2021 in paleobotany

This article records new taxa of fossil plants described during the year 2021, as well as other significant discoveries and events related to paleobotany that occurred in 2021.

Ferns and fern allies

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Arthropitys buritiranensis [1]	Sp. nov	In press	Neregato et al.	Permian	Motuca Formation	Brazil	A member of Calamitales.
Azolla andreisii [2]	Sp. nov	In press	De Benedetti et al.	Late Cretaceous (Maastrichtian)	La Colonia Formation	Argentina	A species of Azolla .
Caulopteris ellipticus [3]	Sp. nov	In press	Wang et al.	Early Permian	Taiyuan Formation	China	A marattialean tree fern belonging to the family Psaroniaceae.
Caulopteris neimengensis [3]	Sp. nov	In press	Wang et al.	Early Permian	Taiyuan Formation	China	A marattialean tree fern belonging to the family Psaroniaceae.
Caulopteris obovatus [3]	Sp. nov	In press	Wang et al.	Early Permian	Taiyuan Formation	China	A marattialean tree fern belonging to the family Psaroniaceae.
Cicatricosisporites pseudograndiosus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the genus Ceratopteris .
Cladarastega [5]	Gen. et sp. nov	Valid	Poinar	Late Cretaceous (Cenomanian)	Burmese amber	Myanmar	A fern belonging to the family Dennstaedtiaceae. Genus includes new species C. burmanica.
Claytosmunda zhangii [6]	Sp. nov	In press	Tian, Wang & Jiang	Late Jurassic	Tiaojishan Formation	China	A fern, a species of Claytosmunda .
Dennstaedtia christophelii [7]	Sp. nov	Valid	Pigg et al.	Early Eocene	Klondike Mountain Formation	United States ( Washington)	A fern, a species of Dennstaedtia .	Dennstaedtia christophelii
Echinosporis densiechinatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Marattiaceae.
Eoangiopteris congestus [8]	Sp. nov	Valid	Sun et al.	Early Permian	Taiyuan Formation	China	A fern belonging to the group Marattiales.
Hymenophyllum axsmithii [7]	Sp. nov	Valid	Pigg et al.	Early Eocene	Klondike Mountain Formation	United States ( Washington)	A fern, a species of Hymenophyllum .
Iberisetum [9]	Gen. et sp. nov	In press	Correia, Šimůnek & Sá	Carboniferous (Gzhelian)	Douro Basin	Portugal	A member of Equisetales. Genus includes new species I. wegeneri.
Laevigatosporites cultellus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Polypodiaceae.
Marsileaceaephyllum ciliatum [10]	Sp. nov	In press	Wang et al.	Cretaceous	Burmese amber	Myanmar	A member of the family Marsileaceae.
Nemejcopteris haiwangii [11]	Sp. nov	In press	Pšenička et al.	Permian (Asselian)	Taiyuan Formation	China	A zygopterid fern.
Neocalamites iranensis [12]	Sp. nov	Valid	Kustatscher, Mazaheri-Johari & Roghi in Mazaheri-Johari et al.	Late Triassic (Carnian)	Miakuhi Formation	Iran	A member of the family Equisetaceae.
Odontosoria marekgaltieri [13]	Sp. nov	In press	Pšenička, Sakala & Dašková	Early Miocene	Most Basin	Czech Republic	A species of Odontosoria .
Oligosporangiopteris [14]	Gen. et sp. nov	In press	Votočková Frojdová et al.	Early Permian	Taiyuan Formation	China	A leptosporangiate fern. Genus includes new species O. zhongxiangii.
Osmunda zhangpuensis [15]	Sp. nov		Wang & Sun in Wang et al.	Miocene	Fotan Group	China	A fern, a species of Osmunda .
Patagoniapteris [16]	Gen. et sp. nov	Valid	Gnaedinger & Zavattieri	Late Triassic (Norian–Rhaetian)	Paso Flores Formation	Argentina	A member of the family Dipteridaceae. Genus includes new species P. artabeae.
Pectinangium xuanweiense [17]	Sp. nov	In press	Zhou et al.	Permian (Lopingian)		China	A fern belonging to the group Marattiales.
Polypodiisporites densus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Polypodiaceae.
Polypodiisporites fossulatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Lomariopsidaceae.
Psilatriletes marginatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the genus Cyathea .
Qasimia yunnanica [18]	Sp. nov	In press	Guo et al.	Permian (Lopingian)	Xuanwei Formation	China	A fern belonging to the group Marattiales.
Reticulosporis diversus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Polypodiaceae.
Rothwellopteris sanjiaoshuensis [19]	Sp. nov	In press	He et al.	Late Permian	Xuanwei Formation	China	A fern belonging to the group Marattiales.
Sphenophyllum fanwanense [20]	Sp. nov	In press	Huang et al.	Late Devonian		China	A sphenophyllalean equisetid
Sphenophyllum parvifolium [21]	Sp. nov	In press	Libertín et al.	Early Permian	Taiyuan Formation	China	A sphenophyllalean equisetid
Tapelrayen [22]	Gen. et sp. nov	In press	Machado et al.	Eocene	Huitrera Formation	Argentina	Fertile remains of a fern comparable with Thelypteridaceae and Dryopteridaceae. Genus includes new species T. helgae.
Thyrsopteris cyathindusia [23]	Sp. nov	In press	Zhang et al.	Cretaceous	Burmese amber	Myanmar	A tree fern, a species of Thyrsopteris .
Woodwardia changchangensis [24]	Sp. nov	In press	Naugolnykh & Song in Song et al.	Middle Eocene	Changchang Formation	China	A fern, a species of Woodwardia .

Bennettitales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Nilssoniopteris jogiana [25]	Sp. nov		Blomenkemper & Abu Hamad in Blomenkemper et al.	Permian (Changhsingian)	Umm Irna Formation	Jordan	A member of Bennettitales.
Nilssoniopteris shanxiensis [25]	Sp. nov		Bäumer, Backer & Wang in Blomenkemper et al.	Permian (Cisuralian)	Upper Shihhotse Formation	China	A member of Bennettitales.
Pterophyllum pottii [25]	Sp. nov		Bomfleur & Kerp in Blomenkemper et al.	Permian (Changhsingian)	Umm Irna Formation	Jordan	A member of Bennettitales.
Weltrichia magna [26]	Sp. nov	Valid	Guzmán-Madrid & Velasco de León	Middle Jurassic (Bajocian)	Zorrillo Formation	Mexico
Weltrichia xochitetlii [27]	Sp. nov	In press	Lozano-Carmona et al.	Middle Jurassic (Callovian)	Tecomazuchil Formation	Mexico	A member of Bennettitales.
Williamsonia sanjuanensis [28]	Sp. nov	In press	Lozano-Carmona & Velasco-de León	Middle Jurassic		Mexico

Cycadales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Becklesia franconica [29]	Sp. nov	In press	Van Konijnenburg-van Cittert et al.	Late Triassic (Rhaetian)	Exter Formation	Germany	A member of Cycadales of uncertain phylogenetic placement.
Iratinia [30]	Gen. et sp. nov	In press	Spiekermann et al.	Permian (Kungurian)	Irati Formation	Brazil	A cycad-like plant. Genus includes new species I. australis.

Ginkgoales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Eretmophyllum hamiensis [31]	Sp. nov	In press	Tang et al.	Middle Jurassic	Xishanyao Formation	China
Ginkgoites villardeseoanii [32]	Sp. nov	In press	Andruchow-Colombo et al.	Late Cretaceous (Maastrichtian)	Lefipán Formation	Argentina
Ginkgoxylon arcticum [33]	Sp. nov	In press	Afonin & Gromyko	Early Cretaceous		Russia ( Arkhangelsk Oblast)	A member of Ginkgoales described on the basis of fossil wood.
Karkenia irkutensis [34]	Sp. nov	In press	Nosova, Crane & Shi	Middle Jurassic (Aalenian)	Prisayan Formation	Russia

Vladimariales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Pseudotorellia doludenkoae [35]	Sp. nov	Valid	Nosova, Kostina & Bugdaeva	Late Jurassic–Early Cretaceous (Oxfordian–Berriasian)	Dublikan Formation Talyndzhan Formation	Russia ( Khabarovsk Krai)
Pseudotorellia irkutensis [36]	Sp. nov	In press	Nosova	Middle Jurassic (Aalenian–Bajocian)	Prisayan Formation	Russia	A Vladimariales foliage species
Umaltolepis irkutensis [36]	Sp. nov	In press	Nosova	Middle Jurassic (Aalenian–Bajocian)	Prisayan Formation	Russia	A Vladimariales reproducive structure species

Conifers

Araucariaceae

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Agathoxylon mendezii [37]	Sp. nov	Valid	Del Fueyo et al.	Early Cretaceous (Berriasian–Valanginian)	Springhill Formation	Argentina	An Araucariaceae fossil wood.
Agathoxylon santanensis [38]	Sp. nov	In press	Dos Santos et al.	Early Cretaceous (Aptian)	Crato Formation	Brazil
Araucaria violetae [39]	Sp. nov	In press	Batista et al.	Early Cretaceous		Brazil	A species of Araucaria .

Cheirolepidiaceae

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Brachyoxylon lalongense [40]	Sp. nov	In press	Yang & Li	Early Cretaceous (Berriasian–Barremian)	Duoni Formation	China
Brachyoxylon patagonicum [41]	Sp. nov	In press	Rombola et al.	Late Cretaceous	Cerro Fortaleza Formation	Argentina	Fossil wood of a member of the family Cheirolepidiaceae.
Watsoniocladus cunhae [42]	Sp. nov	In press	Kvaček & Mendes	Early Cretaceous (Aptian–Albian)	Almargem Formation	Portugal	A member of the family Cheirolepidiaceae.

Cupressaceae

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Cupressinoxylon widdringtonioides [43]	Sp. nov	Valid	De Wit & Bamford	Late Cretaceous		South Africa	Fossil wood of a member or a relative of the family Cupressaceae.
Fokienia tianpingensis [44]	Sp. nov	Valid	Wu & Jin in Wu et al.	Miocene	Erzitang Formation	China	A species of Fokienia .
Nishidastrobus [45]	Gen. et sp. nov	Valid	Atkinson et al.	Late Cretaceous		Japan	A member of the family Cupressaceae belonging to the subfamily Cunninghamioideae. Genus includes new species N. japonicum.
Ohanastrobus [45]	Gen. et sp. nov	Valid	Atkinson et al.	Late Cretaceous		Japan	A member of the family Cupressaceae belonging to the subfamily Cunninghamioideae. Genus includes new species O. hokkaidoensis.
Protaxodioxylon metangulense [46]	Sp. nov	In press	Nhamutole, Bamford & Araújo	Permian (late Capitanian)	K5 Formation	Mozambique	A member of the family Cupressaceae.
Protaxodioxylon verniersii [46]	Sp. nov	In press	Nhamutole & Bamford in Nhamutole, Bamford & Araújo	Permian (late Capitanian)	K5 Formation	Mozambique	A member of the family Cupressaceae.
Thujopsoxylon [47]	Gen. et sp. nov	Valid	Dolezych, LePage & Williams	Oligocene (Chattian)	Korlikov Formation	Russia ( Tomsk Oblast)	Genus includes new species T. schneiderianum.

Pinaceae

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Lepidocasus [48]	Gen. et sp. nov	Valid	Herrera et al.	Early Cretaceous (Aptian–Albian)		Mongolia	A member of the family Pinaceae. Genus includes new species L. mellonae.
Nothotsuga sinogaia [49]	Sp. nov	In press	Ding et al.	Late Miocene		China	A species of Nothotsuga
Piceoxylon nikitinii [47]	Sp. nov	Valid	Dolezych, LePage & Williams	Oligocene (Chattian)	Korlikov Formation	Russia ( Tomsk Oblast)	A Picea relative wood.
Pinus leptokrempfii [50]	Sp. nov	Valid	Zhang et al.	Early Oligocene		China	A pine.
Pinus nongyaplongensis [51]	Sp. nov	In press	Grote in Grote & Srisuk	Oligocene-early Miocene		Thailand	A pine.
Pinus weichangensis [52]	Sp. nov	In press	Li et al.	Early Miocene		China	A pine.
Schizolepidopsis ediae [53]	Sp. nov	Valid	Matsunaga et al.	Early Cretaceous	Huolinhe Formation Tevshiin Govi Formation	China Mongolia	A member or a close relative of the family Pinaceae.

Podocarpaceae

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Podocarpus yunnanensis [54]	Sp. nov	In press	Wu et al.	Early Pliocene		China	A species of Podocarpus .
Protophyllocladoxylon hilarioense [55]	Sp. nov	In press	Vallejos Leiz, Crisafulli & Gnaedinger	Late Triassic (Norian–Rhaetian)	Hilario Formation	Argentina	A member of the family Podocarpaceae.
Protophyllocladoxylon yiwuense [56]	Sp. nov	In press	Gou & Feng in Gou et al.	Middle Jurassic	Xishanyao Formation	China	A conifer of uncertain phylogenetic placement, possibly belonging or related to the family Podocarpaceae.

Other conifers

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Cargalostrobus [57]	Gen. et sp. nov	Valid	Gomankov	Permian		Russia ( Orenburg Oblast)	A member of Pinales belonging to the family Sashiniaceae. Genus includes new species C. demetrii.
Megaporoxylon sinensis [58]	Sp. nov	In press	Wan et al.	Late Triassic (Carnian–Norian)	Huangshanjie Formation	China	A coniferous trunk.
Taxus huolingolensis [59]	Sp. nov	In press	Dong et al.	Early Cretaceous	Huolinhe Formation	China	A species of Taxus .
Voltzia edithae [60]	Sp. nov	Valid	Forte, Kustatscher & Van Konijnenburg-van Cittert	Middle Triassic (Anisian)		Italy	A member of Voltziales.
Xenoxylon utahense [61]	Sp. nov	In press	Xie & Gee in Xie et al.	Late Jurassic	Morrison Formation	United States ( Utah)	Fossil wood of a conifer.
Zhuotingoxylon [62]	Gen. et sp. nov	In press	Wan et al.	Permian (Changhsingian)	Guodikeng Formation	China	A silicified trunk with coniferous affinities. Genus includes new species Z. liaoi.

Flowering plants

Basal angiosperms

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Allonymphaea [63]	Nom. nov	Valid	Doweld	Eocene		Egypt	A replacement name for Thiebaudia Chandler (1954).

Unplaced non-eudicots

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Alcainea [64]	Gen. et sp. nov	Valid	Sender et al.	Early Cretaceous (Albian)	Escucha Formation	Spain	A member of the family Chloranthaceae. Genus includes new species A. eklundiae.
Todziaphyllum [64]	Gen. et sp. nov	Valid	Sender et al.	Early Cretaceous (Albian)	Escucha Formation	Spain	A member of the family Chloranthaceae. Genus includes new species T. elongatum.

Magnoliids

Canellales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Aristolochia macginitieana [65]	Nom. nov	Valid	Freitas & Doweld	Oligocene		United States ( California)	An Aristolochia species; a replacement name for Aristolochia triangularis MacGinitie (1937).
Cryptocaryoxylon grandoleaceum [66]	Sp. nov	Valid	Akkemik	Middle Miocene	Kesmekaya Volcanics	Turkey	A member of the family Lauraceae.
Laurus elliptica [67]	Nom. nov	Valid	Winterscheid in Winterscheid & Kvaček	Oligocene		Germany	A species of Laurus ; a replacement name for Laurus obovata Weber (1852).
Rosarioxylon [68]	Gen. et sp. nov	In press	Cevallos-Ferriz, Catharina & Kneller	Late Cretaceous (Campanian)	Rosario Formation	Mexico	A member of the family Lauraceae. Genus includes new species R. bajacaliforniensis.
Winteroxylon oleiferum [69]	Sp. nov	Valid	Brea et al.	Early Eocene	Huitrera Formation	Argentina	A member of the family Winteraceae.

Monocots

Alismatid monocots

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bognerospadix [70]	Gen. et sp. nov	Valid	Stockey, Hoffman & Rothwell	Paleocene		Canada ( Alberta)	A member of the family Araceae. Genus includes new species B. speirsiae.

Lilioid monocots

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Mirafloris [71]	Gen. et sp. nov	Valid	Poinar	Cretaceous	Burmese amber	Myanmar	A member of the family Liliaceae. Genus includes new species M. burmitis.

Commelinid monocots

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Arecipites invaginatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Arecaceae.
Arecocaryon [63]	Nom. nov	Valid	Doweld	Eocene	Messel pit	Germany	A member of the family Arecaceae; a replacement name for Friedemannia Collinson, Manchester & Wilde (2012).
Cyperaceaepollis wesselinghii [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Cyperaceae.
Eograminis [72]	Gen. et sp. nov	Valid	Poinar & Soreng	Eocene	Baltic amber	Russia ( Kaliningrad Oblast)	Grass belonging to the subfamily Arundinoideae. Genus includes new species E. balticus.
Luminidites amazonicus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Arecaceae.
Orthogonospermum [73]	Gen. et sp. nov	Valid	Smith et al.	Late Cretaceous (Maastrichtian)	Deccan Intertrappean Beds	India	A member of the family Zingiberaceae. Genus includes new species O. patanense.
Sabalites colaniae [74]	Sp. nov	In press	Song, Su, Do & Zhou in Song et al.	Oligocene	Dong Ho Formation	Vietnam	A member of the family Arecaceae belonging to the subfamily Coryphoideae.
Trichotomosulcites normalis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Arecaceae.

Commelinid monocot research

• A study on the evolutionary history of palms throughout the Cenozoic era, aiming to determine the impact of Cenozoic environmental changes on the diversification and biogeography of palms, is published by Lim et al. (2021). ^[75]
• Pollens of member of the family Poaceae preserving the same morphological characteristics as that of modern cereal grains are described from a sedimentary core from Lake Acıgöl (Turkey) by Andrieu-Ponel et al. (2021), who interpret this finding as indicative of the presence of proto-cereals in Anatolia since 2.3 million years ago, likely evolving from wild Poaceae as a result of trampling, nitrogen enrichment of soils and browsing by large mammal herds, and evaluate possible benefits from the availability of these proto-cereals for early hominins. ^[76]

Basal eudicots

Proteales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Platanocarpelia [77]	Gen. et sp. nov	Valid	Maslova, Kodrul & Kachkina	Late Cretaceous (Turonian)		Kazakhstan	A member of the family Platanaceae. Genus includes new species P. kyzyljarica.
Proteacidites pseudodehaanii [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Proteaceae.

Superasterids

Campanulid euasterids

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Pittosporum ettingshausenii [78]	Nom. nov	Valid	Doweld	Miocene		New Zealand	A species of Pittosporum ; a replacement name for Pittosporum elegans (Ettingshausen) W.R.B. Oliver (1950).
Xenopanax [78]	Gen. et comb. nov	Valid	Doweld	Eocene		Russia ( Kamchatka Krai)	A new genus for "Pittosporum" beringianum Chelebaeva & Akhmetiev (1983).

Lamiid euasterids

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Adina vastanenesis [79]	Sp. nov	Valid	Shukla et al.	Early Eocene	Cambay Shale Formation	India	A species of Adina .
Dicolpopollis? costatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Macoubea .
Dolichandra pacei [80]	Sp. nov	Valid	Franco, Brea & Cerdeño	Miocene (Santacrucian)	Mariño Formation	Argentina	A species of Dolichandra .
Fraxinoxylon beypazariense [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Oleaceae.
Fraxinus eoemarginata [81]	Sp. nov	In press	Mathewes, Archibald & Lundgren	Early Eocene	Quilchena site	Canada ( British Columbia)	A species of Fraxinus .
Kapgateophyllum [82]	Nom. nov	Valid	Deshmukh	Late Cretaceous (Maastrichtian) - early Eocene	Deccan Intertrappean Beds	India	A member of the family Acanthaceae; a replacement name for Acanthophyllum Ramteke & Kapgate (2014).
Ladakhipollenites campbellii [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Vitex .
Lymingtonia splendida [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Convolvulaceae.
Maryendressantha [83]	Gen. et sp. nov	Valid	Singh et al.	Early Eocene	Cambay amber	India	A member of the family Apocynaceae. Genus includes new species M. succinifera.
Perfotricolpites hexacolpatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Merremia .
Verrustephanoporites intraverrucosus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Apocynaceae.

Non euasterids

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Anacolosidites reticulatus [84]	Sp. nov	In press	Morley, Huang & Hoorn in Huang et al.	Middle and late Eocene	Yaw Formation		Pollen probably derived from plants belonging to the genus Ptychopetalum .
Camptotheca manchesterii [85]	Sp. nov	In press	Xie et al.	Late Miocene	Bangmai Formation	China	A species of Camptotheca .
Diospyros christensenii [86]	Sp. nov	Valid	Denk & Bouchal	Miocene		Denmark	A species of Diospyros .
Halesia mosbruggeri [87]	Sp. nov	Valid	Kvaček	Early Miocene	Most Basin	Czech Republic	A species of Halesia .
Loranthacites tabatingensis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Struthanthus .
Mecsekispermum [88]	Gen. et sp. nov	Valid	Hably & Erdei	Miocene (Burdigalian)	Feked Formation	Hungary	Possibly a member of the family Theaceae. Genus includes new species M. gordonioides.
Miranthus [89]	Gen. et 2 sp. nov	Valid	Friis, Crane	Late Cretaceous (Campanian-Maastrichtian)		Portugal	A Primulaceae genus. Genus includes new species M. elegans and M. kvacekii.
Nyssa nanningensis [90]	Sp. nov	In press	Xu & Jin in Xu et al.	Late Oligocene	Yongning Formation	China	A tupelo.
Paranyssa [63]	Nom. nov	Valid	Doweld	Paleocene		United States ( Montana)	A member of the family Nyssaceae; a replacement name for Browniea Manchester & Hickey (2007).
Parsonsidites? minibrenacii [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Amaranthaceae.
Ternstroemites klettwitzensis [91]	Sp. nov	Valid	Striegler	Miocene (Tortonian)	Rauno Formation	Germany	A member of the family Theaceae.

Superrosids

Fabids

Fabales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Albizia mahuadanrensis [92]	Sp. nov	In press	Hazra, Hazra & Khan in Hazra et al.	Pliocene	Rajdanda Formation	India	A species of Albizia .
Albizia palaeoprocera [92]	Sp. nov	In press	Hazra, Hazra & Khan in Hazra et al.	Pliocene	Rajdanda Formation	India	A species of Albizia.
Cercis zhangpuensis [93]	Sp. nov	In press	Wang et al.	Miocene	Fotan Group	China	A species of Cercis .
Cladrastis haominiae [94]	Sp. nov	In press	Jia et al.	Paleogene		China	A species of Cladrastis .
Enterolobiumoxylon [95]	Gen. et sp. nov	In press	Pérez-Lara, Estrada-Ruiz & Castañeda-Posadas	Eocene	El Bosque Formation	Mexico	Fossil wood of a member of the family Fabaceae. Genus includes new species E. triserial.
Gleditsioxylon fiambalense [96]	Sp. nov	In press	Baez	Miocene	Tambería Formation	Argentina	A member of Leguminosae.
Kingiodendron mexicanus [95]	Sp. nov	In press	Pérez-Lara, Estrada-Ruiz & Castañeda-Posadas	Eocene	El Bosque Formation	Mexico	Fossil wood of a member of the family Fabaceae.
Ladakhipollenites? pseudocolpiconstrictus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Fabaceae.
Leguminocarpum lottii [97]	Sp. nov	Valid	Li & Manchester in Li et al.	Early Eocene	Tepee Trail Formation	United States ( Wyoming)	A member of the family Fabaceae.
Leguminocarpum olmensis [98]	Sp. nov	Valid	Centeno-González et al.	Late Cretaceous (Campanian)	Olmos Formation	Mexico	A member of the family Fabaceae.
Neopapilionia [99]	Gen. et sp. nov	Valid	Hazra, Hazra & Khan in Hazra et al.	Pliocene	Rajdanda Formation	India	A member of the family Fabaceae. Genus includes new species N. indica.
Ormosia cyclocarpa [100]	Sp. nov	In press	Li et al.	Miocene		China	A species of Ormosia .
Peltophorum asiatica [101]	Sp. nov	In press	Hazra, Hazra & Khan in Hazra et al.	Pliocene	Rajdanda Formation	India	A species of Peltophorum .
Polyadopollenites minimus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Mimosa .
Salpinganthium [102]	Gen. et sp. nov	Valid	Poinar & Chambers	Burdigalian	Dominican amber	Dominican Republic	A member of the family Fabaceae belonging to the tribe Detarieae. Genus includes new species S. hispaniolanum.
Striatopollis crassitectatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Macrolobium .
Syncolporites foveolatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Swartzia .

Fagales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Eucarpinoxylon kayacikii [66]	Sp. nov	Valid	Akkemik	Middle Miocene	Kesmekaya Volcanics	Turkey	A member of the family Betulaceae.
Fagus dodgei [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Fagus species wood.
Lithocarpoxylon ashwillii [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Lithocarpoxylon species wood.
Morella absarokensis [104]	Comb nov	valid	(Wheeler, Scott, & Barghoorn) Wheeler & Manchester	Middle Eocene	Lamar River Formation	USA Wyoming	Moved from Myrica absarokensis (1978)
Morella scalariformis [104]	Comb nov	valid	(Kruse) Wheeler, Baas, & Manchester	Middle Eocene	Eden Valley Formation	USA Wyoming	Moved from Myrica scalariformis (1954)
Myricamentum [105]	Gen. et sp. nov	Valid	Wilde, Frankenhäuser & Lenz	Eocene	Eckfelder Maar	Germany	A catkin-like male inflorescence, probably of myricaceous affinity. Genus includes new species M. eckfeldensis.
Ostryoxylon gokceadaense [66]	Sp. nov	Valid	Akkemik	Middle Miocene	Kesmekaya Volcanics	Turkey	A member of the family Betulaceae.
Palaeocarpinus borealis [106]	Comb nov	Valid	(Heer) Correa & Manchester	Paleocene late Paleocene	Menat Formation	France Puy-de-Dôme	A Coryloideae species. First described as Anchietea borealis (1859). Moved from Atriplex borealis (1912)
Palaeocarpinus parva [106]	Sp. nov	Valid	Manchester & Correa	Middle Eocene	Clarno Formation	USA Oregon	A Coryloideae species.
Palaeocarpinus pterabaratra [106]	Sp. nov	Valid	Correa & Manchester	Paleocene Middle Paleocene	Fort Union Formation "unnamed lower member"	USA Wyoming	A Coryloideae species.
Palaeocarpinus pteravestigia [106]	Sp. nov	Valid	Correa & Manchester	Paleocene Middle Paleocene	Fort Union Formation "unnamed lower member"	USA Wyoming	A Coryloideae species.
Palaeocarya indica [107]	Sp. nov	Valid	Hazra, Hazra & Khan in Hazra et al.	Pliocene	Rajdanda Formation	India	A member of the family Juglandaceae.
Paralnoxylon [63]	Nom. nov	Valid	Doweld	Paleocene		United Kingdom	A member of the family Betulaceae; a replacement name for Cantia Stopes (1915).
Quercoxylon yaltirikii [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Fagaceae.

Fagalean research

• A study on wood anatomy in extant and fossil members of Fagales is published by Wheeler, Baas & Manchester who transfer two Eocene species from Myrica to Morella. ^[104]

Malpighiales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Euphorbiotheca deccanensis [108]	Sp. nov	In press	Reback et al.	Late Cretaceous (Maastrichtian)	Deccan Intertrappean Beds	India	A member of the family Euphorbiaceae.
Passiflora appalachiana [109]	Sp. nov	Valid	Hermsen	Pliocene	Gray Fossil Site	United States ( Tennessee)	A species of Passiflora .
Passifloriidites [4]	Gen. et sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Passifloraceae. Genus includes new species P. pseudoperculatus.
Populoxylon sebenense [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Salicaceae.
Salicoxylon galatianum [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Salicaceae.
Verrustephanoporites circularis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Mascagnia .

Oxalidales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Tropidogyne euthystyla [110]	Sp. nov	Valid	Poinar, Chambers & Vega	Cretaceous	Burmese amber	Myanmar	A possible member of Cunoniaceae.

Rosales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Alloceltidoxylon [63]	Nom. nov	Valid	Doweld	Eocene	Clarno Formation	United States ( Oregon)	A flowering plant with possible affinities with urticalean rosids; a replacement name for Scottoxylon Wheeler & Manchester (2002).
Celtis popsii [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Celtis species wood.
Crataegoxylon sibiricum [47]	Sp. nov	Valid	Dolezych, LePage & Williams	Oligocene (Chattian)	Korlikov Formation	Russia ( Tomsk Oblast)
Morus asiatica [111]	Sp. nov	In press	Patel, Rana & Khan in Patel et al.	Early Eocene		India	A species of Morus .
Prunoidoxylon prunoides [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Rosaceae.
Psilatriporites minimus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Celtis .
Pyracantha pseudococcinea [91]	Sp. nov	Valid	Striegler	Miocene (Tortonian)	Rauno Formation	Germany	A species of Pyracantha .
Ulmoxylon kasapligilii [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Ulmaceae.
Ventilago tibetensis [112]	Sp. nov	Valid	Del Rio et al.	Middle Eocene		China	A species of Ventilago .
Zelkovoxylon crystalliferum [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Ulmaceae.

Malvids

Malvales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Bombacacidites hooghiemstrae [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Malvaceae.
Craigia lincangensis [113]	Sp. nov	In press	Wang & Xie in Wang et al.	Late Miocene		China	A species of Craigia
Dipterocapus fotanensis [114]	Sp. nov	Valid	Chen et al.	Miocene		China	A species of Dipterocarpus
Discoidites angulosus [84]	Sp. nov	In press	Huang, Morley & Hoorn in Huang et al.	Late Eocene	Yaw Formation	Myanmar	Pollen probably derived from plants belonging to the genus Brownlowia .
Thymelipollis amazonicus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Thymelaeaceae.
Tilia asiatica [115]	Sp. nov	In press	Jia & Nam in Jia et al.	Middle Miocene	Pohang Basin	South Korea	A species of Tilia
Wataria kvacekii [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Wataria species wood.

Myrtales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Florschuetzia impostora [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen, possibly of a member of the family Lythraceae.
Lazarocardenasoxylon [116]	Gen. et sp. nov	In press	Estrada-Ruiz & Martínez-Cabrera in Estrada-Ruiz, Martínez-Cabrera & García-Hernández	Late Cretaceous	San Carlos Formation	Mexico	Possibly a member of the family Myrtaceae. Genus includes new species L. aldamense.
Lythrum portugalliense [117]	Sp. nov	In press	Vieira et al.	Pliocene (Piacenzian)		Portugal	A species of Lythrum .
Syzygium guipingensis [118]	Sp. nov	In press	Li et al.	Miocene	Erzitang Formation	China	A species of Syzygium .
Terminalioxylon mozambicense [119]	Sp. nov	Valid	Bamford & Pickford	Probably late Eocene		Mozambique	A member of the family Combretaceae.
Trapa sanyingensis [120]	Sp. nov	In press	Aung et al.	Late Pliocene	Sanying Formation	China	A water caltrop.
Verrutricolporites pusillus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen, possibly of a member of the family Lythraceae.
Xystonia [121]	Gen. et sp. nov	Valid	Carvalho et al.	Paleocene	Bogotá Formation	Colombia	A member of the family Melastomataceae. Genus includes new species X. simonae.

Sapindales

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Aceroxylon aceroides [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Sapindaceae.
Ampelorhiza [122]	Gen. et sp. nov	Valid	Jud et al.	Early Miocene	Cucaracha Formation	Panama	A member of the family Sapindaceae belonging to the subfamily Sapindoideae and the tribe Paullinieae. Genus includes new species A. heteroxylon.
Anacardium gassonii [123]	Sp. nov	Valid	Rodríguez-Reyes, Estrada-Ruiz & Terrazas in Rodríguez-Reyes et al.	Oligocene-Miocene		Panama	A species of Anacardium .
Atalantioxylon thanobolensis [124]	Sp. nov	Valid	Soomro et al.	Miocene	Manchar Formation	Pakistan	Fossil wood of a member of the family Rutaceae.
Melia santangensis [125]	Sp. nov	Valid	Liu, Xu & Jin in Liu et al.	Late Oligocene	Yongning Formation	China	A species of Melia .
Pistacia terrazasae [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Pistacia species wood.
Proteacidites poriscabratus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Sapindaceae.
Sorindeioxylon [119]	Gen. et sp. nov	Valid	Bamford & Pickford	Probably late Eocene	Mazamba Formation	Mozambique	A member of the family Anacardiaceae. Genus includes new species S. gorongosense.
Syncolporites tenuicolpatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the genus Serjania .
Tetradium nanningense [126]	Sp. nov	Valid	Huang et al.	Late Oligocene	Yongning Formation	China	A species of Tetradium .
Verrutricolporites simplex [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Pollen of a member of the family Simaroubaceae.

Non eurosid superrosids

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Hamamelidoxylon crystalliferum [103]	Sp nov	Valid	Wheeler & Manchester	Late Eocene	John Day Formation	USA Oregon	A Hamamelidoxylon species wood.
Liquidambar guipingensis [127]	Sp. nov	In press	Huang et al.	Miocene	Erzitang Formation	China	A species of Liquidambar .
Liquidambaroxylon efeae [66]	Sp. nov	Valid	Akkemik	Early Miocene	Hançili Formation	Turkey	A member of the family Altingiaceae.
Obirafructus [128]	Gen. et sp. nov	Valid	Kajita & Nishida in Kajita, Suzuki & Nishida	Late Cretaceous (Coniacian–Santonian)	Haborogawa Formation	Japan	A member of Saxifragales of uncertain phylogenetic placement. Genus includes new species O. kokubunii.
Paleoaltingia [129]	Gen. nov	Valid	Lai et al.	Late Cretaceous (Turonian)		United States ( New Jersey)	A member of Altingiaceae. Genus includes P. ovum-dinosauri and P. polyodonta.

Other angiosperms

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Araliaephyllum vittenburgii [130]	Sp. nov	Valid	Golovneva & Volynets in Golovneva et al.	Early Cretaceous (Albian)	Galenki Formation	Russia ( Primorsky Krai)	A flowering plant of uncertain phylogenetic placement.
Baderadea [131]	Gen. et sp. nov	Valid	Pessoa, Ribeiro & Jud	Early Cretaceous (Aptian)	Crato Formation	Brazil	A herbaceous eudicot similar to some members of Ranunculales. Genus includes new species B. pinnatissecta.
Byttneripollis rugulatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Dilcherifructus [132]	Gen. et sp. nov	Valid	Wang	Middle Jurassic	Simón Formation	Mexico	Possibly a fruit of an early flowering plant. Genus includes new species D. mexicana.
Farabeipollis deccanensis [133]	Sp. nov	Valid	Sonkusare, Samant & Mohabey	Late Cretaceous (Maastrichtian)	Deccan Intertrappean Beds	India	Pollen of a flowering plant of uncertain affinity.
Florigerminis [134]	Gen. et sp. nov	In press	Cui et al.	Middle-Late Jurassic	Jiulongshan Formation	China	A possible flower bud. The type species is F. jurassica. First announced online 2021, Final article published 2022.	Florigerminis jurassica
Gansufructus [135]	Gen. et sp. nov	In press	Du in Du et al.	Early Cretaceous (late Aptian-early Albian)	Zhonggou Formation	China	A eudicot of uncertain phylogenetic placement. Genus includes new species G. saligna.
Inaperturopollenites tectatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Ladakhipollenites? corvattatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Ladakhipollenites? endoporatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Ladakhipollenites nanus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Ladakhipollenites? sphaericus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Margocolporites bilinearis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Margocolporites incertus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Multiporopollenites intermedius [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Nigericolpites [136]	Nom. nov	Valid	Hernández	Late Cretaceous (Maastrichtian)		Nigeria	Pollen of a flowering plant; a replacement name for Clavatricolpites Hoeken-Klinkenberg (1964).
Psilaperiporites circinatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Psilaperiporites depressus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Psilastephanocolporites ectoporatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Psilastephanocolporites pseudomarinamensis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Ranunculacidites pontoreticulatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retibrevitricolpites pseudoretibolus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retiperiporites retiporatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retistephanocolpites liberalis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retistephanocolporites loxocolpatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitrescolpites benjaminensis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitrescolpites brevicolpatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitrescolpites grossus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitrescolpites kriptoporus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitrescolpites marginatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitriporites discretus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Retitriporites sifonis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites apertus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites? colpiverrucosus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites crassinexinicus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites crassitectatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites grossomurus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites guttatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites lolongatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites protoguttatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites pseudocrassopolaris [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites pseudopilatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites pseudoscabratus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites quantulus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Rhoipites vilis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Tetracolporopollenites nanus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.
Tetracolporopollenites xatanawensis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil pollen.

Other plants

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Adelocladoxis [137]	Gen. et sp. nov	Valid	Durieux et al.	Devonian (Emsian)	Battery Point Formation	Canada ( Quebec)	A member of Cladoxylopsida. Genus includes new species A. praecox.
Baragwanathia brevifolioides [138]	Nom. nov	Valid	Kraft & Kvaček	Silurian (Přídolí)	Požáry Formation	Czech Republic	A member of the family Drepanophycaceae; a replacement name for Baragwanathia brevifolia Kraft & Kvaček (2017).
Camarozonosporites fossulatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Lycopodiaceae.
Cingulatisporites cristatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the genus Selaginella .
Cingulatisporites matisiensis [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the genus Phaeoceros .
Closterium mosbruggeri [139]	Sp. nov	Valid	Ivanov & Belkinova	Miocene (Serravallian)		Bulgaria	A green alga, a species of Closterium .
Colpodexylon mergae [140]	Sp. nov	In press	Harris et al.	Devonian (Famennian)	Witpoort Formation	South Africa	A member of Lycopsida.
Colpodexylon pullumpedes [140]	Sp. nov	In press	Harris et al.	Devonian (Famennian)	Witpoort Formation	South Africa	A member of Lycopsida.
Cordaabaxicutis martii [141]	Sp. nov	Valid	Šimůnek & Lojka	Carboniferous (Pennsylvanian)	Kladno Formation	Czech Republic	Cordaitalean cuticles.
Cordaabaxicutis papillosus [141]	Sp. nov	Valid	Šimůnek & Lojka	Carboniferous (Pennsylvanian)	Kladno Formation	Czech Republic	Cordaitalean cuticles.
Cordaadaxicutis raristomatus [141]	Sp. nov	Valid	Šimůnek & Lojka	Carboniferous (Pennsylvanian)	Kladno Formation	Czech Republic	Cordaitalean cuticles.
Cynodontium luthii [142]	Sp. nov	Valid	Bippus, Rothwell & Stockey	Late Cretaceous		United States ( Alaska)	A moss belonging to the family Rhabdoweisiaceae, a species of Cynodontium .
Dayvaultia [143]	Gen. et sp. nov	Valid	Manchester et al.	Late Jurassic	Morrison Formation	United States ( Utah)	A seed-bearing structure of gnetalean affinity. Genus includes new species D. tetragona.
Distefanopolia [144]	Gen. et comb. nov	Valid	Barattolo, Romano & Conrad	Late Triassic and possibly Early Jurassic		Austria Czech Republic Germany Greece Italy Oman Slovakia	A green alga belonging to the group Dasycladales and the family Bornetelleae. Genus includes "Heteroporella" micropora Di Stefano & Senowbari-Daryan (1985), "Heteroporella" macropora Di Stefano, 1981 ex Di Stefano & Senowbari-Daryan (1985), "Chinianella" zanklii Ott (1967), "Chinianella" crosii Ott (1968) and "Heteroporella" carpatica Bystrický (1967).
Distichophytum mogilatii [145]	Sp. nov	Valid	Naugolnykh	Devonian		Russia ( Krasnoyarsk Krai)	A member of the family Zosterophyllaceae.
Dragastanella [146]	Gen. et sp. et comb. nov	Valid	Barattolo, Bucur & Marian	Early Cretaceous		Italy Romania Spain	A green alga belonging to the group Dasycladales. Genus includes new species D. transylvanica, as well as "Zittelina" hispanica Masse, Arias & Vilas (1993), "Zittelina" massei Bucur, Granier & Săsăran (2010) and "Triploporella" matesina Barattolo (1980).
Elandia [147]	Gen. et sp. nov	Valid	Gess & Prestianni	Devonian (Lochkovian?)	Baviaanskloof Formation	South Africa	An early polysporangiophyte. Genus includes new species E. itshoba.
Flabellopteris [148]	Gen. et sp. nov	In press	Gess & Prestianni	Devonian (Famennian)	Witpoort Formation	South Africa	A fern-like plant of uncertain affinities. Genus includes new species F. lococannensis.
Foraminisporis connexus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the genus Anthoceros .
Frullania kachinensis [149]	Sp. nov	Valid	Li et al.	Cretaceous	Burmese amber	Myanmar	A liverwort, a species of Frullania .
Frullania palaeoafricana [150]	Sp. nov	In press	Bouju et al.	Miocene	Ethiopian amber	Ethiopia	A liverwort, a species of Frullania.
Frullania shewanensis [150]	Sp. nov	In press	Bouju et al.	Miocene	Ethiopian amber	Ethiopia	A liverwort, a species of Frullania.
Gilboaphyton fuyunensis [151]	Sp. nov	In press	Liu et al.	Late Devonian	Kaxiweng Formation	China	A member of Protolepidodendrales.
Guazia [152]	Gen. et sp. nov	In press	Wang et al.	Late Devonian	Wutong Formation	China	A seed plant of uncertain phylogenetic placement. Genus includes new species G. dongzhiensis.
Ixostrobus daohugouensis [153]	Sp. nov	In press	Na & Sun in Na et al.	Middle Jurassic		China	A member of Czekanowskiales.
Kenrickia [154]	Gen. et sp. nov	In press	Toledo et al.	Devonian (Emsian)	Battery Point Formation	Canada	An early euphyllophyte belonging to the group Radiatopses. Genus includes new species K. bivena.
Krommia [147]	Gen. et sp. nov	Valid	Gess & Prestianni	Devonian (Lochkovian?)	Baviaanskloof Formation	Brazil South Africa	An early polysporangiophyte. Genus includes new species K. parvapila.
Lejeunea abyssinicoides [150]	Sp. nov	In press	Bouju et al.	Miocene	Ethiopian amber	Ethiopia	A liverwort, a species of Lejeunea .
Lycaugea [155]	Gen. et sp. nov	Valid	Meyer-Berthaud, Decombeix & Blanchard	Devonian (Famennian)		Australia	A lycopsid. Genus includes new species L. edieae.
Lycopodiumsporites amazonicus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Spores of a member of the family Lycopodiaceae.
Melvillipteris sonidia [156]	Sp. nov	In press	Bai et al.	Devonian (probably Famennian)		China	A member of Rhacophytales.
Mesochara adobensis [157]	Sp. nov	In press	De Sosa Tomas et al.	Early Cretaceous	Los Adobes Formation	Argentina	A member of Charophyta.
Mesochara dobrogeica [158]	Sp. nov	In press	Sanjuan et al.	Early Cretaceous (Berriasian)		Romania	A member of Charophyta.
Mixoxylon [159]	Gen. et sp. nov	In press	Chernomorets & Sakala	Early Cretaceous (Albian)	Whisky Bay Formation	Antarctica	A homoxylous wood of uncertain systematic affinities. Genus includes new species M. australe.
Mtshaelo [147]	Gen. et sp. nov	Valid	Gess & Prestianni	Devonian (Lochkovian?)	Baviaanskloof Formation	South Africa	An early polysporangiophyte. Genus includes new species M. kougaensis.
Omniastrobus [160]	Gen. et sp. nov	Valid	Bonacorsi et al.	Devonian (Emsian)	Campbellton Formation	Canada	A lycophyte. Genus includes new species O. dawsonii.
Palaeonitella trifurcata [161]	Sp. nov	Valid	Martín-Closas et al.	Early Cretaceous (Barremian–Aptian)		Spain	A member of Charophyta belonging to the family Characeae.
Paratingia wuhaia [162]	Sp. nov	Valid	Wang et al.	Permian (Asselian)	Taiyuan Formation	China	A progymnosperm belonging to the group Noeggerathiales and the family Tingiostachyaceae.
Permotheca? musaformis [163]	Sp. nov	Valid	Foraponova & Karasev	Permian		Russia	A pteridosperm.
Radula heinrichsii [164]	Sp. nov	In press	Feldberg et al.	Cretaceous	Burmese amber	Myanmar	A liverwort, a species of Radula .
Rehamnia [165]	Gen. et sp. nov	In press	Oukassou & Naugolnykh	Late Devonian		Morocco	A member of Lycopodiophyta of uncertain phylogenetic placement. Genus includes new species R. michardis.
Ricciopsis baojishanensis [166]	Sp. nov	In press	Han & Yan in Han et al.	Late Triassic	Nanying'er Formation	China	A liverwort.
Salopella laidae [167]	Sp. nov	Valid	McSweeney, Shimeta & Buckeridge	Devonian (Pragian)	Yea Formation	Australia	An early land plant of uncertain affinities.
Skyttegaardia [168]	Gen. et sp. nov	In press	Friis, Crane & Pedersen	Early Cretaceous (Berriasian)		Denmark	A plant of uncertain phylogenetic placement, possibly close to cycads. Genus includes new species S. galtieri.
Thysananthus aethiopicus [150]	Sp. nov	In press	Bouju et al.	Miocene	Ethiopian amber	Ethiopia	A liverwort belonging to the family Lejeuneaceae.
Velascoa [169]	Gen. et sp. nov	Junior homonym	Flores Barragan, Velasco de León & Ortega Chavez	Permian	Matzitzi Formation	Mexico	Fossil leaves of a plant of uncertain phylogenetic placement, with a morphology similar to Ginkgophyta. Genus includes new species V. pueblensis. The generic name is preoccupied by Velascoa Calderón & Rzedowski (1997).
Vitinellopsis [170]	Gen. et sp. nov	In press	Vachard, Bucur & Munnecke	Silurian		Sweden	A green alga belonging to the group Bryopsidales. Genus includes new species V. gotlandica.
Zosterophyllum confertum [171]	Sp. nov	In press	Gossmann et al.	Early Devonian		Germany	A zosterophyll.

Palynology

Name	Novelty	Status	Authors	Age	Type locality	Location	Notes	Images
Camarozonosporites trilobatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	A spore of lycopodialean affinity.
Echinatisporis infantulus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Erlansonisporites duwaensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Erlansonisporites exquisitus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Erlansonisporites perbellus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Erlansonisporites textilis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Ginkgomonocolpites [173]	Nom. nov	Valid	Hernández	Paleogene		India	A gymnosperm pollen; a replacement name for Psilamonocolpites Mathur (1966).
Hamulatisporis bareanus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Henrisporites longibaculiformis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Horstisporites comitus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Horstisporites denticulatus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Horstisporites subtilis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Horstisporites tarimensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Hughesisporites reticulatus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Hughesisporites unicus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Ischyosporites dubius [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	A spore of dicksoniaceous affinity.
Ischyosporites granulatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Laevigatosporites indigestus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Luntaispora [172]	Gen. et sp. nov	Valid	Li et al.	Mesozoic		China	Genus includes new species L. laevigata.
Microfoveolatosporis simplex [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Minerisporites tarimensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Minerisporites triangularis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Narkisporites conicus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Narkisporites densibaculatus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Narkisporites densiconicus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Narkisporites tarimensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Neoraistrickia dubia [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Noniasporites triassicus [174]	Sp. nov	Valid	Ghosh et al.	Early Triassic	Panchet Formation	India	A megaspore.
Otynisporites tarimensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Polypodiisporites discretus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Psilatriletes delicatus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Punctatosporites latrubessei [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Rotverrusporites amazonicus [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.
Stellibacutriletes [172]	Gen. et 4 sp. nov	Valid	Li et al.	Mesozoic		China	Genus includes new species S. capillaris, S. gracilis, S. rarus and S. solidus.
Striatriletes inconspicuus [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Tarimispora [172]	Gen. et 2 sp. nov	Valid	Li et al.	Mesozoic		China	Genus includes new species T. auriculata and T. perfecta.
Tricristatispora trilobata [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Tricristatispora yingmailensis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Trileites plicatilis [172]	Sp. nov	Valid	Li et al.	Mesozoic		China
Verrucatotriletes pseudovirueloides [4]	Sp. nov	Valid	D'Apolito, Jaramillo & Harrington	Miocene	Solimões Formation	Brazil	Fossil spores.

Palynological research

• Strother & Foster (2021) describe an assemblage of fossil spores from the Ordovician (Tremadocian) of Australia, representing a morphology that was intermediate morphology between confirmed land plant spores and earlier forms of uncertain phylogenetic placement, and evaluate the implications of these fossils for the knowledge of the evolution of land plants from their algal ancestors. ^[175]
• A study on the fossil pollen record from New Zealand, dating from 100 million years ago to the present, is published by Prebble et al. (2021), who report evidence indicating that Cretaceous diversification was closely followed by an increase in flowering plants frequency, but their maximum frequency did not occur until the Eocene. ^[176]
• A study on changes of abundance in spores and pollen record from the Danish Basin, and on their implications for the knowledge of the impact of the Triassic–Jurassic extinction event on land plants, is published by Lindström (2021). ^[177]
• A study on the vegetation history in the southwestern Balkans, as indicated by pollen from the sedimentary record in the Lake Ohrid extending to 1.36 million years ago, is published by Donders et al. (2021). ^[178]

Research

• A study on changes of the morphological complexity of reproductive structures of land plants throughout their evolutionary history, based on data from fossil and extant land plants, is published by Leslie, Simpson & Mander (2021). ^[179]
• Revision of Silurian (Wenlock to Přídolí) assemblages of polysporangiophytes with dispersed spores and cryptospores, aiming to determine the relationship between Silurian plant evolution and climate changes linked with perturbations of the global carbon cycle, is published by Pšenička et al. (2021). ^[180]
• Reconstruction of the structure and development of the rooting system of Asteroxylon mackiei is presented by Hetherington et al. (2021). ^[181]
• A study on factors influencing the extent of arboreal vegetation during the Late Paleozoic icehouse is published by Matthaeus et al. (2021), who interpret their findings as indicating that Pangaea could have supported widespread arboreal plant growth and forest cover based on leaf water constraints, but the forest extent was restricted because of impact of freezing on plants, and estimate that contracting forest cover increased net global surface runoff by up to 6.1%. ^[182]
• Description of the reproductive organs of the lycopsids from the Upper Devonian Wutong Formation (China), and a study on the ability of the sporophyll units for wind dispersal, is published by Zhou et al. (2021), who name new form species Lepidophylloides longshanensis and Lepidophylloides changxingensis. ^[183]
• An exceptionally well preserved Brasilodendron -like lycopsid forest containing over 150 upright stumps is described from an early Permian postglacial landscape of western Gondwana (Paraná Basin, Brazil) by Mottin et al. (2021). ^[184]
• A study on the anatomy of Stigmaria asiatica is published by Chen et al. (2021). ^[185]
• Stump casts of Sigillaria , preserving traces of internal anatomy, are described from the earliest Permian Wuda Tuff (China) by D'Antonio et al. (2021). ^[186]
• A study aiming to determine probable causes of the world-wide proliferation of members of Isoetales, particularly Pleuromeia , during and in the aftermath of the Permian–Triassic extinction event, and evaluating the implications of this proliferation for the knowledge of environmental stresses during and in the aftermath of this extinction event, is published by Looy, van Konijnenburg-van Cittert & Duijnstee (2021). ^[187]
• New fossil material of Saportaea salisburioides, providing new information on leaf morphology and growth of this plant, is described from the Permian Umm Irna Formation (Jordan) by Kerp et al. (2021), who interpret their findings as indicating that Saportaea grandifolia and Baiera virginiana were synonyms of S. salisburioides, and possibly indicating that the fructification belonging to the genus Nystroemia is a part of Saportaea. ^[188]
• Description of Geinitzia reichenbachii from its gross morphology to the cellular scale, and a study on the likely ecology of this conifer, is published by Moreau et al. (2021). ^[189]
• A study on the evolutionary history of the family Cycadaceae, based on genomic data and fossil record, is published by Liu et al. (2021). ^[190]
• Well-preserved recurved cupules of seed plants are described from the Lower Cretaceous of China by Shi et al. (2021), who interpret the structure of these cupules as consistent with the recurved form and development of the second integument in the bitegmic anatropous ovules of flowering plants, and evaluate the implications of these fossils for the knowledge of the origin of the flowering plants. ^[191]
• Taxonomically diverse flora from the Seafood Salad locality, found ~65 m below the Cretaceous-Paleogene boundary in the Hell Creek Formation (Montana, United States), is described by Wilson, Wilson Mantilla & Strӧmberg (2021), who study the affinities of plants of this locality and compare them with other Late Cretaceous floras of the Western Interior. ^[192]
• A study on the timing of the origin of the flowering plants, based on data from fossil record and from the diversity of extant members of this group, is published by Silvestro et al. (2021), who interpret their findings as indicating that several flowering plant families originated in the Jurassic. ^[193]
• A study on the diversity of insect damage types in fossil plants from the Cretaceous (Albian to Cenomanian) Dakota Formation (United States), evaluating their implications for the knowledge of the early evolution of angiosperm florivory and associated pollination, is published by Xiao et al. (2021). ^[194]
• New fossil material of Callianthus dilae is described from the Lower Cretaceous Yixian Formation (China) by Wang et al. (2021), who reconstruct the whole plant of Callianthus, interpreting it as an aquatic flowering plant. ^[195]
• A study on the anatomy of the epidermal features of the floating leaves of Quereuxia angulata from the Upper Cretaceous Yong'ancun Formation (China) is published by Liang et al. (2021). ^[196]
• A study on plant extinction and ecological change in tropical forests resulting from the Cretaceous–Paleogene extinction event, based on data from fossil pollen and leaves from Colombia, is published by Carvalho et al. (2021), who report evidence indicative of a long interval of low plant diversity in the Neotropics after the end-Cretaceous extinction, and the emergence of forests with a structure resembling modern Neotropical rainforests, with a closed canopy and multistratal structure dominated by flowering plants, during the Paleocene. ^[197]
• A study on the impact of the mid-Eocene greenhouse warming event on floras from southernmost South America is published by Fernández et al. (2021). ^[198]
• Evidence from middle Eocene-middle Miocene tuffaceous deposits of central and northern Patagonia, indicating that soils, vegetation, insects and mammal herbivores began to record diverse traits related to the presence of grasslands with mosaic vegetation since middle Eocene, is presented by Bellosi et al. (2021). ^[199]
• A study on Middle Miocene microfloral assemblages from ten localities in the Madrid Basin (Spain), providing evidence of prevalence of open habitats with grass-dominated, savannah-like vegetation under a warm and semi-arid climatic regime in the Iberian Peninsula in the Middle Miocene, is published by Casas-Gallego et al. (2021). ^[200]
• Crump et al. (2021) present a record of vegetation from the Last Interglacial based on ancient DNA from lake sediment from the Baffin Island (Canada), and report evidence of major ecosystem changes in the Arctic in response to warmth, including a ~400 km northward range shift of dwarf birch relative to today. ^[201]

Deaths

• Alan Graham (1934–2021), passed away on 8 July 2021. Graham earned his PhD in 1962 under the guidance of Chester A. Arnold, and was noted for a career studying the Cenozoic paleobotany of the Caribbean and Central America. ^[202]

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