Abstract Much progress has been made in the understanding of the phylogeny of Silurian-Early Devonian vascular land plants during the past years. However,no consensus has been reached considering the delimitation and subdivision of polysporangiophytes,rhyniopsids,and zosterophyllopsids;the sister-group relationship between zosterophyllopsids and lycopsids is questioned;detailed studies of basal euphyllophytes are needed. The evolutionary history of the Silurian-Early Devonian vascular plants can be divided into two phases:The pre-Pragian prelude phase and the Early Devonian Pragian explosion(burst)phase. The first phase is characterized by initial emergences of representatives of vascular plant major clades. The second is the main phase which witnessed explosive radiation of vascular plant clades. During the Pragian,typical features of lycopsids such as regular phyllotaxy,sporophylls,and strobilus-like structures appeared;the diversification of euphyllophyte lineages occurred;megaphylls and their homologous structures(branch-leaf complexes)evolved multiple times independently;and developmental interrelationship between sporangia and leafy structures convergently established in different lineages. In South China of the northeastern Gondwanan phytogeographic unit,zosterophyllopsids were dominated by those with spirally arranged sporangia and euphyllophytes showed high diversity and disparity during the Pragian,while in the Laurussian phytogeographic unit,zosterophyllopsids with rowed sporangia were abundant and the appearance of comparable euphyllophytes occurred later than in South China. The cladogenesis of horneophytopsids,cooksonioids,and rhynialeans might have mainly occurred at middle and high paleolatitudes such as southern Laurussia and northwestern Gondwana. The cladogenesis and biogeographic radiation of zosterophyllopsids,lycopsids,and euphyllophytes may accord with the “out of the tropics”model,that is,these groups diversified and rapidly radiated earlier at low paleolatitudes;the explosion of vascular plant clades in Pragian was mainly contributed by the evolutionary novelty of tropical floras.
About author: Xue Jinzhuang,born in 1981,is an associate professor of Peking University. His current research focuses on the evolution of early vascular plants. E-mail: pkuxue@pku.edu.cn.
Cite this article:
Xue Jinzhuang,Hao Shougang. Phylogeny, episodic evolution and geographic distribution of the Silurian-Early Devonian vascular plants: Evidences from plant megafossils[J]. JOPC, 2014, 16(6): 861-877.
Xue Jinzhuang,Hao Shougang. Phylogeny, episodic evolution and geographic distribution of the Silurian-Early Devonian vascular plants: Evidences from plant megafossils[J]. JOPC, 2014, 16(6): 861-877.
布科,陈旭,Scotese C R,等. 2009. 显生宙全球古气候重建[M]. 北京:科学出版社, 1-173. 蔡重阳,王怿. 1995. 泥盆纪植物群[A]. 见:李星学主编. 中国地质时期植物群[M]. 广东广州: 广东科技出版社, 22-57. 郝守刚,王德明. 2003. 中国云南早泥盆世坡松冲植物群:探究早期陆生维管植物演化分异的窗口[J]. 地球科学进展,18(6):877-883. 郝守刚,王德明,王祺. 2002. 陆生植物的起源和维管植物的早期演化[J]. 北京大学学报(自然科学版),38(2):286-293. 李承森. 1982. 徐氏蕨属:中国云南早泥盆世陆地植物的一个新属[J]. 植物分类学报,20(3):331-342. 李星学,蔡重阳. 1977. 中国西南地区早泥盆世工蕨化石:中国西南部早泥盆世植物群研究之一[J]. 古生物学报,16(1):12-36. 耿宝印. 1983. 云南早泥盆世穗蕨属(新属)的发现及意义[J]. 植物学报,25(6):574-579. 耿宝印. 1986. 贵州中志留世羽枝属(新属)的形态和解剖[J]. 植物学报,28(6):664-672. 王德明,熊聪慧. 2014. 中国早期陆生维管植物演化、古地理和古环境研究进展[J]. 古生物学报,53(1):101-107. 王怿,蔡重阳. 1996. 滇东南早泥盆世云南穗蕨(Stachyophyton yunnanense Geng)的再研究[J]. 古生物学报,35(1):99-108. 王怿,徐洪河. 2009. 中国志留纪陆生植物研究综述[J]. 古生物学报,48(3):453-464. 王怿,郝守刚,傅强,等. 2006. 中国志留—泥盆纪早期陆生维管植物多样性[A]. 见:戎嘉余(主编). 生物的起源、辐射与多样性演变:华夏化石记录的启示[M]. 北京:科学出版社, 383-398. 王祺,徐洪河. 2011. 志留纪晚期拟莱尼蕨植物Junggaria(准噶尔蕨属)的命名问题[J]. 古生物学报,50(3):326-329. 徐仁. 1966. 云南泥盆纪植物化石和其在该区泥盆系地层划分上的意义[J]. 植物学报,14(1):50-69. 薛进庄,郝守刚. 2008. 泥盆纪枝蕨纲的分支系统学研究[J]. 北京大学学报(自然科学版),44(2):299-307. 薛进庄,黄璞,王祺. 2014. Monilophytes(广义真蕨类)的概念溯源及中文定名[J]. 植物学研究,3:23-26. 杨楠,李承森. 2009. 中国泥盆纪维管植物的组成与古地理分布[J]. 古地理学报,11(1):91-104. Andrews H N,Gensel P G,Kasper A E. 1975. A new fossil plant of probable intermediate affinities(trimerophyte-progymnosperm)[J]. Canadian Journal of Botany,53:1719-1728. Banks H P. 1968. The early history of land plants[A]. In:Drake E T(ed). Evolution and environment:A symposium presented on the one hundredth anniversary of the foundation of Peabody Museum of Natural History at Yale University[C]. New Haven:Yale University Press. 73-107. Banks H P. 1975. Reclassification of Psilophyta[J]. Taxon,24:401-413. Bateman R M,Crane P R,Dimichele W A, et al. 1998. Early evolution of land plants:Phylogeny,physiology,and ecology of the primary terrestrial radiation[J]. Annual Review of Ecology and Systematics,29:263-292. Beck C B,Stein W E. 1993. Crossia virginiana gen. et sp. nov.,a new member of the Stenokoleales from the Middle Devonian of southwestern Virginia[J]. Palaeontographica Abteilung B-Palaophytologie,229:115-134. Berry C M,Stein W E. 2000. A new iridopteridalean from the Devonian of Venezuela[J]. International Journal of Plant Sciences,161:807-827. Bonamo P M,Banks H P,Grierson J D. 1988. Leclercqia,Haskinsia,and the role of leaves in delineation of Devonian lycopsid genera. Botanical Gazette,149:222-239. Cai C Y,Ouyang S,Wang Y, et al. 1996. An early Silurian vascular plant[J]. Nature,379:592-592. Cascales-Miana B,Meyer-Berthaud B. 2013. Diversity dynamics of Zosterophyllopsida[J]. Lethaia,DOI:10.1111/let.12051. Crane P R,Herendeen P,Friis E M. 2004. Fossils and plant phylogeny[J]. American Journal of Botany,91:1683-1699. Dawson J W. 1859. On fossil plants from the Devonian rocks of Canada[J]. Quarterly Journal of the Geological Society of London,15:477-488. Edwards D. 1990. Constraints on Silurian and early Devonian phytogeographic analysis based on megafossils[A]. In:McKerrow W S,Scotese C R(eds). Palaeozoic Palaeogeography and Biogeography[C]. Geological Society Memoirs 12:233-242. Edwards D. 2003. Xylem in early tracheophytes[J]. Plant,Cell and Environment(2003)26,57-72. Edwards D. 2004. Embryophytic sporophytes in the Rhynie and Windyfield cherts[J]. Transactions of the Royal Society of Edinburgh:Earth Sciences,94: 397-410. Edwards D,Berry C M. 1991. Silurian and Devonian[A]. In: Cleal C J. Plant Fossils in Geological Investigation: The Palaeozoic[C]. New York: Ellis Horwood, 117-153. Edwards D,Davies M S. 1990. Interpretations of early land plant radiations:“facile adaptationist guesswork” or reasoned speculation?[A]. In: Taylor P D,Larwood G P(eds). Major Evolutionary Radiations[C]. Oxford:Clarendon Press, 351-376. Edwards D,Edwards D S. 1986. A reconsideration of the Rhyniophytina Banks[A]. In: Spicer R A,Thomas B A(eds). Systematic and Taxonomic Approaches in Palaeobotany[C]. Oxford: Clarendon Press, 199-220. Edwards D,Feehan J. 1980. Records of Cooksonia-type sporangia from Late Wenlock Strata in Ireland[J]. Nature,287:41-42. Edwards D,Wellman C H. 2001. Embryophytes on land:The Ordovician to Lochkovian(Lower Devonian)record[A]. In: Gensel P G,Edwards D(eds). Plants Invade the Land:Evolutionary and environmental perspectives[C]. New York:Columbia University Press, 3-28. Edwards D,Wellman C H. 2004. Silurian and Lower Devonian plant assemblages from the Anglo-Welsh Basin:A palaeobotanical and palynological synthesis[J]. Geological Journal,39:375-402. Edwards D,Davies K L,Axe L. 1992. A vascular conducting strand in the early land plant Cooksonia[J]. Nature,357:683-685. Edwards D,Wang Y,Bassett M G, et al. 2007. The earliest vascular plant or a later rooting system?Pinnatiramosus qianensis from the marine lower Silurian Xiushan Formation,Guizhou Province,China[J]. Palaios,22:155-165. Edwards D S. 1986. Aglaophyton major,a non-vascular land plant from the Devonian Rhynie Chert[J]. Botanical Journal of the Linnean Society,93:173-204. Gensel P G. 1992. Phylogenetic relationships of the zosterophylls and lycopsids:Evidence from morphology,paleoecology,and cladistic methods of inference[J]. Annals of the Missouri Botanical Garden,79:450-473. Gensel P G. 2008. The earliest land plants[J]. Annual Review of Ecology Evolution and Systematics,39:459-477. Gensel P G,Berry C M. 2001. Early lycophyte evolution[J]. American Fern Journal,91:74-98. Gensel P G,Kasper A E. 2005. A new species of the Devonian lycopod genus,Leclercqia,from New Brunswick,Canada[J]. Review of Palaeobotany and Palynology,136:105-123. Gerrienne P,Dilcher D L,Bergamaschi S, et al. 2006. An exceptional specimen of the early land plant Cooksonia paranensis,and a hypothesis on the life cycle of the earliest eutracheophytes[J]. Review of Palaeobotany and Palynology,142:123-130. Gerrienne P,Gensel P G,Strullu-Derrien C, et al. 2011. A simple type of wood in two Early Devonian plants[J]. Science,333:837-837. Gerrienne P,Meyer-Berthaud B,Fairon-Demaret M,et al. 2004. Runcaria,a Middle Devonian seed plant precursor[J]. Science,306:856-858. Gonez P,Gerrienne P. 2010a. A new definition and a lectotypification of the genus Cooksonia Lang 1937[J]. International Journal of Plant Sciences,171:199-215. Gonez P,Gerrienne P. 2010b. Aberlemnia caledonica gen. et comb. nov.,a new name for Cooksonia caledonica Edwards 1970[J]. Review of Palaeobotany and Palynology,163:64-72. Gonez P,Huu H N,Hoa P T, et al. 2012. The oldest flora of the South China Block,and the stratigraphic bearings of the plant remains from the Ngoc Vung Series,northern Vietnam. Journal of Asian Earth Sciences,43:51-63. Gray J. 1993. Major Paleozoic land plant evolutionary bio-events[J]. Palaeogeography,Palaeoclimatology,Palaeoecology,104:153-169. Hao S G, Beck C B. 1991. Yunia dichotoma, a Lower Devonian plant from Yunnan, China[J]. Review of Palaeobotony and Palynology, 68:181-195. Hao S G,Gensel P G. 1998. Some new plant finds from the Posongchong Formation of Yunnan,and consideration of a phytogeographic similarity between south China and Australia during the Early Devonian[J]. Science in China Series D-Earth Sciences,41:1-13. Hao S G,Gensel P G. 2001. The Posongchong floral assemblages of southeastern Yunnan,China: Diversity and disparity in Early Devonian plant assemblages[A]. In:Gensel P G,Edwards D(eds). Plants Invade the Land: Evolutionary and Environmental Perspectives[C]. New York: Columbia University Press, 103-119. Hao S G,Xue J Z. 2011. A new zosterophyll plant,Ramoferis gen. nov.,from the Posongchong Formation of Lower Devonian(Pragian)of Southeastern Yunnan,China[J]. Acta Geologica Sinica-English Edition,85:765-776. Hao S G,Xue J Z. 2013a. The Early Devonian Posongchong Flora of Yunnan-A Contribution to an Understanding of the Evolution and Early Diversification of Vascular Plants[M]. Beijing: Science Press, 1-366. Hao S G,Xue J Z. 2013b. Earliest record of megaphylls and leafy structures,and their initial diversification. Chinese Science Bulletin,58:2784-2793. Hao S G,Wang D M,Beck C B. 2003. Observations on anatomy of Adoketophyton subverticillatum from the Posongchong Formation(Pragian,Lower Devonian)of Yunnan,China[J]. Review of Palaeobotany and Palynology,127:175-186. Hao S G,Wang D M,Wang Q, et al. 2006. A new lycopsid,Zhenglia radiata gen. et sp. nov.,from the Lower Devonian Posongchong Formation of southeastern Yunnan,China,and its evolutionary significance[J]. Acta Geologica Sinica-English Edition,80:11-19. Hao S G,Xue J Z,Guo D L, et al. 2010. Earliest rooting system and root:Shoot ratio from a new Zosterophyllum plant[J]. New Phytologist,185:217-225. Hao S G,Xue J Z,Liu Z F, et al. 2007. Zosterophyllum Penhallow around the Silurian-Devonian boundary of northeastern Yunnan,China[J]. International Journal of Plant Sciences,168:477-489. Hao S G,Xue J Z,Zhu X, et al. 2012. A new genus of Early Devonian plants with novel strobilar structures and vegetative appendages from the Posongchong Formation of Yunnan,China[J]. Review of Palaeobotany and Palynology,171:73-82. Hillebrand H. 2004. On the generality of the latitudinal diversity gradient[J]. The American Naturalist,163:192-211. Hueber F M. 1983. A new species of Baragwanathia from the Sextant Formation(Emsian),Northern Ontario,Canada[J]. Botanical Journal of the Linnean Society,86:57-79. Hueber F M. 1992. Thoughts on the early lycopsids and zosterophylls[J]. Annals of the Missouri Botanical Garden,79:474-499. Hueber F M,Banks H P. 1979. Serrulacaulis furcatus gen. et sp. nov.,a new zosterophyll from the Lower Upper Devonian of New York State[J]. Review of Palaeobotany and Palynology,28:169-189. Jablonski D. 1993. The tropics as a source of evolutionary novelty through geological time[J]. Nature,364:142-144. Jablonski D,Roy K,Valentine J W. 2006. Out of the Tropics:Evolutionary dynamics of the latitudinal diversity gradient[J]. Science,314:102-106. Kenrick P. 2000. The relationships of vascular plants[J]. Philosophical Transactions of the Royal Society B:Biological Sciences,355:847-855. Kenrick P,Crane P R. 1991. Water-conducting cells in early fossil land plants:Implications for the early evolution of tracheophytes[J]. Botanical Gazette,152:335-356. Kenrick P,Crane P R. 1997. The rigin and Early Diversification of Land Plants,a Cladistic Study[M]. Washington:为Smithsonian Institution Press, 1-441. Kenrick P,Wellman C H,Schneider H, et al. 2012. A timeline for terrestrialization:Consequences for the carbon cycle in the Palaeozoic[J]. Philosophical Transactions of the Royal Society B-Biological Sciences,367:519-536. Kerp H,Hass H,Mosbrugger V. 2001. New data on Nothia aphylla Lyon 1964 ex El-Saadawy et Lacey 1979,a poorly known plant from the Lower Devonian Rhynie Chert[A]. In: Gensel P G,Edwards D(eds). Plants Invade the Land:Evolutionary and Environmental Perspectives[C]. New York: Columbia University Press, 52-82. Kidston R,Lang W H. 1917. On Old Red Sandstone plants showing structure,from the Rhynie Chert Bed,Aberdeenshire. Part 1. Rhynia gwynne-vaughani Kidston and Lang[J]. Transactions of the Royal Society of Edinburgh,51:761-784. Kidston R,Lang W H. 1920. On Old Red Sandstone plants showing structure,from the Rhynie Chert Bed,Aberdeenshire. Part Ⅱ. Additional notes on Rhynia gwynne-vaughani,Kidston and Lang;with descriptions of Rhynia major,n.sp. and Hornea lignieri,n.g.,n.sp.[J]Transactions of the Royal Society of Edinburgh, 52:603-627. Knoll A H,Niklas K J,Gensel P G, et al. 1984. Character diversification and patterns of evolution in early vascular plants[J]. Paleobiology,10:34-47. Kotyk M E,Basinger J F. 2000. The Early Devonian(Pragian)zosterophyll Bathurstia denticulata Hueber[J]. Canadian Journal of Botany,78:193-207. Kotyk M E,Basinger J F,Gensel P G, et al. 2002. Morphologically complex plant macrofossils from the Late Silurian of Arctic Canada[J]. American Journal of Botany,89:1004-1013. Lang W H,Cookson I C. 1935. On a flora,including vascular land plants,associated with Monograptus,in rocks of Silurian Age,from Victoria,Australia[J]. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences,224:421-449. Li C S. 1992. Hsüa robusta,an Early Devonian plant from Yunnan Province,China and its bearing on some structures of early land plants. Review of Palaeobotany and Palynology,71:121-147. Li C S,Edwards D. 1992. A new genus of early land plants with novel strobilar construction from the Lower Devonian Posongchong Formation,Yunnan Province,China[J]. Palaeontology,35:257-272. Li C S,Edwards D. 1995. A reinvestigation of Halles Drepanophycus spinaeformis Gpp. from the Lower Devonian of Yunnan Province,Southern China[J]. Botanical Journal of the Linnean Society,118:163-192. Li C S,Edwards D. 1997. A new microphyllous plant from the Lower Devonian of Yunnan Province,China[J]. American Journal of Botany,84:1441-1448. Lyon A G. 1964. Probable fertile region of Asteroxylon mackiei K. and L.[J]. Nature,203:1082-1083. Morris J L,Richardson J B,Edwards D. 2011. Lower Devonian plant and spore assemblages from Lower Old Red Sandstone strata of Tredomen Quarry,South Wales[J]. Review of Palaeobotany and Palynology,165:183-208. Niklas K J, Banks H P. 1990. A reevaluation of theZosterophyllophytina with comments on the origin of lycopods[J]. American Journal of Botony, 77:274-283. Niklas K J,Tiffney B H,Knoll A H. 1983. Patterns in vascular land plant diversification[J]. Nature,303:614-616. Pryer K M,Schneider H,Smith A R, et al. 2001. Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants[J]. Nature,409:618-622. Pryer K M,Schneider H,Zimmer E A, et al. 2002. Deciding among green plants for whole genome studies[J]. Trends in Plant Science,7:550-554. Raymond A. 1987. Paleogeographic distribution of Early Devonian plant traits[J]. Palaios,2:113-132. Raymond A,Gensel P,Stein W E. 2006. Phytogeography of Late Silurian macrofloras[J]. Review of Palaeobotany and Palynology,142:165-192. Raymond A,Parker W C,Barrett S F. 1985. Early Devonian phytogeography[A]. In: Tiffney B H(ed). Geological Factors and the Evolution of Plants[C]. New Haven,CN: Yale University Press, 129-167. Scheckler S E,Skog J E,Banks H P. 2006. Langoxylon asterochlaenoideum Stockmans:Anatomy and relationships of a fern-like plant from the Middle Devonian of Belgium[J]. Review of Palaeobotany and Palynology,142:193-217. Taylor T N,Taylor E L,Krings M. 2009. Paleobotany: The Biology and Evolution of Fossil Plants(Second edition)[M]. Amsterdam: Academic Press, 1-1230. Tims J D,Chambers T C. 1984. Rhyniophytina and Trimerophytina from the early land flora of Victoria,Australia[J]. Palaeontology,27:265-279. Wang D M. 2007. Two species of Zosterophyllum from south China and application of a biostratigraphic method to an Early Devonian flora[J]. Acta Geologica Sinica-English Edition,81:525-538. Wang D M. 2008. A new iridopteridalean plant from the Middle Devonian of Northwest China[J]. International Journal of Plant Sciences,169:1100-1115. Wang D M,Hao S G. 2002. Guangnania cuneata gen. et sp. nov. from the Lower Devonian of Yunnan Province,China[J]. Review of Palaeobotany and Palynology,122:13-21. Wang D M,Hao S G. 2004. Bracteophyton variatum gen. et sp. nov.,an Early Devonian plant from the Xujiachong Formation of Yunnan,China[J]. International Journal of Plant Sciences,165:337-345. Wang D M,Hao S G,Wang Q. 2003. Tracheid ultrastructure of Hsüa deflexa from the Lower Devonian Xujiachong Formation of Yunnan,China[J]. International Journal of Plant Sciences,164:415-427. Wang Y, Edwards D, Bassett M, et al. 2013. Enigmatic occurrence of Permian plant roots in Lower Silurian rocks, Guizhou Province, China[J]. Palaeontology, 56:679-683. Wang Y,Fu Q,Xu H H, et al. 2007. A new Late Silurian plant with complex branching from Xinjiang,China[J]. Alcheringa,31:111-120. Wang Y,Hao S G,Cai C Y, et al. 2006. A diminutive plant from the Late Silurian of Xinjiang,China[J]. Alcheringa,30:23-31. Wang Y, Wang J, Xu H H, et al. 2010. The evolution of Paleozoic vascular land plant diversity of South China [J]. Science China-Earth Sciences, 53:1828-1835. Xiong C H,Wang D M,Wang Q, et al. 2013. Diversity Dynamics of Silurian-Early Carboniferous Land Plants in South China[J]. PLoS ONE,8:e75706. Xu H H,Berry C M,Wang Y, et al. 2011. A new species of Leclercqia Banks,Bonamo et Grierson(Lycopsida)from the Middle Devonian of North Xinjiang,China,with a possible climbing habit[J]. International Journal of Plant Sciences,172:836-846. Xu H H,Feng J,Jiang Q, et al. 2013. Report of Drepanophycus Gppert(Lycopsida)from the Middle Devonian of Xinjiang,China[J]. Journal of Systematics and Evolution,51:765-772. Xu H H,Xue J Z,Wang Q. 2014. Notes on a fertile rhyniophytoid from the Lower Devonian of Guangxi,southwestern China[ ]. Historical Biology,http:∥dx.doi.org/10.1080/08912963.2014.910205 Xue J Z. 2012. Lochkovian plants from the Xitun Formation of Yunnan,China,and their palaeophytogeographical significance[J]. Geological Magazine,149:333-344. Xue J Z. 2013. New material of Hueberia zhichangensis YANG,LI & EDWARDS,a basal lycopsid from the Early Devonian of Yunnan,China[J]. Neues Jahrbuch für Geologie und Palontologie,Abhandlungen,267:331-339. Xue J Z,Hao S G,Basinger J F. 2010. Anatomy of the Late Devonian Denglongia hubeiensis,with a discussion of the phylogeny of the Cladoxylopsida[J]. International Journal of Plant Sciences,171:107-120. Yang N,Li C S,Edwards D. 2009. Hueberia zhichangensis gen. et sp. nov., an Early Devonian(Pragian)plant from Yunnan,China[J]. Palynology,33:113-124.