Notes on Fossil Leaves

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Fossil material of plants can be difficult to identify as to species, genus, or larger taxonomic units, as usually what is found is individual parts of plants, such as wood, leaves, flowers, fruits, seeds or pollen, and these are often insufficient for identification, particularly for older material which is less closely related to modern material, and may be less well preserved. Consequently, and as fossils of one plant part often cannot be unambigiously associated with those of another plant part, palaeobotanists use form genera to classify parts of plants of uncertain taxonomic position. The suffix -phyllum is often used in generic names, indicating a similarity with the leaves of the modern genus whose name is combined with that suffix. It cannot be assumed that the fossil leaves represent a species particularly close to the modern genus.

I suspect that the genera and species of fossil malvaceous leaves listed below are far from an exhaustive coverage. Furthermore fossil leaves may be ascribed to a living genus, and this will be even harder to track down.

The identification of fossil leaves is particularly difficult, and historically many errors have been made. Early identifications were often made on the basis of the overall shape of the leaf, which is not a reliable guide. (A great many leaf fossils, including some now thought to be malvaceous, were originally identified as Ficus.) More modern work takes account of the details of the venation, the architecture of the stomata, and the nature of the indumentum. However as even living plants can be difficult to identify in the absence of flowers or fruits I retain some skepticism.

An index of fossil leaf names [1, 2, 3, 4], with a terminal date depending on volume of some time in the 21st century, has been used as a source of names in known malvaceous and putatively malvaceaous genera. Not all volumes were available for examination. (Some of the names, e.g. Berrya racemosa and Buettneria perplexans, in that work turn out to be applicable to other plant parts, rather than to leaves.)


Abutilon eakinii Hollick is recorded from the Tertiary of Alaska [5]. This was collected from the Alaska Peninsula, where the Tertiary rocks are of Eocene or Miocene age. Alaska is a surprising location to find find a putative Abutilon; my expectation is that the genus is too young to be found in Palaeocene and Eocene rocks, and too cold intolerant to be present in Alaska in younger rocks. However the specimen has secondary and tertiary veins terminating at the apices of teeth, which fits Malvaceae better than the offered alternative of Populus.

Abutilon acerites Massalongo.

Abutilon sp.


A paper [6] describing a late Eocene flora from northern Bohemia classifies Acherniaephyllum Rásky as belonging to Malvaceae s.l. The type of this genus is Acherniaephyllum kraeuseli Rásky, but the only recognised species is Acherniaephyllum hydrarchos (Unger) Hably, based on the earlier Ficus hydrarchos Unger. Leaves assigned to this species have been previously named Passifloriaephyllum kraeuseli Rásky, Cinnamonum rotundifolium Principi, Dombeyopsis dubia Principi, Ficus populina Heer, Magnolia ovalifolia Principi and Sterculia variabilis Saporta [7]. Other specimens of Sterculia variabilis are referred to Sloanea olmediaefolia (Elaeocarpaceae) [7]. The Eocene material,treated as cf Acherniaephyllum hydrarchos [6], was originally figured as Populus mutabilis Heer.

Acherniaephyllum hydrarchos is recorded from the Oligocene of Italy [7, 8] and Hungary [9].


Actinovena Tanai is a genus introduced for leaves believed to be tiliaceous (perhaps related to Pentace, i.e. brownlowioid). These leaves are nearly symmetrical, ovate to narrowly ovate, 3-nerved and rounded at the base, with entire margins and acuminate apices. The assignment to Tiliaceae is based on details of the secondary and subsequent venation.

The name refers to the actinodromous nature of the primary venation.

Actinovena ishikariensis Tanai is recorded in the Late Eocene Ikushunbetsu Fm of Hokkaido. [10]


Leaf fossils described as Apeiba improvisa are recorded from the Eocene of Wyoming.


Apeibopsis deloesi Heer, Apeibopsis fischeri Heer and Apeibopsis gaudini Heer are listed without comment for the Lower Oligocene of Liguria, Italy. [8]

Some leaves described as Apeibopsis represent Nyssa alata (Ward) R.W.Brown. [11]


Astrapaeites pumicosus Langeron is described from a leaf fossil from the Eocene (Thanetian) of France (Sezanne).


Berryophyllum Jones & Dilcher is an extinct fagaceous genus. The name is not derived, as might be suspected, from the brownlowioid genus Berrya, but from the name of the American palaeobotanist Edward Wilber Berry, i.e. "Berry's leaf", not "Berrya-leaf".


Bombaciphyllum is a form genus representing fossil leaves similar to those of the genus Bombax (or bombacoids in general). However fossil leaves perceived to similar to living Bombax are more commonly placed in that genus, than in Bombaciphyllum.

Bombacophyllum argillaceum is recorded from the Upper Cretaceous. (I am treating Bombacophyllum as an orthographic variant of Bombaciphyllum.) It may well be the same as Bombax argillaceum.

Bombaciphyllum opacum Engelhardt is recorded from the Tertiary of Chile.


Leaf fossils described as Bombacites jacksoniana are recorded from Eocene (Jackson Fm.) of Texas.

Bombacites ceibaoides Ball is recorded from the Eocene (Wilcox Fm.) of Texas and Bombacites formosus Berry from the Eocene (Wilcox Fm.) of Louisiana and Texas

Bombacites orientalis Lakhanpal is described from the Kopili stage of the Middle Eocene of Assam [12]. It is also recorded from the Upper Palaeocene (Tura Fm.) of Assam and Meghalaya.


Bombaxlongifolium and “Bombaxprocaccinii, from the Lower Oligocene of Liguria, Italy, are identified as lauraceous [8]. Bombax oblongifolium is listed without comment in the same work.

Leaves (leaflets) assigned to Bombax sturtii Ettingshausen and Bombax mitchellii Ettingshausen are recorded from the Tertiary of Australia. Leaves assigned to Bombax chorisioides Friedr. and Bombax lepsii Engelhardt are recorded from the Lower Tertiary of Germany


Leaf material ascribed to Brachychiton is reported from the Tertiary of Australia [13] and New Zealand. Brachychiton-like leaves have also been found in South America.


Byttneria iizimae Tanai is recorded from the Late Eocene Ikushunbetsu Fm of Hokkaido. It is similar to Byttneriophyllum tiliaeophyllum, from which it differs in its more nearly symmetric base, and in its sparser tertiary venation. [10] (Buettneria is a variant spelling of Byttneria.)

Buettneria blackii Ball is recorded from Eocene (Wilcox Fm.) of Texas, and Buettneria jacksoniana Berry from the Eocene (Jackson Fm.) of Texas. Buettneria asterotrichiformis and Buettneria lanceolatifolia are recorded from the Eocene or Miocene of Argentina.


Byttneriophyllum is a form genus representing fossil leaves similar to those of the genus Byttneria.[a, 14]

Byttneriophyllum tiliaeofolium is recorded from the Neogene of Slovenia, the Miocene of Poland, the Middle Miocene of Romania, and also from East Asia. It has been confused with Dombeyopsis lobata, from which it differs in possessing a strongly asymetrical leaf base. Whether this represents a malvaceous plant is not clear; on the one hand it is associated with the malpighiaceous fruit morphospecies Banisteriaecarpum giganteum, on the other hand it has been associated with the pollen morphospecies Intratriporopollenites instructus, which is probably tilioid.


Byttneriopsis daphnogenes (Ettingshausen) Kvaček & Wilde, Byttneriopsis steuerii (Engelhardt) Kvaček & Wilde and Byttneriopsis spiegelii (Engelhardt) Kvaček & Wilde [15]


Leaf material ascribed to Cola, under the name Cola amharensis, is recorded from the Late Oligocene of northwestern Ethiopia.



Dombeya novi-mundi Hickey is described from early Tertiary Golden Valley Fm. of western North Dakota.


Worobiec, Worobiec and Kvaček [8] interpret Dombeyopsis as a monotypic genus, with the single morphospecies Dombeyopsis lobata Unger. Dombeyopsis lobata co-occurs with the fruits of Craigia bronnii, and is thought to represent the foliage of that species.

Dombeyopsis lobata has many synonyms, including Bombax dechenii (Weber) Friedrich, Dombeyopsis dechenii C.Weber, Dombeyopsis grandifolia Unger pro parte, Dombeyopsis pentagonalis Weber, Dombeyopsis sidaefolia Unger, Dombeyopsis tridens Ludwig, Firmiana germanica Menzel, Gothan & Sapper, Firmiana lobata (Unger) Kryshtofovich, Firmiana tridens (Ludwig) Kryshtofovich, Sterculia dombeyopsis (Unger) Schimper, Sterculia tridens (Ludwig) Schimper, Tilia expansa Saporta, and a variety of names in non-malvaceous genera. The type is Ficus dombeyopsis Unger.

It is recorded from the Miocene [b, c] and Upper Oligocene [d] of Germany, the Miocene of France [e], and the Early Miocene and Oligocene [f] of Bohemia.

Many other names in the genus have been introduced, including


Empedoclea is a living genus, belonging to Dilleniaceae. However, the fossil Empedoclea repando-serrata Engelhardt, from Chile, is believed to be bombaceous, and perhaps to be placed in Bombax.




Leaf material assigned to "Fremontia" coriacea and "Fremontia" lobata is found in the Miocene of California. Material referred to the former is also found in the Pliocene (Piru Gorge), and to the latter in the Miocene Kinnick and Pliocene Tasssajero Fms. Material referred to "Fremontia" is also found in Eocene deposits in Yellowstone.



Fossil leaves from the Tertiary of North America and Eurasia (including Greenland and Spitzbergen) have in the past been assigned to various species of Grewia, including Grewia auriculata Lesquereux, Grewia crenata (Unger) Heer, Grewia crenulata Heer and Grewia obovata Heer; these four species are now all assigned to the synonym of Trochodendroides crenulata (Cercidiphyllaceae). These should not be confused with living Grewia species with the same names.

A number of leaves from the Tertiary of India have been described as or ascribed to species of Grewia.

Grewia mallotophylla Konomatsu & Awasthi is recorded from the Miocene of Nepal [17]. (This is one of several species described in a paper based on leaf and seed macrofossils; the abstract doesn't indicate on which G. mallotophylla was raised, but the specific name is a strong hint that it is based on leaf material.) Grewia tiliaefolia Vahl and Grewia tisaensis Antal & Prasad are recorded from the Middle Miocene of West Bengal [18].

Other material assigned to Grewia includes


Material originally described under the name Grewiopsis elliptica Andreánszky is assigned to Sloanea olmediaefolia (Elaeocarpaceae) [9].

Other names in Grewiopsis are


Guazuma tertiaria Berry.


Hampea conditionalis Hollick is described from Alaska [5]. This has subsequently been redescribed in Menispermaceae as Paratinomiscium conditionalis (Hollick) Wolfe []. This latter placement was not challenged in a later review of the fossil record of Menispermaceae, which dates it to the Palaeocene [].




Leaves assigned to the species Kleinhovia basitruncata (Oishi & Huzioka) Tanai are recorded from the Late Eocene Ikushunbetsu Fm of Hokkaido. These were previously assigned to Marlea basitruncata Oishi & Huzioka and Alangium basitruncatum (Oishi & Huzioka) Tanai.

They differ from the leaves of the sole living species, Kleinhovia hospita, in being truncate to broadly rounded at the base (rather than cordate). [10]


Kydia kraueseli is described from the Oligocene of Liguria, Italy, and stated to be also be present in the Inner Carpathian region []. (These two locations are part of the Adriatic Block, then separate from Europe).

Kydia hungarica, Kydia palaeocalycina


Laria rueminiana


Luehea roxoii Dolianiti is described from a leaf fossil from the Pliocene of Minas Gerais in Brasil.


Lueheopsis dissymetra Langeron




Malvaciphyllum is a form genus representing fossil leaves similar to those of living mallows.

Malvaciphyllum macondicus is recorded form the Palaeocene Cerrajon Fm of Colombia [19].

Malvaciphyllum quenquiadensis is recorded from the Upper Miocene of Argentina, and fossils assigned to this genus, but to no specific species, from the Pliocene of Brasil [f]. The former is thought to be related to Abutilon [20].


In his monograph on fossil dombeyoids Massalongo introduced a genus, Peltophyllum, with two species, Peltophyllum nelumboides and Peltophyllum lobkowitzianus. The genus name is a later homonym (1854) of Peltophyllum Gardner (1843), and if the genus is to be recognised would need to be replaced.


Penosphyllum cordatum is a replacement name for Pterospermites cordatus.


Plafkeria includes leaf fossils from the Palaeogene of North America and North East Asia, and is considered to be tiliaceous. They were originally named as Willisia[21], but this turned out to be a later homonym of a genus in Podostemonaceae, and Plafkeria was introduced as a replacement name[22].

The type species is Plafkeria rentonensis (Wolfe) Wolfe syn. Willisia rentonensis Wolfe. .... They have asymmetric leaf blades and palmate venation.

Plafkeria rentonensis (Wolfe) Wolfe is recorded from the Middle and Upper Palaeocene of the Renton Fm of Washington[21], from the Lowest Eocene of the Liuqu Fm of the Transhimalaya of Tibet [23].

Plafkeria obliquifolia (Chaney) Wolfe is reported from the Late Oligocene of Oregon (also Eocene John Day beds) and the Eocene of California.

Plafkeria basiobliqua (Oishi & Huzioka) Tanai is reported from the Late Eocene Ikushunbetsu Fm of Hokkaido. Other names for this taxon include Marlea basiobliqua Oishi & Huzioka, Alangium basiobliquum (Oishi & Huzioka) Tanai, Ficus ezoensis Endo, Ficus planicosta Endo and Ficus platanifolioides Endo. Specimens of this taxon have also been assigned to Ficus tiliaefolia (Braun) auct. non Heer and Ficus yubariensis Endo. [10]

Plafkeria, unspecified as to species, is recorded from the Lower Eocene of British Columbia [], and from the Eocene Kushutaka Fm of Alaska [10]. The latter may represent Plafkeria basiobliqua.


Some material of Pterospermites incertus Principi from the Oligocene of Italy is referred to Sloanea olmediaefolia (Elaeocarpaceae) [7].


Pterospermum palaeoheyneanum and Pterospermum siwalicum are recorded from the Neogene of India [24].


Pterygota ezoana Tanai is recorded from the Late Eocene Ikushunbetsu Fm of Hokkaido. These leaves are similar to those of the living Pterygota alata, but differ in being broadly rounded to broadly cordate (as opposed to usuallly broadly cordate), and in finer venation [10].

Pterygota palaeoalataSrivastava and Mehrotra is described from the Late Oligocene Tikat Parbat Fm. of Assam. Similar leaves, also from South Asia, have also been assigned to Pterygota cordata, and to the extant Pterygota alata. [16]


Sphaeralcea exhumata is recorded (as Malvastrum exhumatum Cockerell) from the Florissant Fm of Colorado. An image of a leaf is available at the Florissant Fossil Beds National Monument web site.


Leaf fossils assigned to Sterculia are found in the Eocene to Oligocene of Europe, and in other areas of the world. A palmate lobed form from Dalmatia and Slovenia is known as Sterculia labrusca.

In south Asia, the extant species Sterculia versicolor Wall. and Sterculia villosa Roxb. are recorded from the late Tertiary of Jharkand & Bihar respectively; other names used include Sterculia kathgodamense, Sterculia mioensifolia, Sterculia premontana and Sterculia tertiaria. [16]

A specimen of Sterculia variabilis is identified as Sloanea olmediaefolia [8]; the same specimen was elsewhere included within Kydia kraueseli [7]. Sterculia gaudinii, Sterculia spectablis, Sterculia trilobata are listed in the same work without comment.




Tilia eojaponica Endo and Tilia protojaponica Endo are recorded from the Eocene and Miocene respectively of Hokkaido. Material from the Lower Pleistocene is assigned to the living species Tilia japonica Simonk..


Tiliaephyllum is a form genus representing fossil leaves similar to those of the genus Tilia. As this form genus is recorded from the Upper Cretaceous, and the fossil record of Tilia otherwise extends no earlier than the Palaeocene, it seems to me that these leaves do not belong to Tilia, and may not belong to Malvaceae. (Confusion between fossil Tilia and Populus (Salicaceae) leaves, for example, has occurred.) In particular Tillaephyllum dubium Newberry from the Turonian Raritan Fm is too early to be malvaceous.

Tiliaephyllum tsagajanicum is recorded from Manchuria, Kamchatka, the Amur river region, and northern Alaska. It may be referable to Davidia [25].


  1. J. van der Bergh, Index of Angiosperm leaf species names A-B, 1823-2000, Fossilium Catalogus Plantae 107: 1-143 (2005)
  2. J. van der Bergh, Index of Angiosperm leaf species names D-G, 1823-2006, Fossilium Catalogus Plantae 109: 1-154 (2008)
  3. J. van der Bergh, Index of Angiosperm leaf species names H-L, 1823-2008, Fossilium Catalogus Plantae 110: 1-126 (2010)
  4. J. van der Bergh, Index of Angiosperm leaf species names M-O, 1823-2008, Fossilium Catalogus Plantae 111: 1-116 (2011)
  5. Hollick, Arthur, The Tertiary Floras of Alaska, Geological Survey Professional Paper 182 (1936)
  6. Kvaček, Zlatko & V. Teodoris, The Late Eocene flora of Kuclin near Bilina in north Bohemia revisited, Acta Musei Nationalis Pragae B 67(3-4): 83-144 (2011)
  7. Lilla Hably, The Early Oligocene Flora of Santa Giustina (Liguria, Italy) ? Revision and Comparison with the Flora of the Tard Clay Formation, Rivista italiana di Paleontologia e Stratigrafia 116(3): 405-420 (2010)
  8. Bonci, Maria Cristina, Grazia Vanucci, Simona Tacchino & Michele Piazza, Oligocene fossil leaves of the Perrando Collection: history, preservation, and paleoclimatic meaning, Bollettino della Società Paleontologica Italiana 50(3): 145-164 (2011)
  9. Lilla Hably & Boglárka Erdei, The early Oligocene flora and palaeo-environment of the Tard Clay Formation — latest results, Hantkeniana 10: 113-124 (2015)
  10. Tanai, The Revision of the so-called "Alangium" Leaves from the Paleogene of Hokkaido, Japan, Bull. Natn. Sci. Mus., Tokyo, Ser. C 15(4): 121-149 (1989)
  11. Richard H. Eyde, Fossil Record and Ecology of Nyssa (Cornaceae), Bot. Rev. 63(2): 97-123 (1997)
  12. Lakhanpal, R.N., Recognisable species of Tertiary Plants from Damalgiri in the Garo Hills, Assam, The Palaeobotanist 3: 27-33 (1955)
  13. Christophel, D.C., L. J. Scriven & D. R. Greenwood, An Eocene Megafossil Flora from Nelly Creek, South Australia, Transactions of the Royal Society of S. Aust. 116(2). 65-76 (1992)
  14. Worobiec G., E. Worobiec & Z. Kvaček, Neogene Leaf Morphotaxa of Malvaceae s.l. in Europe, International Journal of Plant Sciences 171(8): 892?914 (2010)
  15. Kvaček, Z. & Wilde, V., Foliage and seeds of Malvalean plants from the Eocene of Europe,, Bulletin of Geosciences 85(1): 103-122 (2010)
  16. Gaurav Srivastava & R C Mehrotra, Further contribution to the low latitude leaf assemblage from the late Oligocene sediments of Assam and its phytogeographical significance, J. Earth Syst. Sci. 122(5): 1341-1357 (2013)
  17. Konomatsu & Awasthi, Plant fossils from Arung Khola and Binai Khola formations of Churia Group (Siwalik), west central Nepal, and their palaeoecological and phytogeographical significance, The Palaeobotanist 48(2): 163-181 (1999)
  18. Antal, J.S. & M. Prasad, Morphotaxonomic study of some more fossil leaves from the lower Siwalik sediments of West Bengal, India, The Palaeobotanist 47: 88-98 (1998)
  19. Carvalho, M.R, et al, Paleocene Malvaceae from northern South America and their biogeographical implications, Am. J. Bot. 98(8): 1337-1355 (2011)
  20. Anzotegui, L. M. & R. Herbst, Megaflora (hojas y frutos) de la Formación San José (Mioceno Medio) en Río Seco, Departamento Santa María, provincia de Catamarca, Argentina
  21. Wolfe, Jack A., Paleogene Biostratigraphy of Nonmarine Rocks in King County, Washington, Geological Survey Professional Paper 571 (1968)
  22. Wolfe, Jack A., Palaeogene Floras from the Gulf of Alaska region, Geological Survey Professional Paper 977 (1977)
  23. Robert A. Spicer, Alexander Farnsworth & Tao Su, Cenozoic topography, monsoons and biodiversity conservation within the Tibetan Region: An evolving story, Plant Diversity 42: 229-254 (2020)
  24. Srivastava. Rashmi, R K Saxena and Gaurav Srivastava, Pterospermumocarpon, a new malvalean fruit from the Sindhudurg Formation (Miocene) of Maharashtra, India, and its phytogeographical significance, J. Earth Syst. Sci. 121(1): 183-193 (2012)
  25. Steven R. Manchester, Leaves and Fruits of Davidia (Cornales) from the Paleocene of North America, Systematic Botany 27(2): 368-382 (2002)


  1. Givulesco, R, Observations nouvelles sur Byttneriophyllum tiliaefolium (al. Braun) Knobloch et Kvaček, Rev. Palaeo. Palyno. 10(3): 233-242 (1972)
  2. Schweigert, Guenter, Die untermiozaene Flora (Karpatium, MN 5) des Suesswasserkalks von Engelswies bei Messkirch (Baden-Wuerttemberg)., Stuttgarter Beitraege zur Naturkunde; Serie B (Geologie und Palaeontologie) 188: 1-55 (1992)
  3. Knobloch, E & Zlatko Kvaček, Miozäne Blätterfloren vom Westrand der böhmischen Masse, Rozpravy Ustredniho Ustavu Geologickeho 42: 1-131 (1976)
  4. Juliana Koehler, Die Fossillagerstätte Enspel; Vegetation Vegetationsdynamik und Klima im Oberoligozän, Ph. D. thesis (Tubingen) (2003)
  5. Roiron, Paul, La macroflore d`age Miocene Superieur des diatomites de Murat, Palaeontographica Abteilung B 223: 169-203 (1991)
  6. Kvaček, Zlatko & Harald Walther, Oligocene flora of Bechlejovice at Decin from the neovolcanic area of the Ceske stredohori Mountains, Czech Republic, Acta Musei Nationalis Pragae B 60(1-2), 9-60 (2004)

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