吉林延边地区早白垩世西山坪组的铁角蕨属
摘要
本文对吉林省东部延边地区早白垩世含煤地层西山坪组的铁角蕨属进行了详细的形态学研究,并结合繁殖器官等特征,鉴定、描述了铁角蕨属化石2种,Asplenium tiefanum Deng和Asplenium popovii Samylina,并建立1新种Asplenium changcaium Sun et Liu (sp. nov.)。
在查阅大量铁角蕨属化石资料基础上,按照不同的时代和层位,对铁角蕨化石进行了地理分布研究,提出铁角蕨属可能起源于早白垩世中期,起源地点为东北亚(我国东北、俄罗斯以及日本)温暖潮湿气候区。首先在欧亚大陆广布,之后经古白令陆桥到现今北美等高纬度地区。随着晚白垩世全球气温降低,温暖潮湿气候区范围扩大,铁角蕨属分布范围也逐渐扩散到南半球。与此同时,在扩散的过程中,铁角蕨属逐渐适应新的环境,演化出不同的种类,分布范围也遍布全球。
化石铁角蕨属中,通过对A. changcaium和A. popovii原位孢子形态和大小的研究,并结合蕨叶形态,与现生铁角蕨属(孢子和叶型)进行了系统比较。在孢子形态和大小方面,A. changcaium孢子形态变化与铁角蕨属铁角蕨组中铁角蕨系的孢子形态变化有共性,其极轴稳定,赤道轴随成熟度增加而变大;在叶型方面,A. changcaium与铁角蕨属铁角蕨组的隆脉系和变异叶系最为相近。A. popovii的孢子比现生铁角蕨属的孢子略大,与铁角蕨属单叶组以及铁角蕨组的隆脉系的孢子大小较为接近;在叶型方面,该种同铁角蕨属铁角蕨组变异叶系中的东海铁角蕨最为相似。因而,推断现生铁角蕨属铁角蕨组与A. changcaium、A. popovii关系最为密切。
The early Cretaceous coal-bearing strata overlying on Tuntianying Formation in Yanbian region of eastern Jilin are composed of two sedimentation cycles, and can be divided into two formations, i.e. Xishangping Formation and Changcai Fomation. The lower sedimentary cycle is named as Xishanping Formation, bearing thin coal seams, while the upper one is called Changcai Formation, bearing thick coal beds. The two formations are divided by the sandy conglomerate on the bottom of the Changcai Formation which shows parallel unconformable contact relationships between them. This suit of coal-bearing strata is distributed mainly in Laotougou and Tuntianying of Yanji County; Changcai, Fudong, Nanyang, Tushanzi, Songxiaping and Dajinchang of Helong County in the eastern Jilin. It is an important coal-bearing strata consisting mainly of array sandstone, sandy conglomerate and gray-green conglomerate intercalated with black shale, carbonaceous mudstone and coal beds, yielding abundant fossil plants.
Since 1950’s, 42 species of 11 genera of ferns have been recorded from the Changcai and Xishanping formations inYanbian region (Yang, 1959; Zhou, et al., 1980; Shang, 1992; Li, 1995, etc.), based on the gross morphology. The present paper deals mainly with the fossil Asplenium found from the Lower Cretaceous Xishanping Formation in Yanbian area.
Based on the gross morphology and the fertile organ and spores in situ which plays an important role in identification and phylogeny, three species of Asplenium, i.e. Asplenium changcaium Sun et Liu (sp. nov.), Asplenium tiefanum Deng and Asplenium popovii Samylina and are recognized for the first time.
On the sterile fronds, the present new species of Asplenium, Asplenium changcaium Sun et Liu (sp. nov.) is different from the known species of fossil Asplenium by having the specialized oval segments on acroscopic side of pinnule and 2-3 times forking lateral veins. For the fertile fronds, Asplenium changcaium Sun et Liu (sp. nov.) is also distinguishing from the known fossil Asplenium by the small elliptical sori attached longintudinally at the middle part of veins on every segments of the pinnules. With regard to the monolete spores in situ, Asplenium changcaium Sun et Liu (sp. nov.) is characterized by having short polar axies and fabaceous shape (lateral view) which is quite different from the spores in situ of Asplenium tiefanum Deng and A. popovii Samylina.
Based upon studies of fossil records of Asplenium and the distribution of fossil and living Asplenium, the place and time of origin and the migration route of Asplenium are discussed preliminarily, concerning on phylogeny, the paleoclimatology and plate tectonic theory. This provides a new basement for the further researches on Asplenium.
The lowest horizon of reliable appearance of Asplenium in the world might be the middle part of the early Cretaceous (i.e. Hauterivian, Barremian and Aptian) in NE China, Siberian and central Honshu of Japan. During that time, the Northeast Asia was in the warm and humid climate zone and might be the original places of Asplenium in the middle of early Cretaceous. Later on, Asplenium dispersed widely on the Eurasian. During the late Cretaceous, Asplenium was already living on the middle Europe, accompanied by the northward migration of warm and humid climate zone. At the same time, the Asplenium occupied the high altitude region of North American (warm and humid climate in the Cretaceous and early Tertiary) via Bering Land Bridge. In the early late Cretaceous, Asplenium might spread to the South Hemisphere due to global climate cooling and the extension of warm and humid climate zone. With the radiation and dispersal, Asplenium diversified, adaped to different environmentsits, and gradually distributed worldwide.
Systemic comparision between the fossil Asplenium changcaium, A. popovii and the living Asplenium is done in the aspects of shape and size of spores in situ and the gross morphology of leaves.
For the shape and size of spores in situ, A. changcaium is resembled to the living Ser. Trichomanoidea of Sect. Asplenium, the length of polar axis changes little, while the equatorial axis varied greatly with the maturity of fossil and living ones. The spores of A. popovii is slightly bigger than those of the living Asplenium and close to those of the living Sect. Holophyllum and Ser. Falcata of Sect. Asplenium in size. On the gross morphology of fronds, A. changcaium is similar to the living Ser. Falcata and Ser. Variantia of Sect. Asplenium, while the only analogue of A. popovii is Asplenium castaneoviride of Ser. Trichomanoidea. This might indicate that the living Sect. Asplenium is close related to A. changcaium and A. popovii.
在查阅大量铁角蕨属化石资料基础上,按照不同的时代和层位,对铁角蕨化石进行了地理分布研究,提出铁角蕨属可能起源于早白垩世中期,起源地点为东北亚(我国东北、俄罗斯以及日本)温暖潮湿气候区。首先在欧亚大陆广布,之后经古白令陆桥到现今北美等高纬度地区。随着晚白垩世全球气温降低,温暖潮湿气候区范围扩大,铁角蕨属分布范围也逐渐扩散到南半球。与此同时,在扩散的过程中,铁角蕨属逐渐适应新的环境,演化出不同的种类,分布范围也遍布全球。
化石铁角蕨属中,通过对A. changcaium和A. popovii原位孢子形态和大小的研究,并结合蕨叶形态,与现生铁角蕨属(孢子和叶型)进行了系统比较。在孢子形态和大小方面,A. changcaium孢子形态变化与铁角蕨属铁角蕨组中铁角蕨系的孢子形态变化有共性,其极轴稳定,赤道轴随成熟度增加而变大;在叶型方面,A. changcaium与铁角蕨属铁角蕨组的隆脉系和变异叶系最为相近。A. popovii的孢子比现生铁角蕨属的孢子略大,与铁角蕨属单叶组以及铁角蕨组的隆脉系的孢子大小较为接近;在叶型方面,该种同铁角蕨属铁角蕨组变异叶系中的东海铁角蕨最为相似。因而,推断现生铁角蕨属铁角蕨组与A. changcaium、A. popovii关系最为密切。
The early Cretaceous coal-bearing strata overlying on Tuntianying Formation in Yanbian region of eastern Jilin are composed of two sedimentation cycles, and can be divided into two formations, i.e. Xishangping Formation and Changcai Fomation. The lower sedimentary cycle is named as Xishanping Formation, bearing thin coal seams, while the upper one is called Changcai Formation, bearing thick coal beds. The two formations are divided by the sandy conglomerate on the bottom of the Changcai Formation which shows parallel unconformable contact relationships between them. This suit of coal-bearing strata is distributed mainly in Laotougou and Tuntianying of Yanji County; Changcai, Fudong, Nanyang, Tushanzi, Songxiaping and Dajinchang of Helong County in the eastern Jilin. It is an important coal-bearing strata consisting mainly of array sandstone, sandy conglomerate and gray-green conglomerate intercalated with black shale, carbonaceous mudstone and coal beds, yielding abundant fossil plants.
Since 1950’s, 42 species of 11 genera of ferns have been recorded from the Changcai and Xishanping formations inYanbian region (Yang, 1959; Zhou, et al., 1980; Shang, 1992; Li, 1995, etc.), based on the gross morphology. The present paper deals mainly with the fossil Asplenium found from the Lower Cretaceous Xishanping Formation in Yanbian area.
Based on the gross morphology and the fertile organ and spores in situ which plays an important role in identification and phylogeny, three species of Asplenium, i.e. Asplenium changcaium Sun et Liu (sp. nov.), Asplenium tiefanum Deng and Asplenium popovii Samylina and are recognized for the first time.
On the sterile fronds, the present new species of Asplenium, Asplenium changcaium Sun et Liu (sp. nov.) is different from the known species of fossil Asplenium by having the specialized oval segments on acroscopic side of pinnule and 2-3 times forking lateral veins. For the fertile fronds, Asplenium changcaium Sun et Liu (sp. nov.) is also distinguishing from the known fossil Asplenium by the small elliptical sori attached longintudinally at the middle part of veins on every segments of the pinnules. With regard to the monolete spores in situ, Asplenium changcaium Sun et Liu (sp. nov.) is characterized by having short polar axies and fabaceous shape (lateral view) which is quite different from the spores in situ of Asplenium tiefanum Deng and A. popovii Samylina.
Based upon studies of fossil records of Asplenium and the distribution of fossil and living Asplenium, the place and time of origin and the migration route of Asplenium are discussed preliminarily, concerning on phylogeny, the paleoclimatology and plate tectonic theory. This provides a new basement for the further researches on Asplenium.
The lowest horizon of reliable appearance of Asplenium in the world might be the middle part of the early Cretaceous (i.e. Hauterivian, Barremian and Aptian) in NE China, Siberian and central Honshu of Japan. During that time, the Northeast Asia was in the warm and humid climate zone and might be the original places of Asplenium in the middle of early Cretaceous. Later on, Asplenium dispersed widely on the Eurasian. During the late Cretaceous, Asplenium was already living on the middle Europe, accompanied by the northward migration of warm and humid climate zone. At the same time, the Asplenium occupied the high altitude region of North American (warm and humid climate in the Cretaceous and early Tertiary) via Bering Land Bridge. In the early late Cretaceous, Asplenium might spread to the South Hemisphere due to global climate cooling and the extension of warm and humid climate zone. With the radiation and dispersal, Asplenium diversified, adaped to different environmentsits, and gradually distributed worldwide.
Systemic comparision between the fossil Asplenium changcaium, A. popovii and the living Asplenium is done in the aspects of shape and size of spores in situ and the gross morphology of leaves.
For the shape and size of spores in situ, A. changcaium is resembled to the living Ser. Trichomanoidea of Sect. Asplenium, the length of polar axis changes little, while the equatorial axis varied greatly with the maturity of fossil and living ones. The spores of A. popovii is slightly bigger than those of the living Asplenium and close to those of the living Sect. Holophyllum and Ser. Falcata of Sect. Asplenium in size. On the gross morphology of fronds, A. changcaium is similar to the living Ser. Falcata and Ser. Variantia of Sect. Asplenium, while the only analogue of A. popovii is Asplenium castaneoviride of Ser. Trichomanoidea. This might indicate that the living Sect. Asplenium is close related to A. changcaium and A. popovii.
引文
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150.Figueiredo E., New findings of pteridophytes from the mountain rainforests of Sao Tome and Principe.Fern Gazette 16 (4) : 191-193 2001.
151.Fischer M. A., Niklfeld H., New floristic findings (51-56).Neilreichia 1 : 237-241 2001.
152.Fontoura S. B., Ganade G., Larocca J., Changes in plant community diversity and composition across an edge between Araucaria forest and pasture in South Brazil.Revista Brasileira de Botanica 29 (1) : 79-91 JAN-MAR 2006.
153.Freiberg M., Turton S. M., Importance of drought on the distribution of the birds nest fern, Asplenium nidus, in the canopy of a lowland tropical rainforest in north-eastern Australia.Austral Ecology 32 (1) : 70-76 FEB 2007.
154.Gabancho L.R., Sanchez C., Prada C., Asplenium veneticolor, a new spleenwort (Aspleniaceae, Pteridophyta) from Cuba.Willdenowia 36 (2) : 871-880 2006.
155.Galera H., Sudnik-Wojcikowska B., The most interesting species in the flora of the Palace of Culture and Science in Warsaw.Fragmenta Floristica et Geobotanica Series Polonica 7 : 117-128 2000.
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160.Giraldo-Canas D., Floristic and phytogeographical analysis of the Andean secondary wet forest, Central Cordillera (Antioquia, Colombia).Darwiniana (San Isidro) 39 (3-4) : 187-199 2001.
161.Glazkova E. A., New floristic records on the islands and the northern coast of the Gulf of Finland.Botanicheskii Zhurnal (St. Petersburg) 89 (7) : 1182-1188 July 2004.
162.Gogoi R., Das M. K., Ethnobotanical studies of some ferns used by the Garo tribals of Meghalaya.Advances in Plant Sciences 15 (2) : 401-405 December 2002.
163.Gray A., Pelembe T., Stroud S., The conservation of the endemic vascular flora of Ascension Island and threats from alien species.Oryx 39 (4) : 449-453 OCT 2005.
164.Griffiths O., Tatayah V., Rapid survey of marine turtles in Agalega, Western Indian Ocean.MARINE TURTLE NEWSLETTER (115) : 14-16 JAN 2007.
165.Grund S. P., Parks J. C., Conservation status of pteridophytes in the 49 continental United States: A preliminary report.Fern Gazette 16 (6-8) : 290-294 2002.
166.Guo Xiaosi , Additional materials for the pteridophytic flora of Qinling Mountains.Xibei Zhiwu Xuebao 19 (5) : 096-102 June 1999.
167.Guo Xiaosi, Li Bin, Li Junchao, Preliminary study on the characteristics of pteridophyte flora in the Loess Plateau.Xibei Zhiwu Xuebao 25 (7) : 1446-1451 JUL 2005.
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173.Hietz P., Briones O., Photosynthesis, chlorophyll fluorescence and within-canopy distribution of epiphytic ferns in a Mexican cloud forest.Plant Biology (Stuttgart) 3 (3) : 279-287 May, 2001.
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179.Jasprica N., Kovacic S., Ruscic M., Flora and vegetation of Sveti Andrija Island, southern Croatia.Natura Croatica 15 (1-2) : 27-42 JUN 30 2006.
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191.Kirn H. S., Pteridophytic flora of Poonch district of Jammu and Kashmir State, north-west Himalayas.Indian Fern Journal 17 (1-2) : 92-105 2000.
192.Kirn H. S., Kapahi B. K., Ethnobotanical notes on some ferns and fern-allies of Jammu and Kashmir State, India.Indian Fern Journal 18 (1-2) : 35-38 2001.
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