Acta Palaeobot. Suppl. 1: 5–30, 1994

TAXONOMICAL REVISION OF SELECTED POLLEN AND SPORES TAXA FROM NEOGENE DEPOSITS

Rewizja taksonomiczna wybranych taksonów spor i ziarn pyłku z osadów Neogenu

MARIA ZIEMBIN´ SKA-TWORZYDŁO1, IRENA GRABOWSKA2, ALEKSANDRA KOHLMAN-ADAMSKA3, KRYSTYNA SKAWIN´ SKA4, BARBARA SŁODKOWSKA5, LEON STUCHLIK4, ANNA SADOWSKA6 and HANNA WAZ˙ YN´ SKA5

1 – Warsaw University, Institute of Geology, Z˙ wirki i Wigury 93, 02–089 Warszawa 2 – 02–508 Warszawa, ul. Puławska 53B 3 – Museum of the Earth, Polish Academy of Sciences, Aleja Na Skarpie 20/26, 00–488 Warszawa 4 – W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31–512 Kraków 5 – Polish Geological Institute, Rakowiecka 4, 00–975 Warszawa 6 – University of Wrocław, Institute of Geological Sciences, Paleobotany Department, Cybulskiego 30, 50–205 Wrocław

ABSTRACT. For unification the taxon determination of the most common pollen and spores from the Neogene of Central Europe a checklist of selected genera and species was elaborated. All taxons are illustrated and ordered in both morphological and botanical systems. For several taxons a systematical revision according to the rules of the International Code of Botanical Nomenclature was carried out. All enumerated taxons were classified to a proper group of geofloristic element.

KEY WORDS: Neogene, Paratethys, Polish Lowland, pollen, spores, taxonomy, botanical affinity, geofloristic element, palaeo- tropical, arctotertiary

INTRODUCTION

International co-operation on correlation of establishing 15 new genera, 7 new species, 50 Neogenic deposits from Paratethys area and combinations and 6 new names. In spite of European Lowland basins, based on results of this, 1 new status was established and 3 diag- palynological investigations (Planderová et al. nosis were emended. In such a way elaborated 1993), was prompted the authors of this paper list, together with illustrations, can provide to make an attempt to unify the taxonomy and the basis for the univocal interpretation the to make revision of some scattered fossil taxa. results of palynological correlations. The next In the first stage of the mentioned co-oper- stage means the elaboration of common data- ation a preliminary list of 108 taxa, based on base for quantitative comparisons of spores the comparison of results of palynological ana- and pollen grains of Neogene deposits in Po- lyses from various localities in Paratethys and land and in neighbouring countries. The data- European Lowland areas, was elaborated. It base was created on the basis of the list pro- might serve as a basis for proper under- posed by the authors of the present paper. standing of lists with percentage values, in The lists of chosen sporomorph taxa from which these species frequently occurred. The Neogene deposits were arranged according to necessity of revision of many taxa has widened two systems. the scope of preliminary plans. 1. Morphological system: spores in trilete- In the effect a list comprises 95 genera and monolete arrangement and pollen grains of 185 taxa of spores and pollen grains was ela- Gymnospermae with morphological types: in- borated. During the work 82 taxa were re- aperturate, saccate, polyplicate; of Angiosper- vised. Also, it became evident the necessity of mae with morphological types: monoporate, 6 triporate, polyporate, monocolpate, tricolpate- fested (e.g. Nelumbo europaea Tarasevich tricolporate, polycolpate-polycolporate, syncol- 1983). This type of nomenclature was com- pate, tetrade. An order of taxa in the particu- monly used by palynologists, among others lar morphological groups is alphabetical. from former Soviet Union. This is not in con- 2. Botanical system in genera and families formity with recommendations by the Interna- arrangements follows: Tryon and Lugardon (1991 tional Code of Botanical Nomenclature (1961, – for Pteridophyta); Szweykowska and Szwey- art. 20, Rec. 20b), according to these in generic kowski (1993 – for Cycadophytina and Pinophy- name the organ, being the base for the descrip- tina); Takhtajan (1987 – for Magnoliophytina). tion, should be shown. The proper name for the fossil species of pollen of Nelumbo genus should be Nelumbopollenites europaeus (Ta- HISTORY OF INVESTIGATIONS rasevich) comb. nov. At the same time when Potonié, Thiergart The differences in taxonomical approaches and others in Europe used Sporites and Polle- among various palynologists from different scien- nites for genus names of spores and pollen tific centres have historical reasons. During grains, in the U.S.A. Wodehouse (1933) added the early stages of introducing the palynologi- the suffix “-pites” for root of the botanical cal methods for investigations of the Tertiary generic name (e.g. Caprifoliipites), stressing in deposits Potonié (1931a–d, 1934) used generic this way its organogenic source. Couper (1953) names Sporites and Pollenites. The differentia- used in palynological taxonomy the suffix tion among morphological types of distin- “-idites” for stressing the fossil origin of sporo- guished sporomorphs was only seen on the morph kinds (e.g. Haloragacidites). level of a fossil taxon (e.g. Pollenites margari- In older papers also the suffix “-idites” may tatus Potonié 1931a). In the next stage of de- be found, what in combination with contem- velopment of the discipline Potonié (1934), porary genus or family names stresses fossil Thiergart (1937), Raatz (1937) attempted to type of an organ and points to its morpholo- attribute species to botanical genera by the ad- gical similarity {e.g. Quercoidites (Potonié, ding generic name before the epithet Pollenites Thomson & Thiergart 1950), Osmundacidites and Sporites, e.g. Ilex-pollenites margaritatus (Couper 1953)}. Since the times of Potonié Raatz 1937, Thiergart 1937. From that time a (1934) in the names of fossil genera of botani- clear division among botanists is dated back: cal origin the suffixes “-pollenites” and on those preferring the botanical systematics “-sporites” are often used. Pflug (1953a) intro- and those favouring only the morphological duced for genus names the suffixes “-pollis” nomenclature. and “-sporis” (e.g. Classopollis). When one was using the natural systema- The other trend in nomenclature of the Ter- tics, a morphological similarity of fossil spro- tiary sporomorphs was based solely on the morphs to present-day spores or pollen grains morphological features. The most famous mor- were frequently taken into consideration, what phological systematics was worked out by was reflected by adding to genera or botanical Thomson and Pflug in 1953. According to it the species names the epithet “type” (e.g. Ilex – generic names must contain the basic informa- type, Pinus haploxylon – type, etc.) This mode tion on a sporomorph structure. The first part of classifying the pollen grains and spores of the name informs about the quantity, type would not have given the possibilities to show- and arrangement of the germination appara- ing the full morphological diversity at genera tus (e.g. Triplano-; Tricolporo), and the second level, if one had not shown previously the part of the generic name defines the type of an species similarity. Mostly such pointing is not organ (-sporites or –pollenites). A clear formu- possible because present-day equivalents of lation of principles of this systematics, the uni- fossil morphological types frequently do not vocal genera diagnoses and very rich fossil ma- exist. In such the case a fossil species name terial elaborated enabled the wide spreading creation is necessary. Among advocates of pure of it, especially while older Tertiary deposits botanical nomenclature an extreme tendency were under the work. to giving the species names for fossil grains, The morphological systematics proposed by classified to contemporary botanical genera, Van der Hammen (1956) was close to that of without showing the type of organ, is mani- Thomson and Pflug’s (1953). The only dif- 7 ference was the stressing in the first part of genera, retaining the rule of priority. The lack generic name the character of a sporomorph of proper recommendations of ICBN regarding superficial relief (e.g. Clavatitricolporites). In the fossil remains forced in effect the necessity Europe this systematics did not find too many of using some rules by the International Code followers. of Zoological Nomenclature (ICZN) 1961 (1963) When Neogene palynological deposits are elaborated for fossil fauna. According to the elaborated a precise using of the morphological Code the concept of the type species is a crite- taxonomy solely is insufficient. The possibility rion for a genus distinction. Thomson and of relating of fossil taxa to contemporary Pflug (1953) and Potonié (1956) used the con- families or genera led palynologists to idea of cept of “Genotypus” for evaluation of the basic displaying this affiliation in generic name. The species to a genus distinction. Nowadays, in greatest contribution in the development of the biological literature the concept of “geno- this systematics had Potonié (Synopsis 1956– type” is referred to as a complex of genetical 1970) and Krutzsch (1959–1971). Both investi- features. ICZN 1961 (1963) in Art. 67, rec. 67a gators attempted to set in order hitherto used denied using the name “genotype” in relation divisions in all the systematics, taking into to the type species. consideration the priorities in genus names de- 2. The authors have adopted the rule that if scribed according to rules by the International it is a possibility to identify the morphological Code of Botanical Nomenclature (1988). How- type of pollen grains and spores to a given bo- ever, in spite of these efforts still exist several tanical taxon of various rank (family, subfam- names of genera similarly sounding, often sy- ily, genus, species) then will be also changed nonimical, based on the same type species (for the morphological taxon, so in effect the bo- example “parmularius” species was typical of tanical rank will be clearly visible: Lycopodia- three genera: Cornoidites, Tricolpopollenites ceaesporis – affinity in the rank of family, Cas- and Psilatricolporites). Also, not all palynolog- taneoideaepollis – affinity in the rank of sub- ists creating the new genus names used the family, Selaginellisporis - affinity in the rank same rules; so in the consequence a complete of genus and Selagosporis – affinity in the chaos arose. To help palynologists in this rank of species. “jungle” of genera names Jansonius and Hills 3. While creating new genera the suffixes (1976 – with current supplements) took up an “-sporis”, “-sporites”, “-pollis” and “-pollenites” elaboration of a checklist of sporomorph gen- were used. era from all the geological periods. In this 4. While creating a new name (nomen work every genus have obtained a catalogue novum), to existing names of valid genera the card with the successive number, diagnose, suffix showing the type of organ is added, to schematic drawing, references to literature which this name is relating. For example, the and type species, and sometimes a short auth- genus Milfordia Erdtman 1960 was com- ors’ comment, regarding the importance of a plemented by suffix “-pollis” (Milfordiapollis genus. Erdtman nom. nov.). In the up to date palynological literature, 5. If there were changed the names im- following Potonié’s instructions (Synopsis I–VI properly suggesting affinity to spores, e.g. Spo- 1956–1970), the changes in genus names were rotrapoidites Klaus 1954 to Trapapollis gen. observed by, among others, Nagy (1969; 1985) nov. and Planderová (1990). At the moment there 6. If it was created previously the genus of is a prevailing tendency to creating genera botanical etymology, then fossil species were names, which indicate the botanical affinity. incorporated into it, which had the same bo- Also, there is the same trend in the present tanical affinity, and their features cover a elaboration. genus diagnosis. For example to Aesculidites Elsik 1968 genus a new species A. hippocasta- neoides sp. nov. was incorporated, correspond- RULES ADOPTED FOR TAXA REVISIONS ing morphologically to present-day species Aesculus hippocastanum. 1. The rules by the International Code of 7. If the morphological type of a fossil sporo- Botanical Nomenclature (ICBN) 1988 have ac- morph represents the botanical genus, how- knowledged only formally existing important ever no precisely comparable to the morpho- 8 logy of a given present-day species, then the phological type of pollen grains of “Tilia type” organ name for the genus should be created in Tiliaceae family presently occurs in two and a fossil species incorporated into it. For subfamilies in different genera (in Brownlo- example, from Butomus genus – Butomuspol- wioideae the genera: Berrya, Brownlowia, and lenites longicolpatus (Doktorowicz-Hrebnicka in Tilioideae: Tilia). In this case it was no 1964) comb. nov. possible to use the genera name of family 8. In the case, when the botanical division origin because there are in Tiliaceae family of a family into subfamilies is also visible in also other morphological types of pollen the morphology of sporomorphs, then morpho- grains. logical differences at the rank of a subfamily 12. Morphological name of a genera is re- were taken into consideration. For the Cas- tained if there is a morphological similarity of tanoideae, a subfamily within the Fagaceae, fossil sporomorphs to spores or pollen grains of the genus Castaneoideaepollis was established more than one botanical taxon, e.g. the name and it includes fossil pollen grains of the struc- Tricolporopollenites megaexactus is retained tural type found in all genera of that subfam- because there is no possibility to asses univo- ily. The fossil pollen grains acknowledged to cally the botanical affinity of this fossil species Quercoideae subfamily could not obtain to a given botanical taxon; in this case in Cyri- generic name pointing at this subfamily be- laceae and Clethraceae families as similar cause admitted genus Quercoidites Potonié, morphological type occurs as in Tricolporopol- Thomson & Thiergart 1950 have existed. Ac- lenites megaexactus. cording to the priorite rule pollen grains of Quercoideae subfamily to this unit were ac- The symbols: knowledged. P – generally palaeotropical element 9. Sometimes changes in botanical nomen- P1 – tropical clature at the genus level recall for the P2 – subtropical necessity of fossil genera to be changed. For A – generally arctotertiary element example, Lycopodiumsporites Thiergart 1937 A1 – warm-temperate genus, in spite of the botanical affinity, could A2 – cool-temperate not be retain because Holub (1983) and Oll- Abbrevations “p.p.” means pro parte gaard (1987) divided contemporary Lycopo- dium genus into new genera (Huperzia, Lyco- podiella, Lepidotis and Diphasiastrum). Ac- SYSTEMATIC PART cording to this, for fossil sporomorphs, which morphological equivalents in Lycopodiaceae Cingulisporis Waz˙yn´ska gen. nov. family may be found, but no in given genera, a Polypodiaceoisporites Potonié 1951a ex Potonié 1956 new genus of Lycopodiaceaesporis was created. p.p. Cingulatisporites Thomson 1953 10. The fossil genera of pollen grains or Bifacialisporites Nagy 1963 spores include in the name botanical affinity Type species. Cingulisporis marxhei- lost validity when their type species was im- mensis (Mürriger & Pflug 1952) comb. nov. properly classified botanically. For example, (= Triradiato – sporites marxheimensis Mür- Juglanspollenites verus Raatz 1937 cannot be riger & Pflug 1952, pl. 11, fig. 2; Marxheim, included into Juglans genus because its mor- Upper Oligocene). phological composition points to affinity to Cel- tis genus. In this case Potonié (1960) rule was D e r i v a t i o n o f n a m e. Presence of a cin- obeyed (p. 135 – “if the type species of a genus gulum on the equator. is transferred to another genus the previous D i a g n o s i s. Equatorial outline more or less one lose his validity”. triangular with rounded apices, sides straight 11. While a botanical affinity of a fossil or slightly concave or convex. On the equator a species is designated to present-day species or smooth cingulum, often more narrow on the api- genus the word “type” is added. It means that ces. Trilete mark distinct, arms nearly reach- above all the morphological similarity of fossil ing the equator. Surface on the distal face ru- spores or pollen grains to the taxon of the gate with irregular distributed rugae, often same rank occurs, what not always points to fused together forming a pseudoreticulate pat- the botanical affinity. For example, the mor- tern with irregular anastomosing muri. Sculp- 9 ture on the proximal face more or less similar Botanical affinity. ? Polypodiaceae, ? Pte- to that of the distal face. Equatorial diameter ridaceae, p.p. Pteris type. μ 27–78 m. Geofloristic element. Palaeotropical (P1). Botanical affinity. P.p. Schizaeaceae, Occurrence. Lower Miocene. p.p. Dicksoniaceae, p.p. Polypodiaceae, p.p. R e m a r k s. A new combination is a result of Cyatheaceae, p.p. Pteridaceae the genus change – see remarks in the diag- Geofloristic element. Palaeotropical (P). nosis of genus. The used combination Polypo- R e m a r k s. A new genus based on the mor- diaceoisporites corrutoratus Nagy suggested phological features was created. It was used the botanical affinity of this fossil species to the names of spores of similar morphology: 1. the extant Polypodiaceae. In this family mor- Cingulatisporites Thomson 1953 – the genus phological variability of spores is very high lost validity because its type species C. laevis- and the morphological type C. corrutoratus is peciosus Pflug 1953 was transferred (with mis- within this variability, however C. corrutora- take in the species name C. levis) by Krutzsch tus is comparable to spores of Pteridaceae fam- (1963b) to Leiotriletes genus. 2. Bifacialis- ily. Hence, the botanical affinity of fossil porites Nagy 1963 – according to Potonié species is unknown and to the morphological (1966) it has not the typical cingulum. 3. Poly- type could be only classified. podiaceoisporites Potonié 1951a – it suggests family the botanical affinity to the Polypodia- Cingulisporis gracillimus (Nagy 1963) Wa- ceae improperly because this type of structure z˙yn´ska comb. nov. is found in other extant families fern, e.g. Pl. 1, fig. 1a-c Pteridaceae, Schizaeaceae and Dicksonia- 1963 Polypodiaceoisporites gracillimus n. sp.; Nagy, p. ceae. 398–400, pl. 1, figs 3–6. Botanical affinity. ? Polypodiaceae, ? Schi- Cingulisporis marxheimensis (Mürriger & zaeaceae (Lygodium), ? Pteridaceae, Pteris type. Pflug 1952) Waz˙yn´ska comb. nov. Pl. 1, figs 2a, b; 3a, b Geofloristic element. Palaeotropical (P1). Occurrence. Lower Miocene. 1952 Triradiato – sporites marxheimensis n. sp.; Mürriger & Pflug, p. 57, 61, 64, pl. 11, figs 2–4. R e m a r k s. The new combination was intro- 1953 Cingulatisporites marxheimensis (Mürriger & duced in the result of the genus change. The Pflug) n. comb.; Thomson & Pflug, p. 58, pl. 1, comparison of the C. gracillimus spores with figs 13–15. present-day spores of Schizaeaeceae family 1959 Polypodiaceoisporites marxheimensis (Mürriger & Pflug) n. comb.; Krutzsch, p. 180. (Lygodium) by Nagy (1963) and of Pteridaceae (Pteris) by Planderová (1990) validated the Botanical affinity. P.p. Schizaeaceae, usage of the morphological generic name. p.p. Dicksoniaceae Geofloristic element. Palaeotropical (P1). Cryptogrammasporis Skawin´ska gen. nov. O c c u r r e n c e. Oligocene – Lower Miocene. p.p. Stereisporites Krutzsch 1963b R e m a r k s. A new combination is a result of Type species. Cryptogrammasporis mag- the genus change – see remarks in the diag- noides (Krutzsch 1963b) comb. nov. {= Ste- nosis of genus. Cingulisporis marxheimensis reisporites (Stereigranisporis) magnoides; was chosen as the type species taking into con- Krutzsch 1963b, p. 90, pl. 26, fig. 1–3, Rütter- sideration the distinctive morphological fea- berg, Pliocene}. tures and relatively common, however not too D e r i v a t i o n o f n a m e. After the recent ge- abundant. nus Cryptogramma. D i a g n o s i s. Spores trilete, equatorial outline Cingulisporis corrutoratus (Nagy 1985) triangular with rounded apices. Exospore 3 μm Waz˙yn´ska comb. nov. thick, two-layered with more or less undulated Pl. 1, fig. 4a–c surface. Trilete mark with long laesura arms. 1985 Polypodiaceoisporites corrutoratus n. sp.; Nagy, Proximal face covered by small densely spaced p. 96, pl. 27, figs 14–16, pl. 28, figs 1–6. verrucae, distal face covered by big verrucae, 10

3–4 μm in diameter, densely spaced. Equator- Type species. Lycopodiaceaesporis (Re- ial diameter about 45 μm. titriletes ex Krutzsch 1963a) pseudoclavatus B o t a n i c a l a f f i n i t y. Pteridaceae, Crypto- (Krutzsch 1963a) comb. nov. (= Retitriletes gramma type. pseudoclavatus Krutzsch; Krutzsch 1963a, p. 110, pl. 36, figs 1–5, Gerstungen/Werra, Upper Geofloristic element. Arctotertiary (A). Pliocene). Genus Cryptogramma occurs at present in the temperate zone of northern hemisphere. Derivation of name. After the Lycopo- diaceae family. R e m a r k s. Diagnosis of genus and new com- bination of species were for the first time given D i a g n o s i s. Spores trilete, equatorial outline in the archival doctoral dissertation (K. Ska- rounded-triangular. Leasura arms reaching win´ ska 1989). In species Stereisporis (Sterei- the equator. Exospore thick, two-layered, the granisporis) magnoides Krutzsch, the name of outher layer thicker than the inner. On the genus and subgenus has been changed into distal face the sculpture is varying between ir- Cryptogrammasporis, because this morpho- regular ridges formed by fusing together muri logical type of fossil spores is approximate to (Camarozonosporites and Hamulatisporis sub- spores of the extant genus Cryptogramma genera) to regular reticulate with polygonal (Pteropsida). The form of spore exine in this lumina (Retitriletes subgenus). On the proxi- genus is different from the spores of genus Stere- mal face the sculpture is more or less similar isporites, comparable to Sphagnum (Bryopsida). to that of the distal face, totaly or sometimes partialy covering the surface. In the leasura arms area muri partialy free, anastomosing Cryptogrammasporis magnoides (Krutzsch 1963) Skawin´ska comb. nov. forming an irregular sculpture. Equatorial μ Pl. 2, fig. 1a-c diameter 24–25 m. Botanical affinity. Lycopodiaceae. 1963b Stereisporites (Stereigranisporis) magnoides n. sp.; Krutzsch, p. 90, pl. 26, figs 1–12. Geofloristic element. Arctotertiary 1980 Cryptogramma crispa (Linne) Braun; Shatilova (A)/Palaeotropical (P), today to cosmopolitan & Mtshedishvili, pl. 3, figs 5, 5a. family Lycopodiaceae belong 400, mainly tropi- 1993 Cryptogramma sp. 1; Kohlman-Adamska p. 107, pl. 6, fig. 5 cal, species. R e m a r k s. The genus Lycopodiumsporites D e s c r i p t i o n. Spores trilete, equatorial out- Thiergart 1937, cannot be maintained because line triangular with rounded apices. Trilete of nomenclatural reasons (Krutzsch 1963). mark with straight, 20–23 μm long arms. Holub (1983) and Ollgaard (1987) divided ex- Exospore about 3 μm thick, two-layered, sur- tant genus Lycopodium into several genera. In face verrucate. Verrucae densely spaced on the consequence, for fossil spores which morpho- distal face, 3–4 μm in diameter, on the proxi- logical equivalents may be found in present- mal face smaller. Equatorial diameter of the day family Lycopodiaceae, the new genus Lyco- spore about 46 μm. podiaceaesporis was created. This genus has B o t a n i c a l c a l a f f i n i t y. Pteridaceae, Cryp- three subgenera: two determined by Krutzsch togramma type. 1963a (Camarozonosporites and Hamulatis- Geofloristic element. Arctotertiary (A1). poris) as subgenera within genus Camarozo- Occurrence. Miocene, Pliocene. nosporites, and third, Retitriletes, described by Krutzsch (1963a) as a separate genus. The R e m a r k s. Change of genus results from morphological features of all subgenera are determination of the morphological similarity within morphological variability of spores of of fossil spores to extant spores of genus Cryp- Lycopodiaceae family. The genus Selagosporis togramma, e.g. species: C. crispa, C. stelleri, C. Krutzsch 1963a was not included into newly raddeana. formed genus. The spores of this taxon are morphologically very similar to spores of Lycopodiaceaesporis Waz˙yn´ska gen. nov. species Huperzia selago and have quite differ- Lycopodiumsporites Thiergart 1937 ex Potonie 1956. ent morphology than other genera of Lycopo- Camarozonosporites Pant 1954 ex Potonié 1956. diaceae. Retitriletes Hammen van der 1956 ex Pierce 1961 emend. Döring, Krutzsch, Mai & Schulz in Krutzsch 1963a. 11

Lycopodiaceaesporis (Retitriletes ex Krutzsch Lycopodiaceaesporis (Camarozonosporites 1963) pseudoclavatus (Krutzsch 1963) Wa- ex Krutzsch 1963a) heskemensis (Pflanzl z˙yn´ska comb. nov. 1955) Waz˙yn´ska comb. nov. Pl. 3, fig. 8a, b Pl. 3, fig. 9a, b

1963a Retitriletes pseudoclavatus n. fsp.; Krutzsch, p. 1955 Cingulisporis heskemensis n. sp.; Pflanzl in 110, pl. 36, figs 1–11. Mürriger & Pflanzl, p. 85, 87, pl. 5, figs 1–3. 1985 Lycopodiumsporites pseudoclavatus (Krutzsch) n. 1959 Camarozonosporites heskemensis (Pflanz) n. comb.; Nagy, p. 65, pl. 6, figs 12–15. comb.; Krutzsch, p. 187–188, pl. 38, figs 413– 421. Botanical affinity. Lycopodium sect. Ly- 1961 Lycopodium forma priva; Doktorowicz-Hrebni- copodium, L. clavatum type. cka, p. 192, pl. 3, fig. 26. 1963a Camarozonosporites (Camarozonosporites) he- Geofloristic element. Arctotertiary (A1), skemensis (Pflanzl) Krutzsch; Krutzsch, p. 122, Lycopodium clavatum occurs in mixed forests pl. 42, figs 1–14. of temperate zone and in montane forests of 1964 Lycopodium cernuum L. type; Stuchlik, p. 12, pl. 2, fig. 3. tropical zone. Botanical affinity. Lycopodiella sect. Occurrence. Miocene – Pliocene. Campylostachys, Lycopodiella cernua (=Lyco- R e m a r k s. Change of genus results from the podium cernuum) type. determination of botanical relations of fossil Geofloristic element. Palaeotropical (P1). spores to extant family Lycopodiaceae and genus Lycopodium s.s. (Lycopodium clavatum O c c u r r e n c e. Oligocene – Lower Miocene. type). In the specific name of new taxon the R e m a r k s. As in the species mentioned pre- former genus Retitriletes Krutzsch 1963a, de- viously. scribing the sculpture of spore surface, was maintained as a subgenus. Lycopodiaceaesporis (Hamulatisporis Krutzsch 1959) helenensis (Krutzsch 1963) Waz˙yn´- ska comb. nov. Lycopodiaceaesporis (Camarozonosporites Pl. 4, fig. 1 ex Krutzsch 1963) decorus (Wolf 1934) Wa- z˙yn´ska comb. nov. 1963a Camarozonosporites (Hamulatisporis) helenen- Pl. 3, fig. 7a, b sis n. fsp.; Krutzsch, p. 128, pl. 45, figs 1–7. 1964 Lycopodium sp. 2; Camarozonosporites (Hamu- 1934 Sporites decorus n. sp.; Wolff, p. 67, pl. 5, fig. latisporis) helenensis Krutzsch; Stuchlik, p. 13, 12. pl. 2, fig. 9. 1959 Camarozonosporites decorus (Wolff) n. comb.; Krutzsch, p. 187. Botanical affinity: Lycopodiella sect. ca- 1963a Camarozonosporites (Camarozonosporites) de- roliniana: p.p. L. caroliniana (=Lycopodium corus (Wolff) Krutzsch; Krutzsch p. 126, pl. 44, carolinianum) type. figs 1–13. Geofloristic element. Palaeotropical 1964 Lycopodium sp. 1 cf. Camarozonosporites (Ca- marozonosporites) decorus (Wolff) Krutzsch; (P2)/Arctotertiary (A). Stuchlik, p. 13, pl. 2, fig. 4, 5. Occurrence. Miocene. Botanical affinity. Lycopodiella sect. R e m a r k s. As in the species mentioned pre- Campylostachys, Lycopodiella cernua (= Lyco- viously. podium cernuum) type. Geofloristic element. Palaeotropical (P1). Lycopodiaceaesporis (Hamulatisporis Krutzsch Lycopodiella cernua occurs today in moist 1959) rarus (Doktorowicz-Hrebnicka 1960) Waz˙yn´ska comb. nov. places in the tropical and subtropical zones. Pl. 4, fig. 2 O c c u r r e n c e. Oligocene – Miocene. 1960 Lycopodium forma rara; Doktorowicz-Hrebni- R e m a r k s. Change of genus results from the cka, p. 74, pl. 16, figs 15, 16. determination of botanical affinity of fossil 1963a Camarozonosporites (Hamulatisporis) rarus spores to extant Lycopodiaceae and to genus (Doktorowicz-Hrebnicka) emend. et nov. comb.; Krutzsch, p. 130, pl. 46, figs 1–16. Lycopodiella (L. cernua type). In the specific name of new taxon the genus Camarozonos- Botanical affinity. Lycopodiella sect. ca- porites, reflecting the relief of spore surface, roliniana, p.p. L. caroliniana (= Lycopodium was preserved. carolinianum) type. 12

Geofloristic element. Palaeotropical R e m a r k s. Introduction of new combination (P2), Lycopodiella caroliniana occurs today in of subspecies Osmundacidites primarius re- temperate and tropical climate along the sults from frequent occurrence of especially Eeast and South shores of the U.S.A., in South large spores of this species. America – Guyana, Brasilia, and in South Af- rica (absent in West America, Europe and tem- Selaginellisporis Waz˙yn´ska gen. nov. perate West Asia). Echinatisporis Krutzsch 1959 O c c u r r e n c e. Oligocene – Miocene. Type species. Selaginellisporis (Echina- R e m a r k s. Change of genus results from tisporis Krutzsch 1959) miocenicus (Krutzsch determination of the botanical affinity of fossil & Sontag 1963) comb. nov. (= Echinatisporis spores to genus Lycopodiella (L. caroliniana miocenicus Krutzsch & Sontag; Krutzsch, type) from the family Lycopodiaceae. In the 1963b, p. 110, pl. 36, figs 6–9, II Lusatian specific name of new taxon subgenus Hamula- Seam). tisporis, reflecting the relief of spore surface, D e r i v a t i o n o f n a m e. After the recent ge- was preserved. nus Selaginella. Lycopodiaceaesporis (Retitriletes ex Krutzsch D i a g n o s i s. Spores trilete, equatorial outline 1963) lusaticus (Krutzsch 1963) Ziembin´- rounded to triangular with rounded apices. Te- ska-Tworzydło comb. nov. trade mark with long arms nearly reaching Pl. 3, fig. 11a–c the equator. Exospore thick, two-layered. Dis- tal face covered by varied shape and size pro- 1963a Retitriletes lusaticus Krutzsch; Krutzsch, p. 96, pl. 29, fig. 1–5. cesses, loosely or densely spaced. Proximal face smooth or with some processes. Equator- Botanical affinity. Lycopodium clava- ial diameter without ridges 19–70 μm. tum type. B o t a n i c a l a f f i n i t y. Selaginellaceae, Se- Geofloristic element. Arctotertiary (A1). laginella type. O c c u r r e n c e. Oligocene – Miocene. Geofloristic element. Arctotertiary (A)/Pa- R e m a r k s. As in the species mentioned pre- laeotropical (P). Today the species of genus Se- viously. laginella (more than 700) occur mainly in tropical and subtropical regions except for S. Osmundacidites Couper 1953 selaginoides, a boreal species. Type species. Osmundacidites wellmanii Remarks. The new genus Selaginellisporis Couper 1953 was formed for hitherto described fossil spores morphologically similar to spores of extant genus Selaginella. The genus Echinatisporis, Osmundacidites primarius major (Raatz 1937) Ziembin´ska-Tworzydło comb. nov. described by Krutzsch (1959), reflecting the re- Pl. 4, fig. 10a, b lief of spore surface was preserved as a subge- nus. 1937 Sporites primarius Wolff f. major n. f.; Raatz, p. 13, pl. 1, fig. 14. 1960 Osmunda – Sporites primarius Wolff forma pilo- Selaginellisporis (Echinatisporis Krutzsch sa major; Doktorowicz-Hrebnicka, p. 74, pl. 15, 1959) miocenicus (Krutzsch & Sontag figs 11, 12. 1963) Waz˙yn´ska comb. nov. 1967 Baculatisporites primarius major Raatz; Pl. 5, fig. 4 Krutzsch, p. 56, pl. 10, figs 1–6. 1985 Osmundacidites primarius (Wolff) n. comb.; Na- 1963b Echinatisporis miocenicus n. fsp. (Krutzsch & gy, p. 74, pl. 12, figs 5–6. Sontag); Krutzsch, p. 110, pl. 36, figs 6–13. Botanical affinity. Osmundaceae, Os- B o t a n i c a l a f f i n i t y. Selaginellaceae, Se- munda type. laginella type. Geofloristic element. Arctotertiary (A1). Geofloristic element. Palaeotropical Genus Osmunda with 14 species occurs today (P2)/Arctotertiary (A1). from temperate to tropical zones of northern Occurrence. Miocene. hemisphere. Remarks. Selaginellisporis (Echinatisporis) Occurrence. Miocene. miocaenicus has been chosen as the type 13 species of genus Selaginellisporis because is Tsugaepollenites spectabilis (Doktorowicz- morphologically most resembling the spores of Hrebnicka 1964) Słodkowska comb. nov. present-day genus Selaginella. It occurs in Ne- Pl. 6, fig. 16 ogene layers more frequently than other 1964 Tsuga typ diversifolia Rudolph forma spectabi- species of this genus. lis; Doktorowicz-Hrebnicka, p. 38, pl. 6, fig. 17, 17a. 1971 Zonalapollenites reuterbergensis n. sp. (= Tsuga Selaginellisporis (Echinatisporis Krutzsch reuterbergensis n. sp.); Krutzsch, p. 156, pl. 45, 1959 echinoides (Krutzsch & Pacltová figs 1–4. 1963) Waz˙yn´ska comb. nov. 1974 Zonalapollenites spectabilis (Doktorowicz-Hreb- Pl. 5, fig. 5a, b nicka) n. comb.; Ziembin´ ska-Tworzydło, p. 354, pl. 12, fig. 3. 1963b Echinatisporis echinoides n. fsp. (Krutzsch & Pacltová); Krutzsch, p. 114, pl. 38, figs 1–5. Botanical affinity. Tsuga diversifolia type. B o t a n i c a l a f f i n i t y. Selaginellaceae, Se- Geofloristic element. Arctotertiary (A1). laginella type. Occurrence. Miocene – Pliocene. Geofloristic element. Arctotertiary R e m a r k s. The change of genus results from (A1)/Palaeotropical (P2). determination of the morphological similarity Occurrence. Miocene. of fossil pollen grains to the grains of extant genus Tsuga (T. diversifolia type), which is R e m a r k s. As in the species mentioned pre- connected with restoration of taxon Tsugaepol- viously. lenites and underlines the priority of the nomenclator of the species. Two species known Selaginellisporis (Echinatisporis Krutzsch from the literature Zonalapollenites spectabilis 1959) longechinus (Krutzsch 1959) Waz˙yn´- (Doktorowicz- Hrebnicka) Ziembin´ska-Tworzy- ska comb. nov. dło and Z. reuterbergensis Krutzsch do not Pl. 5, fig. 3 manifest any differences in the morphological 1959 Echinatisporis longechinus n. fsp.; Krutzsch, p. structure. 133, pl. 20, figs 217, 218.; pl. 21, fig. 219; pl. 22, figs 229–233. Tsugaepollenites spinosus (Doktorowicz- B o t a n i c a l a f f i n i t y. Selaginellaceae, Se- Hrebnicka 1964) Słodkowska comb. nov. laginella type. Pl. 6, fig. 12 Geofloristic element. Palaeotropical (P1). 1964 Tsuga typ diversifolia Rudolph forma spinosa; Occurrence. Eocene – Lower Miocene. Doktorowicz-Hrebnicka, p. 38, pl. 9, fig. 39. 1971 Zonalapollenites spinulosus n. sp. (=Tsuga spi- R e m a r k s. As in the species mentioned pre- nulosa n. sp.); Krutzsch, p. 148, pl. 41, figs 1–10. viously. 1974 Zonalapollenites spinosus (Doktorowicz-Hrebni- cka) n. comb.; Ziembin´ ska-Tworzydło, p. 353, pl. 12, fig. 1a, b. Tsugaepollenites Potonie & Venitz 1934 ex 1985 Tsugaepollenites spinulosus (Krutzsch) n. comb.; Potonié 1958 Nagy, p. 136, pl. 47, fig. 2. Type species. Tsugaepollenites igniculus Botanical affinity. Tsuga diversifolia type. (Potonié 1931) Potonie & Venitz 1934 ex Po- Geofloristic element. Arctotertiary (A1). tonié 1958 Today Tsuga diversifolia occurs in montane Botanical affinity. Pinaceae, Tsuga type forests of Japan. Geofloristic element. Arctotertiary (A1). Occurrence. Miocene – Pliocene. The genus Tsuga with 14 species occurs on R e m a r k s. The change of genus results from northern hemisphere in the Himalayas, Japan the determination of morphological similarity and North America. of fossil pollen grains to pollen grains of extant R e m a r k s. Because of similar morphology of genus Tsuga (T. diversifolia type) which is con- fossil pollen grains to the grains of extant nected with restoration of taxon Tsugaepolle- genus Tsuga the taxon Tsugaepollenites Po- nites and underlines the priority of the author tonié et Venitz 1934 have been restored replac- of the species. Two species known from the lit- ing later morphological genus (younger sy- erature Zonalapollenites spinosus (Doktoro- nonym) Zonalapollenites Pflug 1953. wicz-Hrebnicka 1964) Ziembin´ska-Tworzydło 14

1974 and Z. spinulosus Krutzsch 1971 do not 1970a Aglaoreidia cyclops Erdtman; Krutzsch, p. 88, manifest distinct differences in morphological pl. 15, figs 1–20. structure. Botanical affinity. Unknown, ? Ruppi- aceae, ? Amarylidaceae, ? Liliaceae. Keteleeriapollenites Nagy 1969 Geofloristic element. Palaeotropical (P1). Type species. Keteleeriapollenites komlon- O c c u r r e n c e. Lower Eocene – Lower Oligocene. sis Nagy 1969 R e m a r k s. Nomen novum in the name of species is the result of changes in genus by ad- Keteleeriapollenites dubius (Chlonova 1960) Słodkowska comb. nov. dition of the suffix -pollis to the name Ag- Pl. 7, fig. 6 laoreidia.

1960 Keteleeria dubia n. sp.; Chlonova, p. 59, pl. 9, fig. 5. Graminidites Cookson 1947 ex Potonié 1960 1971 Abiespollenites dubius (Chlonova) n. comb. (= ? Keteleeria dubia Chlonova 1960); Krutzsch, p. Type species. Graminidites media Cook- 98, pl. 21, figs 1–10. son 1947 Botanical affinity. Pinaceae, Keteleeria type. Geofloristic element. Arctotertiary (A1). Graminidites bambusoides Stuchlik sp. nov. The genus Keteleeria (4 species) occurs today Pl. 8, figs 4, 5 in mixed forests of SE China. Occurrence. Miocene. 1964 Gramineae “Bambusa” type; Stuchlik, p. 76, pl. 24, figs 1, 2. R e m a r k s. The change of genus results from 1970a Graminidites sp. A (Bambusa – typus); determination of the botanical affinities of fos- Krutzsch, p. 51, pl. 1, fig. 1. sil pollen grains to pollen grains of genus Kete- 1990 Graminidites cf. sp. A Bambusa type (Stuchlik) leeria. They are very large (> 100 μm), the corpus n. comb.; Planderová, p. 54, pl. 24, figs 1, 2. is round and of relatively thin exine, a reticu- H o l o t y p e. Pl. 8, fig. 5 (= Gramineae “Bam- lum of the same type both on the corpus and busa” type; 1964 Stuchlik, pl. 24, fig. 1) on the sacs (but on sacs with much larger L o c u s t y p i c u s. Borehole Rypin lumina) and there is no crest on the corpus, so typical of the pollen grains of Abies. S t r a t u m t y p i c u m. Middle Miocene D e r i v a t i o n o f n a m e. After the recent Aglaoreidiapollis Grabowska nom. nov. genus Bambusa. Aglaoreidia Erdtman 1960 D i a g n o s i s. Pollen grains oval to spheroidal, μ Type species. Aglaoreidiapollis cyclops polar axis 50–60 m. On the distal pol a pore μ (Erdtman 1960) nom. nov. (= Aglaoreidia cy- about 4 m in diameter with an annulus about μ μ clops Erdtman 1960) 4 m broad. Exine very thin (less than 1 m with secondary falts, surface psilate to finely Botanical affinity. Unknown. granulate. Geofloristic element. Palaeotropical (P). Botanical affinity. Poaceae, Bambusa type. R e m a r k s. Nomen novum has been formed Geofloristic element. Palaeotropical (P2). by addition to the name of taxon Aglaoreidia Today the genus Bambusa (75 species) occurs an suffix -pollis, in order to show the character in tropical and subtropical Asia and America. of organ. Both the diagnosis of genus according to Erdtman 1960, supplemented by Krutzsch Occurrence. Miocene. 1970a, and the type species are left without R e m a r k s. The name given by Stuchlik changes. (1964) and other authors was not valid in the light of regulations of ICBN. Now the present- Aglaoreidiapollis cyclops (Erdtman 1960) day name Bambusa has been replaced by ad- Grabowska nom. nov. jective form in the name of fossil species bam- Pl. 8, fig. 7 busoides. Similar pollen grains have also been 1960 Aglaoreidia cyclops n. sp.; Erdtman, p. 47, pl. 1, observed in other extant grass genera, e.g. figs b, c. Oryza, Clidonia and Dendrocalamus. 15

Milfordiapollis Grabowska nom. nov. 1960 Milfordia hypolaenoides n. sp.; Erdtman, p. 46, pl. 1, fig. a. p.p. Inaperturopollenites Pflug & Thomson 1953 1961 Milfordia incerta (Thomson & Pflug) n. comb.; p.p. Monoporopollenites Meyer 1956 ex Potonie 1960 Krutzsch, p. 325 Milfordia Erdtman 1960 1970a Milfordia incerta (Thomson & Pflug) Krutzsch; Restionidites Elsik 1968 Krutzsch, p. 72, pl. 9, figs 1–26. Centrolepidacites Wu Zuo-ji & Yu Jin-feng 1981 Botanical affinity. P.p. Restionaceae, Type species. Milfordiapollis incertus p.p. Centolepidaceae. (Pflug & Thomson 1953) Krutzsch 1961 nom. nov. (=Inaperturopollenites incertus Pflug & Geofloristic element. Palaeotropical (P1). Thomson 1953 p. 66, pl. 5, fig. 34) Occurrence. Lower Eocene – Middle Miocene. D e r i v a t i o n o f n a m e. After a locality in R e m a r k s. Nomen novum for this species England (Erdtman 1960) suffix pollis indicates with pores of “centrolepidoid” type is con- the plant organ. nected with the genus which does not show Botanical affinity. P.p. Restionaceae, proper botanical affinities because this mor- p.p. Centrolepidaceae, p.p. Flagellariaceae. phological type of pollen grains occurs in the families Restionaceae and Centrolepidaceae. Geofloristic element. Palaeotropical (P). Families Restionaceae, Flagellariaceae and the majority of genera of Centrolepidaceae occur Milfordiapollis hungaricus (Kedves 1965) Krutzsch & Vanhoorne in Krutzsch 1970 in tropical part of southern hemisphere (Africa, Grabowska comb. nov. Australia, New Zealand, Chile). Some genera of Pl. 8, fig. 11 Centroleidaceae grow in SE China and Malesia. R e m a r k s. Nomen novum has been formed 1965 Monoporopollenites hungaricus n. fsp.; Kedves, p. 50, 51, figs 1–6. by addition of the suffix -pollis to the name 1968 Restioniidites hungaricus (Kedves) n. comb.; Milfordia, in order to show the character of Elsik, p. 313, pl. 15, fig. 13. organ. The morphological genera, to which the 1970a Milfordia hungarica (Kedves) Krutzsch & Van- type species M. incertus was included, have hoorn; Krutzsch, p. 74, 75, pl. 10, figs 1–3. been rejected: 1. Inaperturopollenites Pflug & Botanical affinity. P.p. Restionaceae, Thomson because its morphological structure p.p. Centrolepidaceae, p.p. Flagellariaceae did not correspond with morphology of the Geofloristic element. Palaeotropical (P1). type species; 2. Monoporopollenites Meyer be- cause this is collective genus; 3. Genera Restio- Occurrence. Lower Eocene – Lower Oligo- niidites Elsik and Centrolepidacites Wu Zuo-ji cene. & Yu Jin-feng, connected with extant families, R e m a r k s. Nomen novum for this species have not been accepted because the pollen with pores of “graminoid” type is connected morphology in both families is very similar. with new name of genus which does not show Pores are both of “centrolepidoid” and “grami- a proper botanical affinity. This morphological noid” type (Chanda 1966, Ladd 1977), while in type of pollen grains occur in three extant related family Flagellariaceae only “grami- families: Restionaceae, Centrolepidaceae and noid”. Exine has always tectum perforatum. Flagellariaceae. All these features occur in species of genus Milfordiapollis. Hence, this taxon cannot be Milfordiapollis minimus Krutzsch 1970 included into particular family. Grabowska comb. nov. Diagnosis of the genus (Erdtman 1960) with Pl. 8, fig. 10a, b later supplement (Krutzsch 1970a) and the type species M. incerta, chose and described by 1970a Milfordia minima n. sp.; Krutzsch, p. 76, pl. 10, figs 1–34. Krutzsch 1961, have been left without changes. 1974 Restioniidites minimus (Krutzsch) n. comb.; Kedves, p. 26, pl. 11, figs. 1–3. Milfordiapollis incertus (Pflug & Thomson Botanical affinity. P.p. Restionaceae, 1953) Krutzsch 1961 Grabowska nom. nov. p.p. Centrolepidaceae, p.p. Flagelariaceae. Pl. 8, figs 8, 9 Geofloristic element. Palaeotropical (P1). 1953 Inaperturopollenites incertus n. sp. (Pflug & Thomson); Thomson & Pflug, p. 66, pl. 5, figs O c c u r r e n c e. Eocene – Oligocene, Lower Mio- 31–36. cene. 16

R e m a r k s. As for the former species. Botanical affinity. Corylus avellana type. Geofloristic element. Arctotertiary (A2). Corylopollis Ziembin´ska-Tworzydło gen. nov. Occurrence. Oligocene – Pliocene. non Coryli?-pollenites Potonié 1934 R e m a r k s. The change of genus results from non Corylus-pollenites Raatz 1937 determination of the botanical affinities of fos- non Corylus-pollenites Thiergart 1937 (nomen nudum) sil pollen grains to the grains of extant genus Coryloidites Potonié, Thomson & Thiergart 1950 p.p. Triporopollenites Pflug & Thomson 1953 Corylus.

Type species. Corylopollis coryloides Myricipites Wodehouse 1933 (Pflug 1953b) comb. nov. (= Triporopollenites coryloides Pflug 1953b; Thomson & Pflug Type species. Myricipites dubius Wode- 1953, p. 84, pl. 9, fig. 20, Wallensen, Middle house 1933 Pliocene). Myricipites microcoryphaeus (Potonié 1931) D e r i v a t i o n o f n a m e. After the extant ge- Słodkowska comb. nov. nus Corylus. Pl. 8, figs 23, 24 D i a g n o s i s. Pollen grains triporate, equator- 1931a Pollenites microcoryphaeus n. sp.; Potonié, p. ial outline triangular, apertures on the apices 332, pl. 2, fig. 13. circular or slightly lolongate. Equatorial 1950 Engelhardtioidites microcoryphaeus Potonié; diameter 23–35 μm, polar axis about 10 μm Potonié & al., p. 51, pl. B, fig. 8; pl. C, fig.16. 1953 Triatriopollenites coryphaeus (Potonié) subsp. long. Exine tectate, tectum smooth to provided microcoryphaeus (Potonié); Thomson & Pflug, with very fine irregular spaced processes. En- p. 80, 81, pl. 8, figs 38–63. dexine forming a small annulus around the 1990 non Engelhardtioidites microcoryphaeus (Poto- pore. nié) Potonié; Planderová, p. 214, pl. 61, figs 1–19 B o t a n i c a l a f f i n i t y. Betulaceae, Corylus Botanical affinity. Myrica type. type. Geofloristic element. Palaeotropical (P2). Geofloristic element. Arctotertiary (A). Presently Myrica genus (54 species) occurs Today Corylus (15 species) occurs in temperate mostly in subtropical zone, few species in tem- and warm-temperate zones of Europe, North perate zone and moderately warm-temperate America and Asia. zone of northern hemisphere. Remarks. Triporopollenites coryloides Pflug O c c u r r e n c e. Eocene – Middle Miocene. 1953b possessing features similar to pollen Remarks. Myrcipites microcoryphaeus has a grains of Corylus has been taken as type morphological composition pointing at greater species. From very similar morphologically similarity to pollen grain of Myrica genus genus Myricipites it differs in the structure of (clearly visible atrium, the thick exine), than germinal apparatures. Described hitherto gen- to Engelhardtia. In the light microscope, in era Coryli?-pollenites Potonié 1934 and Cory- specimens determined as Myricipites microco- lus-pollenites Raatz 1937 have been admitted ryphaeus, the punctate tectum is not visible, by Potonié (1960) as younger synonyms of so characteristic of Juglandaceae family. There genus Myricipites Wodehouse 1933. Coryli?- is a feature, which served as the base for pollenites corypheus Potonié, later transferred transferring the species from Engelhard- to genus Myriciptes, was the type species for tioidites genus to Myrcipites one. both genera. To the genus Coryloidites its authors Potonié, Thomson & Thiergart (1950) Celtipollenites Nagy 1969 did not included any species. Type species. Celtipollenites komlonsis Nagy 1969 Corylopollis coryloides (Pflug 1953) Ziem- bin´ ska-Tworzydło comb. nov. Celtipollenites verus (Raatz 1937) Ziem- Pl. 8, figs 17, 18 bin´ ska-Tworzydło comb. nov. Pl. 9, figs 11, 12 1953b Triporopollenites coryloides n. sp. (Pflug); Thomson & Pflug, p. 84, pl. 9, figs 20–24. 1937 Juglans-pollenites verus n. sp.; Raatz, p. 18, pl. 1960 Triporopollenites coryloides Pflug emend. 1, fig. 9. 1961 Carpinus forma naturalis; Dokto- Potonié; Potonié, p. 116, 117. rowicz-Hrebnicka, p. 226, pl. 11, fig. 143. 17

1973 Celtis vera (Raatz) n. comb.; Lubomirova & Ti- very short polar axis. Equatorial outline trian- moshkina, p. 14, pl. 1, figs 6–8. gular with slightly thickened pore area. Exine Botanical affinity. Ulmaceae, Celtis type. less than 2 μm thick, verrucate, verrucae about 3 μm in diameter. Equatorial diameter Geofloristic element. Arctotertiary (A1), of pollen grain about 30 μm. the genus Celtis (80 species) occurs presently in subtropical zone, and only one species in Botanical affinity. Haloragaceae, Halo- temperate zone. ragis (=Hallorrhagis) type. Occurrence. Miocene – Pliocene. Geofloristic element. Palaeotropical (P2). R e m a r k s. The common features of fossil pol- Presently Haloragis (60 species) occurs mainly len graines of Juglanspollenites verus and the on southern hemisphere and on northern in grains of Celtis genus (Lubmirova & Timoshki- S-E Asia. na 1973) enable the species to the Celtipolle- Occurrence. Middle Miocene. nites Nagy genus be incorporated. Because the R e m a r k s. The genus Halorrhagoidites type species for the Juglanspollenites genus Stuchlik 1964 is no longer valid because it was was J. verus, when transferred to other ones, not definied, however species features of H. the genus Juglanspollenites was no longer triporatus Stuchlik 1964 cover the diagnosis of valid. the genus Haloragacidites Couper 1953.

Haloragacidites Couper 1953 Juglandipollis Kohlman-Adamska gen. nov. Type species. Haloragacidites trioratus p.p. Juglanspollenites Raatz 1937 Couper 1953 p.p. Multiporopollenites Pflug 1953b non Juglandipites Simpson 1961 Haloragacidites stephanoporatus (Stuchlik Type species. Juglandipollis juglan- 1964) Stuchlik comb. nov. doides sp. nov. Pl. 9, fig. 16 D e r i v a t i o n o f n a m e. After the recent 1964 Halorrhagoidites stephanoporatus n. sp.; Stu- chlik, p. 52–53, pl. 16, figs 7, 8. genus Juglans D i a g n o s i s. Pollen grains polyporate sphe- D e s c r i p t i o n. Pollen grains 6-porate, equa- roidal to oval. Pores circular to oval dis- torial outline hexagonal. Pores in the equator tributed in the equator area and on one hemis- are on apices with an annulus about 5 μm phere of the pollen grain. Exine two-layered, broad. Exine about 3 μm thick, two layered, ectexine thicker than endexine, in the pore verrucate. Verrucae on their tops flatty trun- area somewhat swollen, forming more or less cate, densely and irregularly spaced. Equator- distinct annulus. Surface tectate, tectum pro- ial diameter 25 μm. vided with spinulae. Equatorial diameter 30– Botanical affinity. Haloragaceae, Halo- 50 μm. ragis type. B o t a n i c a l a f f i n i t y. Juglandaceae, Ju- Geofloristic element. Palaeotropical (P2). glans type. Occurrence. Middle Miocene. Geofloristic element. Arctotertiary (A), R e m a r k s. The genus Halorrhagoidites Stu- presently the Juglans genus (15 species) oc- chlik 1964 is no longer valid because it was curs from southern Europe to eastern Asia and not definied, however species features of H. in the South and North America. stephanopratus Stuchlik 1964 cover the diag- R e m a r k s. There are known three fossil pol- nosis of the genus Haloragacidites Couper len genera, which botanical names are con- 1953. nected with Juglans genus. 1. Juglanspolle- nites is no longer valid because the type Haloragacidites triporatus (Stuchlik 1964) species Juglanspollenites verus Raatz 1937 Stuchlik comb. nov. considered as the Celtis (Lubmirova & Timosh- Pl. 9, fig. 17a, b kina 1973) was transferred to the genus Celti- 1964 Halorrhagoidites triporatus n. sp.; Stuchlik, p. pollenites Nagy 1969. 2. Juglandipites with 52, pl. 16, fig. 6. the type species J. magniforamina Simpson D e s c r i p t i o n. Pollen grains triporate with 1961 was created on the basis of a single spe- 18 cimen and according to Potonié (1970) is a sy- Geofloristic element. Arctotertiary (A1). nonym of Juglanspollenites genus. 3. There is Presently Juglans regia occurs from the me- no longer valid morphological genus Multipo- diterranean areas of Europe to Eastern Asia, ropollenites Pflug 1953 because its type species and J. sigillata in China (the Yunnan Prov- M. maculosus morphologically resembles to ince). pollen grains of present-day Juglans genus, O c c u r r e n c e. Oligocene – Miocene. and it was transferred to a new ones of R e m a r k s. Change of genus results from the Juglandipollis. determination of morphological similarity of fossil grains of Multiporopollenites to pollen of Juglandipollis juglandoides Kohlman- Juglans genus. Transferring the type species Adamska sp. nov. Multiporopollenites maculosus to Juglanspol- Pl. 9, figs 18, 19 lenites genus (presently Juglandipollis) by 1993 Juglans Linné – type 1; Kohlman-Adamska, p. Nagy (1985) makes the morphological genus of 133, pl. 17, fig. 11. Multiporopollenites no longer valid. H o l o t y p e. Pl. 9, fig. 19. Butomuspollenites (Doktorowicz-Hrebnicka L o c u s t y p i c u s. Borehole Kosztowo. 1957) Ziembin´ska-Tworzydło emend. T y p e h o r i z o n. Middle Miocene (S´ cinawa p.p. Monocolpopollenites Pflug & Thomson 1953 Formation). p.p. Arecipites Krutzsch 1970a D e r i v a t i o n o f n a m e. After the recent ge- Type species. Butomuspollenites mono- nus Juglans. colpatus Doktorowicz-Hrebnicka 1957 D i a g n o s i s. Pollen grains polyporate, oval to emend. (= Butomus – Pollenites monocolpatus spheroidal, diameter 30–40 μm. Pores circular f. nov.; Doktorowicz-Hrebnicka, 1957 p. 111, pl. 2–4 μm in diameter, distributed in the equator 22, figs 11, 12, Orłowo, Poznan´ Clay, Upper area and on one hemisphere. Exine two- Miocene). layered about 2 μm thick, ectexine thicker D e r i v a t i o n o f n a m e. After the recent than endexine, tectate, in the pore are some- genus Butomus. μ what swollen, up to 2.5 m forming a distinct D i a g n o s i s. Pollen grains monocolpate, annulus. usually asymetric. Equatorial outline in polar Botanical affinity. Juglans type (J. cine- view irregular ellipsoidal. Colpus on the distal rea, J. nigra). side (anacolpate) narrow, sharpened at the en- Geofloristic element. Arctotertiary (A1). dings. Exine two-layered, ectexine semitectate, Extant species Juglans cinerea and J. nigra reticulate, lumina polygonal to more or less occur in warm-temperate zone of the North circular, muri distinctly simplibaculate. Lumi- America. na diameter bigger on the proximal side, de- creasing on the distal side in the colpus area. Occurrence. Miocene – Pliocene. Equatorial diameter 25–45 μm. Remarks. Juglandipollis juglandoides dif- Botanical affinity. Butomaceae, Buto- fers from J. maculosus having less number of mus type. pores and more clearly visible the ring sur- rounding them. Geofloristic element. Arctotertiary (A). Presently a monotype family Butomaceae with Juglandipollis maculosus (Potonié 1931) the species Butomus umbellatus occurs in wet- Kohlman-Adamska comb. nov. lands and watersides in Europe and Asia. Pl. 9, fig. 20 R e m a r k s. The generic name established by Doktorowicz-Hrebnicka (1957) clearly points 1931b Pollenites maculosus n. sp.; Potonié, p. 27, pl. 2, fig. V19d. to the botanical affinity of fossil pollen grains 1953 Multiporopollenites maculosus (Potonié) n. of this type to pollen of Butomus genus. The comb.; Thomson & Pflug, p. 94, pl. 10, fig. 95. generic name Arecipites Wodehouse 1933 pro- 1985 Juglanspollenites maculosus (Potonié) n. comb.; posed by Krutzsch (1970a) for species A. buto- Nagy, p. 204 pl. 115, fig. 15. moides, A. monocolpatus and A. longicolpatus Botanical affinity. Juglans type (J. re- suggests a botanical affinity to palms (Areca- gia, J. sigillata). ceae). Nichols et al. (1973) transferred A. buto- 19 moides to the Monocolpopollenites genus. Tak- Butomuspollenites longicolpatus (Krutzsch ing into consideration the morphological simi- 1970) Ziembin´ska-Tworzydło comb. nov. larity of fossil grains to the grains of present- Pl. 10, fig. 14a, b day Butomus genus, these species were finally 1970a Arecipites longicolpatus n. sp.; Krutzsch, p. 112, included into the Butomuspollenites genus. pl. 25, figs 1–13. The features differing certain species each of Botanical affinity. Butomus type. other are structure of the reticulum, size of the lumina and size of the specimens. Geofloristic element. Arctotertiary (A2). Occurrence. Miocene – Pliocene. Butomuspollenites monocolpatus (Doktoro- R e m a r k s. Change of generic name results wicz-Hrebnicka 1957) Ziembin´ska-Tworzy- from the determination of botanical affinity of dło emend. fossil pollen grains to Butomus genus. Pl. 10, fig. 13a, b

1957 Butomus – Pollenites monocolpatus f. nov.; Dok- Aceripollenites Nagy 1969 torowicz-Hrebnicka, p. 111, pl. 22, figs 11, 12. 1970a Arecipites monocolpatus (Doktorowicz-Hrebni- Type species. Aceripollenites reticulatus cka) n. comb.; Krutzsch, p. 118, pl. 25, figs 14, Nagy 1969 15. Aceripollenites palmatoides Skawin´ska sp. D i a g n o s i s emended. Pollen grains monocol- nov. pate, equatorial outline irregular, 35–45 μm in Pl. 11, fig. 8a–c diameter. Colpus long, overlapping the outline forming a characteristic appendix-like out- 1985 Acer palmatum type; Skawin´ ska p. 108–109, pl. 1, growth. Exine two-layered, ectexine semitec- figs 1–10. tate, reticulate. Reticulum with irregular cir- H o l o t y p e. Pl. 11, fig. 8a–c. cular lumina. On the proximal side lumina L o c u s t y p i c u s. Borehole Ostrzeszów GEO3. diameter biger decreasing towards the colpus. S t r a t u m t y p i c u m. Middle Miocene (Poz- Botanical affinity. Butomus type. nan´ Formation). Geofloristic element. Arctotertiary (A2). D e r i v a t i o n o f n a m e: After the recent O c c u r e n c e. Upper Miocene – Pliocene. species Acer palmatum. R e m a r k s. The species established by Dokto- D i a g n o s i s. Pollen grains tricolporate, pro- rowicz-Hrebnicka (1957) has the morphologi- late-spheroidal with long colpi nearly meeting cal features resemble pollen grains of Butomus each on the pole. Pores lolongate 8 x 3 μm. genus, and it is distinguished as the type Exine two-layered about 1.5 μm thick, ectexine species of Butomuspollenites genus. Krutzsch twice thick as endexine, semitectate striato-re- (1970a) reserved about too large size of pollen ticulate. Striae distinct, narrow, running paral- grains of this species relating to pollen of ex- lel to each other, anastomosing, forming finere- tant Butomus genus. ticulate pattern polar axis 31–33 μm, equatorial diameter 20–30 μm. Butomuspollenites butomoides (Krutzsch Botanical affinity. Aceraceae, Acer pal- 1970) Ziembin´ska-Tworzydło comb. nov. matum type. Pl. 10, fig. 12a, b Geofloristic element. Arctotertiary (A1), 1970a Arecipites butomoides subsp. butomoides; recently Acer palmatum occurs in warm-tem- Krutzsch, p. 112, pl. 24, figs 12–29. perate zone of Japan. 1973 Monocolpopollenites butomoides (Krutzsch Occurrence. Middle Miocene 1970) n. comb.; Nichols & al., p. 252. R e m a r k s. The species name was introduced Botanical affinity. Butomus type. for the first time and the species described in the Geofloristic element. Arctotertiary (A2). archival doctoral dissertation (Skawin´ ska 1989). Occurrence. Miocene – Pliocene. Aesculidites Elsik 1968 R e m a r k s. Change of generic name results from the determination of botanical affinity of Type species. Aesculidites circumcristatus fossil pollen grains to Butomus genus. (Fairchild 1966) Elsik 1968 20

Aesculidites hippocastaneoides Sadowska the Caprifoliaceae family. C. viburnoides is the sp. nov. species sensu lato covers fossil pollen grains Pl. 11, figs 6, 7 resembling several extant species of Viburnum, 1984 Aesculus; Jahn & al., pl. 9, figs 16–26. e.g. V. odoratissimum, V. carlesi, V. lentago.

H o l o t y p e. Pl. 11, fig. 7. Castaneoideaepollis Grabowska gen. nov. L o c u s t y p i c u s. Outcrop Kłodzko. Castanea-pollenites Raatz 1937 Stratum typicum. Pliocene. Castanopsis?-pollenites Thiergart 1937 Castaneoidites Potonié, Thomson, Thiergart 1950 D e r i v a t i o n o f n a m e. After the recent Castaneoipollenites Potonié 1951b species Aesculus hippocastanum Cupuliferoipollenites Potonié 1951 ex Potonié 1960 D i a g n o s i s. Pollen grains tricolporate, isopo- p.p. Tricolporopollenites Pflug & Thomson 1953 lar, in equatorial view oval with narrow Type species. Castaneoideaepollis ovi- μ rounded pole fields. Polar axis 20–30 m long. formis (Potonié 1934) comb. nov. (= Polle- Colpi long, narrow with circular pores in the nites oviformis Potonié 1934, p. 95, pl. 5, fig. equator area. Along the colpi ectexine provided 27, Geiseltal, Eocene) with small spinules. Between the colpi exine D e r i v a t i o n o f n a m e. After the recent psilate or finely granulate. subfamily Castaneoideae (Fagaceae). Botanical affinity. Hippocastaneaceae, D i a g n o s i s. Pollen grains tricolporate pro- Aesculus hipocastanum type. late (polar axis to 25 μm, equatorial diameter Geofloristic element. Arctotertiary (A1), to 12 μm). Outline in equatorial view oval to presently Aesculus hippocastanum occurs in broadly oval. Colpi narrow nearly parallel not montane areas of the Balkan Peninsula. meeting in the pole, with pores in the equator. Occurrence. Miocene – Pliocene Pores more or less circular to slightly lolong- R e m a r k s. The fossil pollen grains are mor- ate. Exine two-layered rather thin, ectexine as phologically related to pollen of extant species thick as endexine, tectate, tectum psilate or so. Aesculus hippocastanum. This is relatively B o t a n i c a l a f f i n i t y. Fagaceae, Castaneoi- often element of Pliocene floras of Europe. deae (Castanea, Castanopsis, Passania, Litho- carpus types) Caprifoliipites Wodehouse 1933 Geofloristic element. Palaeotropical (P). Type species. Caprifoliipites viridiflumi- Recently members of the Castaneoideae sub- nipites Wodehouse 1933 family occur mainly in subtropical and tropical zones of Asia and less often in North America. Caprifoliipites viburnoides (Gruas – Cavag- R e m a r k s. The genera which names were netto 1978) Kohlman-Adamska comb. nov. connected with the botanical affinity were re- Pl. 12, figs 12a, b; 13a, b. fused: 1. Castaneapollenites Raatz 1937 and 1978 Tricolporopollenites viburnoides n. fsp. Gruas – Castaneoidites Potonié, Thomson & Thiergart Cavagnetto, p. 36, pl. 14, figs 16–19. 1950 because for both taxa the type species 1993 Viburnum L. – typ; Kohlman-Adamska, p. 154, was Pollenites exactus Potonié 1931d. 2. The pl. 33, figs 7a,b; 8a,b. Castaneoipollenites Potonié 1934 is nomen B o t a n i c a l a f f i n i t y. Caprifoliaceae, Vibur- nudum and synonym for Castaneoidites Po- num type. tonié, Thomson & Thiergart. 3. Castanopsis? - pollenites Thiergart 1937 was no validated by Geofloristic element. Arctotertiary (A1). Potonié 1960. 4. Cupuliferoipollenites Potonié Presently the genus Viburnum (200 species) 1951 originates from the old name Cupuliferae occurs in the temperate and subtropical zones used for Fagaceae family. This family comprises of northern hemisphere, mainly in Asia and three subfamilies: Castaneoideae, Fagoideae, North America. and Quercoideae, and each of them is charac- Occurrence. Eocene – Pliocene. terized by different morphology of pollen grains. Remarks. Tricolporopollenites viburnoides 5. For fossil grains with structure similar to pol- Gruas-Cavagnetto is included to the Caprifo- len of Castaneoideae subfamily a new genus liipites genus considering its great similarity Castaneoideaepollis was created. As the type to present-day pollen grains of Viburnum of species C. oviformis (Potonié 1934) was chosen. 21

Castaneoideaepollis oviformis (Potonié 1985 Cercidiphyllum sp.; Skawin´ ska, p. 109, pl. 2, figs 1934) Grabowska comb. nov. 1–3. Pl. 12, fig. 8 B o t a n i c a l a f f i n i t y. Cercidiphyllaceae, Cer- 1934 Pollenites oviformis n. sp.; Potonié, p. 95, pl. 5, cidiphyllum type. fig. 27. 1951a Cupuliferoipollenites oviformis Potonié; Poto- Geofloristic element. Arctotertiary (A1). nié, pl. 20, fig. 70. Cercidiphyllum (2 species) occurs in warm- 1953 Tricolporopollenites cingulum (Potonié) n. temperate zone of Eastern Asia. comb. subsp. oviformis (Potonié) n. comb.; Thomson & Pflug, p. 100, pl. 12, figs 42–49. Occurrence. Miocene. Botanical affinity. Castaneoideae, Cas- Remarks. Tricolpollenites minimireticulatus tanea, Castanopsis, Passania and Lithocarpus Trevisan according to the suggestion of Ska- types. win´ ska has been included into Cercidiphyllum genus of Cercidiphyllaceae family. Trevisan Geoflorictic element. Palaeotropical (P2). (1967) improperly established the botanical af- O c c u r r e n c e. Paleocene – Miocene. finity to Dioscoreaceae, the Monocotyledone R e m a r k s. Change of genus results from the class, which do not posses tricolpate pollen determination of morphological similarity of type. pollen grains to the grains of extant subfamily Castaneoideae. Clerodendrumpollenites Skawin´ska gen. nov.

Castaneoideaepollis pusillus (Potonié 1934) Type species. Clerodendrumpollenites Grabowska comb. nov. microechinatus sp. nov. Pl. 12, figs 6, 7 D e r i v a t i o n o f n a m e. After the recent 1934 Pollenites quisqualis pusillus n. f.; Potonié, p. genus Clerodendrum. 71, pl. 3, fig. 21. 1951b Cupuliferoipollenites pusillus Potonié; Potonié, D i a g n o s i s. Pollen grains tricolporate, in p. 227, pl. 1, fig. 21. equatorial view oval to broadly oval, polar axis 1953 Tricolporopollenites cingulum (Potonié) n. 65–68 μm long. Colpi narrow nearly parallel or comb. subsp. pusillus (Potonié) n. comb.; Thom- somewhat arcuated. Exine two-layered, ectex- son & Pflug, p. 100, pl. 12, figs 28–41. ine thicker than endexine, tectate. Tectum Botanical affinity. Castaneoideae, Cas- supported by distinct columellae, provided tanea, Castanopsis, Lithocarpus, Passania with regularly spaced small spines (microechi- types. nate). Tectum between the spines scabrate to granulate. Geofloristic element. Palaeotropical (P2). B o t a n i c a l a f f i n i t y. Verbenaceae, Clero- O c c u r r e n c e. Paleocene – Miocene. dendrum type. R e m a r k s. Change of genus results from the Geofloristic element. Palaeotropical (P). determination of morphological similarity of Present-day genus Clerodendrum (about 150 fossil pollen grains to grains of extant Casta- species) occurs in tropical and subtropical neoideae subfamily. Transfering Cupuliferoi- areas on both hemispheres. pollenites pusillus to a new genus cancels the name Cupuliferoipollenites as C. pusillus had R e m a r k s. For the first time the genus Cle- been the type species of the genus Cupuliferoi- rodendrumpollenites with the typical species pollenites. C. microechinatus has been described in the archival doctoral dissertation (Skawin´ ska 1989). Cercidiphyllites Mtshedishvili 1961 Type species. Cercidiphyllites brevicolpa- Clerodendrumpollenites microechinatus tus Mtshedishvili 1961 Skawin´ska sp. nov. Pl. 12, fig. 14a–c

Cercidiphyllites minimireticulatus (Trevi- 1985 Clerodendrum sp.; Skawin´ ska, p. 110, pl. 3, figs san 1967) Ziembin´ ska-Tworzydło comb. nov. 1, 2. Pl. 13, fig. 1a, b; 2 H o l o t y p e. Pl. 12, fig. 14a–c. 1967 Tricolpopollenites minimireticulatus n. fsp.; Tre- visan, p. 38, pl. 24, figs 10–12. L o c u s t y p i c u s. Borehole Ostrzeszów GEO3. 22

S t r a t u m t y p i c u m. Middle Miocene (Poz- genus Cornoidites Stuchlik 1964 is a homonym nan´ Formation). of genus Cornoidites Thiergart 1951, which ac- Derivation of name. After the sculpture cording to Potonié 1960 and Jansonius et Hills of surface. 1976 (card no. 603) is nomen nudum. The genus Cornaceoipollenites, which also has C. D i a g n o s i s. Pollen grains tricolporate, pro- parmularius as the type species lost its valida- late amb oval with broadly oval poles tion because of reasons mentioned. To new (68x44 μm). Exine two-layered about 2 μm genus Cornaceaepollis the fossil pollen grains thick, ectexine thicker than endexine, tectate. of morphology similar to pollen of today family Tectum supported by indistinct columelle Cornaceae (subfamilies Cornoideae and Mas- about 1 μm, high. Surface of tectum provided tixioideae) have been included. with loosely spaced, up to 1.5 μm long spines. Between them surface scabrate to granulate. Cornaceaepollis major (Stuchlik 1964) Botanical affinity. Clerodendrum type. Stuchlik comb. nov. Geofloristic element. Palaeotropical (P1). Pl. 13, fig. 11a–c Occurrence. Middle Miocene. 1964 Cornoidites major n.sp.; Stuchlik, p. 62, pl. 19, figs 1–4.

Cornaceaepollis Stuchlik gen. nov. B o t a n i c a l a f f i n i t y. Cornoideae, Cornus type. non Cornoidites Thiergart 1951 p.p. Tricolpopollenites Pflug & Thomson 1953 Geofloristic element. Palaeotropical non Cornaceoipollenites Potonié 1951a ex Potonié 1960 (P2). Today genus Cornus (45 species) occurs Cornoidites Stuchlik 1964 in warm temperate zone of northern hemis- Type species. Cornaceaepollis major phere, mainly in Asia and America. (Stuchlik 1964) comb. nov. (= Cornoidites Occurrence. Middle Miocene. major Stuchlik 1964, p. 62, pl. 19, figs 1–4) R e m a r k s. New combination results from es- D e r i v a t i o n o f n a m e. After the recent tablishing the new generic name for fossil pol- family Cornaceae. len grains of morphology similar to pollen of D i a g n o s i s. Pollen grains tricolporate, in extant family Cornaceae. polar view rounded-triangular to triangular, in equatorial view broadly oval. Polar axis 23–60 μm Cornaceaepollis minor (Stuchlik 1964) long. Colpi rather long, running from one pole Stuchlik comb. nov. to the other nearly meeting in the poles. Apo- Pl. 13, figs 12a, b; 13 colpium diameter 5–15 μm. Pores in the 1964 Cornoidites minor n. sp.; Stuchlik, p. 62, pl. 19, middle of the colpi, set transversally to them. figs 1–7. Exine rather thick 2–5 μm, tectate. Surface re- Botanical afiinity. Cornoideae, Cornus ticulate or granulate. type. B o t a n i c a l a f f i n i t y. Cornaceae, Cornoi- Geofloristic element. Palaeotropical (P2). deae, Mastixioideae. Occurrence. Middle Miocene. Geofloristic element. Palaeotropical (P). R e m a r k s. As in previous species. Present day representatives of Cornaceae fam- ily (12 genera) occur from tropical to tem- Cornaceaepollis satzveyensis (Pflug 1953) perate zone of N. America and in temperate Ziembin´ ska-Tworzydło comb. nov. zone of Asia. Pl. 13, figs 8–10a, b R e m a r k s. Stuchlik (1964) described two 1953b Tricolporopollenites satzveyensis n. sp. (Pflug); species: Cornoidites major and C. minor but he Thomson & Pflug, p. 103, pl. 13, figs 10–13. did not describe genus formally. Thiergart Botanical affinity. Mastixioideae. 1951 established genus Cornoidites without description and illustration in connection with Geofloristic element. Palaeotropical (P1). species C. cf. parmularius. It is the reason of Today the subfamily Mastixioideae with one cancellation of genus Cornoidites because the genus Mastixia (25 species) occurs in Indoma- species Cornoidites parmularius has been layan region in palaeotropical zone. transferred to genus Eucommioipollis. So the O c c u r r e n c e. Paleocene – Miocene. 23

R e m a r k s. Because of the similarity of pollen Eucommioipollis Ziembin´ska-Tworzydło gen. grain structure of this species to pollen of Cor- nov. naceae family, and especially to those from Cornaceoipollenites Potonié 1951a subfamily Mastixioideae, the species has been p.p. Tricolpopollenites Pflug & Thomson 1953 excluded from the collective morphological p.p. Tricolporopollenites Pflug & Thomson 1953 genus Tricolporopollenites and transferred to p.p. Psilatricolporites Hammen van der 1956 non Eucommiapollis Menke 1976 genus Cornaceaepollis, pointing at its botani- cal relations. Type species. Eucommioipollis eucom- mius (Planderová 1990) comb. nov. (=Tricol- Diospyrospollenites Skawin´ska gen. nov. poropollenites eucommi Planderová 1990, p. Type species. Diospyrospollenites ovalis 70, 71, pl. 69, figs 22–24, Hontianske Nemce, (Tarasevich 1980) comb. nov. (= Diospyros Lower Sarmatian) ovalis Tarasevich 1980); Tarasevich, p. 381– D e r i v a t i o n o f n a m e. After the recent 382, pl. 1, fig. 4, Plain Oksko – Don, Middle genus Eucommia. Miocene. D i a g n o s i s. Pollen grains tricolporate in D e r i v a t i o n o f n a m e. After the recent equatorial view outline oval with broadly genus Diospyros. rounded to flatted poles. Colpi short not of the D i a g n o s i s. Pollen grains tricolporate, pro- same length, more or less paralle, considerably late (45x40 μm). Colpi long, not always paral- deeply cut into the poles. Pores lolongate often μ lel reaching the poles. Pores lalongate. Exine indistinct. Exine two- layered 3 m thick, ec- tectate, surface of the tectum psilate to sca- texine tectate. Tectum psilate. Polar axis 26– μ μ brate. 48 m long, equatorial diameter 22–36 m. B o t a n i c a l a f f i n i t y. Ebenaceae, Diospy- Botanical affinity. Eucommiaceae, Eu- ros type. commia type. Geofloristic element. Palaeotropical (P). Geofloristic element. Arctotertiary (A). Today genus Diospyros (c. 500 species) occurs In the present days the monotypic genus Eu- in tropical and subtropical zones of both he- commia occurs in mesophytic mountain forests mispheres. of Central China. R e m a r k s. For the first time a diagnosis of R e m a r k s. To the genus Eucommiapollis genus, its description and a new combination beside type species E. eucommius the species of species were given in the archival doctoral E. parmularius has been included which was dissertation (Skawin´ ska 1989). previously recognized as a typical of the fol- lowing genera: Tricolpopollenites, Cornaceoi- Diospyrospollenites ovalis (Tarasevich pollenites, Cornoidites (cf. parmularius), Psila- 1980) Skawin´ska comb. nov. tricolporites. Because of that according to the Pl. 13, figs 3a, b; 4 International Code of Botanical Nomenclature 1980 Diospyros ovalis n. sp.; Tarasevich, p. 381, 382, the genera mentioned are loosing their va- pl. 1, figs 4, 6–8. lidity. The genus Eucommiapollis Menke has 1985 Diospyros sp.; Skawin´ ska, p. 111, 112, pl. 4, figs completely different morphological features 4–9; pl. 5, figs 1, 2. (especially in the structure of colpi) than pol- Botanical affinity. Diospyros type. len grains of the only living species Eucommia ulmoides. Geofloristic element. Palaeotropical (P1). Occurrence. Middle Miocene. Eucommioipollis eucommius (Planderová R e m a r k s. New combination of the species 1990) Ziembin´ska-Tworzydło comb. nov. results from establishing the new genus Dios- Pl. 13, figs 5, 6 pyrospollenites. Fossil pollen grains of D. 1960 Eucommia af. ulmoides n. sp.; Oszast, p. 22, pl. ovalis morphologically are the most similar to 7, figs 17–19, 21; pl. 8, figs 1, 3. grains of Diospyros lotus and D. kaki. 1974 Tricolporopollenites parmularius (Potonié) Krutzsch; Ziembin´ ska-Tworzydło, p. 399, pl. 24, fig. 12. 1977 Tricolporopollenites parmularius (Potonié) Krutzsch; Krutzsch & Vanhoorne, p. 69, pl. 29, figs 1–8. 24

1990 Tricolporopollenites eucommii n. sp.; Plan- H o l o t y p e: Pl. 14, fig. 9. derová, p. 70, 71, pl. 69, figs 22–24. L o c u s t y p i c u s. Borehole Rypin. Botanical affinity. Eucommia type. S t r a t u m t y p i c u m: Middle Miocene. Geofloritic element. Arctotertiary (A1). D e r i v a t i o n o f n a m e. After the recent Occurrence. Miocene. genus Decodon. R e m a r k s. Morphological features are con- D i a g n o s i s. Pollen grains tricolporate. In sistent with the diagnosis by Planderová equatorial view longitudinal ovate (30 x 18 μm) (1990). To this species large pollen grains of with slightly sharpened pole fields, in polar polar axis exceeding 40 μm and equatorial view the outline triangular to rounded-trian- diameter 30–34 μm have been included. The gular with deeply indented colpi. Colpi short species has been recognized as typical of new and straight, running parallel one to the other genus because morphological structure of fos- along lateral sides. Pores rather big lalongate sil grains here included is the most similar to to circular. Exine two-layered, about 1.5 μm that of today species Eucommia ulmoides. thick, ectexine as thick as endexine, tectate, tectum psilate to finely granulate. Eucommioipollis parmularius (Potonié Botanical affinity. Lythraceae, Decodon type. 1934) Ziembin´ska-Tworzydło comb. nov. Pl. 13, fig. 7a, b Geofloristic element. Arctotertiary (A1). 1934 Pollenites parmularius n. sp.; Potonié, p. 52, Today genus Decodon (1 species, Decodon verti- pl. 2, fig. 7. cillata = Lythrum verticillatum) occurs in swamp 1953 Tricolpopollenites parmularius (Potonié) n. regions of eastern parts of USA. comb.; Thomson & Pflug, p. 97, pl. 11, figs 152– 162. Occurrence. Middle Miocene. 1954 Pollenites cf. parmularius Potonié; Doktorowicz- R e m a r k s. It differs from Lythraceaepolle- Hrebnicka, p. 68, fig. 136. nites bavaricus Thiele-Pfeiffer 1980 by more 1960 Tricolporopollenites parmularius (Potonié) n. comb. (Krutzsch); Krutzsch & al., fig. 94. eloganted polar axis (P/E = 1.66), triangular 1977 Tricolporopollenites cf. parmularius (Potonié) equatorial outline, and psilate to finely reticu- Krutzsch; Krutzsch & Vanhoorne, p. 69, pl. 26, late surface of tectum. figs 9–11. 1978 Psilatricolporites parmularius (Potonié) n. comb.; Kedves, p. 32, pl. 5, figs 1–9. Nelumbopollenites Skawin´ska gen. nov. Botanical affinity. Eucommia type. Type species. Nelumbopollenites euro- Geofloritic element. Arctotertiary (A1). paeus (Tarasevich 1983) comb. nov. (= Ne- lumbo europaea Tarasevich 1983; Kuprianova, Occurrence. Miocene – Pliocene. Tarasevich, p. 142, pl. 2, fig. 7, Tambowskaja Remarks. Eucommioipollis parmularius dif- Obłast, Middle Miocene). fers from typical species in smaller dimensions D e r i v a t i o n o f n a m e. After the recent (polar axis 31–34 μm, equatorial diameter 26– genus Nelumbo. 28 μm), more rounded polar outline and nar- row and convex polar field. Change of genus is D i a g n o s i s. Pollen grains tricolpate, prolate, the result of finding a striking similarity of in equatorial view oval in polar view circular. μ fossil pollen grains to those of extant genus Polar axis 77 m long, equatorial diameter μ Eucommia. about 56 m. Colpi not deep, arcuately bended in the equator. Exine two-layered up to 3.5 μm Lythraceaepollenites Thiele-Pfeiffer 1980 thick, ectexine thicker than endexine, semitec- tate. Tectum supported by rather high colu- Type species. Lythraceaepollenites bavari- mellae forming a reticulate – rugulate pattern. cus Thiele-Pfeiffer 1980 B o t a n i c a l a f f i n i t y. Nelumbonaceae, Ne- lumbo type. Lythraceaepollenites decodonensis Stuchlik sp. nov. Geofloristic elemet – Palaeotropical (P). Pl. 14, figs 9, 10, 11 Genus Nelumbo (4 species) is distributed 1964 Decodon cf. globosus (Reid) Nikitin; Stuchlik, p. today in subtropical and tropical regions of SE 49, pl. 15, figs 6, 7. Asia, SE part of N America and in northern 1984 Decodon; Jahn & al., pl. 10, figs 20–30. part of S America. 25

R e m a r k s. First description of genus and R e m a r k s. New combination for species re- new combination for typical species Nelumbo- sults from finding the morphological similarity pollenites europaeus (Tarasevich) comb nov. is of pollen grains of O. microreticulatus to the in the archival doctoral dissertation (Skawin´s- pollen of extant Oleaceae family without possi- ka 1989). bility of pointing at the close similarity to present-day genus. Nelumbopollenites europaeus (Tarasevich 1983) Skawin´ska comb. nov. Platanipollis Grabowska gen. nov. Pl. 14, fig. 12a–d non Platanoidites Potonié, Thomson & Thiergart 1950 1983 Nelumbo europaea n. sp. (Tarasevich); Kupria- non Platanopollenites Potonié 1951 nova & Tarasevich, p. 142, pl. 2, fig. 27, pl. 5, p.p. Tricolporopollenites Pflug & Thomson 1953 figs 1–4 Platanus L. Pacltová 1982 1985 Nelumbo sp.; Skawin´ ska, p. 112, pl. 5, figs 3–5. Type species. Platanipollis ipelensis Botanical affinity. Nelumbo type. (Pacltová 1966) comb. nov. {= Tricolporopolle- Geofloristic element. Palaeotropical (P2). nites ipelensis Pacltová 1966, p. 61, pl. 19, figs Occurrence. Miocene – Pliocene. 14–16, Oberckov (Ipel Basin), South Slovakia, R e m a r k s. Morphology of pollen grains of Ne- Egerian}. lumbopollenites europaeus is similar to mor- D e r i v a t i o n o f n a m e. After the recent phology of pollen grains of four extant species genus Platanus. of Nelumbo (Zetter & Keri 1989) but differen- D i a g n o s i s. Pollen grains tricolporate, pro- ces in structure of the reticule on the surface late (15–25 x 12–18um). In equatorial view of exine are not so distinct that the fossil outline oval to circular with rounded poles. species could be included into one of present Colpi not distinct deep, parallel or nearly par- day taxa. The establishing of new organ genus allel considerably broadly spaced, not meeting Nelumbopollenites results from principles of in the poles, covered by granulate membrane. ICBN and point 7 of revision presented. Endaperture developed as pore-like endocolpi, transversaly ragged. Exine two-layered, tec- Oleoidearumpollenites Nagy 1969 tate, tectum granulate to reticulate. Type species. Oleoidearumpollenites re- B o t a n i c a l a f f i n i t y. Platanaceae, Plata- ticulatus Nagy 1969 nus type.

Oleoidearumpollenites microreticulatus Geofloristic element. Palaeotropical (P). (Pflug & Thomson 1953) Ziembin´ska- The family Platanaceae (1 genus Platanus Tworzydło comb. nov. with 7 species) occurs today in S Europe, E Pl. 14, fig. 18a–c Asia and N America. 1949 cf. Oleaceae; Kremp, p. 76, pl. 9, fig. 131a–c. R e m a r k s. The diagnosis of genus Plata- 1953 Tricolporopollenites microreticulatus n. sp. noidites Potonié, Thomson & Thiergart 1950 (Pflug & Thomson); Thomson & Pflug, p. 106, pl. and Platanoipollenites Potonié 1951a, b (as a 16, figs 35–39. synonym) does not match the morphology of 1960 Oleaceae forma artificosa; Mamczar, p. 53, pl. 13, figs 177–179. pollen grains of present day genus Platanus, 1961 Oleaceae; Romanowicz, p. 354, pl. 21, figs 264– precisely presented by Pacltová (1982). The 266. type species for both genera mentioned is Pol- 1964 cf. Fraxinus sp.; Stuchlik, p. 70, pl. 21, figs 14– lenites gertrudae Potonié 1931, which has only 17. one pore in each of three colpi. Pacltová (1966) 1969 Caprifoliipites gracilis n. sp.; Nagy, p. 423–424, pl. 44, figs 11, 12. distinguished fossil species Tricolporopolle- 1973 Oleaceae; Stachurska & al., pl. 14, figs 10–14. nites ipelensis which she moved to the extant genus Platanus in 1982, emending simultan- Botanical affinity. Oleaceae. ously the diagnosis of species. The botanical Geofloristic element. Arctotertiary (A1). affinity to extant genus Platanus has been Present-day representatives of Oleaceae fam- preserved while forming the new organ genus ily (27 genera) are distributed from tropical to and the ending -pollis has been added. A wide- temperate zones. ly distributed in Younger Palaeogene and Occurrence. Oligocene – Pliocene. Older Neogene Platanipollis ipelensis (Paclto- 26 vá) species has been chosen as the type structure. The genus Quercopollenites Nagy species. 1969 is younger synonym.

Platanipollis ipelensis (Pacltová 1966) Gra- Quercoidites asper (Pflug & Thomson 1953) bowska comb. nov. Słodkowska comb. nov. Pl. 14, fig. 21a–c Pl. 15, fig. 1 1953 Tricolpopollenites asper n. sp. (Pflug & Thom- 1966 Tricolporopollenites ipelensis n. sp.; Pacltová, p. son); Thomson & Pflug, p. 96, pl. 11, figs 43–47. 25, pl. 19, fig. 14–19. 1961a Tricolporopollenites asper (Potonié) n. comb.; 1982 Platanus ipelensis Pacltová n. emend; Pacltová, Krutzsch, p. 322. p. 61, 62, pl. 4, figs 1–7; pl. 5, figs 1–12; text fig. 7. 1990 Tricolporopollenites sp. type “Platanus”; Plan- Botanical affinity. Quercus robur type. derová, p. 70, pl. 69, figs 19, 20. Geofloristic element. Arctotertiary (A1). Botanical affinity. Platanus type. Today Quercus robur occurs in Europe, N Afri- Geofloristic element. Palaeotropical (P1) ca and Asia Minor. Occurrence. Upper Paleocene – Lower Mio- Occurrence. Neogene. cene. Remarks. The genus Tricolpopollenites R e m a r k s. Chosen as the type species Pla- asper has been included into genus Quer- tanipollis ipelensis was precisely described coidites in order to extending of this genus di- and ilustrated by Pacltová (1982). Its morpho- agnosis. logical features are very similar to those of ex- tant pollen of genus Platanus. Quercoidites granulatus (Nagy 1969) Słod- kowska comb. nov. Pl. 15, figs 10, 11 Quercoidites (Potonié, Thomson & Thiergart 1950 ex Potonié 1960) Słodkowska emend. 1969 Quercopollenites granulatus n. sp.; Nagy, p. 223, pl. 52, fig. 21. p.p. Tricolpopollenites Pflug & Thomson 1953 p.p. Tricolporopollenites Pflug & Thomson 1953 Botanical affinity. Quercus type (with Quercopollenites Nagy 1969 rough – granulate pollen grains). Type species. Quercoidites henrici (Po- Geofloristic element. Arctotertiary (A1). tonié 1931a) Potonié, Thomson & Thiergart Occurrence. Miocene. 1950 Remarks. Quercopollenites granulatus was E m e n d e d d i a g n o s i s. Pollen grains tricol- transferred to Quercoidites genus in accord- porate, equatorial outline oval, amb in polar ance to the widening of its diagnosis. The μ view considerable small. Polar axis 20–60 m genus Quercopollenites (Nagy 1969) lost va- long. Colpi running parallel and nearly meet- lidity because its type species Q. granulatus ing in the pole area. In the equator area along was transferred to the other genus. the colpi exine is divided formig a lolongate pore-like aperture. Exine two-layered, ectexine Quercoidites pudicus (Potonié 1934) Słod- thicker than endexine, tectate. Surface of the kowska comb. nov. tectum provided with various irregular gra- Pl. 15 figs 3, 4 nules or verrucae. 1934 Pollenites confinis pudicus n. f.; Potonié, p. 90, B o t a n i c a l a f f i n i t y. Fagaceae, Quercoideae. pl. 5, figs 12, 13. Geofloristic element. Arctotertiary/Pa- 1951b Fraxinoipollenites pudicus Potonié; Potonié, p. laeotropical (A/P). The genus Quercus (600 species) 277, pl. 1, fig. 49. 1953 Tricolpopollenites pudicus (Potonié) n. comb.; occurs today in northern hemisphere in different Thomson & Pflug, p. 95, pl. 11, figs 27–29. climatic zones – from temperate to tropical. 1960 Fraxinoipollenites pudicus (Potonié) Potonié; R e m a r k s. For all fossil pollen grains mor- Potonié p. 94 1961a Tricolporopollenites pudicus (Potonié) n. comb.; phologically similar to pollen of subfamily Krutzsch, p. 322. Quercoideae the priority generic name Quer- coidites, established in 1950, has been used. Botanical affinity. Quercus type (with Emendation of diagnosis extends the use of this fine – granulate pollen grains). genus over all fossil pollen grains of “quercoidal” Geofloristic element. Arctotertiaty (A1). 27

O c c u r e n c e. Miocene. Occurrence. Miocene – Pliocene. R e m a r k s. The transferring to Quercoidites R e m a r k s. Selected as the type species Tra- genus the species Fraxinoipollenites pudicus papollis erdtmanii was described in details (Potonié) Potonié, which was the type species and illustrated by Nagy (1979). Its morpho- for the genus Fraxinoipollenites (Potonié 1960), logical features are in concordance with the gives in the efect the lost of its validity. features of pollen of Trapa genus.

Trapapollis illingensis (Klaus 1954) Kohlman- Trapapollis Kohlman Adamska gen. nov. Adamska comb. nov. Sporotrapoidites Klaus 1954 Pl. 15, fig. 25a, b

Type species. Trapapollis erdtmanii 1954 Sporotrapoidites illingensis n. sp.; Klaus, p. 122, (Nagy 1979) comb. nov. (= Goerboepollenites pl. 1, figs 1–3. erdtmanii Nagy 1979, p. 185, fig. 3 E – N, fig. 4 A – E, Nagygörbö, 1, 911,0 m, Karpatian). Botanical affinity. Trapa type. D e r i v a t i o n o f n a m e. After the recent Geofloristic element. Arctotertiary (A1). genus Trapa. Occurrence. Miocene – Pliocene. D i a g n o s i s. Pollen grains tricolporate in R e m a r k s. The new registration of species equatorial view more or less spheroidal and originates from the genus change. triangular in polar view. The grains are pro- vided with 3 meridional sexinous crests divi- Tricolporopollenites Pflug & Thomson 1953 ding along the short colpi and meeting at the Type species. Tricolporopollenites dolium poles. Exine rather thick, two – layered, tec- (Potonié 1931b) Thomson & Pflug 1953 tate. Surface of the tectum smooth outside the crest and psilate to verrucate on the crest. μ Tricolporopollenites brühlensis (Thomson Polar axis without the crest 40–75 m long, 1950) Grabowska stat. nov. μ equatorial diameter, crest not included 35–40 m. Pl. 16, figs 1–5 B o t a n i c a l a f f i n i t y. Trapaceae (= Hydroc- 1950 Pollenites cingulum brühlensis n. spm.; Potonié aryaceae), Trapa type. & al., p. 56, 63, pl. B, figs 31–33. Geofloristic element. Arctotertiary (A). 1953 Tricolporopollenites megaexactus (Potonié) n. Present-day genus Trapa (3 species) occurrs in comb. subsp. brühlensis (Thomson) n. comb.; Thomson & Pflug, p. 101, pl. 12, figs 51–57. warm-temperate zone of Europe, Asia and 1960 Cyrillaceaepollenites megaexactus subsp. brühl- northern Africa. ensis (Thomson) n. comb.; Potonié, p. 102. R e m a r k s. Klaus (1954) forming the name of 1966 Tricolporopollenites cf. brühlensis (Thomson) Thomson & Pflug; Sontag, p. 38, pl. 50, fig. 13a, b. Sporotrapoidites genus based on the morpho- logy of polar field, where three collars form B o t a n i c a l a f f i n i t y. P.p. Clethraceae, p.p. sign similar to a tetrade mark, characteristic Cyrillaceae, p.p. Rosaceae. of spores. The transforming the Sporotra- Geofloristic element. Palaeotropical (P2). poidites name into Trapapollis was in the ef- Presently a monotype Clethraceae family (Cle- fect of adopted by authors the rule of eliminat- thra genus with 30 species) occurs in South ing from the genus name the suggestion of its and North America, SE Asia in subtropical morphological affinity, i.e. that there is a zone and in the mountains in tropical areas; spore. Cyrillaceae (3 genera and 14 species) – pres- ently in tropical and subtropical America, es- Trapapollis erdtmanii (Nagy 1979) pecially in the Atlantic area, in south-eastern Kohlman-Adamska comb. nov. America. Pl. 15, figs 23a, b; 24 Occurrence. Eocene – Pliocene. 1979 Goerboepollenites erdtmanii sp. n.; Nagy, p. 185, R e m a r k s. The rank of the taxon T. brühlen- figs 2, 3 E- N, 4 A-D. sis was changed from a subspecies into a 1985 Sporotrapoidites erdtmani (Nagy 1979) n. c.; Na- gy, p. 163, pl. 93, figs 18–20; pl. 94, figs 1–8. species. 1. In the first name Pollenites cingu- lum brühlensis the taxon “cingulum” is not Botanical affinity. Trapa type. confirmed in the pollen morphology of this Geofloristic element. Arctotertiary (A1). type. 2. The replacement of the taxon Tricolpo- 28 ropollenites megaexactus brühlensis by the O c c u r r e n c e. Upper Paleocene – Miocene. species Cyrillaceaepollenites megaexactus done R e m a r k s. Named by Thomson & Pflug by Potonié (1960) is not still valid because T. (1953) the species Tricolpopollenites liblarensis brühlensis differs from T. megaexactus, among fallax , T. liblarensis (=quisqualis) and T. reti- other, by more protrounding the equatorial formis were transferred (excluding T. liblaren- part of colpi with pore beyond the contour of a sis) to Tricolporopollenites genus by Krutzsch pollen. In equatorial view it is characterized (1954, 1961), Krutzsch et al. (1960) and by trefoil like callosity. 3. It cannot be main- Krutzsch & Vanhoorne (1977), proved that in tained Cyrillaceaepollenites genus because the species of the former Tricolpopollenites genus grains of similar structure are also found be- more or less developed pores in colpi occur. In yond Cyrillaceae family. 4. Sontag (1966) can- T. liblarensis species, from which was separ- not be the nomeclator of the species because ated by Krutzsch (1954) T. quisqualis, the the taxon in the species rank is in brackets pores are not always visible, however in the and has “cf.” equatorial part of colpi a loosening of exine or curving of a colpus are distinct. The name of Tricolporopollenites exactus (Potonié 1931) Cupuliferoidaepollenites genus was not used, Grabowska comb. nov. for which T. liblarensis was the type species, Pl. 16, figs 8–10 because it may suggest the botanical affinity 1931b Pollenites exactus n. sp.; Potonié, p. 26, pl. 1, only to Fagaceae family, while the similar pol- fig. V49b. len grains are known from various families. 1953 Tricolporopollenites megaexactus (Potonié) n. comb. subsp. exactus (Potonié) n. comb.; Thom- son & Pflug, p. 101, pl. 12, figs 87–92. Tricolporopollenites photinioides Skawin´s- 1960 Cyrillaceaepollenites exactus (Potonié) n. comb.; ka sp. nov. Potonié, p. 102 Pl. 16, figs 27a, b; 28 B o t a n i c a l a f f i n i t y. P.p. Clethraceae, Cle- H o l o t y p e. Pl. 16, fig. 27a, b. thra type; p.p. Cyrillaceae, Cyrilla type. Locus typicus. Borehole Ostrzeszów GEO 3. Geofloristic element. Palaeotropical (P2). Stratum typicum. Middle Miocene, Po- Occurrence. Eocene – Pliocene. znan´ Formation. R e m a r k s. The morphological genus Tricol- Derivatio of name. After the recent poropollenites was maintained instead of the genus Photinia. Cyrillaceaepollenites one (Potonié 1960), sug- D i a g n o s i s. Pollen grains tricolporate, equa- gesting the botanical affinity to the Cyrilla- torial outline ovate, with slightly rounded ceae family because the pollen of similar mor- poles. Colpi slightly arcuate not meeting one phology in the Clethraceae family also occurs. an other at the poles. Pores lolongate often not It was mantained, according to the first ident- distinct. Exine two-layered 1–1.5 μm thick, ec- ification of the species (Potonié 1931b) and texine thicker than endexine, surface finely stri- later changing (Potonié 1960), the taxon “exac- ate. Polar axis about 30 μm long. tus” in the species rank. Botanical affinity. Rosaceae, Photinia, Sorbus types. Tricolporopollenites liblarensis (Thomson Geofloristic element. Arctotertiary (A1). 1950) Grabowska comb. nov. Presently Photinia genus with 60 species oc- Pl. 16, figs 13, 14 curs in tropical S-E Asia and Central America; 1950 Pollenites liblarensis n. spm. (Thomson); Potonié Sorbus genus (more than 100 species) in tem- et al., p. 55. perate zone of northern hemisphere. 1950 Cupuliferoidae-pollenites liblarensis n. spm. (Thomson); Potonié & al., p. 66, pl. B, figs 26, 27. O c c u r e n c e. Miocene. 1953 p.p. Tricolpopollenites liblarensis (Thomson) (= R e m a r k s. The fossil pollen graines are the quisqualis Potonié) n. comb.; Thomson & Pflug, p. 97, pl. 11, figs 111–132. most similar to those of extant Photinia genus, however similar ones may be also found in B o t a n i c a l a f f i n i t y. P.p. Fabaceae (= Le- Sorbus genus. guminosae), p.p. Fagaceae, p.p. Combretaceae, p.p. Verbenaceae. Geofloristic element. Palaeotropical (P2). 29

Sapotaceoidaepollenites Potonié, Thomson Stratum typicum. Middle Miocene, Po- & Thiergart 1950 ex Potonié 1960 znan´ Formation. Type species. Sapotaceoidaepollenites mani- D e r i v a t i o o f n a m e. After the reticulate festus (Potonié 1931d) ex Potonié 1960 sculpture of pollen grains. D i a g n o s i s. Pollen grains 4–5-colporate, pro- Sapotaceoidaepollenites oblongus (Pflug & late, polar outline ovate, equatorial outline cir- Thomson 1953) Grabowska comb. nov. cular to 4–5-lobate. Polar axis 35–42 μm long, Pl. 17, fig. 5 equatorial diameter 23–28 μm. Colpi more or 1953 Tetracolporopollenites oblongus n. sp. (Pflug & less parallel, long and narrow, nearly meeting Thomson); Thomson & Pflug, p. 110, pl. 15, fig. at the poles. Along the colpi endexine swollen. 31. Pore lalongate. Exine two – layered 2.5 μm Botanical affinity. Sapotaceae. thick, ectexine thicker than endexine, semitec- tate. Columellae fused with their capita for- Geofloristic element. Palaeotropical (P1). ming a reticulate pattern. Lumina irregular, Presently Sapotaceae family with 50 genera smaller at the poles, arranged in a striato-re- occurs mostly in tropical and subtropical ticulate pattern. zones, some of them in warm-temperate zone. Botanical affinity. Rutaceae, Skimmia Occurrence. Upper Eocene – Miocene. type (Skimmia laureola) R e m a r k s. Change in genus results from the Geofloristic element. Palaeotropical (P2). similarity of structure of fossil pollen graines of Sapotaceoidaepollenites oblongus to those of Occurrence. Miocene. extant Sapotaceae family. R e m a r k s. As above.

Skimmiaepollenites Skawin´ska gen. nov. Symplocoipollenites (Potonié 1951 ex Po- Type species. Skimmiapollenites reti- tonié 1960) Słodkowska emend. culatus Skawin´ska sp. nov. Symplocospollenites Potonié, Thomson & Thiergart 1950 ex Potonié 1960 D e r i v a t i o n o f n a m e. After the recent p.p. Porocolpopollenites Pflug 1953b genus Skimmia. Type species. Symplocoipollenites ves- D i a g n o s i s. Pollen grains 4–5-colporate, pro- tibulum (Potonié 1931b) Potonié 1951a,b ex late, polar outline ovate, equatorial outline cir- Potonié 1960 cular to 4–5-lobate. Polar axis 35–42 μm long, equatorial diameter 23–28 μm. Ectexine semi- E m e n d e d d i a g n o s i s. Pollen grains 3–5- tectate, striato-reticulate. colporate, mostly 3-colporate oblate. Equator- ial outline triangular, sometimes more circu- Geofloristic element. Palaeotropical (P). lar, polar outline oval to rhomboidal. Colpi Presently Skimmia genus (10 species) occurs very short only slightly longer than pores in warm-temperate to subtropical zones of diameter. Pores circular seldom oval with dis- eastern Asia. tinct vestibulum and small labrum. Ectexine Botanical affinity. Rutaceae, Skimmia tectate, tectum psilate, granulate to baculate. type. Botanical affinity. Symplocaceae. R e m a r k s. The diagnosis of the genus and species was given for the first time in the the Geofloristic element. Palaeotropical (P). archival doctoral dissertation (Skawin´ska Present-day, a monotype Symplocaceae family 1989). with Symplocos genus (300–400 species) oc- curs in tropical and subtropical zones of S-E Asia, only one species in warm-temperate At- Skimmiaepollenites reticulatus Skawin´ska sp. nov. lantic zone of North America. Pl. 17, fig. 6a–c; 7a, b R e m a r k s. Potonié (1960) globose forms in- cluded to Symplocospollenites genus, triangle 1985 Skimmia laureola Hooker – typ; Skawin´ska, p. 113, pl. 6, figs 1–7. to Symplocoipollenites. Pflug (1953) for both forms created morphological genus Porocol- H o l o t y p e. Pl. 17, fig. 6a–c. pollenites. For all species of this genus the Locus typicus. Borehole Ostrzeszów GEO 3. botanical affinity Symplocaceae family is 30 given, and this is the reason the putting these nicka) n. comb.; Ziembin´ ska-Tworzydło, p. 382–383, species into one genus – Symplocoipollenites. pl. 19, figs 5, 6. This genus was chosen because most Neogene D e s c r i p t i o n. Pollen grains tricolporate, species were included by Potonié (1960) to it. equatorial outline triangular with slightly con- vex sides, angulaperturate. Exine two-layered Symplocoipollenites latiporis (Pflug & 1.5–3 μm thick, surface granulate. Thomson 1953) Słodkowska comb. nov. Pl. 18, figs 1a, b; 2 Botanical affinity. Symplocaceae. 1953 Porocolpopollenites latiporis n. sp. (Pflug & Geofloristic element. Palaeotropical (P2). Thomson); Thomson & Pflug, p. 93, pl. 10, figs Occurrence. Miocene. 123–124. R e m a r k s. As in the species metioned pre- Botanical affinity. Symplocos type. viously. Geofloristic element. Palaeotropical (P2). Occurrence. Miocene. Symplocoipollenites orbis (Pflug & Thom- R e m a r k s. Change in genus results from the son 1953) Słodkowska comb. nov. necessity of including the species into the Pl. 18, figs 5, 6 genus pointing at the botanical origin. 1953 Porocolpopollenites orbis n. sp. (Pflug & Thom- son); Thomson & Pflug; p. 92, pl. 10, figs 109, 111. Symplocoipollenites maturus (Doktorowicz- 1970b Symplocospollenites orbis (Thomson & Pflug) Hrebnicka 1960) Ziembin´ska-Tworzydło Potonié; Krutzsch, p. 336, pl. E 79. comb. nov. Pl. 18, figs 3, 4 Botanical affinity. Symplocaceae. 1960 Symplocos-pollenites vestibulum Potonié forma Geofloristic element. Palaeotropical (P1). matura; Doktorowicz-Hrebnicka, p. 111, pl. 43, O c c u r r e n c e. Paleocene. fig. 227. 1974 Porocolpopollenites maturus (Doktorowicz-Hreb- R e m a r k s. As above.