SCIENCE CHINA Earth Sciences • RESEARCH PAPER •
December 2010 Vol.53 No.12: 1828–1835 doi: 10.1007/s11430-010-4079-8
The evolution of Paleozoic vascular land plant diversity of South China WANG Yi*, WANG Jun, XU HongHe & HE XueZhi State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China Received January 22, 2010; accepted June 12, 2010
Data of Paleozoic and some Early Triassic vascular land plant fossils from South China are synthetically analyzed, revealing evolutionary characteristics of Paleozoic vascular land plant diversity. Vascular land plant diversity keeps increasing in the Paleozoic as a whole. The Silurian witnessed the earliest evolution and initial diversification of land plants. From the Early Devonian to the Early Carboniferous (Mississippian), the great development, diversification, and differentiation really occurred in vascular land plants, with fluctuations of diversity, rapid replacement of the plant types, and an all-out takeover of terrestrial ecological niches. From the Early Permian, land plant diversity dramatically increased, and reached a climax in the Middle-Late Permian. Comparisons between late Paleozoic marine and terrestrial biodiversity reveal co-evolution of the late Paleozoic animals and plants as well as the individual evolutionary patterns of sea/land ecosystems. Vascular land plant diversity dramatically declined in the Frasnian as a result of the F/F event, and the end-Permian mass extinctions completely turned over the phytogroups. Paleozoic, vascular land plants, diversity, sea-land co-evolution, South China
Citation:
Wang Y, Wang J, Xu H H, et al. The evolution of Paleozoic vascular land plant diversity of South China. Sci China Earth Sci, 2010, 53: 1828–1835, doi: 10.1007/s11430-010-4079-8
The biodiversity response to global change has been a topic of universal concern. Biodiversity not only counts for the number of organisms but also is a major factor concerning sustainable development and human future. The present biodiversity crisis should be analyzed in the context of its geologic evolution as to recognize biodiversity trends, examine anthropic impact on nature, and make comparisons on a larger scale. More and more attention has been paid to geological biodiversity studies based on regional rather than global data [1]. Regional biodiversity in geological times is more distinctive in ecological environment, taxa, and biogeography than global biodiversities. Global biodiversity summed
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by data from every paleoblock could reflect genuine diversity change. An analysis of the Chinese database [1] indicates that early marine animal biodiversity evolution in the late Protozoic to Mesozoic, except certain geological intervals, of South China is identical to that in the global context, mainly marked by global events such as mass extinctions and bio-radiations. This is because South China was situated in different places at different geological times and was affected by a range of climates, environments, tectonics, and regional depositional conditions. The evolution of the terrestrial ecosystem is an important component of earth ecology, and land biodiversity evolution characteristics and trends are significant. Although significant progress has been achieved in understanding the Paleozoic marine biodiversity of South China, the evolution of terrestrial biodiversity is still not clear. Study of Paleozoic earth.scichina.com
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terrestrial biodiversity would contribute to interpreting its evolution process and recognizing the evolutionary similarities and differences between the marine and land ecosystems. Here we discuss the features of Paleozoic (Silurian to Permian, and some Early Triassic) vascular land plants biodiversity pattern of South China, the succession of land vegetation, and the origins of key plant groups.
1
Study methods and data sources
Plant mega-fossils are the most direct and most effective materials to study plant evolution, and as a result, analyzing the number of plant mega-fossil genera has become one of the main methods of understanding plant diversity through time. The genus is thought to be the most important and the most essential taxonomic level. This is because as most plant mega-fossils were not completely preserved, plant types from many localities are simple and limited; accurate study at the level of species is still difficult to some extent. Among Silurian to Devonian terrestrial vascular plants, the number of higher categories, e.g., family and order, is less significant than that of genera, and furthermore, relationships between plant groups are controversial. The pattern of diversity, at the level of genus, the most important and fundamental taxon [2], could be very indicative to the flora change. Generic diversity varies among plant groups. Diversity patterns of different groups were analyzed. The main plant groups from the Silurian to the Devonian include rhyniophytes, zosterophyllophytes, trimerophytes, lycophytes, sphenophytes, filicophytes (early ferns), and progymnospermophytes/seed plants; those from the Carboniferous to the Permian include lycophytes, sphynophytes, filicophytes, pteridospermophytes, and gymnospermophytes. Note that the accurate correlation is still lacking between Carboniferous-Permian stratigraphic stages in Chinese terrestrial stratigraphy and those in the International Stratigraphy Chart. We have to use local Chinese stages when analyzing plant fossil data. The rough correspondences could be found, such as early Namurian corresponding to Serpukhovian, Chihsian to Artinskian-Kungurian, Maokouan to RoadianWordian, and Luntanian to Capitanian-Wuchiapingian. All data analyzed were sourced from published papers [3–13].
2 Characteristics of vascular land plant diversity evolution from Silurian to Devonian in South China 2.1
General characteristics
The generic diversity of vascular land plants from the pre-Pridoli (late Llandovery) to the Famennian of South China could be summarized as follows (Figure 1): 1) the number of genera keeps increasing from Pridoli to Famennian, except in Emsian+Eifelian and Frasnian; 2) the diver-
Figure 1 Vascular land plant generic diversity from pre-Pridoli to Famennian in South China. P-Pri: Pre-Pridoli (late Telychian, Llandovery); Pri: Pridoli; Loc: Lochkovian; Pra: Pragian; Ems: Emsian; Eif: Eifelian; Giv: Givetian; Frs: Frasnian; Fam: Famennian. The same below.
sity from pre-Pridoli to the earliest Devonian is low, only 1 to 4 genera were counted, and evolves slowly, indicating an early form of early land flora and the potential start of major evolution; 3) in the Pragian, land-plants evolve rapidly and the first peak of the diversity occurs, 24 genera are counted, and most of them are monotypic; 4) the diversity climaxes in the Givetian and the Famennian, 20 and 29 genera respectively, and land plants diversify dramatically in both morphology and anatomy; 5) the diversity declines remarkably in Emsian-Eiflian and Frasnian, to 12/7 and 10 genera. The cause of the first decrease is probably preservation bias and relatively inadequate study, as a result of restricted occurrences of Emsian-Eiflian strata in South China, whereas the Frasnian decrease is probably related to F/F mass extinction event. 2.2
Analysis of flora evolution
Different plant groups have different morphological and anatomical characters as well as different evolution trends. Analysis is carried out on the Silurian-Devonian plant groups from South China (Figure 2). The plant groups are simplified and subdivided into two, i.e., pre-Eiflian and post-Eiflian, evolutionary floras (EF) (EF1 refers to zosterophyllophytes and trimerophytes, and EF2 refers to the rest of plants being analyzed). Only the diversities of the two groups are compared and analyzed (Figure 3). The EF1 contributes greatly to the plants diversity increase from Pridoli to Pragian, being a majority of Early Devonian land vegetation. From Emsian, the EF1 diversity declines, while the EF2 diversifies rapidly, gradually takes the place of the EF1, and becomes a dominant group of the vegetation. 2.3
Comparisons with coeval plant diversity evolution
Our analysis is roughly consistent with generic diversity statistics based on global data [2, 9, 14]. However, the dif-
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3 Characteristics of Carboniferous to Permian plant diversity from South China 3.1
Figure 2 Diagram of main plant groups diversity during the SilurianDevonian in South China. L-lycopsid; R-rhyniophytes; T-trimerophyte; Zzosterophyllophyte; S-sphenopsid; P-progymnosperms/seed plants; Fferns/pre-ferns.
In South China, both the Late Carboniferous (Pennsylvanian) to the Early Permian (about Namurian, late-Westaphalian, Stephanian, Asselian) terrestrial deposits and plant fossil records are lacking, and abundant plant fossils could only be found from the Early Carboniferous and the Permian beds. Plant diversity shows a trend of continuous rising and slight declining in both the Early Carboniferous and the Permian (Figure 4). Both the diversity rises indicate the process of flora replacement and prosperity. In the Early Carboniferous, Pro-Cathaysian flora is evident; in the Permian, the typical Cathaysian flora is shaped. However, the two declines are caused by different reasons. The decline from Visean to the early Namurian is mainly caused by the loss of terrestrial environment, a result of a large transgression in South China, instead of the suitable climatic condition of Tethys as indicated by the well-developed Late Carboniferous flora in the North China Platform. The diversity decrease between Permian Lungtanian and Changhsingian corresponds to the decline of Cathaysian flora. Comparison of occurrences between the stages indicates that extinction rate of the flora is up to 62% [5] and dramatic changes occurred among the members of Cathaysian flora. 3.2
Figure 3 Diagram showing evolution of the two evolutionary floras during the Silurian-Devonian (EF1 and EF2) of South China.
ferences are: 1) the global data indicate that the diversity in Emsian is higher than that in Pragian, but our data show that the first diversity peak occurs in Pragian; 2) from the global data, plant diversity has a slight decline after Eifelian, whereas our data show a diversity drops to the lowest point in Eifelian; and 3) diversity decrease could be seen from the both data, but our statistics indicate a greater decline rate of 50%, instead of 25% based on the global data. We consider that the differences are caused by 1) Emsian, Eifelian, and Frasnian fossil plants from South China are less collected and studied, and potentially could not reflect the general diversity of the flora; 2) South China was an isolated paleoblock and had different climate conditions in the Devonian, whose flora was specialized and distinct; and 3) our data come from a local and endemic land ecosystem, showing individual characteristics; the global data have a much larger source, showing a general trend and common characteristics.
General characteristics
Analysis of flora evolution
In South China, the evolution of diversity of individual Carboniferous-Permian plant groups (Figure 5) could be summarized as below. (1) Lycophytes, spenophytes, filicophytes, and pteridospermophytes are groups occurring from the Carboniferous to the Permian. The diversities of all these groups experience declines in the Late Carboniferous and Early Permian but are soon restored to the previous or even higher level. Although the Carboniferous lycophytes have a rela-
Figure 4 Plant genera diversity evolution curve during Carboniferous-Permian in South China. Tur: Tournaisian; Vis: Visean; Nam: Namurian; Chi: Chihsian; Mao: Maokouan; Lun: Lungtanian; Cha: Changhsingian. The same below.
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Figure 5 Diagram of the diversity of main plant groups during the Carboniferous-Permian in South China. L: Lycophytes; S: Sphenophyte; F: Fillicophytes; P: Pteridosperms; N: Noeggerathiales; S: Seeds; G: Gymnosperms; I: Incertae sedis.
tively high diversity and are dominant members of the whole flora, their tall and arborescent forms are rare. After the Permian, lycophyte diversity has a slight decline, but lycophytes are still quite common, and most of them are arborescent, coal-forming plants. Sphenophyte diversity has an apparent rise in the Permian, being one of the coal-forming plants, and with dramatically increasing plant bodies. Fillicophytes are the group that has the largest increase in diversity. Fillicophytes are rare in the Carboniferous and become the dominant members of the Permian flora, as indicated by the key feature of Cathaysian flora. Pteridosperm diversity keeps a high value in both Carboniferous and Permian. However, their plant members change. Carboniferous pteridosperms are mostly ancient-pteridosperms (e.g., Paripteris and Linopteri), comprising the main members of the flora; and common, advanced Permian ones (e.g., Petaspermales and Corystospermales). At the end of the Permian, pteridosperms become dominant in some high land flora, resembling those in the Triassic flora. (2) Gymnosperms appear from the Early Permian and their diversity rapidly reaches a high level, as is a key characteristic of Permian flora diversity evolution of South China. The seeds refer to dispersed ovules/seeds of gymnosperms. They were counted separately for their uncertain parent plants. These seeds suggest that the diversity of gymnosperms might be higher than our statistics. The transformation from Palaeophytic to Mesophytic flora is also indicated by the diversity trend. Although the diversities of most plant groups have slight declines in the Changhsingian, the latest Permian, gymnosperm diversity is still at a high level. (3) Incertae sedis plants, including noeggerathialeans, play a key role in the Permian flora. Of course, they originate from the Permian plant radiation and could be classified into some known catalogue or new taxonomy, e.g., Nystroemiaceae, which represent a new broad-leaved seed plant [15], and Noeggerathiales, which probably belong to progymnosperms or an independent group [16].
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Comparison of South China Carboniferous and Permian spore plants (lycopsids, sphenopsids, noeggerathialeans, and fern allies) and seed plants (pteridosperms and gymnosperms) (Figure 6) indicates that diversities of both groups are nearly equal except in the Early Carboniferous, in which the diversity of spore plants is higher than that of seed plants. Both the spore plants and the seed plants have experienced a long period of co-existence before the replacement of Palaeophytic with Mesophytic Flora. Comparison has also been made between pteridosperms and gymnosperms (conifers, ginkgo, and cycads，Figure 7). Gymnosperms, the dominant Mesozoic group, originate in the Early Permian and their diversity increases rapidly and exceeds that of the pteridosperms by the Changhsingian, and therefore the Carboniferous landscapes of exclusive pteridosperms were totally changed. 3.3 Comparisons with worldwide coeval plant diversity trends As is well known, the Carboniferous-Permian sees global phytogeographic differentiation of the North Temperate Angaran Realm, South Temperate Gondwanan Realm,
Figure 6 Diagram showing comparative diversities of spore plants and seed plants during the Carboniferous-Permian.
Figure 7 Diagram showing the comparative diversity of gymnosperms and pteridosperms during the Carboniferous-Permian.
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Tropical/subtropical Euramerican Realm and Cathaysian Realm. The different levels of study of individual phytogeographic realms hinder an accurate comparative study of Carboniferous-Permian floral diversities. In Rees’ [17] genera diversity study on every phytogeographic realm, data on South China and North China were counted separately. It suggested that Permian plant diversity of South China was much lower than that of North China, Angara, and Gondwana. Only in the Late Permian is the diversity of South China higher than that of Euramerica. It is worth noting that sources of Rees’ data need further examination and probably are not complete. Our newest statistics indicate that the generic diversity of Middle-Late Permian plant fossils of South China is more or less equal to that of North China, but their species diversities are quite different [5]. Both South China and North China belong to the sub-realm of Cathaysian flora. The Changhsingian flora of North China is comprised of abundant members of the Euramerican flora mixed with a few plants of the Angaran flora and some Middle Permian plants of the Cathaysian flora [18]. However, the Changhsingian flora of South China is still a part of the typical Cathayian flora [11].
4 Paleozoic plant diversity, evolution, and vegetation replacement of South China 4.1
Evolution mode
Based on our statistics on the Silurian-Early Triassic plant genera of South China, the characteristics of diversity trends could be summarized as below (Figure 8, blue curve): (1) Plant diversity keeps increasing from Silurian to Permian as a whole, climaxing in the Late Permian. (2) The Silurian is the earliest stage of terrestrial vascular plant evolution; plant diversity starts to form. (3) The Early Devonian to Early Carboniferous is the stage of early differentiation and diversification of plants, during which plants diversity fluctuates and plant types changes rapidly. This is the fastest development of plant types; as a result, plants can be found from all terrestrial ecosystems. (4) Plant diversity has a dramatic rise from the Early Permian, and reaches a peak in the Middle-Late Permian. (5) From the Late Permian to the Early Triassic, plant diversity decreases to 15% of that in the Late Permian. Significant changes occurred in the composition of the flora. Such a tremendous change was caused by the Permian-Triassic extinction event. 4.2
Pro-Cathaysian flora vegetation replacement
It is significant to analyze floras of different geological times but the same paleogeographic unit in terms of systematics and evolution of floras. Our data of the Silurian-Devonian flora of South China reveal a series of Pro-Cathaysian floras [19], which could be indicated by floras of three key geological periods: 1) the Early Devo-
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nian, when the Posongchong flora develops in Yunnan, characterized by the endemic members, different from coeval floras in terms of species and assemblages; 2) the late Middle Devonian, when the endemic Xichong flora of Yunnan develops with abundant lycopsids; 3) the late Late Devonian, when in the lower Yangtze River area, the development of the specialized and endemic Wutung flora was dominated by lycopsids and sphenopsids, with some taxa leading the late Middle Devonian flora. Although further study on floras of Pridoli, Emsian, Eifelian, and Frasnian are still needed, the Pro-Cathaysian flora could undoubtedly be traced back to the Early Devonian Posongchong flora (or earlier), which transforms through late Middle Devonian Xichong flora and Late Devonian Wutung flora and eventually forms the Carboniferous-Permian Cathaysian flora. 4.3
Evolution of Cathaysian flora
China is the center of origin and focus of the late Paleozoic Cathaysian flora, with representative members of gigantopterids, emplectopterids, lobatannularians, tingialeans, fascipterids, Conchophyllum, taeniopterids, and oriental lepidophytes. According to the origins and developments of these plant groups, it is widely accepted that the age of the typical Cathaysian flora is the late Early Carboniferous (Namurian) to latest Permian [20]. The flora prior to the Carboniferous is therefore dubbed Pro-Cathaysian [21]. Marine deposits develop from the Late Carboniferous to early Early Permian in South China, lacking plant fossil records, but a variety of abundant plant fossils could be found from the remaining sedimentary sequences. The flora evolution can be subdivided into following steps. 1) Tournaisian, lycopsids, and sphenopsids are prosperous, along with a few ferns/pteridosperms. 2) Visean, the main development of the flora; both the abundance and the diversity of every plant group increase. 3) Namurian (Serpukhovian), plant fossil records are few in the early stage, indicating a new rapid development of pteridosperms. However, poor preservation hinders recognition of the whole flora. From the Namurian to early Early Permian, although no records of the Cathaysian flora are found in South China, it probably is the key period to the evolution and the transformation of the Pro-Cathayian flora, because many pioneer members of the Cathaysian flora have been reported from North and Northeast China. 4) Chihsian (Artinskian-Kungurian), the flora becomes gradually dominated by ferns and pteridosperms, with some new members of Noeggerathiales. The members of the Cathaysian realm are dominant among the flora, accompanied by some members of the Euramerican flora. 5) Maokouan (Roadian-Wordian), the flora keeps rich, among which gigantopterids start a great developments and gradually become dominant. 6) Lungtanian (CapitanianWuchiapingian), the flora, enlarging to a large scale, shows a golden time of the Cathaysian flora. Gigantopterids are more common than ever; gymnosperms such as ginkgo,
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cycads and conifers occasionally occur. The flora is still dominated by ferns/pteridosperms. 7) Changhsingian, the composition of the flora is maintained but some typical members of the Cathaysian flora, such as gigantopterids, show a dramatic decline in abundance and diversity. Gymnosperms, including Ginkgo, cycads and conifers, develop to some extent. At the end of this period, the Cathaysian flora is basically finished.
5 Comparative analysis of diversity of late Paleozoic marine animal and vascular land plant 5.1
Comparison of general characteristics of diversity
Comparisons between the trends of diversity in marine animals [1] (Figure 8, red curve) and vascular plants (Figure 8, blue curve), suggest the following points. (1) Lochkovian and Pragian marine deposits are lacking
Figure 8 plants.
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in South China, and therefore the marine animal diversity of these periods is at its lowest point, being different from the coeval trends in a global context. The local and specialized environment of South China is caused by regional regression and provides favorable conditions for vascular land plants. The diversity of the plants soon reaches its second peak. (2) The transgression of South China in the mid-late Early Devonian leads the diversity of marine animals to reach the highest point in the late Paleozoic. Soon after that, there is a slight decline in the Eifelian, and a quick recovery in the Givetian. In the Emsian-Eifelian, the land plants are at the lowest generic diversity of the late Paleozoic, and their diversity is contrary to that of marine animals. In the Givetian, plant diversity soon recovers, resembling that of animal diversity. Although the number of plants is less in Givetian than in Pragian, the Givetian plant groups are diverse and occupy various land, and regional forests first occur.
Diversity of organisms at the generic level from Silurian to the Early Triassic in South China. 1. marine animals (after ref. [1]); 2. vascular land
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(3) Diversity of marine animals drops by 40%–50% in the Frasnian, which is a result of F/F extinction event. In the Famennian, diversity recovers to a high level. Although further evidence is needed in deciphering how much land-plants are affected by the F/F event, available data indicate that the land plant diversity declines by 50%, an extent greater than that of marine animals. (4) Regression may cause the Tournaisian marine animal diversity decline, as well as the larger land area from Famennian to Tournaisian. However, plant diversity of this period also decreases dramatically. We consider that the low diversity of Tournaisian plants is due to fossil record bias and inadequate study. (5) From the Carboniferous to the Permian, marine animal diversity keeps at a low level because of Gondwana glaciers and fluctuating global sea level. In the Middle-Late Permian, animal diversity keeps increasing except in the Capitanian, when a diversity decline is caused by a large scale of regression and the Emei Mountain basalt eruption event. In the Changhsingian, the animal diversity reaches a peak that is slightly lower than that in the Devonian, with a high succession rate. On the other hand, in the Carboniferous-Permian, land plants climax in both diversity and abundance for the whole Paleozoic, and are in a period of rapid development. The occasional diversity declines of plants are probably caused by the fossil record bias affected by transgression events, such as the Late CarboniferousEarly Permian plant fossil record loss caused by extensive transgression from the Serpukhovian. (6) An extinction event occurred at the end of the Permian in South China, in which nearly 90% marine animal genera died out. Such an extensive and severe marine extinction also caused extreme damage to the land ecosystem, vegetation. and plant diversity. Meanwhile, the fern-dominated vegetation was replaced by the gymnosperm/ angiosperm-dominated Mesozoic flora. The above comparisons between Paleozoic diversity of animals and plants suggest the co-evolution of animals and plants as well as the co-evolution of marine/terrestrial ecosystems. 5.2 Characteristics of plant diversity in the Devonian F/F extinction event of South China Land plant diversity decreases up to 50% after the Givetian and to its lower point in the late Frasnian-middle Famennian. At the same time, the differentiation of plants is quite low, until the end of the Famennian [22, 23]. Diversity of dispersed spores in the late Frasnian keeps increasing until the early Famennian [24]. The first plant diversity crisis occurs around Frasnian/Famennian boundary. During the F/F event, despite lower diversity, plants suffered no extinctions as experienced by marine animals. Plants have inner protective mechanisms of resistance to environmental change, such as spores, seeds, and rhizomes
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or roots that could survive partial death [25, 26]. Plants, compared to animals, are less limited by the scale of population in heredity and can easily be restored from a small population [26]. Nevertheless, plant evolution is still deeply affected by F/F event, after which a number of Carboniferous groups, like xeromorph and trailing sphynopsids, early ferns, and seed plants, start to emerge and become prosperous. 5.3 Characteristics of plant diversity at the P/T extinction event of South China Relative to the rest of China, more abundant Late Permian-Early Triassic plant fossils can be found from South China, among which western Guizhou and eastern Yunnan, with well-developed terrestrial Upper Permian deposits and abundant plant fossils, have been thought to be the key areas to study Permian-Triassic plant replacement [27–29]. Fifty-seven genera of plants have been reported from the Changhsingian, Late Permian (Figure 4), nine from the Induan, Early Triassic [30, 31], and twenty-eight from the Olenekian, Early Triassic [32, 33]. This indicates that during the P/T event, plant diversity drops dramatically, following the replacement from the Cathaysian flora of Paleophytic to Mesophytic floras. The former are dominated by ferns and pteridosperms, with accompanying lycopsids, sphenopsids and ginkgo, cycad and conifer gymnosperms. In the latter flora, conifers are mixed with a number of pteridosperms and ferns of new types.
6
Conclusions
(1) Paleozoic plant diversity in South China keeps rising as a whole. The Silurian is the earliest phase of land plant evolution, when plant diversity starts to form. The Early Devonian and the Early Carboniferous witness the fastest development of plant types. During that period, land plants differentiate and diversify, plant diversity fluctuates, rapid replacement of plant types occurs, and every terrestrial ecosystem is eventually occupied. From the Early Permian onwards, the quick development of plant diversity takes place, and reaches the highest point in the Middle Permian for the whole Paleozoic. (2) Comparisons between marine and terrestrial organisms of late Paleozoic indicate the co-evolution of animals and plant, as well as the co-evolution of the marine/terrestrial ecosystem. (3) The F/F event causes vascular land plants diversity to drop by 50%, a greater loss than that of marine animals. (4) The P/T Extinction caused a considerable loss in diversity of marine animals, as well as that of vascular land plants. Subsequently, the fern-dominated vegetation was replaced by the Mesozoic flora dominated by gymnosperms and angiosperms.
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(5) The pro-Cathaysian floral groups originated from the Early Devonian Posongchong Flora (or earlier), developed through the late Middle Devonian Xichong Flora and late Upper Devonian Wutung Flora, and eventually transformed into the Carboniferous-Permian Cathayian Flora peculiar for China. We express our gratitude to Christopher M. Berry with Cardiff Univerisyt (HK) for improving the English version of this manuscript. This study was supported by Chinese Academy of Sciences (Grant Nos. KZCX2-YW-105, KZCX2-YW-Q05-01), National Natural Science Foundation of China (Grant No. 40523004) and National Basic Research Program of China (Grant No. 2006CB806400).
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Rong J Y, Fan J X, Li G X. Patterns of latest Proterozoic to Early Mesozoic marine biodiversity changes in South China. In: Rong J Y, Fang Z J, Zhou Z H, et al, eds, Originations, Radiations and Biodiversity Changes—Evidences from the Chinese Fossil Record (in Chinese with English summary). Beijing: Science Press, 2006. 785–816, 960–962 Knoll A H, Niklas K J, Gensel P G, et al. Character diversification and patterns of evolution in early vascular plants. Paleobiology, 1984, 10: 34–47 Wu X Y. Carboniferous floras. In: Li X X, Zhou Z Y, Cai C Y, et al, eds. Fossil Floras of China Through the Geological Ages. Guangzhou: Guangdong Science and Technology Press, 1995. 78–126 Shen G L. Permian floras. In: Li X X, Zhou Z Y, Cai C Y, et al, eds. Fossil Floras of China Through the Geological Ages. Guangzhou: Guangdong Science and Technology Press, 1995. 127–223 Tian B L, Wang S J, Li C S, et al. An approach on the origin center, evolution center and the mechanics of evolution and extinction of the Late Palaeozoic Cathaysian flora (in Chinese with English summary). Chin Bull Bot, 2000, 17: 21–33 Cai C Y, Li X X. A review of Silurian and Devonian macrofloras in China. Palaeontol Cath, 1994, 6: 167–214 Cai C Y, Wang Y. Devonian floras. In: Li X X, Zhou Z Y, Cai C Y, et al, eds. Fossil Floras of China Through the Geological Ages. Guangzhou: Guangdong Science and Technology Press, 1995. 28–77 Cai C Y. The nonmarine Devonian System. In: Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, ed. Stratigraphical Studies in China (1979–1999) (in Chinese). Hefei: University of Science and Technology of China Press, 2000. 95–127 Hao S G, Gensel P G. The Posongchong floral assemblages of southeastern Yunnan, China—Diversity and disparity in Early Devonian plant assemblages. In: Gensel P G, Edwards D, eds. Plants Invade the Land—Evolutionary & Environmental Perspectives. New York: Columbia University Press, 2001. 103–119 Wang Y, Hao S G, Fu Q, et al. Diversity of Early Land Vascular Plants in the Silurian and Devonian of China. In: Rong J Y, Fang Z J, Zhou Z H, et al, eds. Originations, Radiations and Biodiversity Changes—Evidence from the Chinese Fossil Record (in Chinese with English summary). Beijing: Science Press, 2006. 383–398, 882–884 Liu L J, Yao Z Q. Plant megafossil from the Permian Changhsingian marine deposits of Fusui, Guangxi, China (in Chinese with English summary). Acta Palaeontol Sin, 2007, 46: 195–212 Wu X Y, Wang J, Zhan Y. Characeteristic of Pre-Late Carboniferous flora of China: A note on Pro-Cathaysian flora (in Chinese with English summary). Acta Palaeontol Sin, 2008, 47: 1–20 Wang S J, Sun K Q, Cui J Z, et al. Fossil Flora of China, Volume 1: Fossil Plants from Coal Balls in China (in Chinese with English summary). Beijing: Higher Education Press, 2009. 222 Knoll A H, Niklas K J, Gensel P G, et al. Phanerozoic land plant diversity in North America. Science, 1979, 206: 1400–1402
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22 23
24
25
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28
29
30
31
32
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Wang J, Pfefferkorn H W. Nystroemiaceae, a new family of Permian gymnosperms from China combining primitive with highly advanced features. Proc R Soc B, 2010, 277: 301–309 Wang J, Pfefferkorn H W, Bek J. Paratingia wudensis sp. nov., a whole noeggerathialean plant preserved in an air fall tuff of earliest Permian age (Inner Mongolia, China). Amer J Bot, 2009, 96: 1676–1689 Rees P M. Land-plant diversity and the end-Permian mass extinction. Geology, 2002, 30: 827–830 Wang Z Q, Wang L X. Late Permian fossil plants from the lower part of the Shiqianfeng (Shihchienfeng) Group in North China (in Chinese). Bull Tianjin Inst Geol Min Res Chin Acad Geol Sci, 1986, 15: 1–80 Wang Y, Berry C M, Hao S G, et al. The Xichong flora of Yunnan, China: Diversity in late Middle Devonian plant assemblages. Geol J, 2007, 42: 339–350 Li X X, Shen G L, Tian B L, et al. Some notes on Carboniferous and Permian floras in China. In: Li X X, Zhou Z Y, Cai C Y, et al, eds. Fossil Floras of China Through the Geological Ages. Guangzhou: Guangdong Science and Technology Press, 1995. 244–304 Li X X, Wu X Y, Shen G L, et al. The Namurian and Its Biota in the East Sector of North Qilian Mountain (in Chinese with English summary). Jinan: Shandong Science and Technology Publishing House, 1993. 482 Raymond A, Metz C. Land plants and the Frasian-Famennian extinction event. Geol Soc Am Abs Prog, 1992, 24: A271 Raymond A, Metz C. Laurussian land-plant diversity during the Silurian and Devonian: Mass extinction, sampleing bias, or both? Paleobiology, 1995, 21: 74–91 Raymond A. Terrestrial-marine links in the Frasnian-Famennian extinction event (abstracts). Seattle Annual Meeting (November 2–5, 2003), Paper No.188-8. Seattle, Washington, 2003 Knoll A H. Patterns of extinction in the fossil record of vascular plants. In: Nitecki M H, ed. Extinctions. Chicago: University of Chicago Press, 1984. 21–68 Traverse A. Plant evolution dances to a different beat. Histor Biol, 1988, 1: 277–301 Yao Z Q, Xu J T, Zheng Z G, et al. The Late Permian biostratigraphy and problem of Permo-Triassic boundary in W. Guizhou and E. Yunnan. In: Nanjing Institute of Geology and Palaeontology, Academia Sinica, ed. Stratigraphy and Palaeontology of the Permian Coal Measures of W. Guizhou and E. Yunnan (in Chinese). Beijing: Science Press, 1980. 1–69 Zhao X H, Mo Z G, Zhang S Z, et al. Late Permian flora from W. Guizhou and E. Yunnan. In: Nanjing Institute Geology and Palaeontology, Academia Sinica, ed. Stratigraphy and Palaeontology of Upper Permian Coal Measures W. Guizhou and E. Yunnan (in Chinese). Beijing: Science Press, 1980. 70–99 Wang S Y, Yin H F. Study on Terrestrial Permian-Triassic Boundary in Eastern Yunnan and Western Guizhou (in Chinese with English abstract). Wuhan: China University of Geosciences Press, 2001. 88 Yao Z Q. On the age of “Gigantopteris Caol Series” and Gigantopteris-flora in South China (in Chinese with English abstract). Acta Palaeontol Sin, 1978, 17: 81–89 Yu J X, Huang Q S, Broutin J, et al. The early Early Triassic Annalepis from western Guizhou and eastern Yunnan, South China (in Chinese with English abstract). Acta Palaeontol Sin, 2008, 47: 292–300 Zhou Z Y, Li B X. A preliminary study of the Early Triassic plants from the Qionghai district, Hainan Island (in Chinese with English summary). Acta Palaeontol Sin, 1979, 18: 444–462 Meng F S. The fossil plants of the Badong Formation. In: Meng F S, Xu A W, Zhang Z L, et al, eds. Nonmarine Biota and Sedimentary Facies of the Badong Formation in the Yangtze and Its Neighbouring Areas (in Chinese). Wuhan: China University of Geosciences Press, 1995. 6–27
