A C T A B I O L . S Z E G E D . 3 3 . PP. 3 5 — 4 7 (1987) PALEOPHYTOGEOGRAPHY OF THE ANGIOSPERM P O L L E N GRAINS DURING T H E UPPER C R E T A C E O U S AND THE TERTIARY I M. KEDVES Department of Botany. Attila Jozsef University H—6701 Szeged. P.O B. 657. Hungary (Received: February 20. 1986) Abstract The U p p e r Cretaceous - - Paleogene palcophytogeographical regions are based on the distribution of the pollen grains of Normapolles. Triprojectacites. Oculata G r o u p . Proteaceae. Ulmaceae. and Olacaceae. The pollen g r o u p Postnormapolles was not sufficiently studied from this point of view. This paper deals with the regional distribution during the geological past of one form-species of early type from the genuses Triatriopollenites and Triporopollenites. all species of the genus Platycaryapollenites and Paraalnipollenites. Triatriopollenites roboratus has the most restricted area and occurrence in the geological time units, namely this species occur in the Paleoccne — Eocene of Eurasia. Triporopollenites robustus is also mostly from Paleocene — Eocene age but there are scattered data f r o m the Upper Cretaceous and from the Eocene until the Miocene. This species occur not only in Eurasia, but in North-America and North-Africa too. The pollen grains of the genus Paraalnipollenites were mainly the elements of the province Aquilapollenites. and the intermediate region, its occurrence in the Normapolles province is scarce. This form-genus seems t o be charasteristic for the Upper Cretaceous. Paleocene and Eocene time, but there are d a t a f r o m younger sediments loo. The pollen grains of the genus Platycarvapollenites a p p e a r in the U p p e r Cretaceous (North-Africa). It is very c o m m o n in the Paleocene. but its largest distribution is during the Eocene, important in the Oligocene. During the Neogcne its area become restricted gradually. From the Plioccne it is only a single occurrence f r o m the Southern Hemisphere f r o m a d e e p sea drilling near the Timor Islands.Thc pollen grains analyzed in this paper may be derived from the Normapolles group. But its distribution overstep the boundaries of the Normapolles laxa, in this way its r a d i a t i r n was very rapid during the U p p e r Cretaceous and the lower part of the Paleogene. Key words: Palynology. Palcophytogeography. Cretaceous — Tertiary. Introduction T h e study of the regional distribution of several s p o r o m o r p h taxa d u r i n g the geological past m a y be considered as a m e t h o d in Palynology. SAKS et al., (1973) emphasized that these investigations are i m p o r t a n t in the reconstruction of the paleophytogeographical environments, a n d in the establishment of the paleoclimatic zones a n d the paleopole. As pioneering publications, dealing with the U p p e r C r e t a c e o u s — Paleogene regions based on palynological d a t a the i m p o r t ance of the p a p e r of K.RUTZSCH (1960) a n d ZAKLINSKAYA (1962) m u s t be e m p h a s ized. C o n c e r n i n g the paleophytogeographical units, which were established in these classical publications, later several supplementary d a t a were published by several 36 M KEIJVES a u t h o r s : MTCHEDLISHVILI a n d SAMOILOVICH ( 1 9 6 2 ) . Z A K L I N S K A Y A ( 1 9 6 3 . 1966. 1967a b 1976). SHAKHMOUNDES (1966). SAMOILOVICH (1967). KEDVES a n d KIRALY (1968. 1970). STANLEY (1970). KHLONOVA (1971). WIGGINS (1976). SRIVASTAVA (1978). a n d BATTEN (1981). As c o m p r e h e n s i v e papers, the publications of MEDUS (1973). HERNGREEN and KHLONOVA (1981) and KEDVES and DlNlZ (1983) may be m e n t i o n e d . I m p o r t a n t d a t a were published on this point of view by ROCHE (1974). A new paleophytogeographical synthesis for the U p p e r C r e t a c e o u s was published by the present writer (KEDVES 1985). T h e p a l e o p h y t o geographical provinces and districts for the Paleocene. Eocene a n d O h g o c e n e were e l a b o r a t e d b y BOITSOVA a n d PANOVA ( 1 9 7 3 ) . D u r i n g the Tertiary, the extraordinarily differentiated U p p e r C r e t a c e o u s a n d Paleogene floras were altered, with a m o r e u n i f o r m vegetation. T h e early a n g i o s p e r m s d i s a p p e a r e d , these taxa are represented first in E u r o p e by the Normapolles g r o u p , which has a very complicated germinal exine structure. Its place was occupied by the Postnormapolles. by the developed types of the Longa.xones. a n d other different kinds of angiosperms. T h e pollen g r o u p of the Postnormapolles m a y be derived at least in its greater p a r t f r o m the Normapolles. But these pollen grains occur not only in the Normapolles ( E u r o p e a n — T u r a n i a n ; KHLONOVA. 1971) province, but in such territories where Normapolles have not existed, or their occurrence is very scarce. In the case of some Postnormapolles a h e t e r o g e n e o u s origin m a y be presumed. This Fig. I. Regional distribution of Triporopollenile.% rohustwi Pt. 1953 during lhe Upper Cretaceous and the Tertiary. P A L F O P H Y T O G F O G R A P H Y OF T H E ANGIOSPF.RM P O L L E N G R A I N S 37 p r o b l e m was focussed d u r i n g our present research project, by the way of the regional a n d chronological distribution of several form-species a n d form-genuses. This p a p e r s u m m a r i z e s the first results concerning this subject. Methods The literature d a t a , which were available f o r us was evaluated and documented for cach taxa. During the collection of the data, the following problems arise: I In several cases there are d o u b t s in the determinations of the sporomorphs. 2. In some papers the precision of the geological age of the spore-pollen assemblage is not satisfactory. 3. Sometimes it was not easy t o establish the localities. On the other hand there were given very large territories. on the other so small geographical objects were given, which a r e not indicated on the large world maps. It was impossible to indicate all the collected data, bccause the proportion size of o u r m a p . But the principal goal was. that each of the most important data for the palcophytogeographical evaluation were indicated on the m a p . of the The objects of this paper were the t n p o r a t e . and triatriatc pollen grains as the most primitive types Posinormapolles. Results Fgen.: Triporopollenites PF. et TH. 1953 Triporopolleniies robustus PF. 1953 subfsp. robust us. Betulaceae (Plate I fie 1,2) This kind of pollen grain is a very early t r i p o r a t e type, which p r o b a b l y may be distinguished t a x o n o m i c a l l y f r o m the o t h e r developed pollen g r a i n s of this m o r p h o l o g i c a l type. T E M d a t a a b o u t this species were published by KEDVES a n d PARDUTZ (1970), a n d by KEDVES a n d STANLEY (1976), this latter p a p e r deals with the results of the S E M studies too. T h e a p p e a r a n c e of this form-species is the U p p e r C r e t a c e o u s , we have d a t a f r o m E u r o p e , a n d f r o m the F a r East, in this way this species a p p e a r e d in the Normapolles a n d Aquilapollenites province at the same time (fig. 1). It is w o r t h y to m e n t i o n that it is not yet known f r o m the intermediate zone between the Normapolles a n d Aquilapollenites province in Eurasia, a n d f r o m N o r t h America too. D u r i n g the Paleocene a n d Lower Eocene firstly in the lower Eocene in Eurasia the distribution of this species is c o m m o n . F r o m N o r t h America only Paleocene a n d few Lower Eocene d a t a are known. W e have d a t a f r o m the Oligocene of E u r o p e , a n d f r o m the F a r East, but the re-study of these specimens m a y alter o u r present-day knowledge concerning the occurrence of this species. T h e M i o c e n e d a t a f r o m N o r t h America is in question. As regards the origins of this pollen type the Normapolles g r o u p m a y be taken first into consideration. But on the basis of its occurrence in the Paleocene it m a y not be presumed that this species radiated f r o m the E u r o p e a n gene c e n t r u m to Siberia. O u r earlier results (KEDVES a n d DLNIZ, 1979) reveal that between the Normapolles P A L F O P H Y T O G F O G R A P H Y O F T H E ANGIOSPF.RM POLLEN G R A I N S 39 a n d Postnormapolles g r o u p there are a n u m b e r of intermediate types a n d it may be presumed t h a t it is necessary to create a new g r o u p f o r s o m e early Brevaxones pollen grains, which occur not only in the Normapolles. but in the Aquilapollenites province. O n the o t h e r h a n d , the c o m m o n taxa of the established paleophytogeographical units did not yet c o m e into the limelight, because the subject of the researchers were firstly the segregate angiosperm pollen types. Fgen.: Trialriopolleniles PF. 1953 Trialriopolleniles roboratus PF. 1953, Myricaceae Syn.: I960 SAUER, in POKROVSKAYA el STELMAK. — Myrica (Plate I, fig. 3 — 6 ) mirabUis. sp. nov., p. 412. pi. V. fig. 9a.b. Its characteristic a t r i u m (Plate I, fig. 5,6) a n d the e x o a p e r t u r e elongated in polar direction refer to an early Brevaxones origin. It is regretful t h a t a b o u t this very i m p o r t a n t pollen grain we have not yet transmission electron microscopical d a t a . This species is i m p o r t a n t f r o m stratigraphical point of view of the Paleocene and the Lower Eocene (fig. 2). In contrast to the occurrence in the Paleocene a n d Lower Eocene layers of the F a r East, based on our present day knowledge, this is an element of the Normapolles province as an a t t e n d i n g of the Normapolles taxa. It is n o t e w o r t h y that against the widespread researches it is not yet f o u n d in N o r t h America. In this respect, this species has the identic, regional, a n d p r o p a r t e stratigraphic value, with the genus Stephanoporopollenites, having a decisive i m p o r t a n c e in E u r o p e a n d in the intermediate region in the d e t e r m i n a t i o n of the Paleocene age, a n d similarly not yet d e m o n s t r a t e d f r o m N o r t h America either (KEDVES, 1977). T h e difference between the stratigraphic value of Triatriopollenites roboratus, a n d the f o r m - g e n u s Stephanoporopollenites is, that the taxa of the latter m e n t i o n e d genus occurred never in the sediments younger as T h a n e t i a n in c o n t r a s t t o the Lower Eocene ( S p a r n a t i a n ) occurrence of Triatriopollenites roboratus. Plate I 1,2 3.4 Triporopollenites table n u m b e r : Trialriopolleniles robuslus PF. 1953 subfsp. minor KDS. 1970, cf. Belulaceae. slide: M e n a t - 2 , crossf o l l o w i n g K E D V E S a n d R U S S E L L ( 1 9 8 2 ) , xlOOO roboratus P F . 1 9 5 3 . slide: M e n a t - 8 . cross-table n u m b e r : 1 8 . 3 / 1 2 0 . 9 , following 14.7/116.6, a n d R U S S E L L , (1982) xlOOO Trialriopolleniles roboratus PF. 1953. slide: M e n a t - 2 6 , cross-table n u m b e r : 14.6/103.7. following KEDVES 5,6 a n d R U S S E L L (1982), x2500 Platycaryapolleniies swasticoidus (ELSIK 1974) F R E D , et C H R I S T . Plaiycarya. slide: M e n a t - 3 9 , cross-table n u m b e r : 14.2/111.3. f o l l o w i n g (1982). x2500 KEDVFS 7.8 9.11 Paraalnipolleniles alterniporus 16.9/117.1, xlOOO 10.12 Paraalnipolleniles alterniporus 16.9/117.1. x2500 1978. KEDVES Juglandaceaea. and RUSSELL (SIMPS. 1961) SRIV. 1975. slide: M e n a t - 3 7 . cross-table n u m b e r (SIMPS. 1961) SRIV. 1975. slide: M e n a t - 3 7 . cross-table number M 40 Fgen.: Plaiycaryapolleniles CHRISTOPHER E. KlDVtS NAGY 1969 emend. FREDERIKSEN et 1978 Plaiycaryapolleniles swasticoidus (ELSIK 1974) FRED, et CHRIST. 1978. Juglandaceae. Plaiycarya (Plate I, fig. 7.8) T h e m o r p h o l o g y of these pollen grains is very characteristic. T E M a n d S E M d a t a a b o u t this f o r m - g e n u s were published by KEDVES a n d STANLEY (1976). T h e i m p o r t a n c e of the regional distribution of the pollen grains of this f o r m - g e n u s is s u p p o r t e d by the fact that we have publications f r o m this point of view; LEOPOLD and MACGINITIE (1972). GRUAS-CAVAGNETTO (1977). Following ZAKLINSKAYA (1962) the a p p e a r a n c e of the genus Platycarya was in the U p p e r C r e t a c e o u s . F r o m U p p e r C r e t a c e o u s layers of Egypt. KEDVES (1971) published as a preliminary report pollen grains belonging to this form-genus. KEDVES a n d DLNIZ (1979) f r o m the layers of Aveiro type spore-pollen assemblage ( S a n t o n i a n - C a m p a n i a n , P o r t u g a l ) published cf Plaiycaryapolleniles semicyclus. because its e n d o a p e r t u r e differ f r o m the pollen grains of the Paleocene and Eocene sediments. In this way the a p p e a r a n c e a n d especially the distribution needs f u r t h e r investigations. It is w o r t h y of m e n t i o n i n g in this place t o o . that the studies of the s p o r o m o r p h s of t h e U p p e r C r e t a c e o u s sediments were interested first in the researches of the Normapolles and Aquilapolleniles (Triprojectaciles), which have a peculiar exine structure. T h e regional distribution of the fossil f o r m s of the genus Platycarya was published by LEOPOLD a n d MACGINITIE (1972), this c o n c e p t i o n was later completed by GRUAS-CAVAGNETTO (1977) with F r e n c h d a t a . O n the basis of the o o Fig. 2. Regional distribution of Triairiupollenites rohorutus d u r i n g the Upper Cretaceous a n d the Tertiary. PF. 1953 PALFOPHYTOGFOGRAPHY OF T H E ANGIOSPF.RM POLLEN GRAINS 41 distribution m a p of LEOPOLD a n d MACGINITIE (1972) the acme of this genus is in the Oligocene epoch. Following o u r new c o m p i l a t i o n (fig. 3) the pollen grains of this genus was very wide-spread in the Paleocene a n d the flowering age was in the Eocene. In the Oligocene its importance d i m i n u t e d , but the pollen grains of this genus were r e m a r k a b l y distributed. As regards the M i o c e n e age our results are identical with those of LEOPOLD and MACGINITIE (1972), namely the reduction of this genus is striking. T h e disappear in N o r t h America, but in E u r o p e occur in several localities. Interesting are d a t a f r o m the Pliocene, in contrast that these are scarce, they c a m e mostly f r o m Europe. N o t e w o r t h y is the occurrence, south of the E q u a t o r , f r o m a deep-sea drilling, near the T i m o r Islands. Fgen.: Paraalnipollenites Syn.: 1976, SUNG TZE-CHEN et H I L L S TSAO LIU. — Fgen. Type: Paraalnipollenites (Plate I, fig. 9-12) Syn.: 1961. et W A L L A C E Fushunpollis alterniporus 1969 here emend gen. nov., p. 155. ( S I M P S O N 1961) S R I V A S T A V A 1975 Alnus alternipora. p. 443, pi. 13. fig. 5. Trialriopollenites confusus. p. 232, pi. 34, fig. 7. cf. 6. 8. 1969, H I L L S and W A L L A C E . — Paraalnipollenites confusus ( Z A K L I N S K A Y A ) n. comb., p. 141. pi. 17. figs. 1-8. 1 9 7 6 . S U N G T Z E - C H E N and T S A O L I U . — Fushunpollis arcuatus sp. nov., p. 155, 156. pi. I I . figs. 32-38. SIMPSON. — 1963, ZAKLINSKAYA. — AUCRETACEO APALEOCENE • EOCENE O OLIGOCENE ® MIOCENE 0 PLIOCENE Fig. 3. Regional distribution of Platycaryapolleniles E . N A G Y 1969 emend. F R E D E R I K S E N et C H R I S T O P H E R 1978 d u r i n g the U p p e r Cretaceous a n d the Tertiary. M. 42 KLDVES Emended diagnosis Triatriate pollen grains. O n one of the hemispheres there is a thickening f o r m of a ring, a n d there are arci t o o . T h e form and the thickness of the arci is varied. By L M m e t h o d n o characteristic sculpture may be detected. R e m a r k s . — T h e pollen grains, which may be classed into this f o r m - g e n u s have four i m p o r t a n t m o r p h o l o g i c characteristic features: 1. the a t r i u m . 2. the polar thickening form a ring. 3. the arci. 4. the more o r less s m o o t h surface. Differential diagnosis T h e joint presence of the above e n u m e r a t e d characteristic features well distinguish f r o m the pollen grains of Trialriopolleniies PF. 1953 a n d Plicapollis PF. 1953. O n the basis of the first d a t a , which were reviewed by HILLS a n d WALLACE (1969) it seemed that the pollen grains of this f o r m - g e n u s are the elements of the Aquilapollenites province, a n d because of its restricted. M a e s t r i c h t i a n . a n d Paleocene occurrence, have a stratigraphic i m p o r t a n c e . T h e earliest d a t a were published f r o m the T u r o n i a n of Europe, this is unique, but no m i c r o p h o t o g r a p h s a c c o m p a n i e d this finding (PACLTOVA, 1973) so it was n o taken into c o n s i d e r a t i o n of the time being. In this way the a p p e a r a n c e r e m a i n the u p p e r part of the U p p e r Cretaceous. T h e youngest occurrences are as follows: Eocene/Oligocene: F a r East, Eocene ?Oligocene: C h i n a . Region of Bohai. T h e r e a r e well established occurrences f r o m E u r o p e a n d Siberia. It is interesting that f r o m the Eocene period these pollen grains were not f o u n d f r o m localities studied palynologically in N o r t h America (fig- 4). Fig. 4. Regional distribution of Paraalnipolleniies H I L L S et W A L L A C E d u r i n g the Upper Cretaceous and the Tertiary. 1969 emend, PALEOPHYTOGEOGRAPHY OF THE ANGIOSPERM POLLTN GRAINS 43 T h e plants p r o d u c t i n g of these pollen grains have probably very peculiar ecologic pretensions. In Western E u r o p e there is only o n e occurrence, published f r o m the Paleocene of M e n a t . F r a n c e in contrast that there are studied several localities. which are rich in s p o r o m o r p h s . Similarly. Eocene is also well k n o w n in Europe, but this genus was d e m o n s t r a t e d only f r o m two localities of Poland, a n d f r o m the Black Sea Region. In this way the occurrence in E u r o p e in the Normapolles province is scarce. N o t yet found in the sub-province of the Atlantic Coast of N o r t h America. Conclusions 1. The regional distribution of all Postnormapolles taxa discussed in this paper overstep the b o r d e r s of the Normapolles province. 2. The geographical a n d stratigraphical occurrence of the most ancient types of triporate a n d triatriate pollen grains reveals t o the h e t e r o g e n e o u s origin of the Postnormapolles. 3. The p a l e o p h y t o g e o g r a p h i c elaboration of the Postnormapolles. Longaxones and other kind of pollen grains may give new ideas t o o u r knowledge about the vegetation history. A C K N O W L E D G E M E N T S On my request. SUNG TzE-CHEN ( N a n k i n g Institiule of Geology a n d Palaeontology. Academia Sinica) was kind e n o u g h t o translate the Chinese text into English a b o u t the genus Fushunpollis. 1 express my sincerest t h a n k s f o r this contribution. Appendix T h e d a t a of the following publications were used for the distribution maps. 1 9 7 5 . Bull Soc. France 17. 6 4 1 - 6 5 5 . ; A K H M I H I V . B R A I S I V A Geol. Palcont. Far Easi. 4 6 - 5 0 . ; B A L K W I L L H O P K I N S . 1 9 7 6 . G E O L . Surv. C a n . Paper 7 6 - 1 B . 3 2 9 - 3 3 4 . ; B A I B U L A T O V A . 1 9 6 8 . Palcont. Meth. for Slrat.. 2 2 6 - 2 2 8 . ; B A I B I L A I O V A . 1 9 7 1 . Palynology of K a z a k h s t a n . 1 1 2 - 1 1 3 . ; B A R B A S H I N O V A . 1 9 7 3 . The Palynology of Cenophytic. 1 0 1 - 1 0 4 . ; B L Y A K H O V A . 1 9 6 6 . F o r the II. Int. Palynol. Conf.. 1 2 7 - 1 3 1 . ; B L Y A K H O V A . 1 9 7 1 . Palynology of Kazakhstan. 1 1 4 - 1 2 5 . ; B L Y A K H O V A . 1 9 7 6 . Palynology of K a z a k h s t a n . 6 9 - 7 7 . ; B L Y A K H O V A KALMINIVA P O N O M A R E N K O . 1 9 7 1 . Palynology of K a z a k h s t a n . 9 3 - 1 0 6 . ; B L Y A K H O V A K O V R I / H N Y K H . 1 9 7 1 . Dokl Akad. Nauk SSSR. 200. 1 4 0 1 - 1 4 0 2 . ; B O I T S O V A — P A N O V A . 1967a. Akad Nauk S S S R . 2 6 2 - 2 7 0 . ; B O I T S O V A — P A N O V A . 1973b. The Paly nology of Cenophytic. 4 2 - 4 7 ; B O M S O V A P A N O V A . 1 9 7 6 . Trudy V S E G E I 1 9 5 . 1 4 4 - 1 7 9 ; BOITSOVA. O S H I RKOVA — P A N O V A . 1 9 7 3 . Methodical Problems of Palynology. 8 - 1 2 . ; B O L O T N I K O V A . 1 9 6 4 . Geology and Geophysics. 1 4 0 - 1 4 2 . ; B O L O I N I K O V A . I9<>6. For the II. Int. Palynol. C o n f . . 1 3 1 - 1 3 6 . ; B O N D A R K N K O . 1 9 7 3 . The Palynology of Cenophytic. X - 1 2 . : B R A I S I V A . 1 9 6 5 . Akad. Nauk SSSR. 1 2 9 . 1 - 4 2 . ; B R A T S F V A . 1 9 6 6 . F o r the I I . Int. Palynol. C o n f . . 1 3 6 - 1 4 1 ; B R A I S I V A . AUFFRET - — VAKHRAMEEV. GRUAS-CAVAGNKTTO. 1976. M 44 P a l y n o l . 2 . 1 1 9 - 1 2 6 . : BRATSEVA. 1 9 6 9 . A k a d N a u k S S S R . 2 0 7 . 1 - 5 6 . ; B R E L I E . F o r t s c h r . 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