Notes on Fossil Fruits

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Fossil material of plants can be difficult to identify as to species, genus, or larger taxonomic unit, as usually what is found is individual parts of plants, such as wood, leaves, flowers, fruits 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. For fossil fruits the suffices -carpa, -carpon, -carpum and -carpus are often used in generic names, indicating a similarity with the fruits of the modern genus whose name is combined with the suffix. The suffix -strobus is also used, albeit rarely, for some angiosperm fruit form genera, but more commonly for the cones of lycopods, horsetails, seed ferns and conifers. It cannot be assumed that the fossil fruits represent a species particularly close to the modern genus; for example Craigia bronnii fruits were earlier interpreted as rutaceous or sapindaceous, and Byttneria perplexans, Nordenskioldia borealis and Sterculiocarpus coloradensis have been transferred to Betulaceae, Trochodendraceae and Papaveraceae respectively.

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


Aerofructus dilhoffii Herrera and Manchester is a winged fruit from the Palaeocene Cerrojón formation of Colombia, which is provisionally assigned to Malvaceae. [1]


Banisteriaecarpum giganteum (Goeppert) Kräusel is a fossil fruit consisting of a large samara. It was originally described as an Acer, and subsequently interpreted as malpighiaceous [a]. (Banisteria L. is a name that has been formally rejected in favour of Heteropterys Kunth.) Recent opinion interprets it as sterculiaceous, and associates it with the leaf fossils Byttneriophyllum tiliaefolium [2]. A second species has been described under the name Banisteriaecarpum papilio Andreanszky.


Berrya Knowlton non Roxb. is a nom. inval. (later homonym) coined in honour of the American palaeobotanist Edward Wilber Berry. The two species assigned to Knowlton's genus, Berrya racemosa (Knowlton) Knowlton and Berrya fructifer (Lesquereux) Kirchbeimer, represent racemose infructescences. These were not considered malvaceous, though they have been assigned to a variety of families, including Fabaceae, Cornaceae, Nyssaceae and Cercidophyllaceae. Under the last interpretation they have been redescribed as Jenkinsella knowltonii Golovneva et P. Alekseev.



Byttneria (Buettneria) perplexans Cockerell,from the Florissant Formation of Colorado is represented by nutlets with persistent 5-lobed calyxes. It has also been identified as being verbenaceous, under the name of Petraea perplexans (Cockerell) MacGinitie, and more recently has been made the type of the fossil betulaceous genus Asterocarpinus [3], as Asterocarpinus perplexans (Cockerell) Manchester. Carpolithes macrophyllus Cockerell may represent the same or a related species.


Cantitilia ("Kentish Lime") is a fossil genus known from seeds found in the Eocene London Clay strata of northern Kent (Isle of Sheppey and Herne Bay). On the basis of a detailed microstructural analysis Read and Chandler classify this genus as closely related to Tilia. Cantitilia differs from Tilia in having more (5) ovules in an ovary locule, and in the mature fruit being 2–3-seeded (most ovules not maturing), as opposed to the single-seeded state characteristic of Tilia.

The fruits of Cantitilia are syncarpous capsules. They are (4–)5-locular, many-ovulate, but with few maturing seeds, with axile placentation, and loculicidal dehiscence.

There are two known species (as of 1961), Cantitilia polysperma Reid and Chandler, and Cantitilia lobata Chandler. The fruits of the former are ovoid, 6½–13 mm long, 5–11½ mm wide, with seeds 4 to 5 mm long, 3 to 5½ mm wide. The fruits of the latter are deeply 5-lobed, 6 cm long by 11 mm across, with seeds about 4½ mm long and 3¾ mm broad. [4]


Carpolithus (with Carpolithes and Carpolites) is a catchall for fossil fruits, and species will belong to different higher taxa.

Carpolithus bowerbanki Reid & Chandler is recorded from the Lower Eocene of England. A similar fruit is recorded from the Late Palaeocene Almont and Beicegel floras of North Dakota. These have been interpreted as malvalean, but have more recently been reinterpreted as ranunculaceous, as Paleoactaea bowerbanki (Reid & Chandler) Pigg & DeVore and Paleoactea nagelii Pigg & DeVore [5].


Chitaleocarpon intertrappea is a small (2.2mm x 2 mm diameter) barrel-shaped fruit from the Intertrappean (latest Cretaceous or Lower Palaeocene) rocks of Mohgaon Kalan (Madhya Pradesh). It is a loculicdal capsule, with seven locules, each containing 2-8 oval to rounded seeds, suggesting the presence of around 8 ovules per locule. [b]

The name Chitaleocarpon deccanii was used in a 1984 thesis. It is not known to me how this relates to Chitaleocarpon intertrappea; it might be a manuscript name superceded by Chitaleocarpon intertrappea, or it may be an unrelated name.


Christianacarpum is a form genus representing fossil fruits similar to those of the living genus Christiana,

Christianacarpum quinquelocularis, from the Eocene of France, is interpreted as belonging to a plant which belongs to the genus Christiana.


Daberocarpon is a presumed malvaceous fruit. Daberocarpon gerhardii Chitaley & Sheikh is based on a 10-locular fruit from the uppermost Cretaceous of the Deccan Intratrappean Beds of India [c]. Each locule is single-seeded.

The authors suggested an affinity with tribe Malveae. However the fruit preceeds the appearance of malvoid pollen within the pollen record, which suggests that we should look elsewhere for the true affinities.


Etheridgea subglobosa is a fossil fruit from the Late Cretaceous of Australia, which is assigned to Tiliaceae (Grewioideae? Brownlowiodeae?).


Fruits ascribed to the fossil species Firmiana yunnanensis are reported from the Miocene of Yunnan [d].


Fracastoria is a fruit fossil. It was described, and placed in Sterculiaceae, in the mid 19th century by Massalongo [6] , from specimens of large fruits from Bolca in northern Italy. Massalongo appears to have been a splitter, and I am skeptical that there are as many distinct taxa as he described. Most subsequent opinion treats Fracastoria as the fruits of a palm (e.g. [e]), though it has been treated as related to the much smaller Hightea (see below).

The described species are :-


Grewia mohgaonse ...


"Harrisocarpon sahnii Chitaley and Nambudiri (1973) is a pentalocular, pentaribbed, septicidal capsule resembling that of Abutilon. Each locule contains two seeds arranged one above the other in a row on a central placenta."


Hibiscocarpon mohgaonensis V.D.Kapgate is a a minute fruit (~1 mm long x 1 mm diameter) from the Intertrappean (latest Cretaceous or Lower Palaeocene) rocks of Mohgaon Kalan (Madhya Pradesh). It is ribbed pentalocular capsule. It named from a perceived similarity to the much larger fruits of Hibiscus (Abelmoschus) esculentus. [7] A later study considers its malvaceous affinities questionable. [f]


Hightea is a fruit fossil, originally assigned to Malvaceae, from the Eocene of Britain. Bowerbank [8] described 10 species - Hightea attenuata, Hightea elliptica, Hightea elegans, Hightea fusiformis, Hightea inflata, Hightea minima, Hightea orbicularis, Hightea oviformis, Hightea turbinata and Hightea turgida. He interpreted the fruits ... Reid and Chandler reinterpreted most of these fruits as myrtaceous, and reduced the number of species to 2 [4].

Hightea is named in honour the botanist John Hight [8].


Fruits assigned to Luehea newberryana (Knowlton) MacGinitie are found in the Early and Middle Eocene of Wyoming. These are now considered to belong to Populus cinnamonoides [9].


Malvacarpon clarus


Malvacarpus is a form genus representing fossil fruits similar to those of extant mallows.

The type is Malvacarpus tertiarius Berry, from the Miocene of Argentina. The species Malvacarpus guinazui Berry is recorded from the Palaeocene or Eocene of the Rio Negro region of Argentina. The name Malvacarpus octoloculus has also been used.


Nordenskioldia is a fruit fossil once assigned to Tiliaceae, but now considered as a member of Trochodedraceae, with an extensive fossil record in time (Palaeocene to Miocene) and space (North America, Greenland, Scotland, Spitzbergen, Russia, Inner Mongolia). The leaf genus Zizyphiodes is thought to represent the same plant.


The name Pentaloculocarpon has been applied to a number of capsular fruits from the Intertrappean beds of the Deccan. At least one has been suggested to be malvaceous.

Pentaloculocarpon chitaleii Kapgate & Kapgate is a fruit fossil from the Intertrappean beds of the Deccan, which may be malvaceous. It is a pentalocular loculicidal capsule with axile placentation and single-seeded locules, of ovoid shape measuring 850µ by 1mm. The pericarp is differentiated into a outer thin epidermal zone, followed by middle multilayered parenchymatous zone consisting intercellular spaces and mucilage canals, followed by 1-2 layered compact inner zone. The seed has a membranous coat. The embryo is dicotyledonous and embedded in endosperm tissue. [g]

(This may not be validly published, as I have found no evidence of publication beyond a conference poster.)

No familial assignment was offered for Pentaloculocarpon intertrappea A.S.Khursel and S.D.Narkhede. [10]

Pentaloculocarpon mohgaonse, which is the subject of a Ph.D. thesis, may also not be validly published. A bare mention of it occurs in a list of capsular fruits from the Intertrappean.


Pteleaecarpum is a form genus representing fossil fruit mistakenly interpreted as related to those of Ptelea (Rutaceae). It has also been interpreted as sapindaceous [11]. However Pteleaecarpum bronnii is now recognised as being the fruit of a species of Craigia (Tilieae), of which floral and foliar material is also available. Pteleaecarpum europeum represents the same species [12]. Fossil material of Craigia bronnii is known from several localities in Europe, including the Lower Oligocene of Hungary, the Lower Miocene of Bohemia, and the Upper Miocene of Germany


Reevesia hurnikii



Sphinxia ovalis is a fossil fruit from the Londay Clay interpreted as being Dombeyoid, and most closely allied to Dombeya and Trochetia. (The name Sphinxia is also applied to a Devonian lycopsid; one or other usage of this generic name will be invalid.)


Fruit assigned to Sterculia palaeovillosa on the grounds of its similarity to the living Sterculia villosa is recorded from the Oligocene of Assam [13].


Sterculiocarpus is a form genus representing fossil fruits similar to those of living Sterculia.

The species Sterculiocarpus coloradensis Berry has been reduced to synonyny with Palaeoaster porosa (Papaveraceae) [14]. Leaf material named Sterculia coloradensis consists of deeply three lobed leaves that don't present a papaveraceous aspect, and presumably does not represent the same taxon.

Other names which has been published are Sterculiocarpus eocenicus Berry and Sterculiocarpus sphericus Berry, from the Eocene Wilcox Fm, and Sterculiocarpus sezannelloides Berry from the Eocene Lagrange Fm. I do not know whether these represent genuine sterculioid fruits.

Sterculiocarpus etayoi is described from the Lower Cretaceous (Aptian) of Colombia. The fossils of casts of fruits are similar in gross shape to the follicles of Sterculia, but no other data is known. Given the date I doubt that they are in fact sterculiaceous. [15]


Tarrietia germanica Rüffle and Tarrietia hungarica Rásky are fruit fossils which have more recently been been interpreted as monospermous legumes, the latter under the name Machaerites hungaricus (Rásky) Andreanszky [h].


  1. Fabiany Herrera, Steven R. Manchester, Mónica R. Carvalho, Carlos Jaramillo & Scott L. Wing, Paleocene wind-dispersed fruits and seeds from Colombia and their implications for early Neotropical rainforests, Acta Palaeobotanica 54(2): 197-229 (2014)
  2. Zlatko Kvaček & Lilla Hably, The Whole plant Reconstruction of Banisteriaecarpum giganteum, Folia Musei rerum naturalium Bohemiae occidentalis. Geologica et Paleobiologica 48(1/2): 1-10 (2015)
  3. Manchester, S.R, & Crane, Peter R., A New Genus of Betulaceae from the Oligocene of Western North America, Bot. Gaz. 148(2): 263-273 (1987)
  4. Reid, Elanor Mary & Marjorie Elizabeth Jane Chandler, The London Clay Flora (1933)
  5. Pigg, Kathleen B. & DeVore, Melanie L., Paleoactaea gen. nov. (Ranunculaceae) fruits from the Paleogene of North Dakota and the London Clay, Am. J. Bot. 92(10): 1650-1659 (2005)
  6. Massalongo, D.A.B, Palaeophyta rariora formationis tertiariae Agri Veneti,, Acta Caes. R. Inst. Ven. 3(3): 35-38 (1858)
  7. V.D. Kapgate, Fossil fruit of Hibiscus esculentus L. of family Malvaceae from Deccan Intertrappean cherts of India, The Palaeobotanist 66(2): 211-216 (2017)
  8. Bowerbank, James Scott, A history of the fossil fruits and seeds of the London clay (1840)
  9. Steven R. Manchester et al, Foliage and Fruits of Early Poplars (Salicaceae: Populus) from the Eocene of Utah, Colorado, and Wyoming, International Journal of Plant Sciences 167: 897-908 (2006)
  10. A. S. Khursel & S. D. Narkhede, A new petrified pentalocular capsular fruit from the deccan intertrappean beds of Mohgaonkalan, M.P., India, Int.J.Curr.Microbiol.App.Sci 5(4): 483-487 (2016)
  11. Buzek, Cestmir, Kvaček, Zlatko & Manchester, Steven R., Sapindaceous Affinities of the Pteleaecarpum Fruits from the Tertiary of Eurasia and North America, Bot. Gaz. 150(4): 477-489 (1989)
  12. Kvaček, Z., Early Miocene records of Craigia (Malvaceae s.l.) in the Most Basin, North Bohemia - whole plant approach, Journal of the Czech Geological Society 49(3/4): 161-171 (2004)
  13. Mehrota, Two new fossil fruits from Oligocene sediments of Makum Coalfield, Assam, India, Current Science (IAS) 79(10): 1482-1483 (2000)
  14. Smith, Una R., Revision of the Cretaceous Fossil Genus Palaeoaster (Papaveraceae) and Clarification of Pertinent Species of Eriocaulon, Palaeoaster, and Sterculiocarpus, Novon 11(2): 258-260 (2011)
  15. Huertas, Gustavo, Sertum Florulae Fossiles Villae de Leivae, Caldasia 10(46): 59-75 (1967)


  1. Kräusel, R., Die tertiäre ?Riesenahorn? Banisteriaecarpum nov. gen. Die systematische Stellung von Acer giganteum und Acer otopteris Goepp., Abh. Seckenberg. Naturforsch. Ges. 485: 75-80 (1952)
  2. V.D. Kapgate, D.K. Kapgate & M.T. Sheikh, First record of seven locular dicot fruit from Deccan Intertrappean Beds of Mohgaon Kalan, Chhindwara district, Madhya Pradesh, Gondwana Geological Magazine 21(2): 109-114 (2006)
  3. Chitaley, S.D. & Sheikh, M.T., A ten locular petrified fruit from the Deccan Intertrappean Series of India, The Palaeobotanist 20(3): 297-299 (1973)
  4. Sanping Xie, Steven R.Manchester, Kenan Liu, Yunfeng Wang & Yang Shao, Firmiana (Malvaceae: Sterculioideae) fruits from the Upper Miocene of Yunnan, Southwest China, Geobios 47(4): 271-279 (2014)
  5. Tralau, H., The genus Nypa van Wurmb, Kungliga Svenska Vetenskapsakademiens Handlingar 10: 5-29 (1964)
  6. Steven R. Manchester, Dashrath K. Kapgate, Bandana Samant, Dhananjay M. Mohabey & Anup Dhobale, Fruits and Pollen of Malvoideae (Malvaceae) in the Maastrichtian–Danian Deccan Intertrappean Beds of Central India, International Journal of Plant Sciences 184(1): 68-84 (2023)
  7. Kapgate, V.D. & Kapgate, D.K.., A pentalocular fruit from the Deccan Intertrappen beds of India, poster at XVI International Botanical Congess
  8. Zlatko Kvaček & Lilla Hably, The Whole plant Reconstruction of Banisteriaecarpum giganteum, Folia Musei rerum naturalium Bohemiae occidentalis. Geologica et Paleobiologica 48(1/2): 1-10 (2015)

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