Margaret E. Collinson 1 & Jerry J. Hooker 2 1 Royal Holloway University of London 2 The Natural History Museum, London

Paleogene vegetation of Eurasia: framework for mammalian faunas

Collinson, M.E. & Hooker, J.J. - Paleogene vegetation of Eurasia: framework for mammalian fau- nas - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAM- MALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 41-83 [ISSN 0923- 9308] Published 1 December 2003

Four major categories of Paleogene vegetation are described and their general distribution in Eurasia is plotted on polar projections of paleogeographic maps. Methods and approaches for reconstructing Paleogene and vegetation from fossil floras are briefly outlined. Paleocene and polar, broad-leaved deciduous forests occurred in the high paleolatitudes and up to the edge of the Arctic Ocean. Relative to subsequent paratropical forests these were very low diversi- ty forests with little habitat variation, very few climbers and more open structure. The environment was warm and equable but with up to three months of winter darkness so that most herbivorous browse and the small, dry fruits and would only have been available seasonally. Late Paleocene and Eocene broad-leaved paratropical to subtropical, evergreen forests are variable in time and space. The most evergreen thermophilic forests are documented in and southern China associated with the Eocene thermal maximum. These forests were highly diverse, multi- stratal, rich in climbers, with closed structure and would have provided many varied habitats. They would have provided year round herbivorous browse of medium to large evergreen and a wide variety of fruits, including many fleshy fruits. Late Eocene and broad-leaved mixed deciduous and evergreen forests developed as climate changed following the thermal maxi- mum from the late Middle Eocene and through the Oligocene. These forests had lower diversity, contained fewer lianas and climbers and would have had a more open structure than the paratro- pical forests. They would have provided a mixture of evergreen and deciduous browse and a variety of fruits but with fewer fleshy fruits and more large dry fruits. Vegetation with sclerophyl- lous elements was variably developed in central China, through Kazakhstan and the tethyan islands and into eastern Europe in the Eocene. In the Oligocene it was well-developed in southern Europe and also occurred in Kazakhstan. This vegetation reflects a warm humid temperate clima- te but with a slightly drier interval where sclerophyllous elements become significant in abundan- ce or diversity. This would have been dominantly a forest vegetation (possibly shrubland in places) but with relatively open structure. In the most sclerophyllous belts, such as the Oligocene of sout- hern Europe, herbivorous browse would have consisted mainly of relatively tough small leaves and the fruits and seeds would have been mainly dry with tough outer coverings. This paper paints only a very general picture of Paleogene vegetation. Paleogene floras provide the potential for detailed interpretation of vegetational, and hence biome, response to global change such as warm climate thermal maxima and tectonic events. Realising this potential requires independent dating of more fossil floras, new physiognomic analyses, rigorous re-investigation of nearest living rela- tive assessments and integrated palynological and macrofossil studies which take account of tap- honomic bias.

Keywords: Paleocene, Eocene, Oligocene, vegetation, Eurasia, paleobotany, paleoecology

Correspondence: M.E. Collinson, Geology Department, Royal Holloway University of London, Egham, Surrey, TW20 0EX, England; email: [email protected]; J.J. Hooker, Palaeontology Department, The Natural History Museum, Cromwell Road, London, SW7 5BD, England; email: [email protected]


INTRODUCTION scattered but useful sources are Mai (1995), This article is intended to provide an over- Kunzmann (1999) and Yang & Jin (2000, for view of the nature and distribution of the four Metasequoia [dawn redwood]). The rare major types of Paleogene vegetation in Paleogene cycads are discussed by Kvacek & Eurasia from the perspective of the role of Manchester (1999). vegetation as mammalian habitats and food Paleogene plants are usually represented in resources. Readers interested in the vegeta- the fossil record by various dispersed, isola- tion or flora of a particular region, zone or ted organs. Rarely, especially in the case of site, or in the variety and nature of aquatic herbs, complete plants may be communities, should consult the references encountered. Usually ‘whole’ plants must be cited for more specific assessments of the reconstructed and a variety of approaches are vegetation and composition. The refe- used (Collinson 1990, 2000b). Relatively rare rence list is not intended to be exhaustive but fossils are able to prove conclusively the to provide key examples for entry into the links between organs if they show organic relevant literature. The paper also provides a connection, e.g. leaves and fruits attached to brief introduction to the ways in which plants a woody twig or within stamens of a and vegetation can be reconstructed from fos- . Biological connection is also valu- sils. A more wide-ranging introduction to able, e.g. if pollen survives on a persistent paleobotanical techniques may be found in stigma in a fruit. Recurrent association can Jones & Rowe (1999). also be used if organs exclusively and/or repeatedly co-occur in samples at different RECONSTRUCTING PALEOGENE localities. Anatomical similarity uses the pre- VEGETATION sence of a distinctive character (e.g. a distinc- tive epidermal hair) to link organs to the Paleogene plants same plant. Finally, it is possible to use the A general summary of the Paleogene plant nearest living relative (NLR) of fossils, espe- record is given in Collinson (2000b) with a cially if there are multiple organs (e.g. pollen, more extensive review of Paleogene plant leaves, fruits, seeds, and possibly flo- paleobiology in Collinson (1990). The wers) all indicating the same type of plant. Paleogene flora is dominated by flowering The philosophy applied here would follow plants with associated ferns, conifers, Ginkgo that of Collinson (1986). and rare cycads. Among the flowering plants, the dicotyledon groups (e.g. Paleogene vegetation [], Lauraceae [laurels]), hamamelids (e.g. Betulaceae [birches], In situ plants, sedimentary facies and tap- Juglandaceae [walnuts, wingnuts], honomy After plants have been reconstruc- [planes], Ulmaceae [elms], ted they must be incorporated into recon- Fagaceae [beeches]) and rosids (e.g. structed vegetation. In this task plants preser- [dogwoods, mastic trees], [soap- ved in their position of growth are of excep- berries], Icacinaceae [icacina lianas]) are tional value, including tree stumps; aquatic or abundant and diverse as are the monocotyle- marginal vegetation inundated by floods, and don groups alismatids (e.g. Stratiotes [water plants in peat-forming environments. In other soldier]) and arecids [palms, aroids]). First circumstances the taphonomy, including sedi- occurrences of families may mentary context (facies) and preservational be found in Collinson et al. (1993a). Details state of the plant fossils is pivotal to judging of Cenozoic ferns are given in Collinson the degree of transport and hence the relative (2001b, 2002). Information on Tertiary conif- autochthony of the assemblage. Litter mats ers and Ginkgo (maidenhair tree) is more may preserve the flora essentially in situ

42 COLLINSON & HOOKER: Paleogene vegetation Eurasia

whereas beach deposits may contain a variety unrelated groups of plants, also indicated the of fossils of plants, which did not originally same forest type on the basis of their nearest grow together in the same vegetation. A low living relatives. In order for this approach to degree of fragmentation and abrasion, the co- have value for reconstructing vegetation on a occurrence of organs of different sizes and regional scale it is essential that a single shapes but derived from the same plant, and a taxonomic approach should be employed lack of evidence of concentration by wind or (ideally by one, or a small group) by specia- water would all indicate minimal transport. lists who are familiar with the floras over the The best approach is to study plant fossils entire area of study. In practice this procedure from a variety of sedimentary facies. This is rarely followed. helps to constrain different sources and to ensure that regional (zonal), rather than only Other biota & sedimentological criteria local (azonal), vegetation is being studied. Reconstructions of ancient vegetation can be Experimental (or modern) taphonomy is of checked for consistency against other lines of considerable value in understanding plant evidence such as that from other biota, prima- fossil accumulations and the extent to which rily insects and mammals. As each of these they reflect ancient vegetation. Approaches groups is subject to varied and different tap- are summarised by Ferguson et al. in Jones & honomic bias the approach is not straightfor- Rowe (1999) in which key literature on plant ward. However, one factor which we have taphonomy is also cited. found informative, is the ecological diversity of mammalian faunas (Andrews et al. 1979, Physiognomy The physiognomy of some Collinson & Hooker 1987, Hooker 1998). plant organs, especially leaves and , Indeed, these provide helpful indications of can also contribute to reconstructing vegeta- spacing of plants in a vegetation (i.e. open or tion through inference of a biome type within closed), a factor that is hard to establish a particular climate regime. Further discus- directly for different kinds of ancient forest in sion of the relative roles of different plant the absence of in situ tree stumps. Relevant organs is given in Collinson (1990, 2000b). sedimentological criteria include coals and In particular leaf size and leaf margin, and evaporites (Parrish et al. 1982) and paleosols more recently the multivariate CLAMP ana- (Retallack in Jones & Rowe 1999). lysis (Wolfe & Spicer in Jones & Rowe 1999), provide evidence of climate regimes, Independent dating Reconstructions of as do the presence, absence and nature of ancient vegetation depend on reliable inde- growth rings (Creber & Francis in Jones & pendent dating of the floras. An entirely cir- Rowe 1999). cular argument is created if the nature of the fossil flora is used to estimate its age. Such Nearest living relative Inferences about independent dating may come from absolute ancient vegetation are often based on nearest dates, magnetostratigraphy, from continental living relatives. This approach may be very sequences intercalated with marine strata well-justified in some cases. An example containing standard biostratigraphic markers would be when the fossil record reveals a and from biostratigraphy using mammalian suite of different fossil organs (e.g. wood, faunas, charophytes or less ideally, palynolo- pollen, leaf, fruit) and all indicate an affinity gy (when studying macrofloras). with a particular modern group of plants all of which grow in one particular forest type Criteria used here Where possible the (e.g. examples of the Juglandaceae, information used in this paper is drawn from Manchester 1987). Further support would be recent work in primary literature, or personal provided if other associated fossils, from observations, based on rigorous systematic


studies (or revisions) of floras from indepen- Trochodendroides and Corylites leaf types) dently dated sites. All approaches listed and taxodiaceous conifers. A representative above have been employed, although not leaf assemblage is from Atanikerdluk, equally for all sites. The senior author (MEC) Greenland (Mai 1995, fig. 131). A fern under- is a specialist in fruits and seeds and in wes- story or forest marginal vegetation is associa- tern European floras and does not claim full ted with these floras (Collinson 2001b, 2002). familiarity with leaf floras or with floras in It includes Osmunda, Onoclea, Woodwardia other areas. Nevertheless she has actively and the extinct Coniopteris. participated in conferences and discussions In the Greenland Region (Greenland, involving many of the authors whose work is Spitsbergen, Ellesmere Island, Scotland) the cited here (see especially Boulter & Fisher floras are typified by Trochodendroides, 1994 section 5 on workshop discussions; and Corylites and Metasequoia. They include also Knobloch & Kvacek 1990, Planderova et Nordenskioldia plants () al. 1993). In the cases of sites in China, and Nyssidium plants (Cercidiphyllaceae), Former Soviet Union, Mongolia and other both with Trochodendroides leaves sites where primary literature is not in the (Nordenskioldia also at Tsagayan, east Roman alphabet, recent review articles in Russia, Crane et al. 1991), Macclintockia English have been used to assess vegetation (Incertae Sedis ?Urticaceae, Mai 1995), in combination with illustrations and floral Ushia (?Fagaceae), Platanaceae (Platanus lists in original literature. The use of literatu- and Platanites, Crane et al. 1988); re is no substitute for study of specimens and Fagopsiphyllum (Manchester 1999), the senior author fully acknowledges this Palaeocarpinus (Golovneva 2002), Ginkgo inadequacy. However, as this article deals and relatively rare pinaceous conifers only with broad vegetation categories the (McIver & Basinger 1999, Golovneva 2000, consequences may not be too severe. Where Boulter & Kvacek 1989, Kvacek & Manum heavy reliance has been placed on literature 1993, Kvacek et al. 1994). an attempt has been made to use articles from In far eastern and northern mainland Russia a small number of authors or collaborators in and Sakhalin the floras are also characterised order to reduce the problems of multiple by a Trochodendroides/Corylites/Metasequoia taxonomic approaches. In spite of the above association (in Kamchatka, Rarytkin Ridge attempts at mitigation this paper includes: and northern Sakhalin (Boshnyakovo information based on floral lists with no illus- Formation) and in the Lena River (Budantsev trations; leaf floras which are impression flo- 1994a, 1997; Herman 1993; Herman & ras with few diagnostic characters; floras Spicer 1996; Golovneva 1994, 2000). This known to require major systematic revision contrasts with the Taxodium/Trochoden- and floras which are not independently dated. droides association in the Zaysan area, These and other problems can only be solved Kazakhstan, Tsagayan and Kivda region of by future research. Readers should check the Amur, Russia and Mongolia (Golovneva primary literature for any particular area in 1996). The aquatic plant Quereuxia is a dis- which they are interested. tinctive element in the far eastern Russian assemblages (Golovneva 1996). This plant PALEOCENE AND EOCENE POLAR, also appears in the North American BROAD-LEAVED, DECIDUOUS Paleocene (e.g. in Canada at Ravenscrag FORESTS Butte, Saskatchewan; McIver & Basinger 1993) and on Ellesmere Island (McIver & Floras Basinger 1999), being one of a number of The floras are characterised by deciduous flo- elements indicating dispersal between these wering plants (especially plants with areas (Manchester 1999) across Beringia.

44 COLLINSON & HOOKER: Paleogene vegetation Eurasia

Occasional thermophilic elements are docu- ments from the polar deciduous forest. mented in the Kamchatka floras, e.g. However additional elements include Nyssa Magnolia, Nyssa and Sassafras at Snatol and this flora also represents a somewhat (Maslova 2000, Fotyanova 1995). In eastern mixed assemblage. A flora of late Paleocene Kazakhstan in the Zaysan Basin (Kara- or early Eocene age in South Sakhalin inclu- Beryuk) a short-lived interval contains a des thermophilic elements such as Alchornea, Trochodendroides-rich polar deciduous type which are found in Eocene floras of Japan of flora (Golovneva 1996). (Kodrul 1994). The floras from Mongolia are similar to Paleocene communities within this vegeta- some of those from the Zaysan Basin tion were interpreted by Krassilov (1995). (Makulbekov 1987). Indeed, the flora from They included a) aquatics (with e.g. the Naran Member at Naran Bulak and Quereuxia in eastern Russia and the USA) b) Tsagan Khushu (Makulbekov 1988, table 4) a marginal waterside community (represented is of very low diversity with several by monocotyledonous leaves); c) a swamp Trochodendroides leaf types, Trochoden- forest community (Taxodiaceae and drocarpus and Nordenskioldia fruits, Myrica Trochodendroides-bearing plants); and d) an and two other dicotyledon leaves, the ende- upslope deciduous woodland or more open mic conifer Gobiostrobus Makulbekov forest. (The inference of openness was based (Makulbekov 1995) and other conifers of the on debris flow deposits indicating erosion). Taxodiaceae. The flora from the Naran Middle Eocene communities within this Member at Kaychin Ula I is more diverse vegetation included Metasequoia swamp with Taxodiaceae, many Trochodendroides forests, (with Alnus groves as a successional leaf types, Macclintockia, Betulaceae, stage). Dicotyledonous angiosperms especial- Ulmaceae, and Viburnum-like ly Betulaceae (and the fern Osmunda) are leaves (Badamgarav & Reshetov 1985, table judged to be a significant component of the 2; Makulbekov 1988, table 4). Many ele- swamp forest although they are often over- ments of these floras are typical of the polar looked in the litter assemblages (McIver & deciduous forests and of the floras of eastern Basinger 1999, p. 535). Tree density in these and western Kazakhstan (Makulbekov 1988, swamp forests reached 1100/hectare compa- table 8). These floras may represent the polar rable to modern tropical rain forests deciduous forest or a taphonomically restric- (Basinger et al. 1994, McIver & Basinger ted, but more mixed, vegetation. The presen- 1999, Vann et al. oral comm. 2001). Rarely, ce of Myricaceae (Makulbekov 1988, table 8) swamp leaf litters exhibit a mixture of other distinguishes this flora from the polar deci- conifers including Pinaceae, which were duous forests, as does the long narrow-leaved evergreen. The remains of these evergreen nature of some unidentified dicotyledon conifers also occur in fluvial sands (channel leaves mentioned by Makulbekov (1983) alt- or point bar deposits) suggesting presence of hough these may remain indeterminate a more distant ‘upland’ mixed evergreen (Makulbekov 1988). For these reasons the conifer and deciduous angiosperm forest floras have been assigned here to the mixed (Basinger et al. 1994, Lepage 2001, McIver component of Paleocene vegetation. The & Basinger 1999). The other main communi- Mongolian floras are referred to as a ty was lowland broad-leaved deciduous Taxodium/Trochodendroides association by forests (with deciduous conifers) in better- Makulbekov (1983) and Golovneva 1996). drained sites with remains deposited in over- The Tsagayan flora (Ohana & Kimura bank or fluvial deposits. In the middle 1995, Krassilov 1995, Golovneva 1996) also Eocene a few thermophilic or evergreen ele- exhibits a predominance of Taxodium and ments occur in these lowland forest commu- Trochodendroides and includes many ele- nities (Menispermaceae, Nyssa, and lobed


oaks) alongside the typical deciduous ele- These forests have essentially no lianas or ments (Basinger et al. 1994, McIver & climbers. One leaf type (Archeampelos) in Basinger 1999). Herbaceous wetlands inclu- the assemblages on Axel Heiberg & ded Equisetum and Sparganium (McIver & Ellesmere Islands may belong to the Basinger 1999). and may possibly represent a vine (McIver & Basinger 1999) and a probable menisper- Vegetation overview maceous vine occurs in the middle Eocene These polar deciduous forests are characteri- floras of Axel Heiberg and Ellesmere Islands. sed by a low to very low diversity (single One leaf from Mull may be related to sites with less than 20 leaf species, often only Vitaceae (Boulter & Kvacek 1989). a few (Basinger et al. 1994). There would Understory, or at least marginal vegetation thus have been little habitat variation within included a low diversity fern flora (Collinson the forests. That the nature of the environ- 2001b, 2002). The trees were mainly deci- ment was warm, equable and largely frost- duous (with a few evergreen elements in free, with a long favourable growing season more southern ‘mixed’ floras) with thin, and a sudden shutdown of growth in winter, medium and large leaves. The fruits were is documented by leaf floras, wood physio- characterised by small to very small dry gnomy and associated biota including large seeds in woody follicles (Cercidiphyllum-like land tortoises, varanid lizards and alligators plants) or capsules (Nordenskioldia) or small (Estes & Hutchison 1980, Basinger et al. nuts (Betulaceae, Juglandaceae, Ulmaceae) or 1994, McIver & Basinger 1999). small seeds in conifer cones. Only Ginkgo The plants include a number of well-under- (apparently not an abundant element) would stood genera, which are known from recon- have produced a substantial-sized (al- structed whole plants. These include though the outer flesh of living Ginkgo has Cercidiphyllum-like plants such as Nyssidium an unpleasant smell which may render it rat- and Trochodendrocarpus; Nordenskioldia her unpalatable). These dietary resources plants, Platanaceae and Betulaceae (Crane et would have been almost entirely seasonal in al. 1991; Crane et al. 1988; Boulter & their availability although evergreen conifers Kvacek 1989; Golovneva 2000, 2002; Crane occurred in ‘uplands’ at some sites (e.g. Axel 1981, 1984; Maslova 1997; Krassilov & Heiberg). This polar deciduous forest, a warm Fotyanova 1995; Pigg & Stockey 1991; Chen forest with winter darkness in a mesothermal et al. 1999; Feng et al. 2000). Metasequoia is humid climate, has no modern analogue and also known from reconstructed plants scarce- this biome is extinct. ly distinguishable from modern species (Basinger 1984) and is represented in the LATE PALEOCENE AND EOCENE Eocene of Axel Heiberg by wood, leaves, BROAD-LEAVED, PARATROPICAL pollen cones and seed cones (McIver & TO SUBTROPICAL, EVERGREEN Basinger 1999). All of these plants are char- (RAIN) FOREST acterised by a colonising strategy. This is pro- ven by Paleocene examples from the Joffre Introduction Bridge site in Alberta, Canada. There, the This vegetation is not equivalent simply to Joffrea plant (Cercidiphyllaceae), platanace- ‘tropical rainforest’ for several reasons. (1) ous plants and Metasequoia-like plants Mean annual temperature (MAT) from leaf (Stockey & Crane 1983, Crane & Stockey physiognomy does not usually indicate tem- 1985, Pigg & Stockey 1991, Falder et al. peratures greater than 25oC, which is the 1999) all have small seeds and exhibit simul- lowest for tropical forest (Wolfe 1987). (2) taneous germination proven by fossil seed- Temperate or deciduous elements/indicators lings. often co-occur, resulting in mixtures not

46 COLLINSON & HOOKER: Paleogene vegetation Eurasia

found in modern tropical rain forests. (3) characteristic deciduous elements of the Late There is no direct analogue for the fossil Paleocene/Eocene polar deciduous forests. forests in modern tropical forests. Although These have been characterised as "mixed" on many nearest living relatives occur in the vegetation map. Floras of ‘mixed’ aspect Southeast Asian tropical rain forests today, with some indicators of the paratropical to key elements, especially dipterocarps and subtropical forest vegetation, (e.g. large ever- epiphytes, are rare to absent in Eocene green leaves, members of the Lauraceae, forests. For further discussion see Collinson fruits and seeds with living relatives in subt- (1983a, 2000b, 2001a). ropical to paratropical forests) occur in wes- tern Europe at Sézanne and Menat (Mai Late Paleocene and 1995), in southern England (Collinson 2000a, Paleocene/Eocene transition floras Collinson & Cleal 2001a), at Vervins and in The early late Paleocene Gelinden flora the Soissonnais (Mai 1995). The Sézanne and (which requires detailed systematic revision) Menat floras, like that from Gelinden, also is judged by Mai (1995) to be dominantly require major revision. thermophilic and taken by him as the key In eastern Kazakhstan in the Zaysan Basin Paleocene representative of this vegetation in (Kara-Beryuk) a short-lived interval contains Europe. The typical leaf assemblage is shown a Trochodendroides-rich polar deciduous type in Mai (1995 fig. 133). It does include a few of flora (Golovneva 1996). This is replaced deciduous elements e.g. Ushia but is domina- by subtropical floras (e.g. at the Chakel’mes ted by evergreen Fagaceae and Lauraceae, site), which may include Dryophyllum & esp. Dryophyllum, typical of the Eocene exa- Dewalquea (at the Tayzhuzgen site) and mples of this vegetation type. Mai (1989, sometimes lack Trochodendroides (e.g. at 1991,1995) considered this as an evergreen Kiin-Kerish), but still retain many affinities laurophyllous forest or evergreen notophyl- with boreal northern Asian deciduous floras lous broad-leaved forest (the latter being e.g. Platanaceae (Golovneva 1996, mesothermal rather than megathermal sensu Makulbekov 1983, 1987). Mongolian floras Wolfe (1987)). Woods associated with this have been discussed above under polar deci- flora show only indistinct growth rings duous forests. All these floras are categorised (Gerrienne et al. 1999). as ‘mixed’ on the vegetation map. The later late Paleocene floras of Eisleben The few Paleocene floras from China are and Roda, Germany, are judged by Mai reviewed in Guo (1990) and Liu et al. (1996) (1987, 1995) to be boreotropical (i.e. ther- with an additional flora from Lingbao mophilic) floras but with smaller and more County, Henan (S. Guo pers. comm. 2001). serrated leaves than is a typical for this vege- These floras indicate a mixed vegetation of tation. The floras from Rusava, Rusca and temperate deciduous broad-leaved elements Dobre Sreca are judged to be thermophilic; with warm temperate to subtropical elements. the former two include palms but, atypically They include Trochodendroides and for this vegetation, lack Fagaceae and Cercidiphyllum-type leaves, Ushia, Lauraceae (Mai 1987,1995). The floras of Betulaceae, Platanaceae and Taxodiaceae, but western Kazakhstan and the Volga area also some more thermophilic elements. (Bolytschka, Kamyshin, Ushi, Romankol and Tykbutak) are subtropical floras with many Eocene floras palms and evergreen Lauraceae and Fagaceae (like Gelinden, also including Ushia) (Shilin Strongly thermophilic/evergreen floras in 2000, Makulbekov 1987). Europe Leaf floras are dominated by evergr- However, in the Paleocene some western een Fagaceae and Lauraceae. A typical exa- European floras also include more of the mple of a leaf flora is that from Messel,


Germany, illustrated in Mai (1995 p. 136). Strongly thermophilic/evergreen floras in The leaf morphology (large leaves, thick leat- China Floras in southern China indicate an hery texture, entire margins) is indicative of evergreen broad-leaved forest growing in a evergreen habit for many leaves. Leaf physio- wet and tropical climate. The leaves are leat- gnomy indicates megathermal to very warm hery and some are coriaceous but these are mesothermal vegetation. Key elements inclu- larger than in the central Chinese region (see de Eotrigonobalanus, Trigonobalanopsis vegetation with sclerophyllous elements). (Fagaceae), Steinhauera (Altingiaceae), Evergreen Fagaceae, Lauraceae and palms are Rhodomyrtophyllum (), and Litsea, significant elements (Guo 1990). Lindera, Ocotea, Daphnogene (Lauraceae) species (Mai 1995, Wilde 1995, Walther Other strongly thermophilic floras More 1994). Sterculiaceae, Icacinaceae, isolated records of thermophilic floras occur Sapotaceae, Menispermaceae, Vitaceae, in Japan and Kamchatka. The Japanese floras Nyssaceae, Apocynaceae, Arecaceae and from the Ube coalfield, Honshu are evergreen Symplocaceae are also recorded in leaf floras mainly tropical floras with some deciduous (Mai 1995, Wilde 1989, 1995). Ferns such as elements (Huzioka & Takahashi 1970, Tanai Lygodium are frequent. Doliostrobus is a 1995). Those from the Ishikari coalfield, common conifer and the cycad Eostangeria is Hokkaido are similar with abundant tropical also present (Wilde 1989,1995, Mai 1995, elements, especially Euphorbiaceae, Kvacek & Manchester 1999, Collinson Icacinaceae, Menispermaceae and 2001b, 2002). Juglandaceae (especially Sterculiacaeae, as well as palms and Platycarya and Engelhardia groups) are also Musaceae (Tanai 1990) and evergreen documented from fruits and foliage Fagaceae (Tanai 1990, 1995). Deciduous ele- (Manchester 1987, Mai 1995). Woods show ments also occur in these floras (Tanai little evidence of growth rings. Growth rings 1981,1990, 1994) and become more common are absent in about 70% of twigs from the in the floras of the Kushiro coalfield (Tanai English London Clay flora (Collinson 1983a, 1970, 1990), which is late Eocene in age Poole 1992, 2000). In European Eocene fruit (Tanai 1990) and has been assigned here to and seed floras, taxa whose nearest living the subtropical floras (see below). relatives occur in paratropical forests are The most thermophilic flora in Kamchatka diverse and abundant (Collinson 1983a, is the flora of the Chemurnaut Bay area, 2000b, Collinson & Cleal 2001b, Tiffney which is still under study (Budantsev 1994a, 1994, Mai 1970, 1989,1995). These elements 1997). This flora contains 43-45% entire mar- include members of the Arecaceae, gined leaves, many leaves are large, and Anacardiaceae, Anonaceae, Burseraceae, some have drip tips and there are many ther- mastixioid Cornaceae, Dilleniaceae, mophilic elements with near living relatives Icacinaceae, Lauraceae, Menispermaceae, in the Lauraceae, Arecaceae, Sapindaceae, Sabiaceae, Sapindaceae, and Vitaceae. Styracaceae, Symplocaceae, Apocynaceae, Although a few members of some of these Sabiaceae and a high diversity of legumes taxa extend today into temperate zones, a etc. (Budantsev 1994a, 1997). This is not a high diversity is only found in megathermal sufficiently thermophilic flora to be conside- to warm mesothermal vegetation. Diverse red equivalent to the European or southern palms are particularly important because Chinese floras. However, it seems to be signi- palm functional biology confirms warm con- ficantly more thermophilic than, and very dif- ditions without recourse to living relatives. ferent floristically from, the other (Paleocene Collinson (2000b, 2001a) also discussed this and Eocene) floras of Kamchatka (Budantsev vegetation type. 1994a, 1997). It has therefore been plotted as strongly thermophilic on the map to draw

48 COLLINSON & HOOKER: Paleogene vegetation Eurasia

attention to this fact. It should also be noted re-appraisal of nearest living relatives and that there is debate about the age of the flo- detailed comparisons between floras, is nee- ras. For example the floristically rich ded to explore the temporal and spatial distri- Tkapravayam Formation is argued to be mid- bution of the variations in the Eocene parat- dle Eocene by Budantsev (1994a, 1997) but ropical/subtropical evergreen forests. late Paleocene by Maslova (1997), both of whom took account of marine faunal eviden- Mangroves Further evidence of the humid ce. Krassilov & Fotyanova (1995) also consi- thermophilic aspect of Eocene vegetation is dered the formation to be late Paleocene. afforded by the presence of mangroves domi- Blokhina (1997, 1998) who studied woods, nated by the Nypa palm (represented by fruits and found them to have distinct growth rings and pollen). These ranged from north western (as would be expected at this high paleolati- Europe through central and eastern Europe to tude), also commented on the disparate India, South East Asia and Australasia and dating. Other Eocene floras in Kamchatka into North and South America, but are not contain thermophilic elements such as the recorded from eastern Asia (given that a pol- most northern occurrence of Sterculiaceae in len record from Japan is questionable). the late Eocene Sedanka site, Napana River A map of the distribution of ancient Nypa (Budantsev & Baranova 1995). Further study was produced by Collinson (2000b, Fig. 16.4, and age refinement (e.g. exploiting the fora- p. 233). Localities with Nypa in the minifera documented by Fotyanova & Serova Paleogene of Eurasia are Figure 2 and Table 1994) of the Kamchatkan floras will clearly 1 site numbers 15,19,45,52,92 and 100 be of great significance for the understanding (Collinson 1993, 2000b; Tralau 1964). In of high latitude Paleogene floras. addition Nypa is recorded from Kiev, Ukraine (near site number 98) (Tralau 1964, Mai Floras designated subtropical/with tempe- 1995, p.106); Hruby Regiel, Tatra Mountains, rate elements These floras are distinguished Poland (near site number 29) (Tralau 1964, on the Eocene map as they do not seem to Mai 1995, p.382); Dudar in the Dorog Basin, represent forests that are as strongly thermop- Hungary (near site number 86) (Tralau 1964, hilic or evergreen as those represented by the Mai 1995, p.382) and near Odessa, southern floras discussed above. The distinction is in Ukraine (Tralau 1964). Details of localities in places tenuous and is largely based on com- England, Belgium and France may be found ments in the literature (see references for spe- in Collinson (1983a, 1993, 1996) and Tralau cific sites in Table 1) where these floras are (1964). All these Nypa sites are associated referred to as ‘subtropical’ (rather than parat- with ancient coastlines bordering the strongly ropical or evergreen/thermophilic) and/or evergreen thermophilic vegetation in western where temperate elements are emphasised. and south-eastern Europe and the Ukraine. In The distinction is complicated by the fact that England, members of the there is a temporal transition, in places star- (true mangroves) such as Ceriops (fossils of ting in the late Middle Eocene, with incre- the viviparous embryos) are associated with asing temperate deciduous (and/or sclero- the Nypa, as is an extinct plant, which produ- phyllous) elements in floras. The transition is ced Wetherellia fruit fossils (Collinson 1983a, discussed under late Eocene to Oligocene 1993, 1996, 2000b, 2001a). Fossil pollen mixed deciduous and evergreen forests. The whose nearest living relatives are mangroves dates for some floras are not specific (e.g. today (e.g. Avicennia and ) suggest the pre- Eocene) and for others the dating is based sence of other mangrove taxa in association largely or entirely on the floral composition. with Nypa at some sites (e.g. Helmstedt, site Further research, including independent number 55, Riegel et al. 1999). dating, physiognomic approaches, in depth


Vegetation overview This vegetation had indicate increased seasonality, decreasing very high diversity. More than 300 species temperatures, or both. This vegetation yields are recorded in some of the fruit and seed flo- a wide variety of fruits and seeds with a ras of the London Clay Formation of the range of sizes. Diverse fleshy fruits are pre- London and Hampshire Basins, southern dicted from nearest living relatives of many England (Collinson 1983a; Collinson & Cleal taxa represented by fossil seeds or fruit sto- 2001a,b) though these are exceptionally well- nes. Fleshy fruits are conclusively demonstra- represented owing to various taphonomic and ted by exceptionally preserved specimens at collecting factors. The Messel leaf flora, Messel with leathery flesh and outer fruit sourced only from the catchment of a relati- pericarp (skin) preserved (Schaal & Ziegler vely small lake, is at the opposite extreme, 1992, Collinson 1999, Hooker & Collinson but even this has 65 species based on leaves 2001). (Wilde 1989). Additional taxa are represented in the fruit, seed and pollen floras, so that 58 LATE EOCENE AND OLIGOCENE families of angiosperms are represented over- BROAD-LEAVED MIXED DECI- all at Messel (Schaal & Ziegler 1992). The DUOUS AND EVERGREEN FOREST middle Eocene Clarno flora, representing this vegetation in North America, has 173 species Introduction based on fruits and seeds. Pollen floras as Areas which preserve a temporal sequence of indicators of former biodiversity in this vege- sites e.g.: southern England (Collinson & tation are considered in the section on India Cleal 2001a,b,c), The Weisselster Basin, and Southeast Asia. The high diversity would Germany (Mai & Walther 1978, 1985; Mai mean that these forests provided a richly 1995); the Czech Republic (Mai 1995, Buzek varied habitat. et al. 1990); Kamchatka (Budantsev 1994a, The plants display a wide range of ecologi- 1997); Kazakhstan (Zhilin 1989, Makulbekov cal strategies, from colonisers to those adap- 1987) and Japan (Tanai 1990) all show transi- ted to climax forests. Leaves are medium to tional floras following those of the Eocene large, leathery and entire margined and trees paratropical/subtropical evergreen forests. were dominantly evergreen. Analogy with These may reflect an increase in temperate modern forests would suggest the presence of and deciduous elements, with vegetation two canopy layers, probably with an addition- changing, for example, through evergreen al emergent layer. There are numerous lianas notophyllous forest to mixed mesophytic and climbers based on nearest living relatives forest. Alternatively, they may show an incre- and leaf physiognomy (especially a high ase in sclerophyllous elements (see below diversity of Menispermaceae and Vitaceae under vegetation with sclerophyllous ele- known from fruits and seeds, as well as from ments). The changes may commence as early leaves with cordate shape and drip tips). as the late middle Eocene in some areas and Fragments of twining stem in the London they continue through the late Eocene and Clay floras demonstrate the presence of clim- into the Oligocene, the exact timing being bers. The most strongly thermophilic of these variable and well-constrained in only a few forests would have experienced little (possi- areas. Each area is discussed below. bly in some cases no) seasonality. Growth rings are absent in 70% of twig woods and European floras very rare in trunk woods in the London Clay The transitional, continuous rather than flora (Poole 1992, 2000). Deciduous trees are abrupt, nature of vegetational change in rare. Increased proportions of deciduous ele- Europe was outlined by Collinson (1992) and ments in some of the floras mapped here as Knobloch et al. (1993). In Central Europe the subtropical/with temperate elements may late Eocene Stare Sedlo flora, Czech

50 COLLINSON & HOOKER: Paleogene vegetation Eurasia

Republic (monographed by Knobloch et al. 1996, Kvacek & Walther 1998), the former 1996) is dominantly evergreen containing being mixed mesophytic forests and the latter many elements of subtropical rain forests and notophyllous evergreen forest. In Hungary referred by Mai (1995) to the same floral the deciduous temperate elements only appe- complex as the Zeitz flora (see below). The ared very sporadically in the early Oligocene transition was very well illustrated for but they became significant in the late Central Europe by diagrams in Buzek et al. Oligocene (Hably 1993, Kvacek 1996). (1990) also redrawn in Mai (1995 fig. 138) Kvacek and Walther (2001) further documen- and was discussed by Walther (1994). The ted provinciality in the Oligocene of Europe, paratropical elements do not all disappear at showing that a southern province was distin- once, indeed some persist well into the guished by the persistence of more subtropi- Oligocene. Lauraceae are retained but are cal elements with temperate elements only represented by different species e.g. of Litsea appearing in the late Oligocene. Fluctuations and Laurophyllum. However, incoming taxa in the proportions of leaf physiognomic types (Betulaceae, Ulmaceae, Acer, Liquidambar, and of nearest living relatives offer great Fagus, Carya, Craigia [formerly Ptelea], potential for more precise vegetation and cli- Tiliaceae, Cercidiphyllum) include many mate delimitation during transitional inter- deciduous elements and most of these first vals. However, great care must be taken to appear in the Oligocene. study regional (zonal) rather than local (azon- In western Europe the transition is more al) vegetation and to take account of taphono- gradual with loss of paratropical elements mic bias and paleogeographic settings and incoming of deciduous elements from the (Kvacek & Walther 1998, 2001; Kvacek late middle Eocene (Collinson 1992). Typical 2000). Such detail is unfortunately outside leaf floras of the Late Eocene have been the scope of this general article. interpreted as evergreen to semi-evergreen The Kundratice and Haselbach floras and mesophilic e.g. Célas, France (leaf assembla- floras of the Lausitz region (Mai 1994) are ge illustrated by Mai 1995, fig. 163) or as other examples of European Oligocene vege- evergreen notophyllous vegetation e.g. the tation. The leaf assemblage at Haselbach is Zeitz Complex, Weisselster Basin, Germany illustrated by Mai (1995, fig. 140) and the (Mai 1995, leaf assemblage illustrated in vegetation is interpreted as mixed mesophytic Walther 2000, fig. 1). forest. The leaf assemblage from Fruits belonging to the incoming elements Kleinsaubernitz is illustrated by Walther include larger dry nuts e.g. of Fagaceae, (1999, text-fig. 16) and the assemblage inter- Carya and Betulaceae. The deciduous compo- preted to represent a mixed mesophytic forest nent is floristically distinct from that of the with more than 50% evergreen and thermop- Paleocene polar deciduous forests. For exam- hilic elements. The floral assemblage from ple, it generally lacks extinct plant genera, Kundratice is illustrated by (Kvacek & has smaller leaf sizes and includes a wider Walther 1998, text fig. 13). At Kundratice, variety of fruit types, especially larger dry (volcanic floras preserved in depressions of fruits. fault basins and maar lakes), communities The late Eocene and Oligocene mixed deci- include a broad-leaved deciduous forest in a duous and evergreen forests are truly a mixed humid habitat (e.g. lake margin) with rare vegetation, with the proportions of different evergreens (Platanus neptuni and elements varying through time and in diffe- Daphnogene) and a mixed mesophytic forest rent geographic areas. One excellent example dominated by deciduous broad-leaved trees. of relatively local geographic variation is the Evergreen Lauraceae persist here. MAT is difference between the early Oligocene floras estimated at 9-13oC with a humid (Cfa type) of the Czech Republic and Hungary (Kvacek climate (Kvacek & Walther 1998). Fruit and


seed floras are represented in the Lausitz leaves including members of the Lauraceae region in the Calau Beds (Mai 1998) and in and Fagaceae. A radical change was noted in Saxonian Lausitz (Mai 1997). The Calau the Oligocene with the rise of a warm tempe- flora was also interpreted as representing a rate deciduous flora, termed the Turgai (= mixed mesophytic forest. Turgay/Turgayan) flora or the flora of Turgai ecological type (Shilin 2000, Zhilin 2001). Japanese floras Further discussion of sclerophyllous ele- Deciduous elements become more common ments, which are very important in these flo- in the floras of the Kushiro coalfield (Tanai ras, is given in the section on vegetation with 1970, 1990), which is late Eocene in age sclerophyllous elements. (Tanai 1990). Floras from near Kobi City, In eastern Kazakhstan, in the Zaysan Basin Honshu (Tanai & Uemura 1994, Tanai 1995, floras such as Kershi and Kalmakpay are Fotyanova 1997) and from the Rikubetsu and dated to middle Eocene on the basis of the Kitami coalfields, Hokkaido (Tanai 1981, associated Obayla fauna and they 1990, 1992, 1994, 1995; Tanai & Uemura represent subtropical forests, having large 1994; Fotyanova 1997), are possibly of late leaves, few temperate elements and lacking Eocene or early Oligocene age. They are such groups as birches. Overlying late referred to as warm temperate floras. Eocene floras (e.g. Akkezen [same location as Kalmakpay]) retain some subtropical ele- Floras in Kamchatka ments (e.g. Lindera) but show an increase in The late Eocene floras of Kamchatka have temperate elements, some of which are small- been divided into three floral stages leaved sclerophylls (Iljinskaya 1984a,b; (Budantsev 1997). The oldest two stages are Makulbekov 1977, 1987). The site at Kiin- considered to represent temperate mesophilic Kerish (same Eocene map location as Kershi) floras. They include a proportion of tempera- yields floras assigned to the late Eocene and te deciduous elements and some small-leaved early Oligocene, with the former containing a elements but they also retain some evergreen higher proportion of subtropical elements and thermophilic elements characteristic of earlier the latter more temperate and sclerophyllous floras. First appearances for the area of ther- elements (Zhilin 1989; Makulbekov 1977, mophilic elements such as Firmiana of the 1987). The Oligocene flora of Ashutas is also Sterculiaceae (Budantsev & Baranova 1995, a temperate flora with sclerophyllous ele- Budantsev 1997) as well as Magnolia and ments. Euphorbiaceae are also notable. The latest In northern and central Kazakhstan, floras floral stage (Irgirninian) contains dominantly are generally sclerophyllous earlier, even in broad-leaved deciduous plants, including taxa the middle Eocene, possibly an effect of local not encountered earlier, and lacks the ther- paleogeography (Makulbekov 1987). Palms mophilic elements (Budantsev 1997). Refined and evergreen Fagaceae and Lauraceae are independent dating of these floras will be characteristic of floras assigned to the middle very important for understanding high latitu- Eocene (Karasor) and the late Eocene to early de Paleogene floral change. As far as we can Oligocene (e.g. Zhamantuz, Akhzar, Shiderty) determine none of the floras in this area are but the leaves are dominantly small, leathery considered to be Oligocene in age. sclerophyllous elements (Makulbekov 1972, 1977, 1987). Floras in Kazakhstan In western Kazakhstan, middle Eocene flo- Shilin (2000) emphasised the subtropical ras (e.g. Karakol) show slight indications of nature of the Paleocene and Eocene floras of xeromorphic elements but also show similari- Kazakhstan, noting the presence of palms and ties with earlier Paleocene floras at Tykbutak. evergreen dicotyledons with narrow leathery The late Eocene (partly possibly middle

52 COLLINSON & HOOKER: Paleogene vegetation Eurasia

Eocene) floras including those at Baki, Freshwater wetlands Zharlykap, Karashandak and Kundyzdy, are The changes in the forest vegetation are very different. They lack the dominants of the accompanied by a change in the coastal wet- previous floras, instead being dominated by land vegetation. Best documented in southern small leaved sclerophyllous elements inclu- England, the mangrove is replaced by a rich ding species of Myrica and oaks. Thermo- herbaceous marsh, dominated by Acrostichum philic elements have increased and include and Typha, including lakes with a rich aquatic palms and narrow-leaved laurels such as flora. The vegetation is likely to have Litsea. Late Eocene to early Oligocene transi- resembled the vegetation of the Florida tional floras (such as Shadiniskii/Shana Everglades today. Taxodiaceous trees were Graben, Romanovsk and Martuk) contain a also part of this wetland as in the earlier mixture of subtropical and warm temperate polar swamps but represented now by elements, indicating a mosaic of thermophilic Taxodium or a different extinct genus and mesophilic elements with a moderately Quasisequoia KUNZMANN (Kunzmann 1999, subtropical aspect (Makulbekov 1977, 1987; formerly named Sequoia couttsiae Heer and Zhilin 1989). Subtropical elements persist in common in England, Collinson & Cleal the early Oligocene but are lost in the late 2001b,c). Oligocene (Zhilin 1989) a comparable situ- ation to parts of central Europe. Vegetation overview This vegetation is obviously variable because Strongly thermophilic floras of its transitional nature. However, broadly it Four of the sites on the Oligocene map (Fig. is based upon mixed mesophytic forest vege- 3) seem to contain a rather high proportion of tation. This mixed broad-leaved deciduous evergreen thermophilic elements. Two of and evergreen forest vegetation has medium these, Tim and Pasekovo in central Russia, to high diversity. The Late Eocene Weissel- previously considered as Oligocene, might be ster Basin Zeitz Complex flora has 149 spe- Late Eocene or Eocene/Oligocene transition cies (Mai & Walther 1985) and the Oligocene (Vickulin 1999a, b) but all these ages have Haselbach flora is similar with 132 species been inferred from the floristic composition, (Mai & Walther 1978), both based on a com- which is currently being revised. The other bination of leaf and seed fossils. The late two sites are from the Caucasus and the inter- Eocene flora of Stare Sedlo has 135 species pretation of these has been taken from a based on the list of leaves (Knobloch et al. general article (Akhmetiev 1995) where the 1996). The Oligocene flora of Kundratice has early Oligocene of western Caucasus 89 species (Kvacek & Walther 1998) based (Akhmetiev 1995 p. 175) was described as mostly on leaves. This forest would therefore ‘inhabited by Lauraceae and evergreen contain a wide variety of habitats but fewer Fagaceae’. This area has marine intercala- than the multistratal, paratropical/subtropical tions but specific evidence for the dating is forests. not given. The interpretation is based partly The plants exhibit a variety of ecological on small floras in older literature (including strategies from colonisers to those adapted to woods and leaves) and partly on valuable, but climax forests. This vegetation contains fewer small, new collections. Therefore, all four of lianas and climbers and is inferred to be more these floras, which we have indicated as open than the paratropical/subtropical forests. ‘strongly evergreen’, may be subject to a The leaves vary from deciduous, medium- revised interpretation in future following furt- sized and thin with serrate margins, to ever- her research. green, larger and leathery with entire margins. Some seasonality is present, being induced either by changes in temperature or water


availability or both. The plants would produ- assemblage (Hably & Fernandez Marron ce a variety of fruits and seeds but with rela- 1998; Kvacek 1996; Mai 1994,1998). In con- tively few fleshy fruits and an increase in the trast, some floras e.g. those at Sarreal in component of large dry nuts such as those of Spain and some sites in Romania (Mera bei the Fagaceae, Juglandaceae and Betulaceae. Cluj, Girbou Cluj, associated with gypsum), Croatia (Mt Promina, Socka,) and Macedonia VEGETATION WITH SCLERO- (Pcinja Basin) have many sclerophyllous ele- PHYLLOUS ELEMENTS ments (Mihajlovic 1990, 1992, 1993; Hably & Fernandez Marron 1998). A typical leaf European floras flora rich in sclerophyllous elements from Several of the European late Eocene floras Ljiljanks Reka (Serbia) is illustrated by (e.g. Haring, Austria, leaf assemblage illu- Mihajlovic (1992, fig. 1). Mihajlovic (1992, strated by Mai 1995, fig. 150) and a larger fig. 2) also illustrates the complete range of number of the European Oligocene floras leaves of these sclerophyllous elements from contain elements where the leaves exhibit the western Balkan Peninsula. The propor- xeromorphy (modification to handle reduced tions of these sclerophyllous elements, and water availability). There are distinct taxa the expression of xeromorphy, seem to incre- with small and thick coriaceous leaves. In ase southwards in Europe. addition, representatives of typically meso- Wilde & Frankenhauser (1999) drew atten- morphic taxa may exhibit enhanced xeromor- tion to the risks of using nearest living relati- phy i.e. smaller and thicker leaves compared ve preferences of a relict endemic taxon like to other occurrences (e.g. oaks, laurels, the Comptonia, a deciduous plant which inhabits conifer Tetraclinis). Characteristic elements poor and acid soils in eastern and central are Zizyphus zizyphoides, Comptonia schran- USA, as indicative of dry habitats in the past. kii and Myrica longifolia, a variety of legu- In fact, fossil Comptonia-like leaves are mes and sometimes the conifer Tetraclinis found associated with fossil floras where all (Mihajlovic 1992,1993; Hably & Fernandez other elements indicate high humidity. Marron 1998; Hably et al. 2000; Kvacek Furthermore, xeromorphy is also known to be 1996; Kvacek et al. 2000). The communities associated with nutrient poor acid soils, as have been variously interpreted as: evergreen with modern Comptonia, some mire species laurophyllous forest, oak/laurel/Zizyphus of Myrica and with the southern hemisphere forest; mixed mesic-subhumid forest; xero- family Proteaceae, all of which have a very phytic shrub forest; sclerophyllous subhumid similar leaf morphology. A few fossils with vegetation or subxerophytic vegetation. xeromorphic leaves should certainly not be Palamarev (1989) discussed the nature and used to justify reconstruction of a sclerophyl- origin of this vegetation, recognising lous vegetation, even if some of their nearest Oligocene xerothermic centres on the Iberian, living relatives inhabit dry areas. Apeninian and Balkan peninsulas and in Nevertheless, it is reasonable to argue that some central European and French regions Comptonia schrankii is a sclerophyllous ele- leading ultimately to the evolution of sclero- ment because it characterises leaf assembla- phyllous paleomediterranean vegetation in ges dominated by sclerophyllous xeromorphic the Neogene. forms (Mihajlovic 1992, Hably & Fernandez The European floras seem to exhibit a very Marron 1998). wide variation in proportions of these sclero- Oligocene sites rich in sclerophyllous ele- phyllous elements. Some floras such as those ments occur in the Ebro Basin in Spain in Lausitz and Hungary have only one or two (Sarreal, Cervera and Tarrega – MP21-MP23 elements within an otherwise mixed meso- – Hably & Fernandez Marron 1998). A sedi- phytic forest or notophyllous evergreen forest mentological study (Anadón et al. 1989)

54 COLLINSON & HOOKER: Paleogene vegetation Eurasia

showed that these floras were preserved in a conditions and more open vegetation; and the series of lacustrine depositional sequences abundance of Cupressaceae pollen argued to which include playas and playa lakes with be an indicator of a drier climate. evaporites and carbonates (limestones and Quantitative data for all taxa were not repor- marls) and in which the megasequences may ted but it is clear that the herbaceous plants reflect fluctuating climate or possibly and flowering plant drier climate indicators Pyrenean tectonic activity. Some of the floras were very rare, as the samples were domina- (Sarreal) are preserved in strata interpreted as ted by bisaccate conifer pollen, Cupressaceae more or less perennial stratified lakes. The pollen and pteridophyte spores. Whilst some Ebro Basin was landlocked since the angiosperm pollen are underrepresented Oligocene (Vilena et al. 1996a) so that loss owing to their insect pollination strategy, of oceanic influence could also have affected grass pollen is wind dispersed and would be the local climate and sedimentation. Vilena et abundant if grass plants had been abundant. al. (1996a,b) documented the presence of The Sarral Formation samples also included a both large and small alluvial fans on the high proportion (49%) of taxa with megather- Iberian Margin of the Ebro Basin along with mal or mega/mesothermal affinities and many mud flats, carbonated lakes and saline lakes. of the taxa have nearest living relatives, They argued for dominantly tectonic controls which are trees. on the alluvial fan sedimentation rather than In southern England, sclerophyllous variations between humid and arid climates Palibinia-like foliage (see other areas below) but they did not exclude climate involvement occurs in the Bembridge Limestone in the shift from evaporitic to carbonate lacu- Formation and Bembridge Marls Member strine sedimentation. In a study of the late (), strata, which span the Oligocene to Late Miocene lacustrine sys- Eocene/Oligocene transition (Collinson et al. tems of the central Ebro basin Arenas & 1993b). The genus Raskya (Manchester & Pardo (1999) proposed a model where ephe- Hably 1997) also occurs within the meral rainfall on the Iberian margin resulted Bembridge Marls in the Insect Limestone and in playa lake settings. Laminated limestones it has recently been reported at the early were deposited at high water levels and nodu- Oligocene site of Céreste (Thiebaut 1999) in lar gypsum deposits at low water levels. laminated lacustrine limestones deposited Overall, sedimentological evidence tends to during water level fluctuations. Raskya also support the presence of intervals of water occurs at Kuclin (Czech Republic) and Obuda deficit in the Oligocene Ebro Basin. and Eger-Kiseged in Hungary (Thiebaut Palynological evidence from the Ebro 1999). The Bembridge Marls Insect Basin was interpreted by Cavagnetto & Limestone flora also contains Zizyphus zizyp- Anadón (1996) to indicate a change, in the hoides but otherwise has a wide variety of late Eocene, to a climate with a dry season. wetland elements and no other sclerophyllous In the early Oligocene Sarral Formation elements (Collinson et al. 1993b, Collinson (from which the macrofossils are derived) in progress). The Bembridge Marls floras evidence for this included: the incoming of were interpreted as an extensive marshland pollen whose nearest living relatives are her- (Collinson 1983b). Drier conditions were baceous plants (e.g. grasses, considered as one possible explanation for , Lamiaceae, Portulacaceae subtle differences between the Bembridge and Valerianaceae); the incoming of taxa such Marls and Bembridge Limestone floras, the as Mimosaceae, Mahonia and Daphne, Insect Limestone of the Bembridge Marls alongside those already present including being more comparable to the latter Ephedra and Chenopodiaceae, whose nearest (Collinson et al. 1993b). Insects from the living relatives were argued to suggest drier Insect Limestone (which contains the sclero-


phylls mentioned) were interpreted, after tap- In the early Oligocene they occur rarely in honomic study, by McCobb et al. (1998) as northern Europe and more commonly in the reflecting a primary subtropical to tropical south. In the south they dominate some sites forest subject to significant seasonal rainfall. (e.g. Sarreal, Spain and Macedonia) and they Open vegetation pollen types are rare to are important at others. Taking leaf floras, absent in pollen spectra, Ephedra type pollen palynofloras, and sedimentological evidence is present but no grasses have been reported into account it seems reasonable to regard the (Machin 1971, Collinson 1983b, Collinson et belt with common sclerophylls in southern al. 1993b). In contrast, insects from Céreste, Europe as vegetation adapted to a climate in particular a butterfly of the family with a slightly drier interval. In view of the Satyridae, (one isolated forewing) were sug- proportions of associated deciduous plants, gested to indicate spreading of grasses in the the drier interval may have been in winter early Oligocene of Europe (Pfretzschner (Mai 1998). It should be emphasised that we 1998). The theridomyid rodent are speaking of an overall humid, warm tem- Pseudoltinomys from Céreste, on the basis of perate climate with only slightly drier win- its gerbil-like locomotor adaptation, was sug- ters. gested to indicate a semi-arid open habitat (Schmidt-Kittler & Storch 1985). Kvacek and Floras of other areas Erdei (2001) reported the presence of xero- In China, Guo (1990) recognised a broad belt morphic foliage of Berberis (Berberidaceae) of distinctive vegetation with sclerophyllous in the Oligocene of south France (including elements in the central region in the Eocene. Céreste) but considered the overall vegetation The key floral element, commonly found in to be a mesophytic forest reflecting a warm every locality in this flora, is Palibinia. Other temperate to subtropical, subhumid climate. floral elements represent deciduous hard- Palynospectra from Toru Ajgyr (middle woods, characterised by thick coriaceous and Eocene) in Kyrgyzstan (Fortuna 1979, Erfurt microphyllous leaves with serrate margins, et al. 1999) are said to be dominated by xero- and evergreen trees, mostly with leathery phytes with grass pollen most frequent in all microphyllous leaves with entire margins samples but they also contain many trees (Guo 1990). Sedimentological indicators of including subtropical elements. Xerophytic drying out such as red beds, gypsum and hali- elements may be represented by chenopods, te are associated with the floras (Guo 1990). Artemesia, Myricaceae and Ephedra in the Palibinia also occurs in the Bembridge palynofloral list (percentages not stated). Limestone and Bembridge Marls floras of Complementary sedimentological, palyno- England (Eocene/Oligocene transitional stra- logical and macrofossil (especially foliar phy- ta) (Tao 1965, Zhilin 1989, Collinson et al. siognomic) studies from sites with varied 1993b); in late (?middle in earlier literature) proportions of sclerophyllous elements across Eocene floras at Badchyza in Turkmenistan Europe are necessary to improve our under- (a flora lacking Lauraceae and palms charac- standing of this vegetation. Palynological teristic of less sclerophyllous floras) and in processing and counting methods would need the Pcinja Ovce Polje group Basin on the to be fully standardised (including quantifica- borders of Serbia with the former tion of grains per sediment volume) and com- Yugoslavian Republic of Macedonia and parable facies (including a range of facies) Bulgaria (Mihajlovic 1993) also associated need to be examined for palynomorphs from with other sclerophyllous elements. The all areas. genus is also very important in the early Oligocene Turgai floras of western Summary Sclerophyllous elements occur Kazakhstan (Zhilin 1989) though it does not rarely in southern Europe in the late Eocene. extend into the late Oligocene and it is appa-

56 COLLINSON & HOOKER: Paleogene vegetation Eurasia

rently lacking in eastern Kazakhstan section on late Eocene to Oligocene mixed (Ashutas, Kiin-Kerish, Shiderty, Zhilin 1989, deciduous and evergreen forests. Makulbekov 1987). Mihajlovic (1993) argued that the vegetation containing Palibinia char- Vegetation overview acterised the Tethyan archipelago, with aridi- This vegetation is very variable depending on fication starting in the middle Eocene and the proportion of the sclerophyllous elements. reaching a maximum in the late Eocene and This overview is based on the sclerophyllous early Oligocene. belt in southern Europe. The vegetation has A thorough systematic revision of all medium to low diversity. Floras rich in scler- Palibinia records by a single specialist, exa- ophyllous elements tend to contain relatively mining cuticles where possible, is badly nee- few easily determined key taxa whilst others ded. Palibinia has been variously compared remain unidentified, rendering even a preli- with Proteaceae and Myricaceae. A relations- minary statement about species diversity dif- hip with the former seems unlikely in Europe ficult. Mihajlovic (1992) figured sixty-two on geographical grounds (Proteaceae being a named categories of leaves from all the floras southern hemisphere family) and other putati- in the western Balkan Peninsula. One of these ve European Proteaceae have been reidenti- may include several taxa of legumes. The key fied to other families (e.g. Kvacek & Erdei taxa include Zizyphus zizyphoides, Comptonia 2001). Myricaceae may also be excluded on schrankii, and Myrica longifolia combined cuticle and hair base detail on new collec- with a high proportion of legume leaflets and tions of British Palibinia-like specimens pods and sometimes the conifer Tetraclinis (Collinson et al. 1993b). Tao (1965) distin- and the enigmatic flowering plant Palibinia. guished Palibinia as having pinnatifid These floras are perhaps best characterised by coriaceous leaves, tertiary veins directed their physiognomic features, having large towards the sinus below, and one or two numbers of small (microphyllous or smaller), interstitial veins developed between the later- thick, coriaceous leaves with entire margins, al veins. There is a strong primary vein and a often narrowly elongate in shape (Mihajlovic secondary vein at 45-65o running straight to 1992, fig. 2; Hably & Fernadez Marron the apex with branching tertiary veins to the 1998). sinus below, sometimes with interstitial veins This vegetation is inferred, by comparison between the secondaries. Cuticle characters with modern analogues rich in sclerophyllous were not described. Palibinia is readily dis- elements, and on the basis of palynomorph tinguished from Comptonia, which, in con- evidence for some herbaceous plants, to have trast, has 2-3 secondary veins leaving the been more open forest or patchy forest (by main vein at angles of 60-90o (Wilde & comparison with the mixed mesophytic Frankenhauser 1999, Manchester 1999). forests) ranging to shrubland. We do not Floras with many sclerophyllous elements know of any direct fossil evidence for the sta- (small, narrow and thick leaved oaks and lau- ture of the plants such as might come from rels as well as Myrica) occur in northern and study of paleosols (rooting depth or size) central Kazakhstan from the middle Eocene or from associated wood floras (large or onwards, in western Kazakhstan in the late small diameter axes). Lianas and climbers Eocene (?possibly middle Eocene) and in the were few or lacking in this flora. Some of the Zaysan Basin in eastern Kazakhstan in the legumes may have been climbers on the basis late Eocene. Oligocene floras of Kazakhstan of the habits of some living representatives. also contain sclerophyllous elements inclu- The vegetation was probably mainly evergr- ding Palibinia as discussed earlier in this sec- een. Xeromorphy would reduce water loss tion (Zhilin 1989, Makulbekov 1987). Floras and enable leaves to be retained in drier inter- of Kazakhstan are discussed further in the vals. The climate is suggested to have been a


humid warm temperate climate with a slight- Paleocene of the northern hemisp- ly drier interval. Fruit resources would be here very variable dependent on the dominant (Fig. 1) taxa. Zizyphus (fossil fruit stones known from Calau) probably had thin leathery flesh whilst Vegetation distribution The distribution fossils show that Myricaceae had small dry map (Fig. 1) shows polar broad-leaved deci- nuts and legumes had dry pods with various duous forest in the high northern paleolatitu- sizes of dry seeds. Fruit resources would be des ranging from the northern-most land, at seasonal in their availability. This vegetation about 80o North, south to between 45 and 60o is therefore characterised by relatively tough North depending on the area. Important areas coarse browse of small leaves, at least some and sites include Alaska; Rarytkin Ridge, of which was probably available at low Kamchatka and Sakhalin in Russia; levels, and many dry fruits/seeds with tough Spitsbergen; Greenland; Axel Heiberg and protective coverings. Ellesmere Islands, Canada; Scotland and Ireland. Very similar floras are also known in DISTRIBUTION OF PALEOGENE Alberta and Saskatchewan (Canada) and in VEGETATION AND COMMENTS the Fort Union Formation of Wyoming ON ASSOCIATED MAMMALS (Montana and North Dakota, USA). ‘Mixed’ vegetation with some polar deciduous ele- Introduction ments but also other, often thermophilic, ele- Wolfe (1985) produced vegetation maps that, ments is mapped in southern England and as he emphasised (Wolfe 1985 p. 372), provi- France, Sakhalin, China, Mongolia and the ded only a partial picture and were painted Zaysan Basin (east Kazakhstan). Elsewhere with a broad brush. They showed vegetation in western Europe and eastwards to west belts on a global scale. The maps produced Kazakhstan the vegetation includes a high here plot vegetation at specific sites in proportion of evergreen thermophilic ele- Eurasia using the paleogeographic maps ments. This thermophilic vegetation is the being developed by Paul Markwick forerunner of the Eocene paratropical to subt- (Markwick et al. 2000) as a framework and ropical evergreen vegetation but it also inclu- benefiting from a wide variety of recent lite- des deciduous elements, which dominate furt- rature (See text and references in Table 1). her north. Unfortunately we are not aware of Occasional sites in North America are inclu- Eurasian records south of about 30o North, ded where they are critical for vegetation which would enable assessment of the wider reconstruction (e.g. Axel Heiberg) or to distribution of this thermophilic vegetation in represent phytogeographic similarity. The the Paleocene. picture is still only partial and the brush is A workshop discussion, reported in Boulter still too broad (see ‘criteria used here’) but & Fisher (1994 p. 391), and studies by Mai the data provide a general framework within (1987, 1989, 1991, 1995) indicate paleophy- which mammal faunas may be interpreted. togeographic subdivision into an Arctotertiary The reader should refer to the individual refe- Realm and a Boreotropical Realm consistent rences cited in the locality lists (Table 1) for with the polar deciduous floras and the more details of the vegetation at any one site or evergreen floras respectively. The former can region. More detailed location maps with be subdivided floristically into the Greenland additional sites may be found in Mai (1995), and Beringian Regions and the latter includes Tanai (1992), Takhtajan (1974, 1982), and the Gelinden Region and the Volgo-Mugojar Budantsev (1994b). Region (Mai 1991, fig. 2). The floras of China and Mongolia were not discussed by these authors but may represent another

58 COLLINSON & HOOKER: Paleogene vegetation Eurasia

Figure 1 Distribution map of the Paleogene floras which are listed in Table 1 (Paleocene). Palaeogeography from unpublished maps being produced by Paul Markwick (see Markwick et al. 2000).


realm or region, where some of the floras (e.g. Mongolia) are only slightly different Mammals in the Mongolian late Paleocene from those of the polar deciduous forests. The mammal faunas with the closest associa- These latitudinally controlled vegetation tion with polar deciduous forests in the late distribution realms, with distinct floral Paleocene are those from Mongolia and nort- Regions, seem to have been inherited from hern China (Inner Mongolia). The relevant the late . Late Cretaceous palyno- faunas of the Naran Member at Naran Bulak floral provinces show a northern and of the Gashato and Bayan Ulan sites need Aquilapollenites province and a southern to be summed to provide an adequate sized Normapolles province and also show distinc- assemblage to assess their ecological diversi- tive vegetation in the European area versus ty. Small dominate e.g. multitubercu- the Kamchatka area (Herngreen et al. 1996). lates, mixodontians, arctostylopids and a Latest Cretaceous leaf floras from Kamchatka pseudictopid. The largest were a wolf-sized and the Rarytkin Ridge and other high nort- mesonychid, a pig-sized dinocerate and a hern areas contain many elements also pre- cow-sized coryphodontid pantodont. sent in the Paleocene polar broad-leaved Available postcranial evidence indicates that deciduous floras and reflect the same vegeta- these were terrestrial (Archibald, Lucas in tion type (Herman 1993; Herman & Spicer Janis et al. 1998, Flerov 1967) or, in the case 1996, 1997; Spicer et al. 1994; Golovneva of the smaller animals (not well known), pro- 1996). bably mostly semiterrestrial (Kielan- Jaworowska & Gambaryan 1994, Li & Ting Dispersal routes Patterns of plant distribu- 1993, Cifelli & Schaff in Janis et al. 1998, tion are of three types; circumboreal, North Sulimski 1968). The only evidence of adapta- American/Asian and North American/ tion for tree-climbing ability belongs to the European (Manchester 1999). In the creodont Prolimnocyon, based on skeletal Paleocene there are many circumboreal remains from North America (Gebo & Rose plants. Some of these distributions may have 1993), although Lucas (in Janis et al. 1998) been inherited from the Cretaceous (e.g. for has hinted that the small pantolambdodontoid Nordenskioldia [see Crane et al. 1991 for pantodonts may have been tree-dwelling. A map] and Nyssidium plants). In contrast, significant number of the mammals from this others, e.g. that of the tree Palaeocarpinus of fauna, including the smaller ones, were brow- the Betulaceae (Crane 1981, Chen et al. sing herbivores. This paucity of tree-dwellers 1999, Golovneva 2002) and the aquatic herb and large numbers of herbivores somewhat Haemanthophyllum (Golovneva 1997), seem resembles patterns obtained from modern to have been achieved in the Paleocene. In semi-deciduous forests (Andrews et al. the Paleocene there are a number of taxa (at 1979), largely consistent with reconstructed least 6 genera including small aquatic plants vegetation. like Quereuxia and trees of the Platanaceae and other reconstructed woody plants like Eocene of the northern hemisphere Amersina of the Cornales (Manchester et al. - polar broad-leaved deciduous 1999)) shared between Asia and North forests (Fig. 2) America indicating dispersal via Beringia. There are also taxa, such as other trees of Vegetation distribution In the Eocene the Platanaceae (Platanites) and trees of the polar deciduous forest is still represented at Fagaceae (Fagopsiphyllum), which occur in high northern latitudes including sites on North America and Europe but not in Asia, Spitsbergen, Axel Heiberg and Ellesmere suggesting dispersal via North Atlantic land Islands (Kvacek et al. 1994, Golevneva 2000, bridges (Manchester 1999). McIver & Basinger 1999). The flora of the

60 COLLINSON & HOOKER: Paleogene vegetation Eurasia

Figure 2 Distribution map of the Paleogene floras which are listed in Table 1 (Eocene). Palaeogeography from unpublished maps being produced by Paul Markwick (see Markwick et al. 2000).


Lena River in northern Russia (Budantsev absence of insectivores may represent a col- 1994a) is similar. Rather different floras lecting bias as relatively little screenwashing occur in the Rarytkin range at the Podbazovyi was undertaken. Lower latitude faunas of Creek on the Taljain Creek tributary in the equivalent age in North America have a gre- Pravataljainskaya Formation (Akhmetiev & ater diversity and abundance of frugivores, Samsonenko 1997) and at Tastakh (Budantsev e.g. in the form of hyopsodontid condylarths 1994a). The extent to which these indicate and notharctid primates notable absentees different vegetation requires more detailed from Ellesmere. systematic and physiognomic comparison. Eberle & Storer (1999) suggested that The floral lists do suggest that they indicate a Brontotheres (perissodactyls - as hindgut fer- more thermophilic vegetation. menters) survived in the 2-3 months of winter darkness by feeding on litter (leaves fruits Mammals associated with Eocene polar and seeds etc.) and bark, branches, twigs etc., broad-leaved deciduous forests In the late being able to handle low quality food provi- Early Eocene, mammal faunas are known ded it is available in high quantity. The other from Ellesmere Island and one species from large herbivores & Coryphodon Axel Heiberg (West & Dawson 1977, 1978, would also have been hindgut fermenters but McKenna 1980, Eberle & Storer 1999). The have low-crowned bilophodont teeth compa- faunas so far published contain no truly arbo- rable to those of modern which live in real mammals (e.g. in the form of primates). very low seasonality tropical forests, bulk Tree-dwellers were potentially scansorial feeding on leaves of evergreen trees. As there (post-cranials not known for confirmation, is no modern analogue for a warm dark forest extrapolation based on partial skeletons, ske- (Basinger et al. 1994, McIver & Basinger letal elements elsewhere of related taxa, spe- 1999) it is difficult to predict how deciduous cimens not yet described), including rodents leaves would behave and how much nutrition (Szalay 1985), paromomyids and plagiomen- would remain in the litter. Clearly, the fossili- ids. The last two groups may have been ferous levels prove that huge thicknesses of gliders but this is controversial (Beard 1990, litter accumulated and persisted. The large- Runestad & Ruff 1995, Szalay & Lucas leaved dicotyledons in the flora might then 1996). Scansorial mammals depend less on have been fed upon from the litter by these trees than truly arboreal mammals in that mammals. The ‘upland’ broad-leaved deci- they do not need the aerial walkways (over- duous forests contained evergreen conifers lappping branches, lianas etc.) afforded by a (McIver & Basinger 1999, Basinger et al. closed canopy forest. 1994) and these may have contributed to the There are relatively few frugivores in the food of herbivore browsers during the winter fauna. The multituberculate Neoplagiaulax is darkness. However, there is an issue that the likely to have had a mixed diet of insects and large hoofed browsers would probably not dry fruit/seeds (Krause 1982). Paromomyids have been able to reach much of the evergr- are usually attributed a mixed fruit/exudate een conifer browse if the ancient trees were diet (Beard 1990, McKenna 1980). The of comparable stature to their nearest living rodents would have had a mixed fruit/insect relatives. This browse would, nevertheless, diet and the dawn horse ""? a have been available to the climbing plagio- mixed fruit/leaf diet. On the other hand the menids. Dentally, however, these seem unsui- diverse plagiomenids and the abundant ted to masticating conifer needles and hiber- Hyrachyus (stem rhino) and Coryphodon nation might have been an alternative option (pantodont) and probably Lambdotherium for them. (brontothere) would have had a near exclusi- ve leaf-browsing diet. It is possible that the

62 COLLINSON & HOOKER: Paleogene vegetation Eurasia

typical of dry open habitats today. They have Eocene of the northern hemisphere had relatively high-crowned teeth since their - paratropical/subtropical evergreen oldest undoubted records in the Eocene and forest with sclerophyllous elements this can be attributed to an abrasive herbivo- rous diet. The Eocene vegetation of Mongolia Vegetation distribution There are several has not yet been studied (Makulbekov 1987). areas where paratropical to subtropical vege- However, the middle Eocene northern tation contains sclerophyllous elements at dif- Chinese lagomorph records overlap with flo- ferent times in the Eocene. The first area is in ras indicating sclerophyllous/microphyllous the late Eocene of southern Europe east to forest vegetation. Ukraine and the Caucasus. These sclerophyl- In contrast, typical bunoselenodont anthra- lous elements are those which become much cotheriid artiodactyls are almost entirely res- more common in southern Europe in the tricted in the Irdinmanhan Asian land mam- Oligocene, especially Zizyphus zizyphoides, mal age (Lutetian) to the extreme south of Comptonia schrankii and other Myricaceae. China and Burma with only one record in The second area is in the late (or in places central China. However, more reach the cen- middle and late) Eocene of Kazakhstan, tral and northern regions in the Sharamu- Turkmenistan, central Iran and Kyrgyzstan runian (Bartonian) but were gone again in the (the last from palynological evidence only). late Eocene Ergilian. These are likely to have The third area is in the Eocene of central had a mixed frugivorous/herbivorous diet China where precise dates are lacking. which might have restricted them from Arguably these form a discontinuous belt moving north into the central, possibly drier, between about 30-50o North. It must be emp- region except during the slight late Middle hasised that the mere presence of sclerophyl- Eocene climatic amelioration. Brontotheriid lous elements in these floras may not be suf- perissodactyls occur throughout the area but ficient to justify reconstruction of a drier cli- show a greater diversity in both the north and mate. If the climate was drier we are spe- south than in the central region of China. aking of a slightly drier interval within an Although no account here is taken of tapho- otherwise humid setting. Palibinia, the key nomic/collecting bias, this pattern does per- sclerophyllous element in the Chinese floras, sist through the middle and late Eocene after is also recorded in Turkmenistan and in which extinction intervened. It is difficult to Eocene/Oligocene transitional and Oligocene account for this but inasmuch as they became floras further west (see vegetation with scler- extinct at the end of Eocene the vegetation ophyllous elements). types in the Eocene (e.g. with sclerophyllous elements) that heralded those of the Mammals in China In China a central belt Oligocene may have been suboptimal for with sclerophyllous elements, and hence pos- them. On the basis of the almost total absence sibly a drier interval, is bounded to the south of any primate, plesiadapiform or bat in the by evergreen paratropical rain forest and to Eocene of the Mongolian plateau except the the northeast by subtropical evergreen forest single early Eocene primate species Altanius (Guo 1990). Information on Chinese orlovi, Meng and McKenna (1998) concluded Paleogene mammalian faunas is obtained that this implied the absence of dense forest from the comprehensive review by Russell & in the region. In contrast, a diversity of pri- Zhai (1987). Lagomorphs are almost entirely mates and plesiadapiforms (all assumed tree- restricted to Mongolia and northern Chinese dwelling) are known from the central Chinese localities, very few straying southwards region (Wutu: Beard & Wang 1995, Yuanqu: (Russell & Zhai 1987, Zhang et al. 2001). Beard et al. 1996, Shanghuang fissure fil- Lagomorphs are ground dwellers and are lings: Beard et al. 1994) suggesting treed


habitats consistent with the plant evidence. sites where the literature uses the term ‘subt- Eocene of the northern hemisphere ropical’ rather than paratropical, and empha- - paratropical/subtropical evergreen sises ‘with temperate elements’, have been forests (Fig. 2) distinguished on the map (Fig. 2, hexagons). The extent to which these floras differ, and Vegetation distribution The most strongly the precise nature of the differences, needs evergreen thermophilic category of this vege- further detailed study. In several areas of tation (Fig. 2 stars) is present in a belt Europe, and in Kazakhstan, Kamchatka and through western, central and eastern Europe Japan trends during the Eocene (either and extending into Ukraine. Strongly ther- through independently dated floras or super- mophilic vegetation also occurs as a belt in posed floras) can be recognised with an southern China and as more isolated records increase in the component of temperate deci- in Japan and Kamchatka. The high latitude duous elements (or sclerophyllous elements) record in Kamchatka is matched by a similar with time (see Oligocene floras below). The high latitude record (the Kultieth flora) in subtropical/with temperate elements vegeta- Alaska. The high latitude Tastakh flora in tion on the Eocene map, therefore, encompas- northern Russia contains a number of ther- ses both spatial and temporal variation in mophilic elements (Budantsev 1994a) but the Eocene vegetation, which cannot be more Lena River flora is warm temperate precisely plotted without improved indepen- (Budantsev 1994a) and more similar to the dent dating and further detailed comparisons Eocene polar deciduous forest floras and was of the floras across Eurasia. Furthermore, it discussed earlier. Unfortunately we have not should be noted that the European region, been able to locate an original reference for with strongly evergreen thermophilic vegeta- the flora of Dzilon-Sise. The Rarytkin Range tion in the Early and early Middle Eocene, Podbazovyi Creek flora is described as ther- also contains sites with more deciduous tem- mophilic mesophilic conifer-broadleaved perate elements in the late Middle and Late deciduous forest vegetation (Akhmetiev & Eocene. These could not all be plotted on a Samsonenko 1997). It lacks the Trochoden- single map. droides leaf types characteristic of Eocene polar deciduous forests. Tanai (1992, fig. 5a) Dispersal routes Dispersal routes are likely mapped mixed Coniferous forest at high nort- to have been present via the North Atlantic hern latitudes in the middle Eocene based on and North Pacific. The Middle Eocene the floras of Rarytkin, Tastakh and Dzilon- (44Ma) Clarno flora of western North Sise (Tanai 1992, fig. 1 map). This does not America (Fig. 2) shares 24% genera with seem to be an appropriate interpretation of Europe (20% genera and 10% species with the floral lists. The variations in far eastern London Clay flora alone) (Manchester 1994). and northern Russian high latitude floras may This clearly indicates interchange prior to the be due to age differences but we are not Middle Eocene. A dispersal route via aware of independent dating for these floras. Beringia is possible but would need confir- Further research is required to compare these mation from records in Asia. Only a very few floras more closely with one another and with newly appearing taxa are shared between the Eocene polar deciduous floras, especially Asia and North America (Manchester 1999) from a physiognomic perspective. but fruit and seed floras comparable to those The majority of the dominantly evergreen of Clarno and the London Clay are not vegetation in Europe occurs around the ther- known in Asia. In any case the Turgai seaway mal maximum in the early and early middle may have intervened. Exchange via the North Eocene (evidence from independent dating). Atlantic (via Greenland and Iceland using eit- Elsewhere the dating is not so precise. Those her the Thule or de Geer Route) is more like-

64 COLLINSON & HOOKER: Paleogene vegetation Eurasia

ly (Manchester 1999, Tiffney 2000). However there is a gradual trend from Eocene paratro- the high latitude floras in the North Atlantic pical/subtropical evergreen forest vegetation area give little hint of the presence of ther- through vegetation with an increasing propor- mophilic taxa at the time when dispersal tion of temperate deciduous elements resul- should have been occurring (Manchester ting in the Oligocene in establishment of 1999, McIver & Basinger 1999). Indeed, broad-leaved mixed deciduous and evergreen more thermophilic floras seem to occur in the forests across Eurasia between about paleola- Beringian region (e.g. at Tastakh and at titudes 55 and 40o North (Fig. 3). Chemurnaut Bay, Kamchatka) although these Four sites in central Russia (Vickulin require more precise dating (see earlier). 1999a,b) and Caucasus (Akhmetiev 1995) may reflect more thermophilic vegetation but Associated mammals in Europe The ecolo- these require further study (see earlier). gical diversity patterns of the mammals from Floras in Hungary indicate retention of a Early Eocene sites in Europe match quite clo- strong thermophilic signal into the early sely with those of evergreen forests in Oligocene but its loss by the late Oligocene. Borneo (Hooker 1998). The ecological diver- There seems to be only one flora (Sergeya in sity of mammals suggests that in the middle Russia) from high northern paleolatitudes Eocene, after the thermal maximum, there is (Tanai 1992). We have not been able to locate an increase in the number of larger mammals recent literature on this site and the age is and of herbivore browsers (with the low uncertain (Takhtajan 1982). Three floras in crowned bilophodont tooth type found today China indicate the presence of tropical/subt- in tapirs, Hooker 2000). By the late Lutetian ropical forests in the south and warm tempe- there was a slight increase in hypsodonty in rate deciduous broad-leaved forests in the perissodactyls (Legendre et al. 1991) indica- north (Guo 1990,1993). The two Japanese ting coarser browse and a slight further incre- sites are late Eocene to early Oligocene and ase in larger mammals suggesting slightly are referred to as having warm temperate more open forests; however, many fruit- deciduous floras. eaters persisted. These changes persisted. The The European floras seem to exhibit very bilophodont tooth types were lost by the end wide variation in the proportions of sclero- of the middle Eocene (Collinson & Hooker phyllous elements. Some floras have only one 1987). The late Eocene saw a further increase or two sclerophyllous elements within an in hypsodonty not just in perissodactyls but otherwise mixed mesophytic forest assembla- especially in rodents such as theridomyids ge (e.g. Lausitz) or a notophyllous evergreen (Vianey-Liaud 1991) and a reduction in frugi- forest (e.g. Hungary). In contrast some floras, vorous types and specialisations for seed- e.g. those at Sarreal in Spain and sites in eating in glirids, demonstrated by gnawed Romania, Croatia and the former Yugoslavian Stratiotes seeds (Collinson & Hooker 2000). Republic of Macedonia, have many sclero- There are also more larger mammals and fewer phyllous elements. The proportions seem to arboreal forms. All these changes are consis- increase southwards. Evidence from leaf flo- tent with the gradual change from the evergr- ras, palynology and sedimentology seems to een thermophilic forests to the mixed deci- indicate that the belt with common sclero- duous and evergreen forests of the Oligocene. phylls in southern Europe was, at least in places, a vegetation adapted to a humid warm Oligocene of the northern hemisp- temperate climate but with a slightly drier here - broad-leaved mixed deci- interval, probably in winter. duous and evergreen forests (Fig. 3) In Kazakhstan the floras also contain scler- ophyllous elements. The sclerophyllous ele- Vegetation distribution As noted above ment Palibinia characterises the early


Figure 3 Distribution map of the Paleogene floras which are listed in Table 1 (Oligocene). Palaeogeography from unpublished maps being produced by Paul Markwick (see Markwick et al. 2000).


COLLINSON & HOOKER: Paleogene vegetation Eurasia

Oligocene floras of western Kazakhstan but it (hopping) (Vianey-Liaud 1991). Furthermore, is lacking in floras further east in the Zaysan hindgut-fermenting perissodactyls diminish Basin. In western Kazakhstan subtropical ele- while foregut-fermenting artiodactyls increa- ments persist in the early Oligocene but are se (Meng & McKenna 1998, Janis 1989) sug- lost by the late Oligocene (Zhilin 1989). gesting use of high quality but rare/patchy browse rather than abundant low quality Dispersal routes A number of identical or browse. These trends were beginning in the closely related taxa are shared between European late Eocene when mammalian eco- Europe and North America in the Oligocene logical diversity is indicative of drier more (e.g. species of the conifer Tetraclinis and the open habitats in southern France than in sout- flowering plants Cercidiphyllum, Mahonia, hern England (Hooker 1992, Franzen 1968). Cedrelospemum, Catalpa and Saporta- Moreover, northward dispersal of certain spermum (Hably et al. 2000). These indicate mammalian taxa suggests northward expan- that exchange must have occurred during or sion of such habitats (Hooker et al. 1995). prior to the Oligocene and although some e.g. Cedrelospermum seem to have established India and Southeast Asia this range in the Eocene, others may have done so in the Oligocene (e.g. Mahonia and Vegetation distribution Indian Paleogene Saportaspermum) (Manchester 1999, Hably floras are best known from the Deccan et al. 2000). In addition there are a number of Intertrappean series (latest Cretaceous/ circumboreal genera in the Oligocene (Hably Paleocene transition). These floras, which are et al. 2000), which may have attained this preserved in cherts, are dominated by fossil distribution prior to or during the Oligocene. wood along with some fruits and seeds. Dispersal could have been via Beringia or These are hard to compare with floras from North Atlantic Bridges, though the Turgai/ Eurasia. Some leaf floras are also known but west Siberian seaway intervened in the I am not aware of any work with a thorough Eocene and it was not until the Oligocene systematic comparison of these with Eurasian that there was a full land connection between leaf floras. Study of wood and leaf physio- Europe and Asia. The extent to which the gnomy, combined with evidence from nearest land bridges were available, is discussed by living relatives of woods, leaves and palyno- Tiffney (2000). The North Atlantic land morphs, suggests that warm, humid tropical bridge would seem to have been broken by to subtropical, evergreen to semi-evergreen the beginning of Chron C24N (early Eocene) forest vegetation existed in India through the (Hooker 1998). Further comparisons at spe- Paleogene (e.g. Kar & Sharma 2001; Morley cies level using reconstructed whole plants 2000; Bande & Chandra 1990; Prakash 1960, will be necessary to establish phytogeogra- 1973; Singh & Sarkar 1990; Kumar 1994; phic similarities with more precision. Awasthi & Mehrotra 1995; Guleria & Srivastava 2001). This was probably similar Mammals indicative of drier habitats to wet evergreen forests in the area today Evidence from the paleoecology of mammals (Guleria & Srivastava 2001) e.g. the western for drier more open habitats in the Oligocene Ghats (Banda & Chandra 1990). Similar has been put forward for both Europe and the vegetation is indicated by palynological evi- Mongolian Plateau. This has involved parti- dence from Java for the Middle Eocene cularly rodents, which, in both areas, undergo (Lelono 2000) and generally in Southeast significant increases in molar crown height Asia (Morley 2000) and also by the rather (Vianey-Liaud 1991, Meng & McKenna sparse macrofossils from Southeast Asia 1998) and several European taxa have enlar- (Bande & Prakash 1986). ged auditory bullae and ricochetal locomotion The family and generic affinities (macro-



fossils) of the Southeast Asian Paleogene flo- Lelono (2000) in his study of the Middle to ras (Bande & Prakash 1986) differ from those Late Eocene coastal sequences of Java recor- of the European floras (Mai 1995). It must ded over 300 different palynomorphs with 80 also be emphasised that the European ther- distinct taxa occurring at over 3% i.e. impor- mophilic floras do not represent modern tant in the flora. Collinson et al. (1981) and Southeast Asian rain forest (Collinson 1983a, Boulter & Hubbard (1982) studying the 2000b, 2001a) although many fossils with Eocene coastal sequences of England, recor- near living relatives in these rain forests do ded 80-100 distinctive and important form occur in the European Paleogene. Notably, in genera on which they based their statistical the Paleogene of Europe, dipterocarps are analyses. (The total number of form species represented only in the London Clay flora was not documented). Thiele-Pfeiffer (1988) and only by one twig, and the rest of the documented 173 form species from the early wood flora (Poole 2000) differs from that of Middle Eocene of Messel, Germany, whilst the Paleogene of India and Southeast Asia. It Nickel (1996) documented 190 from the late should, however, be noted that dominance of Middle Eocene of Eckfeld, Germany, both dipterocarps did not characterise even the representing isolated lake basins. Harrington Southeast Asian rainforests until the Neogene & Kemp (2001) and Harrington (2001) docu- (Morley 2000). A further difference is the mented 164 form species in the Paleocene apparent absence of epiphytes (Collinson /Eocene boundary interval of the United 2000b, 2001a) though this may be due to tap- States Gulf Coast paratropical forests whilst honomic factors. At least some lignites in Wing & Harrington (2001) recorded only 93 Java have been shown to originate from in the comparable interval in the continental palm-dominated mires based on palynological interior Bighorn Basin, Wyoming, USA. evidence (Lelono 2000, Morley 2000) and These figures tend to suggest that northern palms are abundant and diverse in the hemisphere Eocene evergreen thermophilic Paleogene floras of both India and Southeast forests had similar levels of diversity to the Asia (references above). Eocene Southeast Asian forests in their ‘com- Modern Southeast Asian tropical rainforests mon’ constituents (around 80-100 form spe- are characterised by very high diversity (e.g. cies) whilst the overall diversity may have Morley 2000) and it is interesting to consider been lower, totalling up to 200 rather than the past situation. The very different nature of over 300 taxa. These figures represent a very plant macrofossils and facies in the Southeast inadequate glimpse into this topic and further Asian and Indian Paleogene floras makes it research by a single individual applying a impossible to compare diversity from this uniform taxonomic approach and using samp- evidence. Palynological data offer the best les from comparable facies (ideally several opportunity for comparison but this is also facies) is essential to provide a more mea- severely hampered by different taxonomic ningful comparison. approaches including different nomenclature systems in Southeast Asia and Europe for his- Dispersal The palynological record provides torical reasons. During the course of PhD strong evidence for dispersal of plants from work by Lelono (2000), Morley, Collinson India into Southeast Asia by the Middle and Lelono noted a number of pollen types Eocene (Morley 1998, 2000, Lelono 2000). that seem to be very similar to, if not the Lelono (2000) and Morley (1998, 2000) same as, those from the European Paleogene reported at least 15 distinctive palynomorphs, (see also Harley & Morley 1995). Further which first appear in the Paleocene or early work is needed by an experienced palynolo- Eocene of India (e.g. Frederiksen 1994, Kar gist making direct comparisons of material & Sharma 2001), and are also found in the from both areas. Middle Eocene of Southeast Asia. These


COLLINSON & HOOKER: Paleogene vegetation Eurasia

include taxa whose near living relatives are 2001). Plant fossil evidence (above) indicates several kinds of palms (pollen of that the mammalian dispersals would have Proxapertites, Longapertites, Quilloni- occurred within humid thermophilic evergr- pollenites, and the peat-former Palmae- een forests, although disturbance to this vege- pollenites kutchensis) and members of the tation should be predicted as a result of tecto- Bombacaceae (Lakiapollis ovatus), Sapin- nic activity (e.g. vulcanicity). daceae and possibly Proteaceae (Beau- preadites), Linaceae, Olacaceae, Polygalaceae ACKNOWLEDGEMENTS and Ctenolophonaceae and one spore type. We would like to thank Hans de Bruijn and Diverse arrays of Spinizonocolpites (= Nypa the conference organisers for the invitation pollen) species are present in the Paleocene and opportunity for MEC to present this key- of Indo-Pakistan but by the Eocene only one note address on the occasion of the celebra- survives (Frederiksen 1994). The occurrence tion of the 70th birthday of Hans de Bruijn. of only one (although variable) type (S. echi- We would also like to thank Paul Markwick natus), in Southeast Asia (Morley 2000, for allowing us to use his unpublished pale- Lelono 2000) suggests that dispersals from ogeographic maps and for his considerable India did not reach Southeast Asia in the help in providing those maps in various for- Paleocene. Frederiksen (1994) also argues mats. Robert Hall provided very helpful dis- that the occurrence of Normapolles pollen in cussion of the India/Southeast Asia region as the late Paleocene of Pakistan may be eviden- well as help with paleogeography in general. ce for sporadic filtered dispersals from We thank James Mcgrow for drafting the fos- Eurasia to Indo-Pakistan at this time. sil flora sites onto the paleogeographic maps The paleogeography of the India and and for his patience during several phases of Southeast Asian areas has been studied by revision. MEC would like to thank numerous Hall (1998, pers comm. 2001) who argues paleobotanists for helpful discussion over that the Indian/Asian collision had taken many years, especially those who participated place by the end of the early Eocene (50Ma) in workshop discussions reported in Boulter implying the earlier close proximity of land. & Fisher (1994) and those with whom she Roger et al. (2000) also argue in favour of regularly exchanges reprints without which India’s initial impact "well before" 48 Ma on this article could never have been written. the basis of overthrusting already in progress Guo Shuangxing, Engin Ünay, Funda Akgun, at that time. Rowley (1996) presented eviden- Alexei Herman, Gary Nichols and Norm ce for well constrained dating of the collision Macloed are thanked for various helpful starting in the late Ypresian in the western advices. Zanskar-Hazara region of the Himalayan region. Areas of land in Greater India and REFERENCES Greater Asia are largely speculative (Hall Akhmetiev, M.A., 1995 – Paleobotanical data on the 1998), but supported, at least as island areas, geological age of the Greater Caucasus – Paleonto- by early Eocene mammalian faunas from logical Journal 29 (1A): 172-181 Pakistan (Gingerich et al. 1997, Clyde & Akhmetiev, M.A. & Samsonenko, V.L., 1997 – New Khan 2000). India has been cited (Krause & plant species from the Eocene of the Central Rarytkin Maas 1990) as one possible source of the Range, Northern Koryakia – Paleontological Journal mammals that spread rapidly across the nort- 31 (2): 215-224 hern hemisphere at the beginning of the Anadón, P., Cabrera, L., Colldeforns, B. & Saez, A., Eocene in what is termed the Mammalian 1989 – Los sistemas lacustres del Eoceno superior y Dispersal Event. Mammalian dispersal from Oligoceno del sector oriental de la Cuenca del Ebro – central Asia into the Indian subcontinent at Acta Geologica Hispanica 24: 205-230 this time has also been proposed (Maas et al. Andrews, P., Lord, J.M. & Nesbit Evans, E.M., 1979 –



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during the Middle Eocene-early Oligocene of the significance in the context of co-eval British Tertiary eastern Ebro Basin, northeast Spain – Review of floras – Tertiary Research 16: 175-202 Palaeobotany and Palynology 92: 281-305 Collinson, M.E., 1999 – Evolution of angiosperm fruit Cernjavska, S., Palamarev, E. & Petkova, A., 1988 – and seed morphology and associated functional biolo Micropaleobotanical and macropalaeobotanical gy: status in the Late Cretaceous and Paleogene –in: characteristics of the Paleogene sediments in Hvojna Kurmann, M.H. & Hemsley, A.R. (eds.) – The evolu- Basin (Central Rhodopes) – Paleontologiya, tion of plant architecture – pp. 331-357, Royal Stratigrafiya I Lithologiya, Sofia 26: 26-36 Botanic Gardens, Kew Chandrasekharam, A., 1974 – Megafossil flora from the Collinson, M.E., 2000a – Fruit and seed floras from the Genesee locality, Alberta, Canada – Palaeonto- Paleocene-Eocene transition and subsequent Eocene graphica B 147:1-41 in southern England: comparison and palaeoenviron Chen Zhi-Duan, Manchester, S.R. & Sun Hai-Ying, 1999 mental implications – GFF 122: 36-37 – Phylogeny and evolution of the Betulaceae as infer Collinson, M.E., 2000b – Cenozoic evolution of modern red from DNA sequences, morphology and palaeo- plant communities and vegetation – in: Culver, S.J. & botany – American Journal of Botany 86: 1168-1181 Rawson, P.F. (eds.) – Biotic response to global chan- Clyde, W.C. & Khan, I.H., 2000 – Tectonic and bio- ge the last 145 million years – pp. 223-243, geographic implications of the Ghazij Formation Cambridge University Press, Cambridge (lower Eocene), Baluchistan Province, Pakistan – Collinson, M.E., 2001a – Rise of modern land plants and GFF 122: 34-35 vegetation – in: Briggs, D.E.G. & Crowther, P.R. Collinson, M.E., 1983a – Fossil plants of the London (eds.) – Palaeobiology II – pp. 113-116, Blackwells, Clay – Palaeontological Association Field Guides to Oxford Fossils Number 1, Palaeontological Association, Collinson, M.E., 2001b – Cainozoic ferns and their London distribution – Brittonia 53: 173-235 Collinson, M.E., 1983b – Palaeofloristic assemblages Collinson, M.E., 2002 - Cainozoic fern ecology – and palaeoecology of the Lower Oligocene Review of Palaeobotany and Palynology 119: 51-68 Bembridge Marls, Hamstead Ledge, Isle of Wight – Collinson, M.E., Boulter, M.C. & Holmes, P.L., 1993a – Botanical Journal of the Linnean Society 86: 177-225 Magnoliophyta (‘Angiospermae’) – in: Benton, M.J. Collinson, M.E., 1986 – The use of modern generic (ed.) – The fossil record 2 - pp.809-841, Chapman & names for plant fossils – in: Spicer, R.A. & Thomas, Hall, London B.A. (eds.) – Systematic and Taxonomic Approaches Collinson, M.E. & Cleal, C.J., 2001a – The palaeobotany in Palaeobotany – pp. 91-104, Systematics of the Paleocene and Paleocene-Eocene transitional Association Special Volume 31, Clarendon Press, strata in Britain – in: Cleal, C.J., Thomas, B.A., Oxford Batten, D.J. & Collinson, M.E. (eds.) – Mesozoic to Collinson, M.E., 1990 – Plant evolution and ecology Tertiary Palaeobotany of Great Britain, Geological during the early Cainozoic diversification – Advances Conservation Review Series No 22 – pp. 157-184, in Botanical Research 17: 1-98 Joint Nature Conservation Committee, Peterborough Collinson, M.E., 1992 – Vegetational and floristic chan- Collinson, M.E. & Cleal, C.J., 2001b – Early and early ges around the Eocene/Oligocene boundary in west- Middle Eocene (Ypresian-Lutetian) palaeobotany of ern and central Europe – in: Prothero, D.R. & Great Britain – in: Cleal, C.J., Thomas, B.A., Batten, Berggren, W.A. (eds.) – Eocene-Oligocene climatic D.J. & Collinson, M.E. (eds.) – Mesozoic to Tertiary and biotic evolution – pp. 437-450, Princeton Palaeobotany of Great Britain, Geological University Press, Princeton Conservation Review Series No 22 – pp. 187-226, Collinson, M.E., 1993 – Taphonomy and fruiting biology Joint Nature Conservation Committee, Peterborough of recent and fossil Nypa – Special Papers in Collinson, M.E. & Cleal, C.J., 2001c – Late middle Palaeontology 49: 165-180 Eocene and early Oligocene (Bartonian-Rupelian) Collinson, M.E., 1996 – Plant macrofossils from the and Miocene palaeobotany in Britain – in: Cleal, C.J., Bracklesham Group (Early and Middle Eocene). Thomas, B.A., Batten, D.J. & Collinson, M.E. (eds.) Bracklesham Bay, West Sussex, England: review and – Mesozoic to Tertiary Palaeobotany of Great Britain,



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myrtaceous affinity from Pasekovo, Middle Russian Wilde, V. & Frankenhauser, H., 1998 – The Middle Upland, southern European Russia – Botanical Eocene plant taphocoenosis from Eckfeld (Eifel, Journal of the Linnean Society 131: 65-98 Germany) – Review of Palaeobotany and Palynology Vickulin, S.V., 1999b – The Eocene and early Oligocene 101: 7-28 floras of the Russian plain and their relation to the Wilde, V. & Frankenhauser, H., 1999 – Comptonia-like palaeofloras of central Europe – Acta Palaeobotanica leaves from the German Middle Eocene – Acta Supplement 2: 429-445 Palaeobotanica Supplement 2: 447-463 Villena, J., Pardo, G., Pérez, A., Munoz, A. & Gonzalez, Wing, S.L. & Harrington, G.J., 2001 – Floral response to A., 1996a – The Tertiary of the Iberian margin of the rapid warming at the Paleocene/Eocene boundary and Ebro basin: sequence stratigraphy – in: Friend, P.F. & implications for concurrent faunal change – Paleo- Dabrio, C.J. (eds.) - Tertiary basins of Spain – pp. 77- biology 27: 539-563 82, Cambridge University Press, Cambridge Wolfe, J.A. 1977 – Paleogene floras from the Gulf of Villena, J., Pardo, G., Pérez, A., Munoz, A. & Gonzalez, Alaska region – Professional Paper of the United A., 1996b – The Tertiary of the Iberian margin of the States Geological Survey 997: 1-108 and plates Ebro basin: paleogeography and tectonic control – in: Wolfe, J.A., 1985 – Distribution of major vegetational Friend, P.F. & Dabrio, C.J. (eds.) - Tertiary basins of types during the Tertiary – in: Sundquist, E.T & Spain – pp. 83-88, Cambridge University Press, Broecker, W.S. (eds.) - the Carbon Cycle and Cambridge Atmospheric CO2: natural Variations Archean to Walther, H., 1990 – the Weisselster Basin (GDR) – an Present – pp. 357-375, American Geophysical Union example of the development and history of Paleogene Geophysical Monograph 32 forest vegetation in central Europe - in: Knobloch, E. Wolfe, J.A., 1987 – Late Cretaceous-Cenozoic history of & Kvacek, Z. (eds.) – Paleofloristic and paleoclimatic deciduousness and the terminal Cretaceous event - changes in the Cretaceous and Tertiary – pp. 149- Paleobiology 13: 215-226 158, Geological Survey Publisher, Prague Wolfe, J.A., 1994 – Alaskan Paleogene climates as infer Walther, H., 1994 – Invasion of Arcto-tertiary elements red from the CLAMP database – in: Boulter, M.C. & in the Paleogene of central Europe - Boulter, M.C. & Fisher, H.C. (eds.) – Cenozoic Plants and climates of Fisher, H.C. (eds.) – Cenozoic Plants and climates of the Arctic – pp. 223-237, Springer-Verlag, Berlin the Arctic – pp. 239-250, Springer-Verlag, Berlin Yang Hong & Jin Jianhua, 2000 – Phytogeographic Walther, H., 1999 – Die Tertiärflora von Kleinsaubernitz history and evolutionary status of Metasequoia: bei Bautzen – Palaeontographica B 249: 63-174 geological and genetic information contrasted – Acta Walther, H., 2000 – Floristic relations between North and Palaeontologica Sinica 39 (Sup.): 288-307 central America and Europe in the Eocene – Acta Zhang Zhao-Qun, Dawson, M.R. & Huang Xue-Shi, Universitatis Carolinae Geologica 44: 51-57 2001 – A new species of Gobiolagus (Lagomorpha, West, R.M. & Dawson, M.R., 1977 – Mammals from the Mammalia) from the middle Eocene of Shanxi Paleogene of the Eureka Sound Formation: Ellesmere Province, China - Annals of the Carnegie Museum Island, Arctic Canada – Géobios Mémoire spécial 1: 70: 257-261 107-124 Zhilin, S.G., 1989 – History of the development of the West, R.M. & Dawson, M.R., 1978 – Vertebrate paleon- temperate forest flora in Kazakhstan, U.S.S.R. from tology and the Cenozoic history of the North Atlantic the Oligocene to the early Miocene – The Botanical region – Polarforschung 48: 103-119 Review 55 (4): 205-330 Wilde, V., 1989 – Untersuchungen zur systematik der Zhilin, S.G., 2001 – Structure of the Turgayan flora in Blattreste aus dem Mitteleozän der grube Messel bei the Oligocene and Miocene and its palaeoclimatic Darmstadt (Hessen, Budesrepublik Deutschland) – features - Acta Palaeobotanica 41: 141-146 Courier Forschungsinstitut Senckenberg 115: 1-213 Wilde, V., 1995 – Die Makroflora aus dem Mitteleozän Received 19 June 2001 des Geiseltalgebietes, kurze Übersicht unde verglei Accepted 10 December 2002 che – Hallesches Jahrbuch Geowissenchaften B 17: 121-138

80 COLLINSON & HOOKER: Paleogene vegetation Eurasia . val stated ined within the inter column * indicates age undef ‘Age’ In the as which are plotted on Figures 1-3. aleogene flor le 1 List of sites the P ab T

81 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 ued) (contin val stated. ined within the inter column * indicates age undef ‘Age’ In the as which are plotted on Figures 1-3. aleogene flor le 1 List of sites the P ab T

82 COLLINSON & HOOKER: Paleogene vegetation Eurasia ued) (contin val stated. ined within the inter column * indicates age undef ‘Age’ In the as which are plotted on Figures 1-3. aleogene flor le 1 List of sites the P ab T