction selectivity and changes in taxonomic, morphological and ecological diversity of diplobathrid crinoids throughout their evolutionary history is published by Cole & Hopkins (2021).

Conodonts

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Ancyrogondolella? bohorensis	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Ancyrogondolella goldingi	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Apsidognathus yanbianensis	Sp. nov	Valid	Yan & Wu	Silurian		China
Caudicriodus anitae	Sp. nov	Valid	Barrick, Sundgren & McAdams	Devonian (Lochkovian)		United States
Caudicriodus murphyi	Sp. nov	Valid	Barrick, Sundgren & McAdams	Devonian (Lochkovian)		United States
Dollymae peregrina	Sp. nov	In press	Świś	Devonian (Famennian)		Poland
Epigondolella buseri	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Epigondolella kozjanskoensis	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Epigondolella ritae	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Austria Slovenia	A member of the family Gondolellidae.
Epigondolella senovoensis	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Epigondolella slovenica	Sp. nov	Valid	Karádi et al.	Late Triassic (Norian)		Slovenia	A member of the family Gondolellidae.
Mosherella longnanensis	Sp. nov	Valid	Li & Lai in Li et al.	Late Triassic (Carnian)	Dengdengqiao Formation	China
Ozarkodina huenickeni	Sp. nov	In press	Gómez et al.	Silurian (Ludfordian) to Devonian (Lochkovian)	Los Espejos Formation	Argentina
Paragondolella ebruae	Sp. nov	Valid	Kılıç	Middle Triassic (Anisian)		Turkey
Paragondolella hirschii	Sp. nov	Valid	Kılıç & Budurov in Kılıç	Middle Triassic (Anisian)		Turkey
Paragondolella praecornuta	Sp. nov	Valid	Kılıç et al. in Kılıç	Middle Triassic (Anisian)		Turkey
Parvigondolella ciarapicae	Sp. nov	Valid	Rigo & Du in Du et al.	Late Triassic (Norian and Rhaetian)	Gabbs Formation San Hipolito Formation Scillato Formation	Hungary Italy Mexico United States (Nevada)
Praeicriodus simpsoni	Sp. nov	Valid	Barrick, Sundgren & McAdams	Silurian (Ludlow–Pridoli)		Australia
Tasmanognathus coronatus	Sp. nov	Valid	Yang et al.	Ordovician (Katian)		China

Fish

New taxa

Jawless vertebrates

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Jiangxialepis	Gen. et sp. nov	Valid	Liu et al.	Silurian (Telychian)	Fentou Formation	China	A member of Galeaspida belonging to the group Eugaleaspidiformes. The type species is J. retrospina.
Qushiaspis	Gen. et sp. nov	In press	Jiang et al.	Early Devonian	Xujiachong Formation	China	A member of Galeaspida. Genus includes new species Q. elaia.

Placoderms

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Leptodontichthys	Gen. et sp. nov		Jobbins et al.	Devonian (Givetian)	Taboumakhlouf Formation	Morocco	A member of Arthrodira belonging to the family Plourdosteidae. The type species is L. ziregensis.

Acanthodians

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Nostolepis digitus	Sp. nov	Valid	Li et al.	Devonian (Lochkovian)	Xitun Formation	China
Nostolepis qujingensis	Sp. nov	Valid	Li et al.	Devonian (Lochkovian)	Xitun Formation	China

Cartilaginous fishes

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Aquilolamna	Gen. et sp. nov		Vullo et al.	Late Cretaceous (Turonian)	Agua Nueva Formation	Mexico	A probable planktivorous shark of uncertain phylogenetic placement, possibly a member of Lamniformes. The type species is A. milarcae.
Dracopristis	Gen. et sp. nov	Valid	Hodnett et al.	Late Carboniferous (Kasimovian)	Atrasado Formation	United States (New Mexico)	A medium-sized ctenacanthiform shark known from a complete skeleton with soft tissue. The type species is D. hoffmanorum.
Durnonovariaodus	Gen. et sp. nov	Valid	Stumpf et al.	Late Jurassic (Tithonian)	Kimmeridge Clay	United Kingdom	A member of the family Hybodontidae. The type species is D. maiseyi.
Nebriimimus	Gen. et sp. nov	Valid	Collareta et al.	Pliocene (Zanclean)		Italy	A member of Rajiformes, possibly a skate. The type species is N. wardi.
Phoebodus curvatus	Sp. nov	Valid	Ivanov	Devonian (Givetian–Frasnian)		Australia Poland Russia
Pseudocorax kindlimanni	Sp. nov	In press	Jambura, Stumpf & Kriwet	Late Cretaceous (Cenomanian)	Sannine Formation	Lebanon

Ray-finned fishes

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Archaemacruroides vanknippenbergi	Sp. nov	In press	Schwarzhans & Jagt	Late Cretaceous (Maastrichtian)	Maastricht Formation	Belgium Netherlands	A member of Gadiformes of uncertain phylogenetic placement.
Austelliscus	Gen. et sp. nov		Figueroa, Weinschütz & Friedman	Middle Devonian or older	Paraná Basin	Brazil	An early ray-finned fish. Genus includes new species A. ferox.
Auxis koreanus	Sp. nov	Valid	Nam, Nazarkin & Bannikov	Middle Miocene	Duho Formation	South Korea	A species of Auxis.
Bardackichthys	Gen. et sp. nov	In press	Hacker & Shimada	Late Cretaceous (Cenomanian)	Woodbine Formation	United States (Texas)	A member of Ichthyodectiformes. Genus includes new species B. carteri.
Bobbitichthys	Gen. et comb. nov	Valid	Schwarzhans, Milàn & Carnevale	Paleocene (Selandian)	Kerteminde Marl	Denmark	A member of the family Macrouridae. The type species is "Hymenocephalus" rosenkrantzi Schwarzhans (2003).
Brauccipycnodus	Gen. et comb. nov	Valid	Taverne & Capasso	Early Cretaceous (Albian)		Italy	A member of the family Pycnodontidae. The type species is "Proscinetes" pillae Capasso (2007).
Cheirolepis jonesi	Sp. nov	In press	Newman et al.	Devonian (Givetian)	Tordalen Formation	Norway
Choichix	Gen. et sp. nov	Valid	Cantalice, Than‐Marchese & Villalobos‐Segura	Late Cretaceous (Cenomanian)		Mexico	A member of Acanthopterygii of uncertain phylogenetic placement. Genus includes new species C. alvaradoi.
Cretaserranus	Gen. et sp. nov	In press	Schwarzhans & Jagt	Late Cretaceous (Maastrichtian)	Maastricht Formation	Belgium Netherlands	A member of Perciformes, possibly belonging to the family Serranidae. Genus includes new species C. maastrichtensis.
Feroxichthys panzhouensis	Sp. nov	Valid	Ma, Xu & Geng	Middle Triassic (Anisian)	Guanling Formation	China	A member of the family Colobodontidae.
Gobiosoma? axsmithi	Sp. nov	In press	Ebersole, Cicimurri & Stringer	Oligocene (Rupelian)	Byram Formation	United States (Alabama)	A member of the family Gobiidae.
Guiclupea	Gen. et sp. nov	Valid	Chen et al.	Oligocene		China	A member of Clupeomorpha belonging to the group Ellimmichthyiformes. The type species is G. superstes.
Pteronisculus changae	Sp. nov	In press	Ren & Xu	Middle Triassic (Anisian)	Guanling Formation	China
Raususetarches	Gen. et sp. nov	Valid	Yabumoto & Nazarkin	Late Miocene	Koshikawa Formation	Japan	A member of the family Scorpaenidae. Genus includes new species R. sakurai.
Rhinocephalus cretaceus	Sp. nov	In press	Schwarzhans & Jagt	Late Cretaceous (Maastrichtian)	Maastricht Formation	Belgium	A member of the family Merlucciidae.
Saurichthys sceltrichensis	Sp. nov	Valid	Renesto, Magnani & Stockar	Middle Triassic (Ladinian)	Meride Limestone	Switzerland
Severnichthys	Gen. et sp. nov	In press	Stringer & Schwarzhans	Late Cretaceous (Maastrichtian)	Severn Formation	United States (Maryland)	Possibly a member of Polymixiiformes. Genus includes new species S. bourdoni.

Research

• A study aiming to determine whether the earliest vertebrates may have swum under various conditions without a clearly-differentiated tail fin, based on data from an abstracted model of Metaspriggina walcotti, is published by Rival, Yang & Caron (2021).
• Miyashita et al. (2021) report larval and juvenile forms of four stem lampreys from the Paleozoic era (Hardistiella, Mayomyzon, Pipiscius and Priscomyzon), including a hatchling-to-adult growth series of Priscomyzon, and report that the studied larvae display features that are otherwise unique to adult modern lampreys, and lack the defining traits of ammocoetes.
• A study on the morphological and functional diversity of osteostracan and galeaspid headshields, and on its implications for the knowledge of the ecology of the immediate jawless relatives of jawed vertebrates, is published by Ferrón et al. (2021).
• A study on the histology of the dermal skeleton in Procephalaspis oeselensis, Aestiaspis viitaensis, Dartmuthia gemmifera and four species of Tremataspis is published by Bremer et al. (2021), who interpret their findings as indicative of the emergence of the complex pore‐canal system in Tremataspis through the modification of the structures already present in other taxa.
• A study on the morphology of the earliest osteocytes in Tremataspis mammillata and Bothriolepis trautscholdi is published by Haridy et al. (2021), who interpret their findings as indicating that the earliest known osteocytes in the fossil record had similar morphology and likely similar physiological capabilities to their modern counterparts, and attempt to determine initial driver favoring evolution of cellular (osteocytic) over acellular (anosteocytic) bones in vertebrates.
• Zhu et al. (2021) use CT scanning to reveal the endocast of Brindabellaspis stensioi, and evaluate the implications of its anatomy for the knowledge of the phylogenetic relationships of early jawed vertebrates.
• Redescription of the anatomy of the headshield of Parayunnanolepis xitunensis is published by Wang & Zhu (2021).
• Description of new fossil material of Palaeacanthaspis vasta from the Devonian (Lochkovian) Chortkiv Formation (Ukraine), and a study on the phylogenetic relationships of this species, is published by Dupret et al. (2021).
• A study on the development of teeth in acanthodians, and on its implications for the knowledge of the evolution of teeth of jawed vertebrates, is published by Rücklin et al. (2021).
• Description of the first known skull remains of Onchopristis numidus from the Cretaceous Kem Kem Group (Morocco), and a study on the anatomy and phylogenetic relationships of this species, is published by Villalobos-Segura et al. (2021), who name a new family Onchopristidae.
• New, exceptionally well‐preserved skeleton of Asteracanthus ornatissimus, preserved with teeth that markedly differ from other teeth referred to Asteracanthus, is described from the Tithonian Altmühltal Formation (Germany) by Stumpf et al. (2021), who interpret this specimen as indicating that Asteracanthus and Strophodus represent two valid genera distinct from all other hybodontiforms.
• A study on the biomechanics of teeth of five species of Otodus, aiming to assess the functional significance of morphological trends in otodontid teeth and to test whether the morphology of otodontid teeth enabled the transition from piscivory to predation on marine mammals and the evolution of titanic body sizes, is published by Ballell & Ferrón (2021)
• A study on a bonebed in the Oligocene Chandler Bridge Formation (South Carolina, United States) with a large sample of Carcharocles angustidens dominated by small teeth is published by Miller, Gibson & Boessenecker (2021), who interpret this bonebed as a nursery area for C. angustidens.
• A study on growth patterns, reproductive biology and likely lifespan of Otodus megalodon is published by Shimada et al. (2021).
• Perez, Leder & Badaut (2021) present a novel method for estimating body size in fossil lamniform sharks, and attempt to determine the body size of Otodus megalodon.
• Revision of the fossil record of the extant tiger shark and the extinct members of the tiger shark lineage is published by Türtscher et al. (2021).
• Redescription of Striatolamia tchelkarnurensis is published by Malyshkina (2021).
• Shark teeth which might represent the first occurrence of the blacknose shark in the Pacific Ocean are described from the Pliocene Upper Onzole Formation (Ecuador) by Collareta et al. (2021), who evaluate the implications of this finding for the knowledge of the evolutionary history of the blacknose shark and the whitenose shark.
• Evidence of a previously unknown major extinction of sharks in the early Miocene, ~19 million years ago, is presented by Sibert & Rubin (2021).
• A platysomid specimen, representing the earliest deep-bodied actinopterygian reported to date, is described from the Carboniferous (Tournaisian) Horton Bluff Formation (Canada) by Wilson, Mansky & Anderson (2021), who evaluate the implications of this findings for the knowledge of the evolution of early ray-finned fishes.
• A review of the fossil record of Early–Middle Triassic marine bony fishes, aiming to determine the implications of poor fossil record from the late Olenekian-early middle Anisian interval on the knowledge of the Triassic radiation of bony fishes, is published by Romano (2021).
• A diverse assemblage of late Maastrichtian and Paleocene ray-finned fishes is described from Evrytania (Greece) by Argyriou & Davesne (2021).
• New fish fauna dating to the Paleocene–Eocene Thermal Maximum, indicating that diverse fish communities thrived in the paleotropics during this time period, is reported from Egypt by El-Sayed et al. (2021).
• A study on the morphological diversity and evolution of pycnodontiforms is published by Cawley et al. (2021).
• A study on fossil crushing dentitions of Pycnodus zeaformis and P. maliensis, providing evidence of a distinct pattern of gap‐filling tooth addition in pycnodonts, with individual large teeth replaced by multiple small teeth, is published by Collins & Underwood (2021).
• A study on the evolutionary history of lanternfishes, primarily based on the fossil record of otoliths, is published by Schwarzhans & Carnevale (2021).
• A study on the phylogenetic relationships of extant and fossil coelacanths is published by Toriño, Soto & Perea (2021).
• A study on the morphology and histology of the scales of Miguashaia bureaui, and on its implications for the knowledge of the evolution of the squamation in coelacanths, is published by Mondéjar‐Fernández et al. (2021).
• New fossil remains representing one of the largest known coelacanths ever reported are described from the Middle Jurassic of Normandy (France) by Cavin et al. (2021), who also compare the relationship between taxic diversity and body size diversity in coelacanths and ray-finned fishes over the Devonian–Paleocene time interval.
• An ossified lung of a mawsoniid coelacanth is described from the Maastrichtian of Oued Zem (Morocco) by Brito et al. (2021), representing the last known record of a Mesozoic coelacanth and the first known occurrence of coelacanths in the phosphate deposits of North Africa.
• A study on the evolution of feeding modes among tetrapodomorphs, as indicated by the anatomy of the skull of Tiktaalik roseae, is published by Lemberg, Daeschler & Shubin (2021), who report the simultaneous occurrence of anatomical modifications of the skull for prey capture through biting, as well as joint morphologies suggestive of cranial kinesis that is also present in suction-feeding fish.

Amphibians

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bermanerpeton	Gen. et sp. nov	Valid	Werneburg, Schneider & Lucas	Carboniferous (Kasimovian)	Atrasado Formation	United States (New Mexico)	A dvinosauroid temnospondyl. The type species is B. kinneyi.
Laosuchus hun	Sp. nov	Valid	Liu & Chen	Late Permian	Naobaogou	China	A chroniosuchian.
Palaeobatrachus codreavladi	Sp. nov	Valid	Roček, Rage & Venczel
Palaeobatrachus minutus	Sp. nov	Valid	Roček, Rage & Venczel
Rocekophryne	Gen. et sp. nov	Valid	Rage et al.	Eocene		Algeria	A frog belonging to the group Ranoidea. The type species is R. ornata.

Research

• A study on the function and evolution of forelimbs of early tetrapods, based on data from three-dimensional models of bones and muscles of forelimbs of Eusthenopteron foordi, Acanthostega gunnari and Pederpes finneyae, is published by Molnar et al. (2021).
• A study on the locomotor capabilities of tetrapods from the earliest Carboniferous Blue Beach site (Nova Scotia, Canada) is published by Lennie et al. (2021).
• A study on the early evolution of long bone elongation and bone marrow in tetrapods, based on data from temnospondyls (Apateon and Metoposaurus) and seymouriamorphs (Seymouria and Discosauriscus), is published by Estefa et al. (2021), who find the terrestrial Permian seymouriamorphs to be the oldest known tetrapods exhibiting a centralized marrow organization of long bones (which allows production of blood cells as in extant amniotes), and argue that the migration of blood-cell production in long bones probably wasn't an exaptation predating the water-to-land transition.
• A study on the skeletal anatomy of the holotype specimen of Ichthyerpeton bradleyae is published by Ó Gogáin & Wyse Jackson (2021).
• Description of the anatomy of the postcranial skeleton of Whatcheeria deltae is published by Otoo et al. (2021).
• A study on the relations between vertebral shape and terrestriality in the evolution of temnospondyls is published by Carter et al. (2021).
• Description of new fossil material of temnospondyls from the Triassic of the Ruhuhu and Luangwa basins (Tanzania and Zambia), providing new information on the diversity of Triassic African temnospondyls and their recovery after the Permian–Triassic extinction event, is published by Steyer et al. (2021).
• A study on the anatomy and phylogenetic relationships of "Cheliderpeton" lellbachae is published by Schoch (2021), who transfers this species to the genus Glanochthon in the family Sclerocephalidae.
• A study on the histology of different-sized femora and vertebra of specimens of Platyoposaurus stuckenbergi is published by Uliakhin, Skutschas & Saburov (2021).
• A study on the anatomy and phylogenetic relationships of Tertrema acuta is published by Slodownik, Mörs & Kear (2021).
• Redescription of the metoposaurid fossil material from the Upper Triassic Zions View locality (New Oxford Formation; Pennsylvania, United States) is published by Gee & Jasinski (2021), who assign this material to the species Anaschisma browni, expanding known geographic range of this taxon.
• Redescription of the holotype specimens of Borborophagus wyomingensis and Koskinonodon princeps, and a reassessment of their synonymy with Anaschisma browni, is published by Kufner & Gee (2021).
• A study on the anatomy and phylogenetic relationships of Timonya anneae and Procuhy nazariensis is published by Marsicano et al. (2021).
• A study on the anatomy and phylogenetic relationships of Macrerpeton huxleyi is published by Schoch & Milner (2021).
• Description of a new specimen of Conjunctio from the Permian Cutler Formation (Colorado, United States), and a study on the phylogenetic relationships of this genus, is published by Gee et al. (2021).
• New fossil material of Micropholis stowi, expanding known geographic range of this species, is described from the lower Fremouw Formation (Halfmoon Bluff, Antarctica) by Gee & Sidor (2021).
• New early adult specimen of Milnererpeton huberi, providing new information on the ontogenetic development of amphibamiform temnospondyls, is described from the Carboniferous (Kasimovian) Atrasado Formation (New Mexico, United States) by Werneburg, Schneider & Lucas (2021).
• An early Campanian assemblage of anuran bones, suggestive of high local species richness of frogs, is described from the Aguja Formation (Texas, United States) by Wick (2021).
• Description of new fossil material of Hungarobatrachus szukacsi from the Upper Cretaceous (Santonian) Csehbánya Formation (Hungary), and a study on the anatomy and phylogenetic relationships of this species, is published by Venczel, Szentesi & Gardner (2021).
• Revision of the fossil record of the family Ceratophryidae is published by Gómez & Turazzini (2021).
• Revision of the fossil material of Mesozoic temnospondyls and anurans housed in the collections of the Sirindhorn Museum and the Palaeontological Research and Education Centre of Mahasarakham University (Thailand), including fossils of brachyopids resembling the Chinese forms, is published by Nonsrirach, Manitkoon & Lauprasert (2021).

Reptiles

Main articles: 2021 in reptile paleontology and 2021 in archosaur paleontology

Synapsids

Non-mammalian synapsids

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Acratophorus	Gen. et comb. nov	Valid	Kammerer & Ordoñez	Middle Triassic (Anisian)?	Río Seco de la Quebrada	Argentina	A kannemeyeriid dicynodont, the type species is "Kannemeyeria" argentinensis.
Borealestes cuillinensis	Sp. nov	Valid	Panciroli et al.	Middle Jurassic (Bathonian)	Kilmaluag Formation	United Kingdom	A docodont.
Dobunnodon	Gen. et comb. nov	Valid	Panciroli et al.	Middle Jurassic (Bathonian)	Forest Marble Formation	United Kingdom	A docodont; a new genus for "Borealestes" mussettae Sigogneau−Russell (2003).
Fossiomanus	Gen. et sp. nov	Valid	Mao et al.	Early Cretaceous (Aptian)	Jiufotang Formation	China	A cynodont belonging to the family Tritylodontidae. Genus includes new species F. sinensis.
Isengops	Gen. et sp. nov	In press	Sidor, Tabor & Smith	Late Permian	Madumabisa Mudstone	Zambia	A burnetiamorph biarmosuchian. Genus includes new species I. luangwensis.
Kannemeyeria aganosteus	Sp. nov	Valid	Kammerer & Ordoñez	Middle Triassic (Anisian)?	Quebrada de los Fósiles	Argentina	A species of Kannemeyeria.
Mobaceras	Gen. et sp. nov	Valid	Kammerer & Sidor	Middle Permian	Madumabisa Mudstone	Zambia	A burnetiid therapsid. The type species is G. zambeziense.
Turfanodon jiufengensis	Sp. nov	Valid	Liu	Late Permian	Naobaogou Formation	China	A dicynodontoid dicynodont.

Research

• A study on the evolution of the vertebral column in synapsids is published by Jones et al. (2021), who interpret their findings as refuting the idea that the transition from non-mammalian synapsids to mammals involved a shift from reptile-like lateral bending of the backbone to sagittal bending, and argue that non-mammalian synapsids were characterized by their own unique functional regime of the vertebral column, distinct from that of extant reptiles and amphibians.
• A study comparing the forelimb morphology in extant mammals and fossil non-mammalian synapsids, aiming to determine whether extant mammals are good ecomorphological analogues for extinct synapsids, whether examples of ecomorphological convergence can be found among synapsids, and whether evolutionary history determined available functional solutions in synapsid forelimbs, is published by Lungmus & Angielczyk (2021).
• A study comparing the morphology of the maxillary canal of Heleosaurus scholtzi, Varanosaurus acutrostris, Orovenator mayorum and Prolacerta broomi, and evaluating the implications of the morphology of the maxillary canal for the knowledge of the phylogenetic placement of varanopids, is published by Benoit et al. (2021).
• A study on the skeletal anatomy and phylogenetic relationships of Raranimus dashankouensis is published by Duhamel et al. (2021).
• A study on the paleoneurology and likely paleobiology of Anteosaurus magnificus is published by Benoit et al. (2021).
• New specimen of Lanthanostegus mohoii, providing new information on the anatomy of the skull of this dicynodont and providing the first direct correlation between the lower Abrahamskraal Formation at Jansenville on the eastern side of the Karoo Basin and the southwestern part of this basin, is described by Rubidge, Day & Benoit (2021).
• New burrow casts containing skeletons of Diictodon, including associated remains of adult and infant specimens, are described by Smith et al. (2021), who consider it likely that portions of underground burrows produced Diictodon by were facultatively used as brood chambers.
• Redescription and a study on the phylogenetic relationships of Kunpania scopulusa is published by Angielczyk, Liu & Yang (2021).
• A study on the bone histology and likely life history of specimens of Lystrosaurus from the Lower Triassic Turpan Basin (Xinjiang, China), comparing them with specimens from South Africa, is published by Han, Zhao & Liu (2021).
• A new postcranial specimen of a stahleckeriid dicynodont, possibly of Stahleckeria, is described from the Chañares Formation, representing the oldest record of stahleckeriine dicynodonts from the Ischigualasto-Villa Unión Basin in Argentina.
• A study on the quality of the early cynodont fossil record in time and space, and on its implications for the understanding of the group's evolutionary history, is published by Varnham, Mannion & Kammerer (2021).
• A study on the anatomy and variation of the stapes in Thrinaxodon and Galesaurus is published by Gaetano & Abdala (2021).
• A study on the morphology of the nasal cavity of Exaeretodon riograndensis and Siriusgnathus niemeyerorum is published by Franco et al. (2021).
• A study on the morphology of the endocast of a specimen of Riograndia guaibensis from the Linha São Luiz site (Candelária Sequence of the Santa Maria Supersequence, Brazil) is published by Kerber et al. (2021).
• New specimen of the Middle Jurassic haramiyidan Vilevolodon diplomylos with well-preserved malleus, incus and ectotympanic is described by Wang et al. (2021).

Mammals

Main article: 2021 in mammal paleontology

Other animals

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Country	Notes	Images
Anulitubus	Gen. et sp. nov	Valid	Moczydłowska in Moczydłowska et al.	Ediacaran	Stáhpogieddi Formation	Norway	A member of Eumetazoa of uncertain phylogenetic placement. The type species is A. formosus.
Arienigraptus balticus	Sp. nov	Valid	Maletz & Ahlberg	Ordovician (Darriwilian)		Sweden	A graptolite.
Arienigraptus delicatus	Sp. nov	Valid	Maletz & Ahlberg	Ordovician (Darriwilian)		Sweden	A graptolite.
Arienigraptus robustus	Sp. nov	Valid	Maletz & Ahlberg	Ordovician (Dapingian)		Sweden	A graptolite.
Blastochaetetes reitneri	Sp. nov	In press	Sánchez-Beristain & García-Barrera in Sánchez-Beristain, García-Barrera & Juárez-Aguilar	Late Cretaceous	Tamasopo Formation	Mexico	A chaetetid demosponge.
Coniculus	Gen. et sp. nov	Valid	Moczydłowska in Moczydłowska et al.	Ediacaran	Stáhpogieddi Formation	Norway	A member of Eumetazoa of uncertain phylogenetic placement. The type species is C. elegantis.
Cornulites spinosus	Sp. nov	Valid	Vinn & Eyzenga	Late Ordovician		Netherlands	A cornulitid tubeworm.
Dailyatia icari	Sp. nov	Valid	Claybourn et al.	Cambrian Series 2		Antarctica	A camenellan tommotiid.
Fistula	Gen. et sp. nov	Valid	Moczydłowska in Moczydłowska et al.	Ediacaran	Stáhpogieddi Formation	Norway	A member of Eumetazoa of uncertain phylogenetic placement. The type species is F. crenulata.
Palaeocorvospongilla	Gen. et sp. nov	In press	Samant et al.	Late Cretaceous (Maastrichtian)	Deccan Intertrappean Beds	India	A sponge belonging to the family Palaeospongillidae. Genus includes new species P. cretacea.
Palaeosaccus minus	Sp. nov	Valid	Luo et al.	Cambrian	Shuijingtuo Formation	China	A sponge.
Papiliograptus retimarginatus	Sp. nov	Valid	Kozłowska & Bates	Silurian (Homerian)		Germany Poland	A graptolite belonging to the family Retiolitidae.
Saetaspongia jianhensis	Sp. nov	Valid	Ling et al.	Cambrian Stage 4	Balang Formation	China	A sponge of uncertain phylogenetic placement, possibly with protomonaxonid affinities.
Triticispongia giganta	Sp. nov	Valid	Luo et al.	Cambrian	Shuijingtuo Formation	China	A sponge.
Turriserpula	Gen. et sp. nov	In press	Dieni & Massari	Early Cretaceous (Berriasian)		Italy	A microserpulid. Genus includes new species T. coralliophila.
Vauxia paraleioia	Sp. nov	In press	Wei et al.	Cambrian Stage 3		China	A vauxiid sponge.
Vauxia pregracilenta	Sp. nov	In press	Wei et al.	Cambrian Stage 3		China	A vauxiid sponge.

Research

• A study aiming to identify characters of Kimberella, Ikaria, Dickinsonia and Tribrachidium controlled by conserved developmental processes, as well as genetic elements likely responsible for their expression, is published by Evans, Droser & Erwin (2021), who also attempt to determine phylogenetic positions of these taxa relative to extant animals.
• Structures interpreted as traces of motor activity of Dickinsonia are reported by Ivantsov & Zakrevskaya (2021), who interpret the studied traces as indicating that Dickinsonia was capable of both attachment and mobility.
• A study aiming to determine the feeding mode of Arkarua adami is published by Cracknell et al. (2021).
• Shore et al. (2021) report the first three-dimensional, pyritized preservation of soft tissue in Namacalathus hermanastes from the Nama Group (Namibia), and evaluate the implications of this finding for the knowledge of the phylogenetic relationships of this animal.
• A new assemblage of fossil eggs, embryos attributable to the early scalidophoran Markuelia, and early post-embryonic developmental stages of camenellans is described from the Cambrian Stage 3 Salanygol Formation (Mongolia) by Steiner et al. (2021).
• Redescription of Stanleycaris hirpex, and a study on the phylogenetic relationships of this species and on the functional specialization of the frontal appendages of this and other stem euarthropods, is published by Moysiuk & Caron (2021).

Other organisms

New taxa

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bicellum	Gen. et sp. nov		Strother & Wellman in Strother et al.	Torridonian	Diabaig Formation	United Kingdom	An organism of uncertain phylogenetic placement, possibly an early member of Holozoa. Genus includes new species B. brasieri. Appears to have differentiated multicellularity.
Gigarimaneta	Gen. et sp. nov	Valid	Taylor et al.	Ediacaran	Mistaken Point Formation	Canada (Newfoundland and Labrador)	A organism growing on the seafloor in a manner similar to Fractofusus and Beothukis. Genus includes new species G. samsoni.
Quadrimurus	Gen. et sp. nov	Valid	Miao, Moczydłowska & Zhu	Early Mesoproterozoic	Xiamaling Formation	China	An organic-walled microfossil. Genus includes new species Q. clavatus.

Research

• Delarue et al. (2021) describe 3.4 billion years old microfossils preserved with a tail-like structure from the Strelley Pool Formation (Australia), and interpret the tail-like appendage as likely providing early microorganisms with movement capabilities.
• Tang et al. (2021) describe dark discoidal, semicircular, or ovate structures preserved on fossil of early Neoproterozoic eukaryotes Tawuia and Sinosabellidites from North China, and interpret these structures as fossils of eukaryotic epibionts that lived on the surface of and may have benefited from an association with their Tawuia and Sinosabellidites hosts.
• Well-preserved communities of large unbranched filamentous microorganisms, bearing morphological and ecological similarities with large sulfide-oxidizing bacteria such as Beggiatoa, are described from the Ediacaran Itajaí Basin (Brazil) by Becker-Kerber et al. (2021).
• Zacaï et al. (2021) attempt to determine the potential timing of establishment of the latitudinal diversity gradient for early Paleozoic acritarchs and its evolution through time .

History of life in general

• A study on the taphonomy of eukaryotic organelles, assessing the basis of the view that organelles decay too rapidly to be fossilized and evaluating the plausibility of the claims of organelles preserved in Proterozoic fossils, is published by Carlisle et al. (2021).
• Evidence of the presence of significant populations of both red and green algae ca. 1.4 billion years ago (600 million years earlier than previously recognized) is reported from the Xiamaling Formation (China) by Zhang et al. (2021).
• A study on the major biotic transitions in the Phanerozoic fossil record of the benthic marine faunas is published by Rojas et al. (2021), who report evidence of three major biotic transitions (across the end-Cambrian, end-Permian, and mid-Cretaceous boundaries).
• A study on changes of diversity of skeletonized marine invertebrates in the fossil record, evaluating the impact of dead clades walking on broader trends in Phanerozoic biodiversity, is published by Barnes, Sclafani & Zaffos (2021), who identify 70 invertebrate orders that experienced major diversity losses without recovery, but note that most of these taxa had a long duration after the drop in diversity, and many drops in diversity without recovery were not associated with mass extinction events.
• Geyer & Landing (2021) report a hitherto unknown Cambrian Stage 3 Lagerstätte from the Amouslek Formation (Morocco), preserving the first relatively abundant fossils with exceptional preservation from the Cambrian of Morocco (and Africa).
• A study on Carboniferous and early Permian tetrapod tracks, and on their implications for the knowledge of evolutionary changes in the anatomy of the trackmakers in and locomotion style close to the origin of amniotes, is published by Buchwitz et al. (2021).
• A study on the impact of Permian mass extinctions on continental invertebrate infauna, based on data from the Iberian Basin (central Spain), is published by Buatois et al. (2021), who report evidence of a dramatic decrease in bioturbation intensity on land by the end of the Capitanian, coinciding with an increase in weathering intensity and acidic conditions, and a collapse in plant communities spanning the late Permian–Early Triassic in the Iberian Basin.
• A review of the state of research on the Capitanian mass extinction event in the Karoo Basin (South Africa) is published by Day & Rubidge (2021).
• Evidence from tetrapod fossil record from the Karoo Basin (South Africa) indicative of a protracted (∼1 Ma) extinction on land during the Permian-Triassic transition is presented by Viglietti et al. (2021).
• Evidence of two pulses of extinction at the Permian–Triassic boundary caused by different environmental triggers is reported from the Liangfengya section in the South China Block by Li et al. (2021).
• Revision of the Triassic record of tetrapod tracks is published by Klein & Lucas (2021).
• A study on the diversity dynamics and evolution of the functional morphology of tetrapod herbivores throughout the Triassic and Early Jurassic is published by Singh et al. (2021).
• Marchetti et al. (2021) revise the tetrapod (including dinosauromorph) footprint assemblage from the Quarziti del Monte Serra Formation (Ladinian of Italy), and interpret this assemblage and other findings of Ladinian dinosauromorph footprints as evidence of wide dispersal of dinosauromorphs as early as the Middle Triassic.
• Zouhri et al. (2021) describe a diverse vertebrate fauna from the Eocene (Bartonian) Aridal Formation (Western Sahara), including 12 species of cartilaginous fishes, at least three species of turtles, at least two longirostrine crocodylian taxa, the oldest record of Pelagornis reported to date, and a proboscidean possibly belonging to the genus Barytherium.
• Description of non-mammal vertebrate fauna from the Miocene (Messinian) Monticino Quarry (Italy), including the oldest known records of the javelin sand boa or a related species, snakes belonging to the genus Malpolon and an unambiguous bustard reported to date, is published by Villa et al. (2021).
• A study on the age of escorias (glassy rock fragments similar to volcanic scoriae, likely products of extraterrestrial impacts) collected along the Pampean Atlantic coast from the "Irene" and Chapadmalal Formations (Argentina), and on their implications for the knowledge of the timing of late Miocene–Pliocene faunal succession in the Pampean Region, is published by Prevosti et al. (2021).
• A study on the age of the most recent Pleistocene megafaunal specimens from Cloggs Cave (Australia), and on its implications for the knowledge of the timing and causes of Late Pleistocene extinctions of Australian megafauna, is published by David et al. (2021).
• A study aiming to determine whether a significant relationship can be detected between demographic susceptibility to extinction of members of Quaternary megafauna of Sahul and their extinction chronology inferred from their fossil record is published by Bradshaw et al. (2021).
• A study aiming to determine whether the fossil record indicates that the arrival of hominins on islands in the Pleistocene was coincident with the disappearance of insular taxa is published by Louys et al. (2021).
• A study on the extinction dynamics of the elephant birds and Malagasy hippos is published by Hansford et al. (2021), who interpret their findings as indicating that these animals persisted for millennia after first human arrival on Madagascar, that their communities collapsed suddenly ∼1200-900 BP, and that their extinctions were closely correlated in time with intensive conversion of forests to grassland, probably resulting from human shift to agro-pastoralism.
• A study aiming to determine how observed extinctions in the geological past can be predicted from the interaction of long-term temperature trends with short-term climate change is published by Mathes et al. (2021).
• A study on the impact of the Capitanian mass extinction event, Permian–Triassic extinction event and Triassic–Jurassic extinction event on terrestrial and freshwater ecosystems, aiming to quantify community resistance during the extinction events and to determine ecological dynamics of communities before and after these extinctions, is published by Huang et al. (2021).
• A study on correlations between fossilization potential and food web features, aiming to determine how fossilization impacts inferences of ancient community structure, is published by Shaw et al. (2021).
• A study on the drilling predation pressure on sea urchins across the Mesozoic and Cenozoic is published by Petsios et al. (2021), who present evidence indicative of the Cenozoic intensification of this predation, and argue that the Mesozoic marine revolution was more likely a series of asynchronous processes with variable significance across different groups of predators and preys, rather than a single synchronized ecosystem-wide event.
• A study on the spatial biodiversity dynamics of unicellular marine plankton throughouth the Cenozoic, aiming to test the generality of the ‘out of the tropics’ hypothesis (positing that the tropics are both a cradle and source of biodiversity for extratropical regions), is published by Raja & Kiessling (2021).
• A study on the evolution of ecophysiological adaptations to life in the sea in extant and fossil marine tetrapods (excluding birds) is published by Motani & Vermeij (2021).

Other research

• Mißbach et al. (2021) report the existence of indigenous organic molecules and gases in primary fluid inclusions in c. 3.5-billion-year-old barites from the Dresser Formation (Pilbara Craton, Australia), providing evidence of the organic composition of primordial fluids that were available for the early microbes.
• A study on the 3.4-billion-year old organic films from the Buck Reef Chert (Kaapvaal Craton, South Africa) is published by Alleon et al. (2021), who interpret their findings as indicating that early Archean organic films carry chemical information directly related to their original molecular compositions, and evaluate the implications of their finding for the knowledge of the initial chemical nature of organic microfossils found in ancient rocks.
• Evidence of prolonged and repeated oxygen stress in the Appalachian Basin associated with the Late Devonian extinctions is presented by Boyer et al. (2021).
• Rakociński et al. (2021) report very large anomalous mercury spikes from the south-western part of Tian Shan (Uzbekistan), and interpret this finding as evidence of intensive volcanic activity both predating and occurring during the Hangenberg Crisis.
• Evidence from the southern Karoo Basin of South Africa indicative of at least four atmospheric carbon dioxide spikes coinciding with extinctions on land and at sea from the Late Permian to the Middle Triassic is presented by Retallack (2021).
• A study evaluating whether fuel-driven changes to fire activity during the Cretaceous period had the ability to counteract rising atmospheric oxygen at this time is published by Belcher et al. (2021), who argue that alteration of fire feedbacks driven by the rise of the flowering plants likely lowered atmospheric oxygen levels from ~30% to 25% by the end of the Cretaceous.
• White & Campione (2021) describe a workflow in which three-dimensional surface profiles of fragmentary fossils can be quantitatively compared to better-known exemplars in order to identify fragmentary fossils, and apply this workflow to megaraptorid theropod unguals from the Cretaceous of Australia.
• A study aiming to test whether histological characters can be used to assign bones to individuals within a quarry, using sauropod dinosaur material from two adjacent Morrison quarries in the Bighorn Basin (Wyoming, United States) as a case study, is published by Wiersma-Weyand et al. (2021).
• A study on diverse amniotic eggshells from the Wido Volcanics (Upper Cretaceous, South Korea), evaluating their utility for assessments of the paleothermometry of the sedimentary deposits, is published by Choi et al. (2021).
• A study on the age and duration of the Lower Cretaceous Yixian Formation (China) is published by Zhong et al. (2021).
• Goderis et al. (2021) report new data revealing a positive iridium anomaly within the peak-ring sequence of the Chicxulub impact structure, and interpret this finding as conclusively tying Chicxulub to the global iridium layer and Cretaceous-Paleogene boundary sections worldwide, confirming the link between crater formation and the iridium peak detected in these sections.
• A study aiming to determine whether a strong link can be established between stable carbon isotopes of tooth enamel of herbivores and vegetation structure in present African ecosystems, and whether enamel stable carbon isotopes of fossil herbivores are useful for making inferences about Plio-Pleistocene vegetation structure in Africa and the environmental context of hominin evolution, is published by Robinson et al. (2021).
• A study on environmental changes in East Africa at the time of the extinction of Paranthropus boisei is published by Quinn & Lepre (2021), who report evidence of a significant reduction in C₄ grasslands during Mid-Pleistocene Transition, and argue that this reduction might have escalated dietary competition amongst the abundant C₄-feeders and influenced P. boisei’s demise.
• Evidence from Chitimwe Beds (northern Malawi), indicating that in the late Pleistocene early modern humans fundamentally altered local landscapes and ecology using fire, is presented by Thompson et al. (2021).
• Ellis et al. (2021) examine current biodiversity patterns in relation to distribution of human populations and land use over the past 12,000 years, and argue that as early as 12,000 years ago nearly three quarters of Earth’s land was inhabited and shaped by human societies.
• Alleon et al. (2021) revise reports of organic molecules in animal fossils, and argue that purported signatures of organic molecules are in reality instrumental artefacts resulting from intense background luminescence.

Paleoclimate

• Scotese et al. (2021) estimate how global temperatures have changed during the last 540 million years.
• A high-resolution proxy record of Late Cambrian and Ordovician climate is presented by Goldberg et al. (2021).
• A study on changes in weathering intensity and temperature along a temperate to subpolar southeastern margin of Gondwana (eastern margin of present-day Australia) across the end-Permian extinction is published by Frank et al. (2021).
• A study on the atmospheric CO₂ levels during the Permian–Triassic transition, based on data from fossil plant remains from sedimentary successions in southwestern China, is published by Wu et al. (2021), who present evidence of a six-fold increase of atmospheric pCO₂ during the Permian–Triassic mass extinction.
• A study on the climate of the Lufeng area (China) during the Early Jurassic, and on the relationship between the global distribution of dinosaur fossils and climate during the Jurassic, is published by Shen et al. (2021).
• Evidence of the presence of a terrestrial climate barrier in the Western Interior Basin of North America during the final 15 million years of the Cretaceous, dividing the Western Interior Basin into warm southern and cool northern biomes, is presented by Burgener et al. (2021), who also report evidence indicating that the biogeographical distribution of plants was heavily influenced by the presence of this temperature transition zone.
• De Winter et al. (2021) present reconstructions of monthly sea surface temperatures at around 50 °N latitude about 78 million years ago, based on data from oyster and rudist shells from the Kristianstad Basin (Sweden).
• A study on CO₂ contents of early Deccan Traps lavas, aiming to determine whether early Deccan magmatism triggered the warming event during the latest Maastrichtian, is published by Hernandez Nava et al. (2021).
• Vento et al. (2021) estimate parameters of the Paleogene to Neogene climate on the basis of data from fossil leaves from the Río Turbio and Río Guillermo formations in southern South America (Argentina).
• 10-million-year long proxy record of Arabian climate is developed by Böhme et al. (2021), who report evidence indicative of a sustained period of hyperaridity in the Pliocene and a number of transient periods of hyperaridity in northern Arabia during the late Miocene which were out of phase with those in North Africa, and argue that these desert dynamics had a strong control on large-scale mammalian dispersals between Africa and Eurasia.
• A study aiming to reconstruct summer and winter temperatures in the Late Pleistocene when Neanderthals were using the site of La Ferrassie (France), based on data from oxygen isotope measurements of bovid tooth enamel, is published by Pederzani et al. (2021).
• Data from analyses and modelling of noble gases in groundwater, indicating that the low-altitude, low-to-mid-latitude land surface (45 degrees south to 35 degrees north) was about 6 °C cooler during the Last Glacial Maximum than during the Late Holocene, is presented by Seltzer et al. (2021).