A summary of fossil records for Arecaceae

Of all monocotyledons the Arecaceae displays by far the richest fossil record, and there is an extensive literature. The earliest unequivocal fossil palm material probably dates from the early to mid Late Cretaceous (Turonian > Coniacian > Santonian). The records are geographically widespread and comprise a wide range of organs: leaves, cuticles, stems, rhizomes, roots, fruits, seeds, endocarps, rachillae, peduncles, inflorescences, individual flowers and pollen. For some of these organs records are rare while for others, such as leaves, stems and pollen, records are abundant. However, fossil material often lacks sufficient diagnostic detail to allow reasonable association with living palm taxa beyond, or even to, subfamilial level. Nevertheless, many fossil genera and numerous species have been described. A brief survey of palm fossil records is presented, and their taxonomy and morphological limitations are considered. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 39–67.     

中国第四纪晚期孢粉记录整理

孢粉数据库是重建过去植被格局以及研究古气候变化和大气圈-生物圈-人类活动相互关系的基础,而孢粉取样信息的记录和整理是建立孢粉数据库的重要前提。该文在收集我国1960–2008年发表的孢粉研究文献的基础上,整理分析了第四纪晚期,尤其是2万年以来(全新世为主)全国孢粉采样点的信息,包括采样地点名称、详细采样位置、省份、采样点经纬度和海拔高度、样品类型、取样深度、孢粉样品数量、14C测年数量及年代记录、覆盖的时间段和参考文献。总结发现,我国目前共有2324个表土/湖泊表层花粉采样点和987个第四纪晚期的地层沉积剖面和钻孔,其中高质量的地层孢粉采样点714个。虽然我国以及部分国际上从事第四纪研究的科学家尽了最大努力开展中国疆域的孢粉学研究,但由于人力、物力以及地形条件的限制,在我国仍然存在部分孢粉采样的"地理空隙",比如北方和西北荒漠地区、青藏高原无人区、中南部山区和东部人类活动频繁区域。然而,该数据信息库的不断完善,将为中国第四纪孢粉数据库的建立奠定良好基础。     

Diverse fossil Onagraceae pollen from a Miocene palynoflora of north-east China: early steps in resolving the phytogeographic history of the family

The origin and evolution of angiosperms can be unravelled by using fossil records to determine first occurrences and phytogeographic histories of plant families and genera. Many angiosperm families, for example the Onagraceae, have a poor macrofossil record, but are more common in palynological records. Modern Onagraceae produce pollen clearly distinct from that of other angiosperms. Combined morphological features obtained by use of light and scanning electron microscopy have enabled assignment of fossil Onagraceae pollen to extant genera, and therefore tracing of the origin and past distributions of extant Onagraceae lineages. We studied a Miocene palynoflora from the Daotaiqiao Formation of north-east China. Using the single-grain technique, we examined individual Onagraceae pollen/tetrads using both light and scanning electron microscopy. Fossil Onagraceae pollen is more frequent than macrofossil remains, but is still rare, and usually represented by a single taxon in palynological samples. Remarkably, samples from the Miocene of north-east China contain five different species: two of Circaea, one of Epilobium, and two of Ludwigia. Such a large number of Onagraceae taxa from a single palynoflora is unknown elsewhere. Whereas Ludwigia pollen is known from Cenozoic sediments of the northern hemisphere, the Circaea pollen is the first fossil pollen assignable to this extant genus. This is also the first fossil record of Epilobium from China. Although the young geological age of this sample does not enable consideration of time of origin for the genera encountered, the co-occurrence of Circaea, Epilobium, and Ludwigia in the mid to late-Miocene of East Asia sheds some light on their phytogeographic histories.     

Past grazing habitats for Svalbard reindeer indicated by the pollen content of 3300‐year‐old faeces from Edgeøya,Svalbard

Sedentary populations, like the Svalbard reindeer, tend to select the most nutritious and easily available plants during the different seasons. It has been shown that plants selected during grazing season are reflected in the pollen content of modern faeces from Svalbard reindeer. In this study the pollen and spore content of fossil reindeer faeces and peat from Raddedalen, western Edgeøya, Svalbard, are presented. The pollen content of the fossil faeces is first compared to published pollen data from modern Svalbard reindeer faeces so as to determine which season they were deposited, and second the pollen in the fossil faeces are compared to the pollen content of the peat in which they were found. The lower part of the peat section is dated to 3435±105?cal?yr BP. The faeces are dominated by pollen from early flowering species such as Saxifraga oppositifolia‐type, S. nivalis‐type, Brassicaceae, and Pedicularis spp. thereby indicating that grazing and faeces deposition took place in spring and early summer. The peat and faeces have some similarities in their pollen and spore content, but different methods of deposition and concentration of pollen in faeces and peat result in different fossil assemblages. The results suggests that the climate of Raddedalen was warmer and moister than today, allowing peat to form in the past.     

Elatides sandaolingensis n. sp. (Cupressaceae sensu lato) — A new fossil conifer with cones from the Middle Jurassic of Xinjiang,northwestern China

Here we describe a new conifer fossil, Elatides sandaolingensis Z.X. Wang and B.N. Sun n. sp., from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. The materials consist of compressions represented by well-preserved leafy shoots, pollen cones, and seed cone. Leaves are characterized by long triangular shapes, with straight apex and entire margins, and two stomatal bands on the abaxial surface. Pollen cones are terminally disposed on the ultimate leafy shoots, borne singly or in clusters. Pollen sacs are long-oval shaped, with three pollen sacs fused together. Pollen grains are spherical and have small germinal papilla and few wrinkles. The seed cone is oblong, with more than 35 helically arranged bract-scale complexes, which are characterized by long-oval shape and triangular apex. Compared with the extant nine genera of Taxodiaceous Cupressaceae in the morphology of seed cone and pollen cones, the present fossil consistently shares many characteristics with the extant genus Cunninghamia, but differs in other aspects. After being compared with the reported fossil records of the fossil plants, the current species is found to be different from any known species; thus, the present fossil is referred to as a new species of Elatides. From the similarity between the present fossil and Cunninghamia, it can be inferred that there may be a genetic relationship between these two genera. Additionally, the new species has thin cuticles and slightly sunken stomata, which can provide evidence indicating that the climate of the Turpan-Hami Basin in Middle Jurassic might have been warm and humid. By studying the geological history and geographical distribution of Elatides, it can be inferred that this genus may have originated in Switzerland, and it was migrated from Xinjiang to North and South China through the Ural Mountains.     

A survey of species assigned to the fossil pollen genus Anacolosidites

Anacolosidites Cookson & Pike, a fossil pollen genus recorded since the Campanian, is peculiar in its morphology – six‐porate with three apertures on each hemisphere, located away from the equator, and with the distal and proximal apertures positioned over each other. Representatives of this fossil genus are widely considered to represent extant Olacaceae from tribe Anacoloseae. Olacaceae is an exclusively tropical angiosperm family with a pantropical distribution; consequently the fossils are often used to suggest a tropical climate and in addition are frequently used as a stratigraphic marker. Fossil species assigned to Anacolosidites are quite variable and may not all represent Olacaceae, in which case they may not indicate tropical climate.

The present study is a morphological survey of fossil pollen assigned to Anacolosidites; it identifies the published reports of the fossil species that probably represent positive occurrences of Olacaceae pollen in the fossil record. Within Olacaceae, Anacolosidites‐type pollen is usually compared with pollen of genera in tribe Anacoloseae, in particular: Cathedra, Anacolosa and Phanerodiscus, but never with the pollen of Ptychopetalum, a genus from tribe Olaceae with closely similar pollen to the other three genera, but with a reticulate tectum and very small circular apertures located near the equator. Nevertheless, the records of reticulate Anacolosidites species which have been excluded from the emended diagnosis are unlikely to be related to Ptychopetalum. The earliest accepted record of Anacolosidites is from the Late Cretaceous (Maastrichtian) of Germany. However, most Late Cretaceous records, and later Russian and Chinese occurrences referred to Anacolosidites, consist mainly of Normapolles‐type pollen, whereas many of the Cenozoic records assigned to Anacolosidites have a much clearer affinity with the pollen of Anacolosa, Cathedra and Phanerodiscus (tribe Anacoloseae). The newly emended genus Anacolosidites may be used as a stratigraphic marker for tropical or megathermal climatic conditions.     

The representation of rainfall and fire intensity in fossil pollen and charcoal records from a South African savanna

Savanna ecosystems of southern Africa are strongly influenced by water availability and fire intensity, and this study aimed to show whether these two specific environmental variables are reflected in fossil pollen and charcoal records. Palaeoecological records of charcoal concentration from three short sedimentary sequences were used to reconstruct fire intensity (the rate of energy released along a fire front) over 50 yrs in the Kruger National Park (KNP), South Africa. Fossil pollen percentages from surface and core-sediment samples taken from water bodies were compared with the reconstructed fire intensity over space and time. Higher fire intensity led to increased herbaceous cover and decreased woody plant growth. Fossil pollen percentages and charcoal concentrations were also compared with rainfall records. Increased macroscopic charcoal abundances and percentages of Cyperaceae pollen corresponded to periods of increased rainfall. The results of this study have shown that fossil pollen and charcoal records from savanna environments can be used to reconstruct past fire intensity and its impact on terrestrial vegetation, as well as changes in rainfall.     

Ancient DNA from pollen: a genetic record of population history in Scots pine

Assessments of plant population dynamics in space and time have depended on dated records of fossil pollen synthesized on a subcontinental scale. Genetic analyses of extant populations have revealed spatial relationships that are indicative of past spatial dynamics, but lack an explicit timescale. Synthesis of these data requires genetic analyses from abundant dated fossil material, and this has hitherto been lacking. Fossil pollen is the most abundant material with which to fill this data gap. Here we report genetic analyses of fossil pollen retrieved from Holtjärnen postglacial lake sediment in Sweden and show that plastid DNA is recoverable from Scots Pine and Norway spruce pollen grains that are 100 and 10 000 years old. By sequencing clones from two short plastid PCR products and by using multiple controls we show that the ancient sequences were endogenous to the fossil grains. Comparison of ancient sequences and those obtained from an extant population of Scots pine establishes the first genetic link between extant and fossil samples in this species, providing genetic continuity through time. The finding of one common haplotype present in modern, 100-year old and 10 000-year old samples suggests that it may have persisted near Holtjärnen throughout the postglacial period. This retrieval of ancient DNA from pollen has major implications for plant palaeoecology in conifer species by allowing direct estimates of population dynamics in space and time.     

Revisiting tree-migration rates: Abies alba (Mill.), a case study

At northern temperate latitudes trees have adjusted their ranges substantially in response to changing climates during the Holocene. Results from dispersal model simulations suggest that postglacial migration rates may have been over-estimated from fossil pollen data. As a contribution to this debate, we infer the migration rates of Abies alba (Mill.), silver fir, as a case-study species, by using a spatially explicit approach based on fossil pollen but taking into account its modern genetic diversity pattern. Maximum estimates of migration rates from fossil pollen data alone are higher than 700 m yr^?1 during the Holocene. Considering the potential refugia as suggested from all the fossil data but restricting the area over which silver fir expanded from each glacial refugium using data on the current haplotype distribution, the estimated maximum migration rates of silver fir are less than 250 m yr^?1. Genetic information may allow for (1) the exclusion of those refugial areas where the species may have survived during the last glacial period but from which it did not spread or spread only very locally and (2) the delineation of the areas over which the species spread from each glacial refugium. The estimated rates in the present study are generally consistent with rates suggested from modelling approaches. This study shows that integrating fossil pollen records can improve simulations of dispersal processes and, thus, allow for better predictions of future changes in tree species’ ranges.     

Early Oligocene fruits and leaves of Burretiodendron (Malvaceae s.l.) from South China

The genus Burretiodendron Rehder is currently endemic to an area near the China–Vietnam border and the limestone mountains of Thailand and Myanmar. The fossil records of this genus were previously found only from the Miocene of Yunnan, Southwest China, and the Oligocene of Guangxi, South China. Here, we describe fossil fruits and associated leaves of Burretiodendron, which were discovered in the lower Oligocene Shangcun Formation of the Maoming Basin, Guangdong, South China. Morphological comparison with extant and fossil Burretiodendron taxa indicates that fruit fossils belong to the species Burretiodendron parvifructum J. Lebreton Anberrée & Z. K. Zhou. This is one of the earliest fossil records of the genus, providing additional evidence for the early biogeographic history of this genus and supporting the inference that the genus originated in South China. According to the habitat conditions of modern species, we speculate that there were limestone mountains around the Maoming Basin in the early Oligocene.     

Modern vegetation–pollen–climate relationships for the Pampa grasslands of Argentina

Aim To analyse the relationships between potential natural vegetation, pollen and climate in order to improve the interpretation of fossil pollen records and provide the background for future quantitative palaeoclimatic reconstructions. Location Pampa grasslands of Argentina, between 33–41° S and 56–67° W. Methods Modern pollen data were obtained from a pollen data base developed by the Grupo de Investigación de Paleoecología y Palinología, Universidad Nacional de Mar del Plata, Argentina (143 surface samples and 17 pollen types). Analysis of pollen and climate data involved multivariate statistics (cluster analysis and principal components analysis), scatter diagrams, Pearson’s correlation and isopoll mapping. Results Vegetation patterns at regional scales (grasslands and xerophytic woodlands) and local scales (edaphic communities) were identified by cluster analysis of pollen surface samples. The main climatic variables that appear to constrain the vegetation distribution and abundance of taxa are mean annual precipitation, annual effective precipitation and summer temperature. Individual pollen types such as Chenopodiaceae, Apiaceae, Cyperaceae, Prosopis, Schinus, Condalia microphylla and other xerophytic taxa are good indicators of moisture regime. Many pollen types are significantly correlated with summer temperature. The modern vegetation–pollen–climate relationships vary in a broadly predictable manner, supporting the contention that fossil pollen assemblages can be related to particular climatic characteristics. Main conclusions An expanded suite of modern analogues facilitated new insights into vegetation–pollen–climate relationships at the regional scale in Pampa grasslands. Relationships between individual pollen types and climate are appraised at a regional scale and new modern analogues are presented. The results provide the basis for improved vegetation and climate reconstruction from fossil records of the study area.     

The first Loranthaceae fossils from Africa

An ongoing re-investigation of the early Miocene Saldanha Bay (South Africa) palynoflora, using combined light and scanning electron microscopy (single grain method), is revealing several pollen types new to the African fossil record. One of the elements identified is Loranthaceae pollen. These grains represent the first and only fossil record of Loranthaceae in Africa. The fossil pollen grains resemble those produced by the core Lorantheae and are comparable to recent Asian as well as some African taxa/lineages. Molecular and fossil signals indicate that Loranthaceae dispersed into Africa via Asia sometime during the Eocene. The present host range of African Loranthaceae and the composition of the palynoflora suggest that the fossil had a range of potential host taxa to parasitise during the early Miocene in the Saldanha Bay region.     

The history of early cereal cultivation in northernmost Fennoscandia as indicated by palynological research

The age of the introduction of cereal cultivation in northern Europe has long been debated by researchers from many disciplines, in particular archaeology and palaeoecology. Over the past 40 years extensive palynological data have been collected concerning pre-industrial land use in northern Fennoscandia. This paper reviews palynological studies that include records of fossil cereal pollen from northernmost Sweden, Finland and Norway at latitudes north of 63°N. The geographical extent of known early cultivation sites is constantly expanding, with more than 100 records of cereal pollen pre-dating ad 1700. The oldest records of scattered cereal pollen derive from Neolithic times. Periods of continuous cultivation, indicated by cereal pollen recorded recurrently in the sediment profiles, derive from the Late Neolithic and Early Bronze Age. Collectively, the reviewed pollen records indicate that cereal cultivation was first introduced into areas close to the coast and later to the interior, and that it may have been practiced locally long before sedentary settlements based on intensive cultivation were established during medieval times. The data do not indicate a latitudinal spread of cultivation from south to north. However, methodological problems relating to pollen morphology of cereals, site characteristics and lack of connections to archaeologically excavated sites imply that the value of many early cereal pollen finds remains unclear. To increase our understanding of the context in which cereal cultivation was introduced in northernmost Fennoscandia, multidisciplinary studies integrating palaeoecology, archaeology and history are needed.     

Fossil pollen as a record of past biodiversity

Quaternary pollen records may contribute uniquely to the understanding of present plant diversity. Pollen assemblages can reflect diversity at community and landscape scales but the time resolution of most studies does not match that of modern ecological studies. Because of the complicating effects of differential pollen productivity and dispersal, pollen records do not directly reflect equitability aspects of vegetation diversity. Vegetation diversity indices other than S (the total number of taxa) are therefore not appropriate for pollen assemblages. As a measure of the species richness palynological richness is biased by the lack of taxonomic precision, by a possible interference on pollen dispersal from vegetation structure and by pollen representation. The nonlinear relationship between species richness and pollen-taxa richness may be used in attempts to estimate past floristic richness from fossil pollen assemblages. Using a hypothetical example the strong effect of cover shifts in the vegetation affecting taxa with different representation (R_rel) values on observed palynological richness is demonstrated. It is suggested that estimates of relative pollen productivity should be used to guide the pollen sum on which pollen-type richness is estimated by rarefaction techniques and this approach is illustrated using a paired site study of late Holocene diversity dynamics. The need for a modern training set relating pollen-type richness to species richness, pollen productivity and vegetation structure is emphasized.     

The mangrove palm Nypa in the geologic past of the New World

The paleontological history of Nypa, known today as the mangrovepalm, is traced through geological time back to the Late Cretaceous. Emphasis is laid on the New World occurrences, especially in NorthAmerica, where Nypa is known from fossil fruits and pollen. In SouthAmerica, the stratigraphic range of this palm extends from theMaastrichtian to the late Eocene, whereas in North America, Nypa isrestricted to only the Eocene. Nypa occurs as pollen all along theAmerican Gulf Coast from the early Eocene (Ypresian) to the late Eocene(Priabonian), while fruit records come from the early and middle Eoceneof Maryland and Texas, respectively. The floristics of these Eocenemangroves, including possible mangrove associates, and the developmentof mangrove vegetation in the neotropics through the Tertiary andQuaternary, are discussed. New paleobotanical evidence from a middleEocene faunal and floral assemblage in Texas, the Casa Blanca flora (LaredoFormation, Claiborne Group), which contains fossil Nypa pollen andfruits, is described in detail.     

Vegetation and environment changes inferred from pollen records since 3000 cal. yr BP in Kanas wetland,Xinjiang

Aims Climate change can significantly affect the vegetation worldwide. Thus, paleovegetation and paleoclimate reconstruction should consider the quantitative relationship between modern vegetation and climate. The specific objectives of this study were (i) to assess the influence of environmental variables on pollen assemblages in the Kanas region, (ii) to reconstruct the evolution of vegetation over the past 3000 years using pollen records and (iii) to quantify historical climate change (including mean annual temperature and total annual precipitation) using a weighted averaging partial least squares regression method (WAPLS) applied to fossil pollen data from the Kanas wetland in Xinjiang, China.     

Transport of airborne Picea schrenkiana pollen on the northern slope of Tianshan Mountains (Xinjiang, China) and its implication for paleoenvironmental reconstruction

The understanding of airborne pollen transportation is crucial for the reconstruction of the paleoenvironment. Under favorable conditions, a considerable amount of long-distance-transported pollen can be deposited far from its place of origin. In extreme arid regions, in most cases, such situations occur and increase the difficulty to interpret fossil pollen records. In this study, three sets of Cour airborne pollen trap were installed on the northern slope of Tianshan Mountains to collect airborne Picea schrenkiana (spruce) pollen grains from July 2001 to July 2006. The results indicate that Picea pollen disperses extensively and transports widely in the lower atmosphere far away from spruce forest. The airborne Picea pollen dispersal period is mainly concentrated between mid-May and July. In desert area, weekly Picea pollen began to increase and peaked suddenly in concentration. Also, annual pollen indices do not decline even when the distance increased was probably related to the strong wind may pick up the deposited pollen grains from the topsoil into the air stream, leading to an increase of pollen concentration in the air that is irrelevant to the normal and natural course of pollen transport and deposition. This, in turn, may lead to erroneous interpretations of the pollen data in the arid region. This study provided insight into the shift in the Picea pollen season regarding climate change in arid areas. It is recorded that the pollen pollination period starts earlier and the duration became longer. The results also showed that the temperature of May and June was positively correlated with the Picea pollen production. Furthermore, the transport of airborne Picea pollen data is useful for interpreting fossil pollen records from extreme arid regions.     

Contribution to the flora history of the genus Polemonium with special reference to North Norway

A key for the identification of pollen of the European species of Polemonium is presented. Fossil pollen records from Norway are surveyed and discussed. The records available show that pollen is derived both from natural stands close to the shoreline, and from plants established as apophytes in grazed meadows. It is suggested that P. caeruleum is a Late Glacial element, which arrived from the south at least, but probably also from the northeast. Pollen of P. boreale -type has a wider distribution in the Late Weichselian, at least in northern Russia. A pollen diagram from Bodø illustrates the fossil record occurrence of Polemonium in a Late Holocene sequence.     

Palaeopalynological Evidence of Phylogeny in Hamamelidaceae

This paper deals with evolution, classification and pollen morphology of the Hamamelidaceae, an important family in phylogeny of angiosperms. I. Pollen morphology and systematics of modern Hamamelidaceae. The pollen morphology of the family may be divided into the following four types: (1) Tricolpate: Hamamelis, Loropetalum, Mytilaria, Corylopsis, Sysopsis, and Distylum etc.; (2) Tricolpate with operculum: Disanthus; (3) Tricolporate: Rhodoleia; (4) Pantoporate: Liquidambar. The tricolpate pollen of the Hamamelidaceae is a primitive type in angiosperms, but the most ancient type is monocolpate pollen. Therefore, the family might have evolved from the Magnoliaceae of the monocolpate pollen. The pantoporate pollen is an evolutionary type in the family. It might have evolved from the tricolpate pollen. II.The fossil pollen of the Hamamelidaceae 1 .General introduction of the fossil pollen of the family Hamamelidaceae The most’ancient fossil pollen belonging to the family was found in the middle-late Early Cretaceous. Palynologists call the fossil pollen of the Hamamelidaceae Retitricolpites, which consists of three genera: Hamamelis L.,Corylopsis Sieb. et Zucc and Fothergilla Murr. Liquidambar is of an advanced type in the fossil pollen of the Hamamelidaceae. It was found in the period from the Palaeogene to the Neogene in China. 2. The geological history of the Hamamelidaceae may be divided into the following four stages: (A) The Early Cretaceous stage or origination stage. The family may be evolved from Magnoliales in the middle-late Early Cretaceous. (B)the Late Cretaceous stage or formation stage. The family is much developed in the period. (C) The Tertiary stage or development stage. The family was a much developed one among angiosperms. (D)The Neogene to modern stage or perfection stage. The evolutionary type, the Liquidambar type of the Hamamelidaceae, was much developed in the Neogene. III. The palaeopalynological evidence of evolution of the Hamamelidaceae The earliest fossil pollen of angiosperms was found in the Barremian (Early Cretaceous) in England, Israel, the United States of America etc., and was named as Clavatipollenites by Couper (1953). In recent years, Clavatipollenites was also found in the middle-late Early Cretaceous in Nei Monggol and Jiangxi Province of China. We also found Retitricolpites in the middle-late Early Cretaceous in Nei Monggol and Jiangxi Province. Retitricolpites, belonging to the Hamamelidaceae, is a primitive type among angiosperms, but it is younger than Clavatipollenites. Therefore, the pollen of Hamamelidaceae may have evoloved fromClavatipollenites, which may have evoloved in turn from that of Magnoliales.