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Fossil Index
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 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 suffices -pollis, -pollenites, -pites, and -dites are sometimes used with generic (or higher taxa) names, indicating a similarity with the pollen of the modern taxon whose name is combined with that suffix. (-pites is an abbreviated form of -pollenites.) It cannot be assumed that fossil pollen represents a species particularly close to the modern genus; for example much Malvacipollis is euphorbiaceous rather than malvaceous.
Material on fossil pollen often omits the taxonomic position of the material, and I have probably omitted some genera which are considered malvaceous; for example, Thomsonipollis is a segregate from Intratriporopollenites, but I don't whether the segregator placed the former in the same taxon (Malvaceae sensu lato) as the latter. (Species of Thomsonipollis include T. magnificus, T. palaeocenicus, T. magnificoides, T. sabinetownensis, T. expolitus and T. gracilis.) The botanical affinities of Suprapollis variabilis are said to be unknown, but it looks rather grewioid (i.e. malvaceous) to my untutored eye.
In some cases pollen may be variously assigned to a living genus, or to a pollen form genus. For example the pollen of Aguiaria, Bernouillia, Bombax, Ceiba and Pseudobombax has been recorded under those names and as species of the form genus Bombacacidites.
The fossil record of pollen is sufficiently rich that I am in the process of dividing my notes into separate pages on echinate, prolate and oblate pollen. Echinate pollen characterises Malvoideae and some Bombacoideae, but is also prevalent in a number of other families such as Asteraceae, Convolvulaceae and Euphorbiaceae. Consequently there have been a number of mistaken assignments of pollen to Malvoideae (formerly Malvaceae) and Bombacoideae (formerly Bombacaceae).
Pollen comparable to that of Aguiaria, which is currently restricted to the Amazon basin, is recorded from Miocene sites in Panama and southern Mexico (Chiapas) [i].
Pollen ascribed to Bernouillia is recorded from the Middle Eocene of Wyoming [ii], the Middle Oligocene of Puerto Rico [iii] and Miocene of Colombia [] and Panama [i].
The name Bernouillia has also been applied to Triassic ferns.
Bombacacidites has a long pollen record extending from the Late Cretaceous to the Holocene. It is generally recognised as representing taxa belonging to Bombacoideae, but B. paulus and others may be referable to Brownlowia or an allied genus. The earliest records are from North America, but in the Lower Tertiary it is broadly distributed. In the Upper Tertiary records become restricted to South America and, oddly, New Zealand. In the Eocene of Colombia at least 20 morphospecies of this pollen genus are known [iv].
Bombacacipites (type B. nacimientoensis Anderson) would appear to be a synonym of Bombacacidites.
Bombacacidites africanus is recorded from the Palaeogene of south west Nigeria.
Bombacacidites annae (Van der Hammen) Germeraad et al. is recorded from the Late Palaeocene of Colombia [v] and the Eocene of Venezuela [vi].
Bombacacidites baculatus is recorded from the Late Miocene of Colombia [15], the Upper Eocene and Oligocene [vii] and lower Miocene of Venezuela [viii], and from the Miocene and Pliocene of Mexico [17, 48] and Brasil [iv ]
Bombacacidites baumfalki and Bombacacidites zuatensis are recorded from the La Rosa and Lagunillas Formations of the Lower Miocene of the Maracaibo Basin of Venezuela [viii]. Bombacacidites baumfalki is also recorded from the Upper Eocene and Oligocene of Venezuela [vii].
Bombacacidites bellus is recorded from the Lower Miocene of Venezuela [viii] and Pliocene or Pleistocene of Colombia [ix]. Similar material is recorded from the Late Oligocene to Middle Miocene [20].
Bombacacidites bombaxoides Couper is recorded from the Late Oligocene to Miocene of New Zealand [3, 34]
Bombacacidites brevis (Dueñas) Muller et al is recorded from the La Rosa and Lagunillas Formations of the Lower Miocene of the Maracaibo Basin of Venezuela [viii], the Upper Eocene and Oligocene of Venezuela [vii], and from the Middle Eocene of Colombia [x].
Bombacacidites ciriloensis is recorded from the Middle Miocene of Colombia [12], and also recorded from the Pliocene or Pleistocene of Colombia [ix].
Bombacacidites fereparilis Frederiksen is recorded from the United States. Similar material is known from the Middle Eocene of Cuba [vi].
Bombacacidites fossureticulatus Jaramillo and Dilcher is recorded from the Middle Eocene of Columbia [x].
Bombacacidites foveoreticulatus Muller et al is recorded from the Middle Eocene of Columbia [11, 19].and the Upper Eocene of Venezuela [vii].
Bombacacidites gonzalezii Jaramillo and Dilcher is recorded from the Middle Eocene of Columbia [x].
Bombacacidites isoreticulatus McIntyre is recorded from the Palaeogene of New Zealand [34].
Bombacacidites kettigensis is recorded from the Eocene [xi] or Miocene [x] of Germany.
Bombacacidites noremmi is recorded from the Oligocene or Miocene of the Gulf of Mexico [48].
Bombacacidites nacimientoensis (Anderson) Elsik is recorded from the Cretaceous [64], Palaeocene [7] and Eocene [vi] of the United States, and from Colombia.
Bombacacidites nanobrochatus Frederiksen is recorded from the United States, and from the Middle Miocene of Argentina. Similar material is known from the Middle Eocene of Cuba [vi].
Bombacacidites paulus is recorded from the Middle Eocene of Texas [x]. A relationship to Brownlowia, or to one of several genera of Bombacoideae, has been proposed.
Bombacacidites protofoveoreticulatus Jaramillo and Dilcher is recorded from from the Palaeocene of Colombia.
Bombacacidites qaidamensis is recorded from the Oligocene Upper Ganchaigou Formation of China and the Miocene Hanjiang Formation of the South China Sea. [xii]
Bombacacidites reticulatus Krutzsch is recorded from the Cretaceous, Palaeocene and earliest Eocene of the United States [4, 5, 6, 7, 64]
Bombacacidites simplireticulatus Jaramillo and Dilcher is recorded from from the Palaeocene of Colombia
Bombacacidites soleaformis is recorded from the Eocene of Colombia [13, 14, 19], the Middle Eocene of Venezuela, and the Upper Eocene of Venezuela [vii].
Bombacacidites tilioides is recorded from the Eocene [xi] or Miocene [x].
Bombacacidites ciriloensis is recorded from the Middle Miocene of Colombia [12], and also recorded from the Pliocene or Pleistocene of Colombia [ix] of Germany. Similar material (Bombacacidites "mirabilis") is found in the Middle Eocene of Cuba [vi]. It is also recorded from the United States [vi].
Bombacacidites triangulatus is recorded from the Oligocene Tikat Parbat Formation of Assam [].
Bombacacidites, unassigned to species, is recorded from Holocene of Maharashtra (India) [45], which presumably reflects either of both of the living genera Bombax and Ceiba. It is also recorded from the Miocene of Peru [a] and the Eocene of Louisiana [x].
The names Bombacacidites psilatus and Bombacacidites pseudosimplireticulatus have also been used.
Named for a general similarity to Bombacacidites, but this genus also shows similarity to the pollen of Triumfetta and that of Coris (Primulaceae) [24]. In particular B. hectorii is prolate, rather than oblate, and hence is more likely grewioid or byttnerioid that bombacoid.
Bombapollis hectorii is recorded from the Late Oligocene of New Zealand [24]. Bombapollis texensis Elsik is recorded from the Middle Eocene of Texas. [27].
Bombacipites is orthographic variant of Bombacacipites.
Pollen ascribed to Bombax is recorded from the Middle Eocene of Wyoming [ii].
Bombax ceiba-type pollen is found in the Palaeocene of South America. []
Pollen ascribed to Cavanillesia is recorded from the Middle Eocene of Wyoming [ii], and the Mio-Pliocene of Bolivia [38].
Pollen ascribed to Ceiba is recorded from the Miocene of Panama [i].
Dermatobrevicolporites Kar
Discoidites represents tricolpate disc-shaped pollen grains. It differs from Tilliaepollenites in possessing short colpi and lacking any indication of vestibulate pores. The type species, Discoidites borneensis is similar to the pollen of living Brownlowia and Pentace []. Other species from Indonesia are Discoidites robustus, Discoidites eflatus, Discoidites maximus, Discoidites novaguinensis and Discoidites pilosus.
Discoidites parvistriatus is recorded from the Lower Palaeocene of North Dakota [28].
Euphorbiacidites is a form genus for pollen similar to that of living Euphorbiaceae. However Euphorbiacidites microreticulatus is comparable to Grewioideae (Grewia and Triumfetta). [xii]
This form genus represents an extinct bombacoid pollen type [b]. It is common enough to be an important index fossil in the north Tethyan floral belt. [b]. The type species is Friedrichipollis duploreticulatus.
Friedrichipollis claibornensis is recorded from the Middle Eocene of Texas [27]. Friedrichipollis is also recorded from Georgia (USA).
This form genus represents pollen similar to that of the living genus Grewia. However it is tricolpate, whereas Grewia has tricolporate pollen, and may not be closely related (or even malvalean). [xii] Grewiapollenites canadensis is recorded from the Cretaceous of North America [33].
This form genus covers pollen morphospecies associated with Tilioideae, Brownlowioideae and perhaps also Bombacoideae. (A possible association with Sterculiaceae is also mentioned in at least one source.) It is divided into 5 subgenera, one of which contains pollen similar that of Tilia and Cragia, a second which contains pollen similar to that of Mortoniodendron, and a third (Insculptipollenites) which contains pollen similar to that of some Brownlowidoideae; the other two subgenera may also related to Brownlowioideae [b]. Modern Brownlowioideae has two types of pollen, which might correspond to the tribes Brownlowieae and Berryeae [].
Intratriporopollenites ambiguus and Intratriporopollenites tiliaceus are described from Japan.
Intratriporopollenites (or Tiliaepollenites) ceciliensis is recorded from the Eocene of Germany [xi]. Intratriporollenites ceciensis (is this the same taxon?) is recorded from the Middle Eocene of North Africa, and is hypothesised to represent a form of brownlowioid pollen [56].
Tiliaepollenites danei is recorded from the Upper Palaeocene of interior Canada [c, d, e] under that name and as Tilia danei.
Intratriporopollenites insculptus (Potonié) Thomson & Pflug is recorded from the Oligocene of Ireland [f] and Germany . It may be the pollen of Byttneriophyllum tiliaefolium (Al. Braun) Knobloch & Z. Kvacek. It may be brownlowioid. It might also represent Craigia. [xii]
Intratriporopollenites kettingensis Pflug is Bombacacidites kettingensis (Pflug) Krutzsch.
Intratriporopollenites magnoporatus Pflug. & Thomson is said to be juglandaceous [58]. Elsewhere it is transferred to the form genus Subtriporopollenites, of unknown affinity.
Material similar to Intratriporopollenites maxoides is recorded from the Eocene of Germany [xi].
Intratriporopollenites microinstructus is recorded from the Lower and Middle Miocene of Germany.
Intratriporopollenites (or Tiliaepollenites) notabilis (Harris) Stover is recorded from the Palaeocene [39], late Eocene [54], early Oligocene [54] and late Oligocene [24] of New Zealand, from the early Eocene of Tasmania [53], from the Palaeocene of Victoria [39], and from South Australia [55]. Similar pollen is recorded from Borneo []
Intratriporopollenites ollivierae is recorded from the Lower Eocene of France, and from the Palaeogene of the southeastern United States.
Intratriporopollenites rhizophorus is recorded from China.
Intratriporopollenites stavensis is recorded from the from the Eocene of Texas [56], Lower Palaeocene (Danian) of South Carolina, and from the Eocene/Oligocene boundary of Alabama and Missisippi.
Intratriporopollenites tetraforaminipites and Intratriporopollenites vescipites are recorded from the Eocene of Wyoming [29]. The latter is also recorded from the Palaeocene of North Dakota. [28] and (as Tilia vescipites) from the Upper Palaeocene of Canada [c]
Intratriporopollenites precrassipites and Intratriporopollentites prevescipites (Google search under Tilia) ...
13 morphospecies of Tiliaepollenites are recorded from China.
Tiliaepollenites formosensis Shaw, Tiliaepollenites taiwanensis Huang, Tiliaepollenites pengchiahsuensis Shaw and Tiliaepollenites speciosus Shaw are recorded from Eocene sediments of the China Sea north of Taiwan. [50], as is Tiliaepollenites zonatus Shaw [51].
Tiliaepollenites is recorded from Indonesia and New Zealand. (I am skeptical of this identification.) []
Tiliaepollenites is a form genus for pollen similar to that of Tilia. More often than not either Tilia or Intratriporopollenites are used instead. Furthermore the genus name is a nomen superfluum (a junior synonym of Tilia), due to a contaminating modern Tilia pollen grain being used as the type. [] Therefore records of Tiliaepollenites are presented here, rather than separately. (Tiliaepollenites may correspond to one of the subgenera of Intratriporopollenites.) The type of Tiliaepollenites, T. indubitalis Potonié, is the same as Intratriporopollenites indubitalis. There are two groups of Tiliaepollenites species. One is smaller (<30μm), with less developed exine and undeveloped pore chambers, and corresponds to Craigia. The other is larger, with thicker exine, coarser ornamentation and developed pore chambers, and corresponds to Tilia. [xii]
Tiliaepollenites (Craigia group)
Intratriporopollenites indubitabilis (Potonié) Thomson & Pflug is recorded from the Eocene [58] and Miocene of Anatolia, from the Miocene of the Gulf of Mexico, and from China. [xii]
Tiliaepollenites lingfengensis is recorded from China. [xii] It agrees with Craigia rather than Tilia in size, and may represent the former. [xii]
Intratriporopollenites (or Tiliaepollenites) microreticulatus is recorded is recorded from Late Eocene of Guangxi [52], from the Early Eocene of Belgium, , and from the Miocene of Bohemia, from Greenland, and from China. It agrees with Craigia rather than Tilia in size, and may represent the former. [xii]
Intratriporopollenites (or Tiliaepollenites) minimus is recorded from the early Palaeocene of Canada, from the Eocene of Germany [xi], and from China. [xii]
Tiliaepollenites retigranulatus is recorded from China. [xii] It agrees with Craigia rather than Tilia in size, and may represent the former. [xii]
Tiliaepollenites (Tilia group)
Intratriporopollenites (or Tiliaepollenites) cordataeformis is a name for pollen similar to that of the living species Tilia cordata. It is recorded from the Upper Miocene of Romania [57], and from China. [xii]
Tiliaepollenites hunchensis is recorded from China. [xii]
Intratriporopollenites (or Tiliaepollenites) instructus (Potonié) Thomson & Pflug is recorded from the Eocene of Wyoming [29], North Dakota [g] and Missisippi [60], from the Eocene [58] and Oligocene-Miocene [59] of Anatolia, from the Miocene of Germany [61], from the Miocene of Denmark [x].
Bombacacidites ciriloensis is recorded from the Middle Miocene of Colombia [12], and also recorded from the Pliocene or Pleistocene of Colombia [ix], and from Eocene sediments of the China Sea north of Taiwan [51].
Intratriporopollenites (or Tiliaepollenites)pseudinstructus Mai is recorded from the Lower Tertiary of the United States [7], and the Eocene of Germany, and from China. [xii]
Lakiapollis is a form genus for pollen similar to that of Durio and related genera, Lakiapollis ovatus represents pollen similar to that of Durio in particular, and Lakiapollis microreticulatus pollen similar to that of Neesia. Lakiapollis is recorded from the Palaeocene and Eocene of India [41], and also from Indonesia
Lakiapollis ovatus is recorded from the Oligocene Tikat Parbat Formation of Assam [].
Lakiapollis matanomadhensis is not correctly placed in this genus, and may not be malvaceous. It is correctly placed in Tricolporopollis (of which Retitribrevicolporites is a junior synonym), and is said to represent pollen of Bombacaceae (Durio?). [Retitribrevicolporites Is a Synonym of Tricolporopollis (Tertiary Pollen from India), Taxon, Vol. 48, No. 3 (Aug., 1999), pp. 493-496 ]
Mansoniaepollis is a form genus for pollen similar to that of Mansonia. It is common enough to be an important index fossil in the north Tethyan floral belt. [b]
Pollen of Mortoniodendron is recorded from the Miocene of Mexico and Panama [], the Mio-Pliocene of Guatemala [36], and the Pliocene of Panama [x].
Bombacacidites ciriloensis is recorded from the Middle Miocene of Colombia [12], and also recorded from the Pliocene or Pleistocene of Colombia [ix].
Parsonsidites psilatus is recorded from the Palaeogene and Neogene of New Zealand. Some or all of this material may represent pollen of Hoheria and Plagianthus (with the spines missing.) However Parsonsidites multiporus is said to be balanophoraceous. [34]. The names P. conspicuus and P. minor have also been used, but I have no reason to suspect that they represent malvaceous taxa.
Pollen ascribed to Psuedobombax is recorded from the Miocene of Panama [37].
Reevesiapollis is a form genus for pollen similar to that of Reevesia. It has a long pollen record in Europe, extending into the Pliocene, and is also recorded from North America. [b ]
Reevesiapollis reticulatus is recorded from northwest Australia (Pilbara) [30], South Australia (Flinders Island) [32] and New Zealand [31, 34]. This pollen species is associated with the living genus Ungeria [30], which is now confined to Norfolk Island.
Reevesiapollis siameniformis is recorded from the Eocene Shahejie Formation of China, and in greater abundance from the Miocene. [xii]
Reevesiapollis arslanensis and Reevesiapollis microreticulatus are recorded from Europe.
The name Reevesiapollenites triangulus has also been used. I am unaware of the distinction, if any, between Reevesiapollis and Reevesiapollenites. As the specific epithet triangulus is used with both generic names, I suspect that they are synonyms, the latter perhaps being an orthographic variant.
Retistephanocolpites is a pollen form genus for reticulate(?) stephanocolpate pollen. Pollen of this form, from the Upper Miocene of Argentina is ascribed to Bombacaceae. [35, 47] Other pollen species of this form genera are assigned to other families. Retistephanocolpites crassiannulatus Lorente is assigned to Quararibea.
Retitribrevicolporites Kar; Retibrevitricolpites macroreticulatus Kumar et Takahashi
Retitricolporites firmianoides pollen is compared to that of the living species Firmiana simplex. [xii] It is recorded from the Eocene Shahejie Formation of China [xii]
Retitricolporites shaoboensis is recorded from the Senonian (Late Cretaceous) Taizhou Formation of China. [xii] It is thought to be grewioid [xii].
Rhoipites guianensis (Van der Hammen & Wymstra) Jaramillo & Dilcher, from the Neogene of western Amazonia, is attributed to Sterculioideae (Firmiana/Hildegardia) or Grewioideae (Glossostemon/Trichospermum or Vasivaea/Trichospermum). It was previously placed in Retitricolporites. There are other species of Rhoipites, but their taxonomic affinities are thought to lie elsewhere (e.g. Araliaceae).
Sandiegopollis elkeae is recorded from the Eocene of California.
Tahitiaoipollis is a form genus for pollen similar to that of Christiana (Tahitia is a synonym of Christiana). It is common enough to be an important index fossil in the north Tethyan floral belt. [b]
Tricolporocolumellites Kar
Trochetiopsis-like pollen is recorded from the late Miocene of St. Helena [40].
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