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Fossil pollen records of extant angiosperms in China.

 I. Abstract
 II. Introduction
III. Fossil Pollen Records in China
 IV. Comparisons of the Oldest Pollen Records Related to
 Extant Angiosperms in and outside China
 V. Acknowledgments
 VI. Literature Cited


I. Abstract

About 250 fossil pollen types related to 113 extant angiospermous families from the Cretaceous and the Tertiary in China are discussed in the first part of this article. In the second part, comparisons of early records of fossil pollen in and outside China are made to trace the oldest fossil records. Based on these data, we conclude that many plants may have originated in China, a finding that may be important for the study of the origin and early development of angiosperms.

II. Introduction

For a discussion of the fossil pollen records of extant angiosperms, Muller (1981) gathered fossil pollen records related to about 139 extant angiospermous families from most parts of the world and published before August 1, 1980. Because many important Chinese contributions to Cretaceous and Tertiary palynology were mainly published after 1980, Chinese data were not included in Muller's article. Song (2000) reported about 12 types of early fossil pollen records and their development in Chinese strata and gave his view on the origin of the steppe and on their development in China. In the present article we enlarge the studied circle to include all fossil pollen records from China that bear some reliable relation to extant angiosperms. This article can thus be considered an addendum to Muller's. The data in it will be important and interesting for study of the origin and development of angiosperms as well as of the phytogeographical divisions of the world, when the fossil data are really related to extant angiosperms.

We used several pollen floras in China to check these fossil pollen data. Among them were "Pollen Flora of China" (Wang et al., 1995), "Angiosperm Pollen Flora of Tropical and Subtropical China" (Institute of Botany and South China Institute of Botany, Academic Sinica, 1982), and "Pollen Flora of Taiwan" (Huang, 1972).

For easy reference to these fossil records, taxonomic sequence in the text is alphabetical by plant family name. Unless otherwise noted, the plate and figure numbers in parentheses after the fossil names are from Fossil Spores' and Pollen of China, vol. 1, Late Cretaceous and Tertiary Spores and Pollen, by Song et al. (1999), which readers will find useful.

III. Fossil Pollen Records in China

1. ACANTHACEAE

Pollen fossils in this family are commonly found in the Pliocene in southeastern China, and they are often labeled as Acanthus, Restellularia, etc. However, Hypoestespollenites taiwanensisi Li et Huang (1990: pl. 14, figs. 1-2), related to Hypoestes cumingima Benth. et Hook (pl. 4, figs. 15-18), occurs in the early Pliocene Lower Erhchiu Formation in northern Taiwan. Muller (1981) reported some pollen types in this family from the lower Miocene in the Tropics.

2. ACERACEAE

Pollen in this family is tricolp(or)ate with microstriate ornamentation. Fossil pollen, first described in the Miocene in Hungary, is called Aceripollenites Nagy 1969. In China, about 12 species in this genus occur in the Late Cretaceous to the Pliocene, most commonly in the Miocene. Among them are: Aceripollenites microstriatus (pl. 85, figs. 9-11), in the Senonian (the Taizhou Formation in northern Jiangsu and the Mengyejing Formation in Yunnan Province) and the Eocene (the Huayong Formation in Guangdong Province); A. paraneus (pl. 85, figs. 21-22), in the Zhoujiadian Formation (Upper Cretaceous) in Jiangxi Province; A. reticulatus (pl. 85, figs. 18-20), in the upper part of the Lingfeng Formation (Paleocene) in the continental shelf of the East China Sea; A. nanhaiensis (pl. 85, figs. 14-16), in the Buxin Formation (Eocene) in Guangdong Province; and A. tener (pl. 85, figs. 23-25), in the Geojia Formation (Eocene) in the Xizang area. The earliest records of Aceripollenites are Late Cretaceous, possibly Senonian. They developed gradually during the Palaeogene and became widely distributed in the Northern Hemisphere during the Miocene. Aceripollis Huang 1980 differs from Aceripollenites Nagy 1969 in being tricolporate, with the latter being tricolpate. Aceripollis taiwanensis Huang, from the Miocene and the Pliocene (Huang, 1980; Li & Huang, 1990) in the Taiwan area, is related to extant Dipteronia. Muller (1981) indicated the Oligocene record of Acer compestre type in Canada; the Upper Cretaceous datum lacks in convincing evidence.

3. AIZOACEAE

Huang (1980) described two species of Tricolporopollenites--T. ellipticus and T. elongatus--from the Miocene in the Taiwan area and pointed out that they resemble recent Aizoaceae pollen (Huang, 1972: pl. 9, figs. 17, 20-26). However, no special characters differentiate them from pollen types in other tricolporate families.

4. ALANGIACEAE

Fossil pollen showing a relationship to Alangium was recorded in the Tertiary: Alangiopollis eocenicus (early Eocene) and A. barghoornianus (Oligocene) in Europe; Alangium havilandi (Miocene), Alangiopollis javanicoides (Pliocene), etc., in Asia (Muller, 1981). In China there are also some early records of the fossil types; for example, Lanagiopollis (al. Alangiopollis) javanicoides, in the Kongdian-Shahejie Formation (Paleocene-Eocene) in the coastal region of the Bohai Sea and the Liushagang Formation (Eocene) in the northern continental shelf of the South China Sea. Others, such as Alangiopollis verruretinus, Lanagiopollis emarginutus, L. minor, and L. ruguloverruretinus, were discovered from the late Eocene to the Oligocene or to the Pliocene. Thus, fossil Alangium pollen in Asia and Europe occur during approximately the same period.

5. ALTINGIACEAE

Fossil pollen in this family is named Liquidambarpollenites Raatz 1938 ex Potonie 1960. It occurs during the Paleocene in southern Europe and in the Rocky Mountains in North America and is widely distributed in Japan and Europe during the Eocene (Muller, 1981). In China the early recorded Liquidambarpollenites sp., dating from the Late Cretaceous, were found in the Sifangtai Formation in the Songliao Basin (Gao et al., 1999), in the upper part of the Furao Formation in Heilongjiang Province (Liu, 1983), and in the Kukebai Formation in the Tarim Basin (Wang et al., 1990). In addition, many species in this genus were discovered in the Tertiary, especially in the Miocene in the continental shelf of the China Sea (Huang & Song, 2002). Liquidambarpollenites pachydermus Ke et Shi (pl. 199, figs. 20-21), resembling pollen types of extant Altingia based on thicker exine and fewer (8-15) pores, occurs during the Eocene in the coastal region of the Bohai Sea and the Upper Tertiary in the continental shelf of the China Sea. In sum, the earliest records of this family were in China: the Late Cretaceous for Liquidambar; the Eocene for the Altingia type.

6. AMARANTHACEAE

Palynologists abroad often combined pollen data of Chenopodiaceae and Amaranthaceae. The oldest record is from the Maestrichtian in Canada (Muller, 1981). Fossil Vaclavipollis Krutzsch 1966 may be comparable to extant Celosia, owing to its fewer pores and clear polar annulus. Vaclavipollis qaidamensis (pl. 203, figs. 10-11), from the Lower Ganchaigou Formation, is Eocene in age. Other species, such as V. pacltovae and V. radiatus, occur during the Neogene in eastern China.

7. ANACARDIACEAE

The fossil type in this family is commonly represented by Rhoipites Wodehouse 1933 and may bear some relation to extant Rhus. However, it is tricolporate, as are many other families with similar grains. Tricolporopollenites pseudocingulum Potonie resembles pollen in extant Cotinus. Its early records are from the Paleocene and the early Eocene in western Europe; later, perhaps during the Oligocene, it occurs in North America (Muller, 1981). In China similar types, Senonian in age, occur in the Taizhou Formation in northern Jiangsu (Song et al., 1981) and in the Nenjiang Formation in the Songliao Basin (Gao et al., 1999). They were widely distributed in the Tertiary, mainly in the Eocene.

8. APOCYNACEAE

Pollen grains in this family are of many types. Fossil Alyxiaepollenites Sun 1989 (pl. 184, figs. 2-16), with two pores that resemble those of extant Alyxia, occurs in the Lingfeng Formation (Paleocene) in the continental shelf of the East China Sea. Similar forms, such as Psilodiporites wolfendeni and Diporites iskaszentgyoryii, were discovered in the Paleocene in Borneo and the Eocene in western Europe, respectively (Muller, 1981). Banksieacidites insularis Huang (1980) was found in the Miocene in Taiwan.

Multiporate types, such as Parsonsidites minor (pl. 203, fig. 15) from the Mengyejing Formation in Yunnan Province, are late Senonian in age. A similar fossil record was reported in the Oligocene in New Zealand (Muller, 1981).

Fossil pollen similar to extant Micrechites lachnocarpa Tsiang and Ecdysanthera rosea Hook et Arn. was found in the Pliocene and the Pleistocene in Zhaotong, Yunnan Province (Song, 1988). Fossil Trachelospermumpollenites taiwanensis Huang (1980), bearing a relation to extant Trachelospermum or Micrechites, was found in the Miocene in the Taiwan area. Fossil Crassimarginpollenites Wang et Zhao 1980 (pl. 151, figs. 25-36), related to extant Rauwolfia, was reported in the Upper Cretaceous in southern China. Outside China, similar fossils occur only during the Eocene. Tricolporites taiwanensis Huang (1980: pl. 12, fig. 1), from the Miocene in the Taiwan area, bears a relation to extant Plumeria rather than to Rauwolfia because of its smaller size. The pollen morphologies of Rauwolfia and Plumeria are similar, and both possess thickened exine along the colpi.

9. AQUIFOLIACEAE

Pollen grains in this family are characterized by pilae or clavae ornamentation, which is easy to determine. Early records were reported in the Touronian in Australia, the Coniacian in Africa, the Senonian in Asia, the Maestrichtian in North America, and the Paleocene in South America. Although Bolchovitina (1953) described Ilex uralensis in the Cenomanian in Russia, it needs further confirmation (Muller, 1981). In China the oldest record of this family is Ilexpollenites microiliacus (pl. 132, figs. 1-2), from the Quantou and Qingshankou Formations and possibly Albian-Cenomanian in age (Gao et al., 1999). If the assigned age is correct, it is the earliest record of this family. More recently, in the Upper Cretaceous and the Tertiary, fossils in this family occur all over China and are widely distributed in the Upper Tertiary in eastern China.

10. ARALIACEAE

Pollen grains in this family are mainly tricolporate, and their botanical relationships are very difficult to confirm. But fossil Araliaceoipollenites (Tricolporopollenites) edmundii and A. euphorii are similar to extant Aralia in pollen morphology, which may indicate a botanical relationship. They occur in the Paleocene in France and are widely distributed in the Tertiary in Europe (Muller, 1981). Similar fossils were discovered in the Taizhou Formation in northern Jiangsu in the Senonian, and they are also widely distributed in the Tertiary in China. Fossil Rhoipites sp. 1 (Gao et al., 1999: pl. 88, figs. 9, 13), found in the Nenjiang Formation (early Senonian) in the Songliao Basin, is similar to Araliaceoipollenites in morphology.

11. BALSAMINACEAE

Fossil Impatiensidites brevicolpus (pl. 99, fig. 5), bearing a relation to extant Impatiens, was found in the Santan Formation (Pliocene) in the continental shelf of the East China Sea. A similar fossil, I. taiwanensis Li et Huang (1990: pl. 14, figs. 10-11), is from the Lower Erhchiu Formation (Pliocene) in the Taiwan area.

12. BETULACEAE

Early records of fossil Alnipollenites, showing a relationship with extant Alnus, occur during the Santonian and the Campanian in Japan, the Maestrichtian in North America (Canada and the United States), and the Paleocene in Europe (Muller, 1981). They were also found in the Nenjiang Formation (Santonian) in the Songliao Basin (Gao et al., 1999), the Furao Formation (Senonian) in Heilongjiang Province (Liu, 1983), and the Lower Tertiary in northeastern China. Fossil Betulaceoipollenites and Betulaepollenites, both related to extant Betula, have the following early records: the Coniacian-Santonian in Canada, the Campanian-Maestrichtian in Japan, and the Paleocene in Europe (Muller, 1981); and the late Senonian (the Sifangtai and Furao Formations; Liu, 1983; Gao et al., 1999) in northeastern China. Fossil pollen related to extant Corylus, Carpinus, and Ostrya all occurs early in the Senonian (the Furao and Taizhou Formations) and are widely distributed in the Tertiary in China. This is consistent with the records outside China.

13. BOMBACACEAE

Fossil pollen in this family are very scarce in China, and only Bombacacidites qaidamensis (pl. 135, fig. 23) and B. sp. (pl. 141, fig. 14) were found, in the Upper Ganchaigou Formation (Oligocene) in the Qaidam Basin and in the Hanjiang Formation (Miocene) in the continental shelf of the South China Sea. The oldest records of the family in the world are from the Maestrichtian in southeastern America and the Paleocene in Texas (Muller, 1981).

14. BORAGINACEAE

Fossil pollen in this family occurs in the Oligocene outside China (Muller, 1981). Tricolporopollenites equiprojectus (pl. 106, fig. 10), with pollen morphology similar to that of this family owing to its equatorial projection, was discovered in the Youshashan Formation (Miocene) in the Qaidam Basin.

15. BUXACEAE

This family comprises two fossil types. One is Buxapollis, related to extant Buxus. Only Buxapollis fushunensis (pl. 198, figs. 24-25) and B. jianghanensis (pl. 202, figs. 12-14) occur in the Paleocene in the Fushun Basin and in the Eocene in the Jianghan Basin. Similar fossil pollen grains occur only in the Miocene in Europe (Muller, 1981). The other type, Erdtmanipollis, which shows a relationship with extant Pachysandra and Sarcococcus, was found only in the Late Cretaceous abroad.

16. CAPRIFOLIACEAE

Fossil pollen bearing a relation to extant Viburnum and Sambaccus may be included in Caprifollipites or Retitricolporites. The latter is a complex, including many retitricolporate types, and determining its botanical relationships is very difficult. Caprifollipites microratus (pl. 116, figs. 1-4) was found in the late Senonian-Eocene in China in places such as the Mengyejing Formation in Yunnan and the Buxin Formation in Guangdong. Similar fossils occur only in the middle Eocene in Europe (Muller, 1981).

Fossil Lonicerapollis and Diervillapollenites. related to extant Lonicera, Diervilla, etc., occur in the Oligocene in Canada and Europe. But in China, Lonicerapollis simplex (pl. 136, figs. 1-6), Senonian in age, occurs in the Taizhou Formation. Diervillapollenites, such as D. echinatus (pl. 187, figs. 15-18) and D. major (pl. 187, figs. 19-21), are from the Funing and Buxin Formations and are Paleocene and Eocene in age.

Lonicerapollis may be divided into three groups. The first, of which L. pachydermus (pl. 137, fig. 26) is an example, has thick exine and microspinate ornamentation and may be related to extant Abelia chinensis. The second group has thin exine with microspinate and/or granulate ornamentation. Examples include L. gellwitzii (pl. 147, fig. 17), related to extant Dipelta yunnanensis, and L. granulatus (pl. 137, figs. 27-29) and L. intrabaculus (pl. 136, figs. 7-9), related to extant Abelia dielsii. The third group has smooth exine, no clear ornamentation, and no clear relationships.

Diervillapollenites echinatus may be related to extant Weigela coraeensis; D. major (pl. 187, figs. 19-21), to extant W. japonica var. sinica. Echitriporites, though triporate in type, may also be related to this family. This is because the colpus in this family is very short, which may be unclear under fossil conditions. Extant Triosteum pinnatifidum is an herb; its pollen grain is triporate, spinate, and 55.7 x 4.4 [micro]m in size, and some types of Echitriporites may be related to it.

17. CARYOPHYLLACEAE

Fossil pollen in this family was found in the Lower Ganchaigou Formation (Eocene) in the Qaidam Basin. Caryophyllidites minutus (pl. 198, figs. 7-8) and C. tenius (pl. 198, figs. 15-17) resemble extant Dianthus. Similar fossils occur in the Oligocene in other parts of the world (Muller, 1981). Drymariapollenites taiwanensis Huang (1980: pl. 2, fig. 12), from the Miocene in the Taiwan area, can be compared with Drymaria, based on its oblong pollen with oblong pores.

18. CASUARINACEAE

Fossil Casuarinidites Cookson et Pike 1954, showing a relationship with this family, occurs early in the Lower Paleocene in New Zealand, in the middle Paleocene in Australia, in the Paleocene in India and Argentina (Muller, 1981), and only in the Upper Tertiary in China. But fossil Casurinaepollenites Sun 1989 (Sun et al., 1989: pl. 12, figs. 3-16), possibly related to extant Casuarina equisetifolia, was found in the Lingfeng Formation (Paleocene) in the continental shelf of the East China Sea. This genus differs from Casuarinidites in its lack of aspis in the pore area and its granulate or microstriate ornamentation: The latter has aspis structure in the pore area and has psilate or intragranulate ornamentation.

19. CELASTRACEAE

Fossil pollen like that of extant plants such as Micratropis, Peritassa, Campylostemon, Hippocraten, etc., commonly occurs in the Oligocene in other parts of the world (Muller, 1981) but was not found in China. In China, Tricolporopollenites microtopiformis (pl. 101, figs. 12-14) has transverse long pores rather than the rounded pores found in extant Micratopis. It is not related to this family. Fossil Evonymoipites Zheng 1985 (pl. 115, figs. 1-6), related to extant Evonymus and Celastrus, was discovered in the Miocene in the continental shelf of the East China Sea. In addition, fossil leaves of Celastrus mioangulata and Evonymus protobungeanus were reported in the Shanwang Formation (Miocene) in Shandong Province.

20. CENTROLEPIDACEAE

Fossil pollen in this family, such as Milfordia incerta = M. hypolaenoides, was found in the Paleocene in Europe, the lower Eocene to Miocene in France, the Oligocene in New Zealand, and the Pliocene in Australia (Muller, 1981). In China, fossil Centrolepidacidites scrobiculatus (pl. 204, figs. 9-13) and C. typicus (pl. 204, figs. 14-15), possibly related to this family, occurs in the Paleocene Nongshan Formation.

21. CERCIDIPHYLLACEAE

Fossil Cercidiphyllites tuberculatus Song (1996: pl. 2, fig. 7), from the Naomugen Formation (Paleocene) in the Inner Mongol area, is tribrevicolpate, with wide (18-25 [micro]m) colpi and rounded terminal ends. These characters are comparable to those of extant Cercidiphyllum, but the fossil pollen has tuberculate and punctate ornamentation, whereas the recent pollen has microreticulate ornamentation. In addition, recent Euptelea pollen has microreticulate ornamentation and tribrevicolpate aperture. A Cercidiphyllum arcticum fossil leaf was collected in the Eocene Fushun Formation. Outside China, both the fossil leaf (Cercidiphyllum ellipticum) and the pollen (Cercidiphyllidites brevicolpatus) were recorded in the Upper Cretaceous in America (Muller, 1981).

22. CHENOPODIACEAE

Pollen types of Chenopodiaceae, Amaranthaceae, and Caryaphyllaceae are all multiporate and distinguishable from each other by exine structure and ornamentation. Early records of Chenopodipollis (fossil Chenopodiaceae pollen) are from the Upper Paleocene (Leopold, 1969), the Eocene (Muller, 1970), and the Maestrichtian-Danian in Kazakhstan (Zaklinskaya, 1963) and Canada (Muller, 1981). In China the earliest records of Chenopodipollis spp. are from the Qingshankou Formation (Cenonmanian) in the Songliao Basin (Gao et al., 1999), the Taizhou Formation (Coniacian-Santonian) in northern Jiangsu (Song et al., 1981), and the Furao Formation (Maestrichtian-Danian; Liu, 1983) in Heilongjiang Province. They were well developed during the Tertiary in northwestern China, occupying about 10% of the assemblage in the Upper Ganchaigou Formation (Oligocene) and 20% in the Youshashan Formation (Miocene) in the Qaidam Basin.

23. CHLORANTHACEAE

Clavatipollenites, related to extant Ascarina in this family, is the oldest real monocolpate angiosperm pollen. According to Muller (1981), Clavatipollenites-Ascarina-type pollen were found: in Central Africa from the Aptian-Albian; in South America from the Aptian; in North America from the Aptian, the Albian, the Cenonmanian, and the Maestrichtian; in Europe from the Aptian, the Albian, and the Cenonmanian; in Austrialia from the Albian, the Cenonmanian, and the Turonian; and in New Zealand from the Maestrichtian to the Pliocene since the Barremian (Muller, 1984). Outside China, however, fossil pollen of the Hedyosmum type were found only from the late Miocene.

According to Hua (1991) and Zhou (1999), the following fossil types belong to this family: Clavatipollenites hughesii, related to extant Acarina and occurring in the Potomac Group (Late Barremian--Early Aptian) in America and the Barremian-Aptian in northern and northeastern China; Asteropollis asteroides, related to extant Hedyosmum and occurring in the same strata as the above species and also in the Saihan Tal and Hadat Formations, in the Eaming Formation in the Inner Mongol area, in the Zhoujiadian Formation in Jiangxi Province, and in the Lumuwun Group in Heinan Province, where they are Late Aptian to Late Cretaceous; Hammenia (Stephanocolpites) fredericksburgensis, possbly comparable to extant Chloranthus, from the Albian in America and the Hadet Formation (Albian) in China; and Chloranthistemon endresii, possibly related to this family from the Late Santonian in Sweden (no data in China). In addition, Chloranthacearumpollenites, which may be related to this family, were found in the Upper Tertiary in Hungary (Nagy, 1969) and China, including the Youshashan and Shizigou Formations in the Qaidam Basin in China, originating in the Miocene-Pliocene.

24. COMPOSITAE

This family is an advanced group of angiosperms, most of which are herbs. Megafossils in this family were first discovered in the Oligocene (Muller, 1970); pollen fossils are from the Oligocene (Muller, 1981), the late Oligocene (Leopold, 1969), the early Miocene (Muller, 1970; Nagy, 1985), and the Pliocene (Krutzsch, 1957; Germeraad et al., 1968) in various parts of the world. The family comprises four pollen fossil genera: Artemisiaepollenites Nagy 1969, Cichoreacidites Sah 1967, Echitricolporites Van der Hammen ex Germeraad et al. 1968, and Tubulifloridites Cookson 1947 ex R. Potonie 1960. Their occurrence and time of flourishing in China are:

Artemisiaepollenites, related to extant Artemisia, was first discovered in the Upper Gancaigou Formation (Oligocene) in the Qaidam Basin and was designated A. minor. It was sparsely distributed during this period but widely distributed during the Upper Tertiary, sometimes with a high frequency. For example, in the Qaidam Basin it occupied about 3% in the Lower Youshashan Formation, about 5% in the Upper Youshashan Formation (Miocene), and up to 8-26% in the Shizigou Formation (Pliocene). In sum, this fossil first occurred during the Oligocene and flourished during the Late Tertiary, especially the Pliocene and the Pleistocene. This fossil pollen was occasionally found in the late Eocene to the middle Oligocene in Kazakhstan (Zaklinskaja, 1957: 22, 28); it was determined to be Artemisia and also occurs abroad in considerable numbers during the Pliocene and the Pleistocene.

Cichoreacidites. Cichoriaearumpollenites Nagy 1969 is a synonym of this genus, and Fenestrites Van der Hammen 1956 is similar to this genus in morphology. Its distribution is synchronous to the former genus. It was discovered in the Upper Ganchaigou Formation (Oligocene) in the Qaidam Basin and determined to be Cichoriaearumpollenites sp., but most species--such as C. donghaiensis (pl. 133, figs. 31-33), C. gracilis (pl. 133, figs. 21-23), C. ixeriformis (pl. 133, figs. 15-16), and C. magnus (pl. 133, fig. 30)--were distributed in the Upper Tertiary.

Echitricolporites is related to the Astereae (Germeraad et al., 1968). Fossil pollen in this genus was first discovered in the upper part of the Lulehe Formation (Paleocene--early Eocene) and called E. conicus (pl. 133, figs. 28-29). In the Lower Ganchaigou Formation (Eocene) this genus contains two or three species and has a frequency of about 1%; in the Upper Ganchaigou Formation, about three or four species and 2.5% frequency; in the Lower Youshashan Formation, about four species and 3.5% frequency; and in the Upper Youshashan Formation (Miocene), about eight species and 15% frequency; its frequency decreases to 3% in the Shizigou Formation (Pliocene). The fossil pollen is widely distributed during the Tertiary in China but was often included in Compositae rather than individually. Therefore, details of its appearance and distribution are not clear.

Tubulilifloridites is related to the Tubuliflorae of the Compositae. Here we also use data from the Qaidaim Basin to describe the appearance and time of flourishing of this fossil. The earliest record of this genus is from the lower part of the Lower Ganchaigou Formation (early--middle Eocene), named T anthemidearum (pl. 134, fig. 7) (about 1%). The genus has about nine species and 2.5% content in the upper part of the same formation, about 10 species and 4.0% in the Upper Ganchaigou Formation (Oligocene), and about 7% and 6% contents in the Youshashan and Shizigou Formations, respectively.

In sum, fossil Compositae pollen first occurred in the Paleocene--early Eocene (e.g., Echitricolporites), the early middle Eocene (e.g., Tubulifloridites), and the Oligocene (e.g., Cichoreacidites and Artemisiaepollenites). In the Upper Tertiary all four genera were distributed widely and in great numbers throughout China, especially the latter two genera, which developed greatly in some areas of China during the Pliocene and the Pleistocene (Song, 2000; Wang, 2004).

25. CONVOLVULACEAE

Fossil pollen in this family occur early in the lower Eocene in Cameroon and during the middle Eocene in South America (Muller, 1981). In China these fossils, first determined to be Comvolulum but later transfered to Fupingopollenites, show some similarity to the Hoplestigmataceae (Wang, 2000) but no relation to the Convolvulaceae. Recent research has shown that Fupingopollenites represents a highly successful but extinct Tertiary taxon whose relationships remain uncertain (Wang & Harley, 2004).

Tricolpites erycibeformis (pl. 96, fig. 11), from the Upper Ganchaigou Formation (Oligocene) in the Qaidam Basin, may be related to extant Erycibe myriantha. Evolvuluspollenites taiwanensis Li et Huang (1990: pl. 15, fig. 14), from the Pliocene in the Taiwan area, resembles extant Evolvulus alsinoides.

26. CORNACEAE

It is difficult to relate Cornaceoipollenites Potonie 1951 ex 1960 to this family. Fossil pollen Cornus bremanoirensis Simpson 1961 was reported from the Maestrichtian in Scotland. A similar fossil, Cornaceoipollenites bremanoirensis (pl. 103, figs. 26-27), was found in the Louhutan Formation (Paleocene) in the Fushun Basin, and low values of C. sp. were found in the Late Cretaceous in sites such as the Taizhou and Furao Formations. About eight species in this genus have been discovered in the Tertiary in China.

27. CRUCIFERAE

Fossil Tricolpites capsellaformis (pl. 96, fig. 8) and T. hesperisformis (pl. 96, figs. 12-13), which may be related to extant Capsella bursa-pastoris and Hesperis matromalis, respectively, were discovered in the Lulehe (Paleo-Eocene) and Youshashan (Miocene) Formations in the Qaidam Basin.

28. CUCURBIATACEAE

Fossil Rousea taiwanensis Li et Huang (1990: pl. 15, figs. 16-17), from the Pliocene in the Taiwan area, can be compared with Thladiantha species.

29. CYPERACEAE

Fossil pollen in this family is called Cyperaceaepollis Krutzsch 1970. It occurs in the Upper Ganchaigou Formation (Oligocene) in the Qaidam Basin and includes two species, C. neogenicus (pl. 196, fig. 38) and C. scholitzensis (pl. 196, fig. 39). Other strata, such as the Santan and Lower Yancheng Formations and the Fotan Group, also contain these fossils. In the upper part of the Fotan Group the fossils in this genus occupy about 4.7% of the assemblage (Zheng et al., 1994). Outside China, these fossils were discovered in the middle Eocene (Muller, 1981) and the Miocene in Germany (Jansonius & Hill, 1976).

30. CYRILLACEAE

Based on pollen morphology, fossil Cyrillaceaepollenites Murrigen et Pflug ex Potonie 1960 is likely to be Cupuliferoipollenites. The former differs from the latter only in its broader body and transverse pores. The oldest record is C. cf. megaexactus, found in the Qingshankou Formation (Cenomanian) in the Songliao Basin in northeastern China (Gao et al., 1999: pl. 88, figs. 10-11) and in the Maestrichtian in America (Muller, 1981).

31. DAPHNIPHYLLACEAE

Fossil Daphniphyllumpollenites ignotus and D. aldhamii Huang (1980: pl. 9, figs. 1-4), from the Miocene in the Taiwan area, resemble extant Daphniphyllum.

32. DIPSACACEAE

Fossil types in this family, such as Dipsacacus, occur in the Miocene in Europe (Muller, 1981); Scabiosaepollenites, in the Upper Tertiary in Hungary (Nagy, 1969). Fossil Scabiosapollis in China differs from Scabiosaepollenites in being tricolpate, whereas the latter is colporate and very large. The former is occasionally found in the latest Cretaceous--e.g., S. minutus (pl. 92, figs. 7-8), in the Mengyejing Formation in Yunnan Province--but is mainly distributed in the Palaeogene--e.g., S. haianensis (pl. 92, figs. 3-4), S. fushunensis (pl. 92, figs. 1-2), and S. intrabaculus (pl. 92, figs. 5-6).

Jianghanpollis and Morinoipollenites Wang et Zhao 1979 may be related to extant Morina sect. Dictocalyx. Muller (1981) proposed that "their detailed description does not match entirely with the data provided by Erdtmann (1952).... It may represent an extinct group of the plants possibly related to Dipsacaceae." But Farabee (1990) thought that they were related to extant Morina. About 15 species of Morinoipollenites and 14 species of Jianghanpollis were discovered, mainly in the Upper Cretaceous--e.g., M. normalis (pl. 163, figs. 7-8, 15-16) and J. ringens (pl. 162, figs. 13-15, 26)--but occasionally in the Paleocene. Jianghanpollis henancrisis (pl. 162, figs. 6, 12, 16-17) is the earliest record in the Qingshankou Formation (Cenomanian) in the Songliao Basin (Gao et al., 1999).

33. DIPTEROCARPACEAE

Fossil Hopeoipollis hopeoides, H. shoreoides, and H. parashoreoides Zheng 2000, from the Fotan Group (Miocene), may be matched to extant Hopea, Shorea, and Parashorea. Outside China, fossil pollen related to Dipterocarpus and Shorea albida have been discovered in the Oligocene and the Upper Miocene in Borneo (Muller, 1981).

34. DROSERACEAE

Fossil Droserapites clavatus and Droserapollis gemmatus Huang (1980: pl. 17, figs. 4-6, 9-10) are tetrad, inaperturate, with clavate, baculate and gemmate processes. They match extant Drosera and were discovered in the Miocene in the Taiwan area.

35. ELAEAGNACEAE

Elaeangnacites Ke et Shi 1978 and Slowakipollis Krutzsch 1960 are the representative of the fossil types in this family. The former--e.g., E. sp. (Song & Qian, 1989)--was discovered in the Taizhou Formation and dates from the Senonian; others--e.g., E. huanghuaensis (pl. 138, figs. 1-5)--are mainly from the Tertiary. The latter, early record is from the Chijiang Formation in Jiangxi Province (Sun & He, 1980) and is Paleocene in age, but S. elaeagnoides (pl. 138, fig. 10), S. neogenicus (pl. 138, fig. 17), etc., were sparsely distributed in the Tertiary. Fossil pollen in this family, determined to be Elaeagnus, was also discovered in the middle-upper Eocene in Kazakhstan (Zaklinskaya 1963: 128) and in the Oligocene in Europe (Song, 2000),

36. ELAEOCARPACEAE

Fossil Elaeocarpollenites taiwanensis Huang (1980: pl. 16, fig. 1) and E. formosonus Huang (1980: pl. 5, figs. 19-20), found in the Miocene in the Taiwan area, have been matched to extant Elaeocarpus. They are tricolporate and small.

37. ERICACEAE

Fossil types in this family have many names, such as Ericipites Wodehouse 1933, Ericaceaepollenites Thiergart 1937, 1938, and Ericaceoipollenites Potonie 1951 ex 1960. The oldest record is from the Green River Formation (Eocene) in America. In China there are two species, Ericipites callidus (pl. 207, figs. 10, 14) and E. ericius (pl. 207, figs. 8-9, 11-13), which occur commonly in the Upper Tertiary. A few specimens of this genus were also found in the Dannan Formation (also Eocene) in northern Jiangsu (Song et al., 1981).

38. EUPHORBIACEAE

Fossil pollen in this family is called Euphorbiacites Zaklinskaja 1965 ex Li, Sung et Li 1978. As a tricolporate type, it includes about 17 species. The oldest species, E. reticulatus (pl. 117, figs. 11, 15) and E. majorporus (pl. 122, figs. 12-13), are from the Upper Cretaceous (the Taizhou, Paomagang, and Mengyejing Formations); the other species are widely distributed in the Tertiary in China. In addition, Planotricolporites (al. Euphorbiacites) euphorbioides (pl. 140, fig. 6), from the Lower Ganchaigou Formation (Eocene) in the Qaidam Basin, may be related to extant Euphorbia marginata, because their tricolporate structure is similar.

39. EUPTELEACEAE

This family has only one genus and two species. In China, pollen Euptelea pleiosperma grains are characterized by their oblong body, tricolpate aperture, and short, broad colpi with rounded ends. Eupteleapollis germanicus Krutzsch 1966, found in the upper Eocene in Germany, differs from recent Euptelea in its periporate aperture, but Muller (1981: 16) nonetheless confirmed their botanical relationship. In China, Eupteleapites elegans and E. euptelanformis Zheng et al. 1994, from the Fotan Group (Miocene) in Fujian Province, are matched to recent Euptelea.

40. FAGACEAE

About four types of pollen grains belong to this family. The Castaneoideae subfamily includes extant Castanea, Castanopsis, and Lithocarpus; the fossil name is Cupuliferoipollenites Potonie 1951 ex 1960. It is small, with tricolporate aperture and smooth exine. The earliest record in China is from the Qingshankou Formation (Cenomanian) in the Songliao Basin (Gao et al., 1999). Pollen fossils are widely distributed in the Upper Cretaceous and the Tertiary, chiefly in the Lower Tertiary, and they are dominant in some assemblages. The fossils are also found in the Santonian/Campanian in Canada, in the Campanian in Europe, and in the Maestrichtian in America (Muller, 1981).

Fossil pollen in the Quercoideae subfamily is named Quercoidites Potonie, Thomson et Theirgart ex Potonie 1960. It is tricolpate, with granulate ornamentation. The small group, such as Q. minutus (pl. 88, figs. 3-5) and Q. microhenrici (pl. 88, figs. 12-15), from the late stage of the Early Cretaceous to the early stage of the Late Cretaceous, were found early in the Denglouku, Quanton, and Qingshankou Formations in the Songliao Basin. Further study will be needed before they can be differentiated from Tricolpites. The large group, including Q. asper (pl. 88, figs. 18-20) and Q. henrici (pl. 88, figs. 15-17), was found early in the Qingshankou Formation (Cenomanian) (Gao et al., 1999).

The Fagoideae fossil type is called Faguspollenites Raatz 1937. It is large and tricolporate pollen with punctate to granulate ornamentation. The oldest record in China is from the Taizhou Formation (Senonian). The pollen is sparsely distributed in the Tertiary.

Fossil Nothofagoideae pollen, called Nothofagidites, is widely distributed in the Late Cretaceous and the Tertiary in the Southern Hemisphere. A few fossils (?) in this genus have also been found in the Mengyejing Formation (late Senonian) in Yunnan Province (Song et al., 1976: pl. 8, fig. 18, Nothofagidites? sp.) in the Northern Hemisphere.

41. GENTIANACEAE

Intragranulitricolporites pachydermus (pl. 113, fig. 15) resembles recent Gentinana in pollen morphology, so the two may be botanically related. It was found in the Miocene Youshashan Formation in the Qaidam Basin. Nymphoidaipites striatus (pl. 140, fig. 8) is a type with 3-4 parasyncolporate, and its pollen morphology is similar to that of extant Nymphoides coreana. Fossils were found in the Miocene Heilongjing and Yuquan Formations in the continental shelf in the East China Sea. In addition, Dermatobrevicolporites dermatus (Song, 1996: pl. 2, figs. 19-20), comparable to extant Biebersteinia heterostemon in pollen morphology, occurs in the Naomugen Formation (Paleocene) in the Inner Mongol area. Fossil pollen related to this family also occurs in the Maestrichtian in Canada and the Paleocene in America (Muller, 1981).

42. GERANIACEAE

Fossil Geraniapollis compactilis (pl. 95, fig. 32), from the Upper Ganchaigou Formation (Oligocene) in the Qaidam Basin, is characterized by its large size, its short, wide, tricolpate aperture, and its solid exine, which matches the pollen morphology of extant Geranium. Fossil types in this family, such as Geranium phaeus, Erodium, and Pelargonium echinatum, all occur in the Late Tertiary in the other parts of the world (Muller, 1981).

43. GRAMINEAE

Fossil pollen in this family is called Graminidites Cookson 1947 ex Potonie 1960. It is monoporate with annulate margin. Its earliest records are from the Paleocene (Muller, 1981) and the early Eocene (Leopold, 1969). In China, fossil pollen in this genus was discovered early in the Kongdian Formation (early Eocene); that of Graminidites major (pl. 182, fig. 23), in the Lower Ganchaigou Formation (early-middle Eocene). It is also sparsely but widely distributed in the Lower and Upper Tertiary. In addition, Monoporopollenites sp. (Gao et al., 1999: pl. 82, fig. 39), which is large (ca. 50 [micro]m), and has thin exine with many folds and annulate pores, may be related to this family. This fossil was discovered in the Denglouku Formation (Albian) in the Songliao Basin (Gao et al., 1999).

44. HALORAGACEAE

Fossil types in this family are commonly Myriophyllum and Haloragis types. Outside China, early records are from the Paleocene for the former and from the early Eocene for the latter (Muller, 1981). In China, Haloragacidites punctatus (pl. 195, figs. 8-9) occurs in the Sandou Formation (Eocene) in northern Jiangsu; Myriophyllumpollenites minimus (pl. 195, figs. 6-7), in the Minghuazhen Formation (Miocene) in northern China.

45. HAMAMELIDACEAE

Fossil Altingia and Liquidambar pollen was mentioned in the section on Altingiaceae. Other fossils in this family are mainly tricolpate or tricolporate, and it is very difficult to determine their botanical relationships. Fossil Parrotia and Hamamelis pollen was found in the Miocene and the Pliocene in France; Fothergilla, possibly in the Eocene in Russia (Muller, 1981).

46. HIPPOCRATEACEAE

Hippocrateaceaedites leizhouensis (pl. 141, figs. 15-18), which resembles Hippocratea in pollen morphology, was found in the Wanglugang Formation (Pliocene) in the northern continental shelf of the South China Sea; similar fossils were also found in the Miocene in southern India (Ramanujam, 1966). Other fossils, comparable to extant Hippocratea volubilis and H. myrianthe, occur in the Oligocene to the Lower Miocene in Cameroon (Muller, 1981).

47. HYDROCARYACEAE

Sporotrapoidites, the fossil pollen type in this family, has special morphology and is easily distinguishable. Sporotrapoidites rotundiporus (pl. 192, figs. 14-16) is found early in the Paleocene Lingfeng Formation and commonly in the Upper Tertiary in China. In addition, this species differs from recent Trapa pollen in its lack of exine rugae, and for some time it was included in the genus Hemitrapa (Liu, 1986).

48. ICACINACEAE

Pollen of the lodes type is tri-pantoporate. Kedves (1970) recognized Compositoipollenites rhizhophorus, which belongs to this type. These fossils occur in the lower Paleocene-upper Eocene in western Europe, the middle Eocene in America, and the lower Eocene-Miocene in Africa (Muller, 1981). In China, similar fossils, determined to be Intratriporopollenites rhizophorus, were found in the Funing and Huayong Formations (Paleocene-Eocene).

Spinotriporites Song 1996 was widely distributed during the Late Cretaceous and the early Tertiary in northwestern China. Pollen in this genus is more subtriangular than rounded, and pores are commonly situated on the equator. It is thus distinguishable from Compositioipollenites.

Pollen of the Platea type is tricolporate. Commonly called Favitricolporites baculoferum, it occurs in the Paleocene in America and Europe (Muller, 1981). Favitricolporites funshunensis (pl. 103, figs. 23-24), F. helmstedtensis (pl. 112, figs. 19, 24), F. minifoveolatus (pl. 112, figs. 20-22), F. oblatus (pl. 132, figs. 18-19), F. retifoveolatus (pl. 112, figs. 25-26), and other fossils have come from the Eocene Guchengzhi Formation in the Fushun Basin.

49. JUGLANDACEAE

Pollen fossils of the Engelhardtia type occur early in the Maestrichtian and Paleocene in western America and in the Paleocene in Europe (Muller, 1981). In China, Paleocene Engelhardtioidites and Engelhardtioipollenites fossils, including E. concavus (pl. 188, figs. 7-8), E. paleocenicus (pl. 188, figs. 10-13), and E. punctatus (pl. 188, figs. 14-18), were found in the Lingfeng Formation.

Early records of fossil Caryapollenites were reported from the middle--late Paleocene in Central America. More recent fossils are distributed in the Tertiary in America and Europe. Nichols believed that the species Momipites leffingwellii might be the ancestral link to the modern Carya type (Muller, 1981). Although some fossils from the Campanian-Maestrichtian in Canada are similar to Carya, their morphology needs further study.

In China, Caryapollenites granulatus (pl. 192, figs. 8-10), found in the Taizhou Formation (Senonian), resembles that of extant Carya tonkinensis in morphology and can also be compared with fossils from the Campanian in Canada with granulate to tuberculate ornamentation. Paleocene fossils, such as C. juxtaporites (pl. 193, figs. 1-2), C. minor (pl. 192, figs. 18-21), C. triangulus (pl. 193, figs. 25-28), and C. polarannulus (pl. 193, figs. 21-24), found in the Lingfeng Formation, can be compared with modern Annamocarya sinensis pollen. Caryapollenites simplex, which resembles modern Carya cathayensis pollen, is widely distributed in the Upper Tertiary in China (Huang & Song, 2002).

Early fossil pollen records of Juglans, such as Juglanspollenites minutus (pl. 198, fig. 8), from the Late Cretaceous and the Eocene in America and from the early--middle Eocene in Europe (Muller, 1981), were also found in the Taizhou Formation (Senonian) in China. These fossils are commonly distributed in the Paleocene and Eocene, even to the Pliocene, in China.

Fossil Platycaryapollenites occurs early in the upper Paleocene in America and Europe (Muller, 1981). In China, similar fossils, such as P. dongyingensis (pl. 190, figs. 1-2), were found in the Lingfeng and Shahejie Formations (Paleocene-Eocene). But a few specimens of P. sp., discovered in the Furao Formation in Heilongjiang Province, are Maestrichtian in age (Liu, 1983).

Palynologists outside China always grouped Juglans and Pterocarya types together, but the former differs from the latter in its pore structures. Pterocaryapollenites stellatus occurs early in the upper Paleocene-Eocene in America and the upper Eocene in Europe (Muller, 1981). Similar fossils (pl. 196, figs. 34-37) were found in the Taizhou Formation (Senonian) and are commonly distributed in the Tertiary in China.

50. LABIATAE

Modern pollen in this family is of two types: tricolpate and hexacolpate. Fossils of the former are called Labitricolpites. Early records, such as L. microgranulatus (pl. 87, figs. 26-28), comparable to extant Leonurus in pollen morphology, were found in the Eocene Kongdian Formation; and other species are distributed in Tertiary, mainly after the Oligocene. Fossil Retihexacolpites taiwanensis, from the Miocene in the Taiwan area, is hexacolpate.

51. LAURACEAE

Pollen exine in this family is too thin for preservation, although Inaperturopollenites palaeogenicus and I. spicatus fossils from the upper Paleocene in France and Belgium are comparable (but not convincingly so) to extant Cinnamomum. Similarly, the real botanical relationships of fossils, including Peltandripites davisii (pl. 204, figs. 21-22), P. lauruciformis (pl. 204, figs. 1-2), P. minor (pl. 204, fig. 23), and P. weixianensis (pl. 204, figs. 25-27), sparsely distributed in the Upper Cretaceous and Lower Tertiary in China, cannot be determined, even though they were compared with this family.

52. LECYTHIDACEAE

Fossil pollen in this family, called Marginipollis Clarke et Froderiksen 1968, is related to extant Berringtonia. Early records are from the lower Eocene in India and the middle Eocene in Cameroon (Muller, 1981). In China, M. elegans (pl. 88, figs. 28-29), M. robustus (pl. 88, figs. 37-38), and other fossils were found only in the Miocene Fotan and Hanjiang Formations.

53. LEGUMINOSAE

Fossil Margocolporites, which show a relationship with extant Caesalpinia and Pterolobium, is characterized by 3-zoni-margocolporate apertures. This pollen type often occurs during the Eocene in equatorial areas, especially in Assam. Muller (1981) mentioned that Assam "may have been the cradle of that part of Caesaloiniaceae." However, M. conspicuus (pl. 117, figs. 67), which occurs in the Taizhou and Furao Formations in China, is Senonian in age and thus older than the Indian pollen.

Tricolporopollenites caraganoides (pl. 105, figs. 1-3), comparable to extant Caragana in pollen morphology, was found in the Late Cretaceous Paomagang Formation and is sparsely distributed in the Tertiary. T. chijiangensis (pl. 108, figs. 1-2) is comparable to extant Lotus in pollen morphology and occurs in the same strata as the former species.

54. LILIACEAE

Fossil Liliacidites may be related to this family. It was recorded in the Upper Lower Cretaceous and Upper Cretaceous in the Songliao Basin, represented by Liliacidites sp. 2 (in the Denglouku Formation), L. sp. 1, L. sp. 3, L. cf. microreticulatus (the Qingshankou Formation), and L. creicus (the Quanton Formation) (Gao et al., 1999). There are certain differences in their morphology compared with the recent pollen in this family. Some fossils, including L. minor (pl. 81, fig. 10) and L. microreticulatus (pl. 81, fig. 15), from the Senonian (in the Taizhou and Furao Formations) may be roughly compared with the recent pollen in this family. Fossil Lilipollis lilioides (pl. 81, fig. 28), which bears a relation to extant Lilium, occurs in the Pliocene in Central Europe and the early Miocene in China.

55. LOGANIACEAE

Fossil Assamialetes taiwanensis Huang (Huang, 1980: pl. 17, figs. 8-9), compared with extant Fagraea sasakii in pollen morphology, has inaperturate pollen with reticulate ornamentation and was discovered in the Miocene in the Taiwan area.

56. LORANTHACEAE

Fossil Gothanipollis may be comparable to extant Loranthus, Taxillus and Amylothera in pollen morphology. Its earliest records were from the early and middle Eocene in Germany and the middle and late Eocene in America. It also occurs in the Oligocene and Miocene in some areas (Muller, 1981). Some species, including Gothanipollis? elegans (pl. 150, figs. 4, 7-8), G. gothani subsp. plicus (pl. 150, figs. 1-3), and G? latiproctus (pl. 130, figs. 16-18), from the Upper Cretaceous in the Songliao Basin, differ in some ways from extant Loranthus, and there is no real relationship with this genus. But G. bassensis (pl. 129, figs. 1-2) can be matched to extant Loranthus in pollen morphology. It was found in the Eocene Liushagang Formation and other Tertiary areas in China. Fossil Loranthacites scabratus (pl. 94, fig. 45), found in the Eocene Huayong Formation, may be compared with extant Elytranthe in pollen morphology. Fossils of Elytranthe Couper (1960), as well as the Cranwellia-Scollardia group, which are mainly distributed in the Upper Cretaceous, may be the ancient representatives of this family.

57. LYTHRACEAE

Fossil Planotricolporites lagerstromiaformis (pl. 120, fig. 1), found in the Guantou Formation (Miocene) in China, resembles extant Lagerstromia subcostata in pollen morphology. Similar pollen grains were found in the Miocene in Korea (Yamanoi, 1992: fig. 8, 16). Fossil Lythraites Yu, Gao et Mao (pl. 153, figs. 10-18, 22-23; pl. 158, figs. 10-12), including about seven species, are all distributed in the Upper Cretaceous in China. They all differ from recent Cuphea in pollen morphology, although original authors assigned their relationship to them.

58. MAGNOLIACEAE

Pollen grains in this family are monosulcate, and it is very difficult to differentiate them from the Mesozoic monosulcites pollen. Fossil leaves in this family occur in the Late Cretaceous Dakota Formation in America, as well as in the Oligocene in Jinggu and the Miocene in Shanwang in China. Magnolipollis magnolioides (pl. 83, figs. 9-10) occurs in the latest Early Cretaceous Baihedong Formation in the Sanshui Basin in China and in the Eocene outside China. The other genus in this family is Liriodendron. It differs from Magnolia in its sparsely tuberculate ornamentation: The latter has only punctate or granulate ornamentation. Fossil Liriodendronpollis sp. (pl. 82, fig. 1), found in the Eocene Xibu Formation in the Sanshui Basin, is earlier than the Oligocene record in Siberia.

59. MALVACEAE

Pollen in this family is periporate, with echinate ornamentation. Early records of Echiperiporites were discovered in the upper Eocene in America and Brazil, the Oligocene in Africa, and the Miocene in other areas (Muller, 1981). In China, Malvacearumpollis magnus (pl. 201, fig. 16) and Malvacipollis dispinus (pl. 201, fig. 15) were found in the Pliocene Santan Formation; Malvacipollis minor (pl. 201, fig. 14), in the Miocene in the Yunnan and Xinjiang areas. Abutilonacidites bohaiensis (pl. 185, figs. 34-36), found in the Miocene Minghuazhen Formation in northern China, resembles extant Abutilon in pollen morphology.

60. MELASTOMACEAE

Fossil pollen Bredeliapollis oldhamii Huang (1980: pl. 10, figs. 28-29) which is tricolporate and small (11-15 [micro]m), resembles that of extant Bredelia oldhemii. It is from the Miocene in the Taiwan area.

61. MELIACEAE

Fossil pollen in this family is called Meliaceoidites Wang 1978 ex Wang 1980. It is tricolporate and resembles extant Cipadeosa in pollen morphology. Early records of M. huaianensis (pl. 109, figs. 3-5) were reported in the Taizhou Formation (Senonian) and the Lulehe and Lower Ganchaigou Formations (Paleocene-Eocene). Tetracolporites sp. (Gao et al., 1999: pl. 89, fig. 39), found in the Mingshui Formation (late Senonian) in the Songliao Basin, may also be compared with extant Toona in pollen morphology. Another species, T. melioides (pl. 140, fig. 5), found in the Eocene Shahejie and Buxin Formations, resembles extant Toona too.

62. MENISPERMACEAE

Fossil Pericampyluspollenites formosensis Huang (1980: pl. 11, figs. 1-5) pollen is tricolporate and small (17 x 5 [micro]m), and its colpi are as long as its polar axis. Discovered in the Miocene in the Taiwan area, it is similar to that of extant Pericampylus formosanus.

63. MORACEAE

Fossil Ficus-type pollen was found in the middle Eocene in America and in the Miocene in Indonisia and Spain (Muller, 1981). In China, a few specimens determined to Engelhardtioidites levis (pl. 188, figs. 2-3), found in the Miocene Wanlugang and Zhujiang Formations in the continental shelf of the South China Sea, may be comparable with extant Ficus altissima in pollen morphology. Moraceoipollenites hubeiensis (pl. 194, figs. 14-15) occurs in the Eocene Qianjiang Formation and resembles extant Morus in pollen morphology, but this cannot invalidate its relationship to Urticaceae.

64. MYOPORACEAE

Myoporumpollenites striatus (pl. 125, figs. 16-18) and M. variabilis (pl. 125, figs. 13-15) fossils are 3-colpate and 6-porate and have two pores per colpus. Recorded in the Upper Cretaceous in the Jianghan Basin, they match extant Myoporum.

65. MYRICACEAE

Triatriopollenites may bear some relation to this family. It is widely distributed in the Upper Cretaceous and early Tertiary in the Northern Hemisphere, and chiefly in the Palaeogene in China. Myricaceoipollenites and Myricipites (pl. 194) may also be related to this family, which is distributed mainly in the Paleocene in the Lingfeng and Luhutai Formations but also in the Upper Cretaceous in the Furao (Liu, 1983) and Kukaban (Wang et al., 1990) Formations in China.

66. MYRSINACEAE

Fossil Tricolporopollenites maesaformis (pl. 101, figs. 9-11), more or less comparable with extant Maesa, was found in the Paleocene Lingfeng Formation.

67. MYRTACEAE

Pollen grains in this family are mainly syncolporate. Early fossils, called Myrtaceidites lisamae (= Syncolporites lisamae), occur in the Santonian in Gabon. Later it was commonly discovered in the Maestrichtian and the Tertiary throughout the world (Muller, 1981). In China, Myrtaceidites granulatus (pl. 139, figs. 4-7) fossils occur in the upper Wuyitake Formation (Turonian-early Senonian) (Wang et al., 1990) in the Xinjiang area. Similar fossils often occur in the Upper Cretaceous in the Songliao Basin, in northern Jiangsu and Jiangxi Provinces, as well as in the Tertiary in China. Fossil M. cajupitiformis (pl. 127, fig. 5), found in the Taizhou Formation (Senonian), resembles extant Eucalyptus in pollen morphology. In addition, Syncolporites fossils, similar to this family in pollen morphology, are distributed in the Upper Cretaceous and Tertiary in China. Other fossils related to the extant Decaspernum and Psidium types are discussed in the section on Sapindaceae.

68. NYMPHAEACEAE

Fossil pollen in this family, comparable to Nymphaea zanzibarensis, was found in the Maestrichtian in Canada; comparable to Nymphaea alba, in the Oligocene in Romania; comparable to Nuphar, in the Pliocene in Turkey; and comparable to Nelumbo, in the Oligocene in Siberia (Muller, 1981). In China, zonicolpate Nymphaeacidites echinatus (pl. 84, figs. 21-22), found in the Latest Cretaceous Mengyejing Formation in Yunnan Province, may be compared with extant Nymphaea in pollen morphology. Tricolpate Retitrescolpites pellucidus (pl. 89, figs. 28-29), comparable to extant Nelumbo in pollen morphology, occurs in the Paleocene-Eocene Funing and Kongdian Formations. Another species, R. nelumboides (pl. 89, figs. 22-23), was found in the Oligocene Dongying Formation in the coastal region of the Bohai Sea.

69. NYSSACEAE

Fossil Nyssa-type pollen, such as Tricolporites kruschi, occurs early in the Paleocene in Spain, France, and America; Nyssapollenites and Nyssoidites, in the Maestrichtian in Canada; N. squamosus and N. lanosus, in the Cenomanian-Turonian in Australia (Muller, 1981). About 17 species of Nyssapollenites have been described in China. The oldest of them, N. analapticus (pl. 129, fig. 26), was found in the Kukebai Formation (Cenomanian-Turonian) in the southern Xinjiang area (Wang et al., 1990). Others, including N. albertensis (pl. 144, figs. 7-8), N. cretaceous, N. margocolpites, and N. pseudolaesus, also occur in the Upper Cretaceous. In addition, fossil N. sinensiformis (pl. 138, figs. 1, 6-8, 18), resembling that of extant Nyssa sinensis in pollen morphology, was discovered in the Paleocene-Eocene in eastern China. This plant may have been growing in this area since the early Tertiary.

70. OLACACEAE

About six Anacolosidites species, including A. luteoides (pl. 197, fig. 1), A. primigenius (pl. 197, figs. 3-4), A. pseudoefflatus (pl. 197, fig. 2), A. substrudens (pl. 197, figs. 9-11), A. supplingensis (pl. 197, figs. 12-15), and A. tenuiplicatus (pl. 197, figs. 37, 39), have been found in the Lingfeng Formation (Paleocene) in eastern China, and they are also distributed in the Eocene in the same areas. Similar fossils occur in the Late Cretaceous (Campanian-Maestrichtian) to Eocene outside China (Muller, 1981).

71. OLEACEAE

Two fossil types, Fraxinoipollenites and Oleoidearumpollenites, may be related to this family. The former, characterized by its oblong body, tricolpate aperture, and reticulate ornamentation, is comparable to extant Jasminum, the latter, with a spheroid body, tricolporate or tricolporoidate aperture, and reticulate ornamentation, is related to extant Osmunthus and others in this family. About 18 Fraxinoipollenites species collected in China, such as F. angustus (pl. 118, figs. 13), A. fusiformis (pl. 123, figs. 3-5), F. granulatus (pl. 116, figs. 11-12), and F. variabilis (pl. 144, figs. 33-34), occur in the Upper Cretaceous. They are also distributed in the Tertiary. Fossils of Oleoidearumpollenites species are commonly distributed in the Upper Tertiary in China, but O. chinensis (pl. 131, figs. 7-9), resembling extant Ligustrum delavayanum in pollen morphology, was found in the Eocene Lower Ganchaigou Formation. Oleoidearumpollenites ligustiformis (pl. 131, figs. 12-13) and O. forsythiaformis (pl. 131, figs. 10-11), related to extant Ligustrum sinensis and Forsythia suspensa, respectively, were found in the Oligocene Upper Ganchaigou Formation.

72. ONAGRACEAE

Early fossil pollen in this family includes Jussitriporites and Trivestibulopollenites, from the Maestrichtian in America and Brazil (Muller, 1981); Onagracites anularis, from the Paleocene in the Ural area in Russia (Pokrovskaja & Stelmak, 1960: 390, pl. 3, fig. 30); Oculis nectis, from the Eocene in Europe (Krutzsch, 1957: tab. 95); and Jussiena or Onagraceae, from the Oligocene in Kazakhstan (Zaklinskaja, 1963: pls. 12-13). In China, fossil Ludwigia trilobapollenites pollen, first collected in the Paleocene Lizigou Formation, increased in frequency to 1% in the middle Eocene Guchengzi Formation in the Fushun Basin (Sun et al., 1980: 58, 63). Corsinipollenites spp. are sparsely distributed in Tertiary, but in the upper part of the Huxian Group (Oligocene), where they are called Jussitriporites champlainensis (Song, 2000), their numbers constitute 5-10% of the total assemblage. In addition, fossil Pseudotrudopollis basoides and Parapecakipollis tuberculatus (Song, 1996: pl. 11, figs. 9-10; pl. 16, figs. 1-2), from the Paleocene Naomugen Formation in the Inner Mongolia area, may be comparable to extant Chamaenerion angustifolium and Circara quadrisulcata in pollen morphology, but the fossil pollen is smaller than the recent pollen.

73. PALMAE

Fossil Dicolpopollis koekelii (pl. 84, figs. 10-12), comparable to extant Calamus in pollen morphology, occurs in the Lingfeng Formation (Paleocene) and is distributed in the Oligocene, especially in the upper Tertiary in the continental shelf of the South China Sea. Sabelpollenites areolatus (pl. 84, figs. 7-9), which may be related to extant Phoenix, occurs in the Mengyejing, Kongdian, and Yangxi Formations, from the latest Cretaceous to the Eocene. Fossil Spinizonocolpites cf. baculatus (pl. 84, figs. 15-16) and S. granulatus (pl. 84, figs. 13-14), bearing a relation to extant Nipa, were reported in the Paleocene Lingfeng Formation.

74. PANDANACEAE

Fossil Pandaniidites shiabensis (pl. 183, figs. 1-3) and P. verruspinus (pl. 183, figs. 6-7) were discovered in the Miocene Guantou Formation in the Bohai region of China. Similar fossils, called Spinamonoporites typicus (Muller, 1981), were found in the latest Cretaceous to late Tertiary outside China.

75. PELLICIERACEAE

Fossil pollen similar to Pelliciera occurs in the Eocene in South America (called Verrutricolporites crassus), the middle Eocene in Brazil (called Psilatricolporites crassus), and the Olig-Miocene in Mexico (Pelliciera type) (Muller, 1981). In China, only Pellicieroipollis langeheimii (pl. 141, fig. 32) occurs in the Pliocene Wanglugang Formation in the continental shelf of the South China Sea.

76. PIPERACEAE

Pollen grains in this family are monosulcites, with small bodies (< 20 [micro]m) similar in pollen morphology to those of the Saururaceae. They are always neglected in Tertiary palynological studies of southern China because of their small size and because they have no geological significance.

77. PLATANACEAE

Fossil pollen in this family, such as Plantanus occidentaloides, occurs in the upper Eocene-Oligocene and the Miocene in southeastern America; Platanoidites gertrudae, in the lower Miocene in Europe and other areas (Muller, 1981). Similar fossils of Platanoidites gertrudae (pl. 94, figs. 17-19) were discovered in the Miocene Minghuazhen Formation in northern China; of P. palaeocenicus (pl. 94, figs. 8-10), in the Paleocene Lingfeng Formation in eastern China. The latter lack colpate membranes and have a clear colpus, unlike recent Platanus pollen. This is possibly a fossil character.

78. POLYGALACEAE

Recent pollen in this family is polycolporate and synorate. Early records of the fossil pollen are Psilastephanocolporites fissilis, from the Paleocene in Chile and the lower Eocene in Guyana, and Polygalacidites clacus, from the Eocene in India; cf. Securidaca, from the middle Eocene in America and Bredomeyera, from the late Miocene in Mexico (Muller, 1981). In China, fossil Tetracolporopollenites zonoratus (pl. 103, fig. 25), found in the Eocene Changhe Formation in northern Zhejiang Province, may be related to this family. Unfortunately, it is only 4-colporate, but Polygalacidites taiwanensis Huang (1980: pl. 11, figs. 6-7), from the Miocene in Taiwan, is more than 5-colpate and synorate.

79. POLYGONACEAE

Persicarioipollis Krutzsch 1962 resembles extant Persicaria and Polygonum in pollen morphology. Persicarioipollis communis (pl. 201, figs. 9-10) was found early in the Eocene in the Shahejie, Oujiang, Pinghu, Honggou, Ganchaigou and Liushagang Formations. They are sparsely distributed in the Palaeogene but become more developed in the Neogene. For example, they occupy about 24.4% in the early-middle Miocene Heilongjing Formation, about 1.8-30.6% in the late Miocene Yuquan Formation, and about 3.3-57.1% in the Pliocene lower part of the Santan Formation in the continental shelf of the East China Sea. They also attain high percentages in the Pliocene Upper Yancheng Formation and the Pliocene upper part of the Minghuazhen Formation: 30% and 20.8-71.3%, respectively (Song, 2000). Fossil pollen in this family outside China are commonly distributed in Neogene but also have old records from the middle Paleocene in central Europe and the upper Paleocene in France (Muller, 1981).

80. PORTULACEAE

Fossil pollen like extant Portulaca were found in the latest Miocene in Spain. Rugaepollis fragilis occurs in the Mio-Pliocene in Alaska. Rugaepollis kachemakensis (pl. 100, fig. 20) was discovered in the Miocene Youshashan Formation in the Qaidam Basin in China. It is mainly rugae-colpate, possibly with 30 rugae, and the rugae are simple and very short (5-6 [micro]m long). Portulacapollenites taiwanensis Li et Huang (1990: pl. 20, fig. 127), from the Pliocene in the Taiwan area, is similar to extant Portulaca pilosa.

81. POTAMOGETONACEAE

Reliable records of this pollen, which is inaperturate and microreticulate, come from the late Miocene in Europe, although records from the Oligocene or even the Paleocene have also been proposed (Muller, 1981). In China, fossil Potamogetonacidites natanoides (pl. 204, figs. 16-18) was found in the Paleocene--early Eocene Lulehe Formation; it is commonly distributed in the Tertiary and increases in occurrence in the upper Tertiary.

82. PROTEACEAE

Fossil pollen in this family occurs commonly in the Maestrichtian or Paleocene in the Northern Hemisphere and developed in the Eocene in Australia and New Zealand. In China, the records are as follows:

Beaupreadites, possibly related to extant Beauprea, is a tricolpate pollen without pores or an atria structure. Species such as B. libitus (pl. 154, figs. 18-19) and B. radiatus (pl. 154, figs. 22-24) are commonly recorded from the Late Cretaceous, but the earliest record is from the Cenomanian in the Songliao Basin (Gao et al., 1999).

Borealipollis, perhaps beating some relation to this family, was differentiated from Proteacidites by its short colpus. It is a tribrevicolporate type with a trilete scar structure on one surface of the pollen. These fossils were discovered only in the middle-late Late Cretaceous and include B. bratzevae (pl. 153, figs. 5, 9, 19), B. songliaoensis (pl. 153, figs. 6-8), and B. yaojianica (pl. 153, figs. 1-4).

Proteacidites was discovered mainly in the latest Cretaceous and earliest Tertiary in China. Song (1996) suggested that Chinese species of this genus may be related to the Normapolles group rather than to this family and proposed another genus, Pseudoproteacidites, to include them.

83. RANUNCULACEAE

Fossil pollen similar to Ranunculus was found in the upper Miocene in France; and fossil pollen similar to Thalictrum, designated Punctioratipollis ludwigi (Muller, 1981), was found in the lower Miocene and the Pliocene in central Europe. In China, tricolpate fossil pollen in this family, called Ranunculacidites Sah 1967, was found in the Eocene to the Pliocene and is more often distributed in the upper Tertiary. The relationship of the fossil species to extant plants is: R. verus (pl. 89, figs. 26-27), found in the Oligocene Dongying Formation, may be related to extant Pulsatilla chinensis; R. minor (pl. 89, figs. 1-3), from the Eocene Kongdian Formation, to Paraquilagia anamonoides, R. vulgaris (pl. 89, figs. 14-16), also from the Kongdian Formation, to Aquilaegia and Clematis; R. pachydermus (pl. 89, figs. 9-10), from the Miocene Minghuazhen Formation, to Adonis amurensis; and R. heilongjingensis (pl. 89, figs. 6-8), from the Miocene Heilongjing Formation, to Kingdonia uniflora. In addition, some fossil Retimultiporopollenites minor (pl. 203, figs. 7, 9) specimens, found in the upper Tertiary, may be compared with the extant Thalictrum type.

84. RESTIONACEAE

Fossil Aglaoreidia cyclops (pl. 205, fig. 19) and Restioniidites menglaensis (pl. 183, figs. 26-27) were found in the latest Cretaceous Mengyejing Formation. Other species, including R. geiseltalensis (pl. 183, fig. 30) and R. minor (pl. 183, figs. 4-5), were found in the Paleocene-Eocene Nongshan and Buxin Formations. Fossil pollen in this family, such as Restio type, early occurs in the Maestrichtian in Senegal, the Paleocene in Argentina, and the Oligocene and Miocene in Australia and New Zealand (Muller, 1981).

85. RHAMNACEAE

Fossil Rhamnus brandonensis pollen was discovered in the Oligocene in America; that of the Ziziphus type, in the upper Miocene in Spain and the Pliocene in northern Africa; and that of the Frangula type, in the Pliocene in Germany (Muller, 1981). Fossil pollen in this family is called Rhamnacidites Chitaley 1951 ex R. Potonie 1960 in China. It was discovered mainly in the latest Cretaceous to the Eocene, but a few examples were found in the Miocene. Fossil species compare with with extant plants thus: Rhamnacidites donghaiensis (pl. 140, figs. 1-2) and R. minor (pl. 127, figs. 7-9) are related to extant Sageretia thea," R. nanhaiensis (pl. 142, fig. 1), to Paliurus hemsloyanus," R. reticulatus (pl. 127, fig. 38), to Rhamnus esquirolii," and R. triangulus (pl. 127, figs. 10-12), to Rhamnus crenata.

86. RHIZOPHORACEAE

Fossil Zonocostatites ramonae in this family occurs in the upper Eocene in the Caribbean area and Brazil, the Eocene in India, and the Oligocene in Australia (Muller, 1981). Zonocostatites cf. ramonae (pl. 130, figs. l-4) was found in the Oligocene-Pliocene Weizhou and Yuehai Formations in the continental shelf of the South China Sea.

87. RHOIPTELEACEAE

Fossil pollen in this family is called Rhoipteapollis Zheng 1985 in China. Two species, R. chilianthoides (pl. 140, fig. 10) and R. triletoides (pl. 140, fig. 9), were discovered in the Pliocene Santan Formation in the continental shelf of the East China Sea.

88. ROSACEAE

Many fossil pollen types related to this family have been discovered in China. Among them is fossil Dendriopoteriumpollis scabratus (pl. 95, figs. 29-30), which bears some relation to extant Dendriopoterium and was found in the Eocene-Oligocene Ganchaigou Formation in the Qaidam Basin. The extent plants in this genus grow in southern Africa.

Grielumpollis microreticulatus (pl. 203, figs. 23-24), which bears an affinity with extant Grielum and Neurada, occurs in the Miocene Lower Youshashan Formation in the Qaidam Basin. Recent plants also grow in Africa. A similar fossil, Syndemicolpites typicus, which was found in the Maestrichtian in Nigeria, was rejected by Muller (1981: 54). Tricolporopollenites chamaerhodesoides (pl. 106, figs. 6-7), T. geumiformis (pl. 105, figs. 14-16), and T. prunusiformis (pl. 106, figs. 12-13), resembling extant Chamaerhode, Geum striatum, and Prunus, respectively, were discovered in the Eocene-Oligocene Ganchaigou Formation. Another form, T. rosaeformis (pl. 107, figs. 20-22), bearing an affinity with extant Rosa, was found in the Miocene Heilongjing Formation. In addition, Rubuspollenites formosensis Huang (1980: pl. 11, figs. 8-9), from the Miocene in the Taiwan area, resembles extant Rubus rosaefolius. Outside China, the fossils in this family occur only from the Oligocene to the Miocene (Muller, 1981).

89. RUBIACEAE

Fossil pollen Triporotetradites nachterstedtensis was found in the upper Eocene in Germany, and T. letouzeyi, from the lower Miocene in Cameroon, may be related to extant Gardenia, which is mainly distributed in the upper Tertiary. Other fossils related to this family occur in the Oligocene outside China (Muller, 1981).

The following fossils were recorded in China: Gardeniapites taiwanensis Huang (1980: pl. 17, figs. 7-8), resembling extant Gardenia, occurs in the Miocene in the Taiwan area. It is tetrad and triporate, with rounded pores and pore margins. Randiapollis Ke et Shi 1978, related to extant Randia, is oblate-spheroid in form, 3-5 porate, and has rounded pores and reticulate ornamentation. Fossil R. reticulatus (pl. 203, figs. 17-20) and R. microreticulatus (pl. 203, figs. 4-5) were discovered in the Eocene Kongdian Formation. These fossils also resemble extant Saprosma ternatum. Their pollen grains have 3-4 pores with rugu-reticulate ornamentation, similar to R. microreticulatus in ornamentation. In addition, fossil Retitriorites aspidatus (pl. 191, fig. 26) and R. megapores (pl. 191, figs. 24-25), which occur in the Taizhou Formation and are Senonian in age, resemble extant Canthium horridum in pollen morphology.

90. RUTACEAE

Reliable fossil pollen in this family, such as Phellodendron, were found in the Pliocene in Western Europe, whereas Tricolporopollenites cingulum fusus, comparable to extant Ptelea, Zanthoxylon, or Hoplophyllum, was discovered in the Eocene-Oligocene in Europe (Muller, 1981). However, without further study of tri-tetracolporate pollen in the family, their relationship is very difficult to determine.

Fossil pollen in this family in China is called Rutaceoipollis Sung et Tsao 1978 ex 1980. It is tricolporate and has transversely elongated pore as rugae that intersect with the colpi in a cross shape. The earliest record of it is R. granulatus (pl. 122, figs. 1-3), which was found in the Taizhou Formation and is Senonian in age, but its main distribution is in the Tertiary. Fossil Rutaceoipollenites He et Sun 1977 differs from the former in its narrow rugae, but its colpi and rugae are also cross shaped. They too were discovered mainly in the Tertiary. Horniella Traverse 1955 emend. Song 1999 differs from the former two genera only in its well-developed reticulate ornamentation, and it may resemble extant Evodia. It is often discovered in the Tertiary, but some species, such as H. baculogranus (pl. 142, figs. 14-16) and H. fusiformis (pl. 122, figs. 4-6), were also found in the Late Cretaceous (Senonian). The fossils are related to extant plants thus: R. cingulatus (pl. 104, figs. 8-9), to extant Clausena excavata; R. granulatus, to Clausena lansium; R. citrusaeformis (pl. 115, figs. 15-16), to Citrus," R. lentiporus (pl. 102, figs. 26-28), to Murraya paniculata; R. oblongatus (pl. 102, figs. 12-13), to Micromelum folcatum; R. minireticulatus (pl. 119, figs. 1-3), to Boenninghousenia albiflora; and Gemmatricolporites acutopollis and G. pedunculata Huang (1980: pl. 12, figs. 4-6, 7-9), to Bryophyllum pinnatum and Acronychia pedunculata, respectively.

91. SALICACEAE

Salixipollenites Srivastava 1966 may chiefly match extant Salix. Early records in China are S. elegans (pl. 91, figs. 6-8), S. trochuensis (pl. 91, figs. 1-4), and S. hians (pl. 91, figs. 1214), which were discovered in the upper Quanton Formation in the Songliao Basin and are Cenonmanian in age (Gao et al., 1999). They are also distributed in the late Upper Cretaceous and Tertiary. As for fossil Populus pollen, although there was a report in the lower Miocene abroad, there is no reliable datum in China.

92. SANTALACEAE

Fossil pollen in this family is called Santalumidites. About eight species in this genus have been discovered in the Upper Cretaceous to the Eocene in China. Late Cretaceous species are S. annulatus (pl. 143, figs. 18-19), S. pseudodolium (pl. 143, fig. 27), and S. wuhuensis (pl. 143, figs. 25-26), possibly Senonian in age. All others are distributed mainly in the Eocene. It must be pointed out that these fossils are all tricolporate, with clear and developed colpi and that they differ from recent Santalum pollen and may be related to Sapotaceae. The Aquilapollenites group may be related to Arjona. The earliest record in China is early Late Cretaceous, similar to the date in the other parts of the world.

93. SAPINDACEAE

Pollen grains in this family are of three types: triporate, tricolporate, and syntricolporate. Triporate-type fossils related to extant Pometia were found in the upper Miocene in Borneo (Muller, 1981) and in the Pliocene in Yunnan Province in China (Song, 1988). In addition, Proteacidites microverrucatus (pl. 181, figs. 4-8), which more or less resemble extant Pometia in pollen morphology, were found in the Paleocene to the Eocene in China.

Tricolporate-type fossils like extant Dodonaea viscosa were found in the Pliocene and the Pleistocene in China (Song, 1988: pl. 37, figs. 15-16, 20-21) and in the Pliocene in New Zealand. Sapindaceidites Wang ex Sun et Zhang 1979, from the Upper Cretaceous and the Paleogene in China, has a triangular amb and is tricolporate with long but not syncolpate colpi. Fossils in this genus may have different relationships: For example, S. concavus (pl. 139, figs. 17-20), S. lobatus (pl. 159, figs. 16-19), S. triangulus (pl. 139, figs. 14-16), and S. vestibulum (pl. 158, figs. 4-6) may be related to extant Sapindus; whereas S. asper (pl. 139, figs. 21-24), S. hunanensis (pl. 118, figs. 7-8), S. liaoningensis (pl. 139, figs. 30-32), S. microrugosus (pl. 144, figs. 15-17), and S. xiningensis (pl. 158, figs. 16-17) are comparable to extant Decaspermum. The others, including S. cangxianensis (pl. 139, figs. 27-29) and S. tetrorisus (pl. 139, figs. 28-29, 33), which are tetracolporate, are comparable to extant Decaspermum combodianum and were recorded in the late Senonian. Fossil Senipites striatus (pl. 152, figs. 9-12, 19), found in the Taizhou Formation, resembles extant Koelreuteria bipinnata and K. paniculata in pollen morphology. Talisiipites Wodehouse 1933 differs from Sapindaceidites only in its rounded arab. The fossils included in this genus show various relationships; for example, T. longicolpus (pl. 142, figs. 2-3), S. nanhaiensis (pl. 127, fig. 25), and T. subtriangulus (pl. 127, figs. 33-35) with extant Sapindus; T. megorites (pl. 149, figs. 22-23, 27) to Koelreuteria bipinnata; T. minor (pl. 135, figs. 24-25) and T. promptus (pl. 129, figs. 23-24) to Delevaya yunnanensis; and T. qaidamensis (pl. 138, figs. 19, 22) to Dodonaea. Except for T. nanhaiensis and T. subtriangulus, which were found in the Taizhou Formation, all are distributed mainly in the Tertiary. Fossil Siberiapollis Tschudy 1971 is also related to different extant plants in pollen morphology: for example, S. angulatus (pl. 128, figs. 27, 30), S. bellus (pl. 154, figs. 10, 16-17), S. bullatus (pl. 180, figs. 10-12), and S. crassiporus (pl. 150, fig. 27) to Decaspermum; and S. formosus (pl. 181, figs. 11-12), S. mollis (pl. 180, figs. 15-17), and S. tumidiporus (pl. 128, fig. 28), to Pometia. However, the recent pollen is triporate, whereas all of the fossil pollen is tricolporate. All of the fossils occur in the Upper Cretaceous.

Syntricolporate-type fossil Sapindaceaepites Biswas 1962 emend. Song 1999 belongs to this type. All of these fossils were discovered in the Upper Cretaceous. Among them, S. laevigatus (pl. 115, fig. 11) and S. punctatus (pl. 144, figs. 29-30) may be compared with extant Amesiodendron and Paranephelium, and S. microstriatus (pl. 155, fig. 15) with Cardiospermum in pollen morphology. Parasyncolporate-type Cupanieidites reticulatus fossils were found in the Upper Cretaceous outside China. Cupanieidites reticulatus (pl. 157, figs. 1-3, 12), C. major (pl. 118, fig. 6), C. projectus (pl. 159, figs. 9-12), S. triangulus (pl. 143, figs. 8-10), and others were found in the Upper Cretaceous and possibly also in the Tertiary in China.

94. SAPOTACEAE

About 10 Sapotaceoidaepollenites species have been collected in China. The earliest species, including S. occultus (pl. 121, figs. 6-7) and S. manifestus (pl. 121, figs. 4-5; pl. 130, figs. 10-11), were found in the Taizhou Formation (Senonian), whereas the others were distributed in the Tertiary. The comparison of fossils to extant plants in pollen morphology is: S. manifestus, S. neyvelii (pl. 103, fig. 7; pl. 106, fig. 2), and S. sapotoides (pl. 103, figs. 14-15) to Chrysophyllum and Marstichodendron; S. kirchheimeri (pl. 130, figs. 15, 20), to Manikara; S. granulatus (pl. 114, figs. 19-20), to Pouteria; S. megaporus (pl. 114, fig. 14); and S. obscurus (pl. 106, fig. 31), to Xantolis. The oldest fossil records outside China are from the Senonian and Paleocene in Borneo, the lower Eocene in Australia (called Sapotaceoidaepollenites robustus), the upper Eocene in Nigeria (Psilastephanocolporites malacanthoides), the Paleocene and the Eocene in Europe (Tetracolporopollenites), the lower Eocene in America, and the Tertiary in other areas (Muller, 1981).

95. SAXIFRAGACEAE

Pollen Tricolporopollenites asper Huang (1980: pl. 18, figs. 11-12) grains from the Upper Tertiary in the Taiwan area are reticulate tricolporate and small. Although Huang included them in this family, their relationships need further study to confirm the placement.

96. SCROPHULARIACEAE

Fossil Tricolpites pedicularidus (pl. 97, figs. 12-14), comparable to extant Pedicularis, is distinguished by its long colpi, which almost forms a syncolpate type; T. tenuicolpus (pl. 97, figs. 7-9) and T. trapeziformis (pl. 98, figs. 4-6) fossils can also be compared with this extant plant. They occur in the Eocene to the Pliocene in eastern China.

97. SIMAROUBACEAE

Fossil Retitricolporites maximus (pl. 121, figs. 21-22) may be compared with extant Harrisonia perfosata in pollen morphology. Recent pollen is retitricolporate, 51.3 x 38 [micro]m in size, and 2.5-3.0 [micro]m in exine thickness. Fossil pollen was discovered in the Taizhou Formation and is Senonian in age. Wilsonipites micireticus (pl. 121, figs. 8-9), also found in the Taizhou Formation, resembles extant Ailanthus in pollen morphology. Paranyssapollenites changheensis (pl. 132, fig. 22), which resembles extant Suriana maritina in pollen morphology, is tricolporate with long colpi and square-shaped pores, 30 x 40 [micro]m in size, and 1.5-2.0 [micro]m in exine thickness, with microreticulate ornamentation. The fossil was found in the Eocene Changhe Formation.

98. SOLANACEAE

The pollen morphology of Tricolporopollenites solanumiformis (pl. 106, fig. 17) is similar to that of some plants in this family. It was discovered in the Pliocene Shizigou Formation in the Qaidam Basin.

99. SONNERATIACEAE

Pollen fossils in this family commonly occur in the Miocene, but Florschetzia trilobata was discovered in the upper Eocene, was developed in the Oligocene, and was extinct in the Miocene; F. semilobata occurs in the lower Miocene; and F. levepoli appeared in the lower Miocene and developed since the middle Miocene outside China (Muller, 1981). In China, F. trilobata (pl. 188, figs. 27-31) was discovered in the Eocene Liushagang and Weizhou Formations, and another two species were found, mainly in the Miocene in the continental shelf of the South China Sea.

100. SPARGANIACEAE

Pollen fossils in this family occur early in the Paleocene in Europe and are widely distributed in the Tertiary (Muller, 1981). Sparganiaceaepollenites neogenicus (pl. 183, figs. 13-16) and S. sparganioides (pl. 183, figs. 17-21) were found in the Paleo-Eocene Lulehe Formation in the China's Qaidam Basin and were distributed throughout China in the Tertiary.

101. STERCULIACEAE

The following pollen fossils may be related to this family in China. Retitricolporites firmianoides (pl. 148, figs. 9-10), which resembles extant Firmiana simplex in pollen morphology, was found in the Eocene Shahejie Formation. Reevesiapollis siameniformis (pl. 120, figs. 2-4), related to extant Reevesia, was found occasionally in the Shahejie Formation (Eocene) and with increased values in the Miocene. Similar fossils occur in the Paleocene and the Eocene in Europe (Muller, 1981). Recent Craigia yunnanensis pollen looks like Tilia of the Tiliaceae in terms of pollen morphology. The former differs from the latter by its smaller size, thinner exine, unclear pore chamber, and annulus, as well as its micireticulate ornamentation. All of these characters are recognized in the fossils Tiliaepollenites lingfengensis (pl. 145, figs. 1-5), T. microreticulatus (pl. 145, figs. 11-13), and T. retigranulatus (pl. 145, figs. 10, 14). These fossils may be related to extant Craigia rather than to Tilia. The early record is from the Paleocene Lingfeng Formation.

102. SYMPLOCACEAE

Early fossil pollen in this family occurs in the Maestrichtian in America and is widely distributed in the Tertiary in many areas in the world after the Eocene (Muller, 1981). Pollen fossils in this family are called Symplocoipollenites and Symplocospollenites in China. Early records, such as S. crassus (pl. 130, figs. 13-15) and S. songliaoensis (pl. 128, figs. 1-4), were found in the early Senonian Nenjiang Formation in the Songliao Basin; S. baculatus (pl. 151, figs. 37-38) and S. oblatus (pl. 152, fig. 5), in the Senonian Taizhou Formation. The others are widely distributed in the Tertiary. Some Late Cretaceous fossils may be also compared with extant plants in this family in pollen morphology: Borealipollis songliaoensis (pl. 153, figs. 68), with Symplocos paniculata; Siberiapollis crassiporus (pl. 150, fig. 27), with Symplocos chinensis; S. formosus (pl. 181, figs. 11-12), with Symplocos crassifolia; S. mollis (pl. 180, figs. 15-17), with Symplocos confusa; and Proteacidites marginus (pl. 180, figs. 1-2, 13), with Symplocos angustifolia. These fossils are about 40 [micro]m in size, just equivalent to recent Symplocos pollen.

103. THEACEAE

Recent pollen grains in this family are characterized by oblate to spheroid bodies, tricolporate apertures with broad colpi and large pores, thick exines, and coarse reticulate ornamentation. The colpi membranes are preserved on recent pollen but are lacking in fossils because of the fossilization. Retitricolporites magnus (60 x 55 [micro]m) (pl. 136, fig. 26; pl. 137, fig. 30) has similar characters and may be compared with extant Polyspora axillaris. Fossils were found in the Eocene Shahejie Formation. Retitricolporites maximus (pl. 121, figs. 21-22) and R. megaporus (pl. 121, figs. 10-11) may also be related to this family, and they all occur in the Senonian Taizhou and Dalongshan Formations. Outside China, Tricolporopollenites srivastavai, found in the lower Eocene in France, may be related to extant Gordonia axillaris (Muller, 1981).

104. THYMELAEACEAE

Fossil Retimultiporopollenites qiongbeiensis (pl. 203, fig. 21) may be compared with extant Wikstroemia indica, but its exine is thinner than that of recent pollen. Fossil pollen was discovered in the Eocene Liushagang Formation.

105. TILIACEAE

About 13 Tiliaepollenites species have been discovered in China. They may be divided into two groups. One group is small (<30 [micro]m), with undeveloped pore chambers, thinner exine, and ornamentation. It includes T. indubitabilis, T. insculptus, T. lingfengensis, T. minimus, and T. microreticulatus and may be related to extant Craigia yunnanensis of the Sterculiaceae. The other group is large (>40 [micro]m), with developed pore chambers, thick exine, and coarse ornamentation. It includes T. cordataeformis (pl. 146, figs. 9-10), T. hunchunensis (pl. 148, figs. 11-12), T. instructus (pl. 146, figs. 13-15), T. polonicus (pl. 146, figs. 17-18), and T. pseudinstructus (pl. 146, figs. 11-12, 16) and may be related to Tilia. The early records are from the Paleocene Lingfeng Formation. In addition, fossil Euphorbiacidites microreticulatus (pl. 118, figs. 18-19), roughly comparable to extant Grewia and Triumfetta, was found in the Paleocene Wuyun Formation. Retitricolporites shaoboensis (pl. 121, figs. 20, 23), found in the Taizhou Formation (Senonian), is also related to the extant plants named above, although its pores are larger and clearer. Grewipollenites Srivastava 1969 is tricolpate and has no relation to extant Grewia, because the latter is tricolporate.

106. TROCHODENDRACEAE

Fossil Eupteleapites eupteleaformis and E. elegans (Zheng et al., 1994: pl. 2, figs. 7-9, 22, 26), related to extant Euptelea, were found in the Miocene Fotan Group in Fujian Province. Operculumpollis operculatus (pl. 94, figs. 20-22), which occurs in the Eocene Liushagang Formation, resembles extant Trochodendron aralioites in pollen morphology.

107. TYPHACEAE

There are two pollen types in this family, tetrad and monads. The two types may be produced in the same species. Tetrad fossils are called Tetradomonoporites Chitaley 1951 ex Krutzsch 1970, and the early record is T. porolatus (pl. 183, fig. 33), found in the Mengyejing Formation (Senonian to Paleocene in age) and commonly distributed in the Tertiary, mainly the upper Tertiary. Monad fossils are always included in Sparganiaceaepollenites. They are similar to Typha in pollen morphology and are also distributed in the Tertiary, mainly the upper Tertiary.

108. ULMACEAE

Pollen grains in this family may be roughly divided into two types. One, which is stephanoporate with rugulate ornamentation, includes the genera Ulmus, Planera, Zelkova, Ampelocera, and Hemiptelea. The other, which is triporate with granulate or smooth ornamentation, includes such as genera as Celtis, Aphanenthe, Gironniera, Pteroceltis, and Trema. The latter type is also found in the Urticaceae and the Moraceae. In China, Ulmus-type fossils, such as Ulmipollenites minor (pl. 196, figs. 13-14, 16) and Ulmoideipites krempii (pl. 197, figs. 27-29), occur early in the Upper Cretaceous, whereas Zelkovaepollenites, including Z. potonie (pl. 197, figs. 34-36) and Z. thiergarti (pl. 197, figs. 31-33), were discovered early in the Eocene Ganchaigou and Anjihaihe Formations in northwestern China and are mainly distributed in the upper Tertiary. Fossil Celtispollenites, such as C. shanghuensis (pl. 198, fig. 37), occurs early in the Paleocene Shanghu Formation but is widely distributed in the Eocene and the rest of the Tertiary.

109. UMBELLIFERAE

The following fossils may be compared with some extant plants in this family: Hydrocostaepites pachydermus (pl. 115, figs. 21-24) and H. reticulatus (pl. 115, figs. 19-20), from the Eo-Oligocene Weizhou Formation, to Hydrocostyle; Umbelliferaepites bupleurumiformis (pl. 107, figs. 10-11), from the Pliocene Santan Formation, to Bupleurum; Nanlingpollis aspidoporatus (pl. 108, figs. 12-15), from the late Senonian-Paleocene Taizhou, Lingfeng, Kongdian, and Xialuisi Formations, to Pleurospermum austricum; and Retitricolporites oblongatus (pl. 104, figs. 26-27, 29, 32), from the Paleocene and Eocene Buxin and Xinggouzui Formations, to Angelica, Carum, and Deucus.

110. URTICACEAE

Pollen grains in this family are 2-3-porate types, and fossils in the family may be included in Celtispollenites. An example is Celtispollenites triporatus (pl. 198, figs. 9-12), which was discovered in the Paleocene Lingfeng Formation. Because of its smaller size and thinner exine, it may be related to this family.

111. VERBENACEAE

Pollen grains in this family are tricolpate or tricolporate (or, occasionally, 4-5-colporate) types with microreticulate, verrucate, and spinulate ornamentation. The following fossils may be related to extant plants in this family: Rutaceoipollis zoniporus (pl. 115, figs. 10-12), found in the Miocene Heilongjing Formation, resemble extant Phyla nodiflora in their cross-shaped aperture; Callicarpapollis firmexinus (pl. 86, figs. 1-3), found in the Miocene, may be comparable to extant Callicarpa; Echitricolpites loniceraformis (pl. 116, figs. 30-31) and E. microechinatus (pl. 86, fig. 26), also found in the Miocene, to extant Clerodendron. Scabiosapollis jilingensis (pl. 92, figs. 12-13), discovered in the Eocene Huichun Formation, is more or less comparable to extant Clerodendron hainanensis.

112. VITACEAE

Vitispollenites scabratus (pl. 106, figs. 3-5), possibly related to extant Vitis, was discovered in the Paleo-Eocene Lulehe Formation in the Qaidam Basin. Fossils related to Vitis and to Parthenocissus occur in the Oligocene outside China (Muller, 1981).

113. ZYGOPHYLLACEAE

Fossil Pokrovskaja Boitzova 1979 emend. Zhu is related to extant Nitraria. Nitraradites Zhu et Xi 1985 and Nitraripollis Xi et Sun 1986 are the synonyms of this genus. The earliest fossils in this genus occur in the upper part of the late Paleocene Qijiachuan Formation in the Xining-Minghe Basin (Wang et al., 1990), and they are widely distributed in the Eocene in China. Here we use the Tertiary palynological data on the Qaidam Basin as an example to show the flourishing of this genus: In the upper Lulehe Formation (Paleocene--early Eocene) this genus occupies about 4% of total assemblage; in the lower part of the Lower Ganchaigou Formation (early Eocene), 5%, in the middle part of this formation, 6%(35%)70%, and in the upper part of this formation, 2.5%(20%)58%; in the Upper Ganchaigou Formation (Oligocene), 0%(5.47%)30%; in the lower part of the Lower Youshashan Formation (Miocene), 1%(2.5%)6% and in the upper part of this formation, 0%(1.17%)1.5%; in the Upper Youshashan Formation (late Miocene), about 1.5%(6.76%)7.5%; and in the Shizigou Formation (Pliocene), about 2.4%(4.81%)8%. Outside China, this fossil pollen was first discovered in the middle Eocene in Kazakhstan (Zaklinskaja, 1963). The flourishing of this fossil in Central Asia may be similar to that in northwestern China (Song, 2000).

IV. Comparisons of the Oldest Pollen Records Related to Extant Angiosperms in and outside China

The comparisons shown in Table I reveal that the earliest records in China of some Early Cretaceous elements, including Clavatipollenites, Asteropollis, and Hammenia of the Chloranthaceae family, are the same as those in the other parts of the world but that the records of other elements, such as Graminidites and Chenopodipollis, may be earlier outside China. These facts may be useful in discussing the origin and early evolution of angiosperms. We hope to discuss some related problems in another publication on early angiosperm pollen in the Early Cretaceous in China.

V. Acknowledgments

This project is supported by the Major Basic Research Projects of Ministry of Science and Technology, China (No. G2000077700), and the National Science Foundation of China (No. 49971076).
Table I

Comparisons of the oldest pollen records related to extant
angiosperm in and outside China.

Plant taxa In China Outside China

 1. Acanthaceae Pliocene Miocene
 2. Aceraceae
 Acertype Senonian Late Cretaceous?
 Dipteronia type Miocene
 3. Aizoaceae Miocene
 4. Alangiaceae Paleo-Eocene Early Eocene
 5. Altingiaceae
 Altingia type Eocene
 Liquidambar type Late Senonian Paleocene
 6. Amaranthaceae
 Vaclavipollis Eocene Maestrichtian
 7. Anacardiaceae
 Rhus type Senonian Paleocene
 8. Apocynaceae
 Alyxia type Paleocene Paleocene
 Ecdysanthera type Pliocene
 Micrechites type Pliocene
 Parsonsidites Late Senonian Oligocene
 Rauwolfia type Late Senonian Eocene
 Trachelospermum type Miocene
 9. Aquifoliaceae
 Ilex type Albian? Cenomanian
 10. Araliaceae Senonian Paleocene
 11. Balsaminaceae Pliocene
 12. Betulaceae
 Alnus type Santonian Santonian
 Betula type Late Senonian Santonian
 Carpinus type Late Senonian Paleocene
 Corylus type Late Senonian Maestrichtian
 Ostrya type Paleo-Eocene Eocene
 13. Bombacaceae Oligocene Maestrichtian
 14. Boraginaceae Miocene
 15. Buxaceae Paleocene Campanian
 16. Caprifoliaceae
 Lonicera type Late Senonian Oligocene
 Diervilla type Paleo-Eocene Oligocene
 Viburnum type Late Senonian Middle Eocene
 17. Caryophyllaceae Eocene Oligocene
 18. Casuarinaceae Paleocene Paleocene
 19. Celastraceae Miocene Oligocene
 20. Centrolepidaceae Paleocene Paleocene
 21. Cercidiphyllaceae Paleo-Eocene Campanian
 22. Chenopodiaceae Cenomanian Maestrichtian
 23. Chloranthaceae
 Clavatipollenites Barremian- Barremian-
 Aptian Aptian
 Asteropollis Aptian Aptian
 Hammania Albian Albian
 Chloranthacearunpollenites Miocene Miocene
 24. Compositae
 Artemisiaepollenites Oligocene Eo-Oligocene
 Cichoreacidites Oligocene Miocene
 Echitricolporites Paleo-Eocene Oligocene
 Tubulifloridites Eocene Oligocene
 25. Convolvulaceae Eocene Eocene
 Erycibe type Oligocene
 Evolvulus Pliocene
 26. Cornaceae Late Senonian Maestrichtian
 27. Cruciferae
 Capsalla type Eocene
 Hesperis type Miocene
 28. Cucurbitaceae
 Thladiantha type Pliocene Oligocene
 29. Cyperaceae Oligocene Eocene
 30. Cyrillaceae Cenomanian Maestrichtian
 31. Daphniphyllaceae Miocene
 32. Dipsacaceae
 Scabiosa type Late Senonian Miocene
 Morina type Cenomanian
 33. Dipterocarpaceae Miocene Oligocene
 34. Droseraceae Miocene Miocene
 35. Elaeagnaceae Late Senonian Eocene
 36. Elaeocarpaceae Miocene Oligocene
 37. Ericaceae Eocene Eocene
 38. Euphorbiaceae Late Senonian Paleocene
 Euphorbia type Eocene
 39. Eupteleaceae Miocene Eocene?
 40. Fagaceae
 Castanea type Cenomanian Sanmtonian
 Fagus type Senonian Oligocene
 Quercus type Late Oligocene
 Cretaceous
 Nothofagus type Late Senonian? Late Cretaceous
 41. Gentianaceae
 Gentiana type Miocene Pliocene
 Nymphoides type Miocene Paleocene?
 42. Geraniaceae Oligocene Miocene
 43. Gramineae Albian-Eocene? Eocene
 44. Haloragaceae
 Haloragis type Eocene Eocene
 Myriophyllum type Miocene Paleocene
 45. Hamamelidaceae Miocene
 46. Hippocrateaceae Pliocene Oligocene
 47. Hydrocaryaceae (Trapaceae) Paleocene- Miocene
 Miocene?
 48. leacinaceae
 Iodes type Paleo-Eocene Paleocene
 Platea type Eocene Paleocene
 49. Juglandaceae
 Annamocorya type Paleocene
 Engelhardtia type Paleocene Maestrichtian
 Carya type Senonian Campanian
 Juglans type Senonian Paleocene
 Platycarya type Maestrichtian Paleocene
 Pterocarya type Senonian Paleocene
 50. Labiatae Eocene
 51. Lauraceae Senonian? Paleocene?
 52. Lecythidaceae Miocene Eocene
 53. Leguminosae
 Caesalpinia type Late Senonian Eocene
 Caragana type Late
 Cretaceous
 Lotus type Late
 54. Liliaccae Cretaceous
 Liliacidites Latest Early- Late Cretaceous
 Late
 Cretaceous
 Lilipollis Miocene Pliocene

 55. Loganiaceae Miocene
 56. Loranthaceae
 Gothanipollis Late Eocene
 Cretaceous,
 Eocene
 Loranthacites Eocene
 Cranwellia-Scollardia Late Late Cretaceous
 Cretaceous
 57. Lythraceae
 Lagerstromia type Miocene Miocene
 Lythraites Late
 Cretaceous
 58. Magnoliaceae
 Liriodendron type Eocene Oligocene
 Magnolia type latest Early Eocene
 Cretaceous
 59. Malvaceae
 Abutilon type Miocene
 Hibiscus type Miocene Eocene
 60. Melastomaceae Miocene ?Oligocene
 61. Meliaceae Late Senonian Paleocene
 62. Menispermaceae Miocene
 63. Moraceae
 Ficus type Miocene? Eocene
 Morus type Eocene
 64. Myoporaceae Late
 Cretaceous
 65. Myricaceae Late Late Cretaceous
 Cretaceous
 66. Myrsinaceae Paleocene Oligocene
 67. Myrtaceae Late Santonian
 Cretaceous
 Eucalyptus type Late Senonian

 Decaspermurn type Late Santonian
 Cretaceous
 Psidium type Late
 Cretaceous
 68. Nymphaeaceae
 Nymphaea type Late Senonian Maestrichtian
 Nelumbo type Eocene Oligocene
 69. Nyssaceae Late Late Cretaceous
 Cretaceous
 Nyssa sinensis type Eocene
 70. Olacaceae Paleocene Campanian
 71. Oleaceae
 Fraxinoipollenites Late Senonian Paleocene
 (Jasminum type)
 Oleo idearumpoll. Eocene Eo-Oligocene
 (Osmanthus type)
 72. Onagraceae Paleocene Maestrichtian
 73. Palmae
 Calamus type Paleocene
 Phoenix type Late Senonian
 Nipa type Paleocene
 74. Pandanaceae Miocene Maestrichtian
 75. Pellicieraceae Pliocene Eocene
 76. Piperaceae Late Tertiary
 77. Platanaceae Paleocene Eocene
 78. Polygalaceae Eocene Paleocene
 79. Polygonaceae
 Persicaria & Polygonum Eocene Paleocene
 (Aguilapollenites)
 type
 80. Portulaceae Miocene Miocene
 81. Potomagetonaceae Paleo-Eocene Eo-Oligocene
 82. Proteaceae Senonian Maestrichtian
 83. Ranunculaceae
 Ranunculus type Eocene Miocene
 Thalictrum type Miocene Miocene
 84. Restionaceae Late Senonian Maestrichtian
 85. Rhamnaceae Late Senonian Oligocene
 86. Rhizophoraceae Oligocene Eocene
 87. Rhoipteleacease Pliocene Campanian?
 88. Rosaceae
 Dendriopoterium type Eo-Oligocene
 Grielum type Miocene Maestrichtian?
 Chamarrhode type Eocene
 Geum type Eocene
 Prunus type Oligocene
 Rosa type Miocene
 Rubus type Miocene
 89. Rubiaceae Eocene Eo-Oligocene
 Gardenia type Miocene Late Tertiary
 Randia type Eocene
 Canthium type Senonian
 90. Rutaceae Late Senonian Eocene
 Clausena type Late Senonian
 Micromelum type Late Senonian
 Murraya type Eocene
 Boenninghousenia type Miocene
 Citrus type Miocene
 Acronychia type Miocene
 Bryophyllum type Miocene
 91. Salicaceae
 Salix type Cenomanian Maestrichtian
 92. Santalaceae
 Santalum type Late Senonian Eocene
 Arjona type Cenomanian- Cenomanian
 Touronian
 93. Sapindaceae
 Dodonaea type Pliocene Pliocene
 Pometia type Late
 Cretaceous
 Sapindus type Senonian
 Koelreuteria type Senonian
 Amesiodendron type Senonian
 Cupania type Senonian Late Cretaceous
 Cardiospermum Senonian
 94. Sapotaceae Senonian Senonian
 95. Saxifragaceae Late Tertiary?
 96. Scrophulariaceae
 Pedicularis type Eocene
 97. Simaroubaceae
 Ailanthus type Senonian Eocene
 Harrisonia type Senonian
 Suriana type Eocene
 98. Solonaceae Pliocene
 99. Sonneratiaceae Eo-Oligocene Eocene
100. Sparganiaceae Eocene Paleocene
101. Sterculiaceae
 Reevesia type Eocene Paleocene
 Firmiana type Eocene
 Craigia type Paleocene
102. Symplocaceae Senonian Maestrichtian
103. Theaceae ?Senonian Eocene
104. Thymelaeceae Eocene Eocene
105. Tiliaceae
 Tilia type Paleocene Paleocene
 Grewia type Senonian Eocene
106. Trochodendraceae
 Euptelea type Miocene
 Trochodendron type Eocene
107. Typhaceae Late Senonian Paleocene
108. Ulmaceae
 Ulmus type Senonian Turonian
 Zalkova type Eocene Miocene?
 Celtis type Paleocene Turonian
109. Umbelliferae
 Hydrocostyle type Oligocene
 Bupleurum type Pliocene
 Pleurospermum type Senonian-
 Paleocene
 Angelica type Paleocene

110. Urticaceae Paleocene Oligocene
111. Verbenaceae Eocene, Miocene
 Miocene?
112. Vitaceae Eocene Oligocene
113. Zygophyllaceae
 Nitraria type Late Paleo- Middle Eocene
 Eocene


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SONG ZHI--CHEN, WANG WEI--MING, AND HUANG FEI

Nanjing Institute of Geology and Palaeontology

Chinese Academy of Sciences

Nanjing 210008, China
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Author:Zhi-Chen, Song; Wei-Ming, Wang; Fei, Huang
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Date:Oct 1, 2004
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