新生代早期鱼类演化的热点区域:环北部湾陆上盆地古近纪鱼化石研究成果与展望
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摘要
环北部湾新生代的鱼化石在20世纪曾有过少量研究.自2002年以来,本文作者等在广西宁明、南宁、百色、广东茂名、三水及海南长昌等环北部湾陆上盆地的古近纪地层中采到了一批鱼化石,并对其中一些做了初步的研究.本文旨在对已有的成果和进展做概览,为进一步深入研究的导向做些探讨.总的来说,本区域古近纪鱼群包括软骨鱼类的鲨鱼类(Selachii,Elasmobranchii),硬骨鱼类的骨舌鱼目(Osteoglossiformes)、鲱超目(Clupeomorpha)、鲤形目(Cypriniformes)、鲇形目(Siluriformes)和鲈超目(Percomorpha)等.鱼化石数量和种类丰富,中生代鱼类的典型代表的孑遗类群(埃笠姆鲱目Ellimmichthyiformes)与新生代新兴、分异的主要类群(鲤形目)并存,几乎涵盖了新生代非海相鱼类所有主要类群的早期代表.值得注意的是,本区域且区域内各盆地的鱼群均以鲤形目为主,是目前世界上古近纪鲤形目化石科一级类群最集中、最丰富的区域.三水盆地古新统心组的江汉鱼科Jianghanichthyidae三水江汉鱼Jianghanichthys sanshuiensis、茂名盆地始新统油柑窝组的鲤科Cyprinidae茂名始原鲤Eoprocypris maomingensis、和南宁盆地渐新统邕宁组的鳅科Cobitidae南宁鳅Cobitis nanningensis分别是各科最早期的化石记录,代表着鲤形目主要分支的分异时间和早期形态.再者,含鱼化石地层的年代从古新世晚期延续至渐新世,除鲤形目和骨舌鱼目外的其他几类鱼可生活于海水和淡水,这几类鱼化石的研究将为判断当时是否存在过海侵提供鱼化石证据,为研究鱼类区系与环境协同演化提供理想的条件.综上,环北部湾陆上盆地无疑是新生代淡水鱼类区系起源与演化的热点区域(evolutionary hotspot),深入研究本区域古近纪鱼群将为诠释现代鱼类区系的形成、有关类群的起源演化以及与之相关的古动物地理学和古环境提供重要的化石证据.
Paleogene sediments are well developed in the onshore basins, including Ningming, Nanning, and Baise Basins of Guangxi, Sanshui and Maoming Basins of Guangdong, and Changchang Basin of Hainan, around the Beibu Gulf(Gulf of Tonkin). A large number of fish fossils have been recovered from these sediments. As our fieldwork and research activities progressed in recent years, the picture of the Paleogene ichthyofauna is gradually unveiled. Here we review the systematic paleontology of the fossil fishes and their implications on paleoecology and paleobiogeography. Known Paleogene fossil fishes from this region include cypriniforms, osteoglossiforms, siluriforms, and perciforms from Sanshui Basin; cypriniforms, siluriforms and sharks from Maoming Basin; cypriniforms from Changchang Basin; cypriniforms, siluriforms, and clupeomorphs from Nanning Basin; cypriniforms, ellimmichthyiforms, siluriforms, and gobiiforms from Ningming Basin; and cypriniforms from Baise Basin. Among all these fossil fishes, cypriniforms, a group of primary freshwater fishes, were the dominant fishes in terms of specimen numbers and taxonomic diversity, which spanned from the Paleocene to the Oligocene and occurred in every basin of this region. Described cypriniforms from this region include(1) Cyprinidae: Eoprocypris maomingensis from late Eocene Youganwo Formation of Maoming Basin; Huashancyprinus robustispina and Ecoparpia ningmingensis from Oligocene Ningming Formation of Ningming Basin, and Nanningocyprinus wui from Oligocene Yongning Formation of Naning Basin;(2) Jianghanichthyidae(the only fossil family of the order Cypriniformes and only occurred in South China): Jianghanichthys sanshuiensis from Paleocene Buxin Formation of Sanshui Basin; and(3) Cobitidae: Cobitis nanningensis from Oligocene Yongning Formation of Nanning Basin. These fishes are among the oldest cypriniform fossils, which provide critical information on the origin and evolution of the largest extant freshwater fish order Cypriniformes. Moreover, Oligocene ellimmichthyiforms(Clupeomorpha) and gobiiforms(Percomorpha) are important discoveries from this region. The former represents the first discovery of ellimmichthyiform from the Oligocene worldwide which is younger than its known fossil records. The latter is the first occurrence of fossil gobiiforms in China. Furthermore, the most abundant Paleogene fishes of this region are cypriniforms, which are strict freshwater fish. Meanwhile, shark materials are also discovered along with possibly marine ellimmichthyiform and siluriforms. The combination of above fishes indicates that the freshwater bodies of this region were invaded occasionally by marine water during the Paleogene. To conclude, the Paleogene ichthyofauna from onshore basins of Beibu Gulf consists of oldest fossils of major clades of Cenozoic fishes and a relict of typical Mesozoic fish taxa, which suggest that this region is an evolutionary hotspot of Cenozoic freshwater fishes, especially of cypriniforms. The vertebrate paleontology, systematics, taxonomy, and paleoecology of the fish assemblages from this region are pending further study. To understand the origin and divergence of the major clades of the Paleogene fish, as well as hypothesize on the development of the recent fish fauna of East and South Asia, we suggest conducting more comprehensive fieldwork to collect scientifically valuable specimens in the onshore basins around Beibu Gulf, and performing in-depth laboratory research using traditional paleontological methods, 2-D and 3 D computational tools, and molecular data of related extant fishes.
Paleogene sediments are well developed in the onshore basins, including Ningming, Nanning, and Baise Basins of Guangxi, Sanshui and Maoming Basins of Guangdong, and Changchang Basin of Hainan, around the Beibu Gulf(Gulf of Tonkin). A large number of fish fossils have been recovered from these sediments. As our fieldwork and research activities progressed in recent years, the picture of the Paleogene ichthyofauna is gradually unveiled. Here we review the systematic paleontology of the fossil fishes and their implications on paleoecology and paleobiogeography. Known Paleogene fossil fishes from this region include cypriniforms, osteoglossiforms, siluriforms, and perciforms from Sanshui Basin; cypriniforms, siluriforms and sharks from Maoming Basin; cypriniforms from Changchang Basin; cypriniforms, siluriforms, and clupeomorphs from Nanning Basin; cypriniforms, ellimmichthyiforms, siluriforms, and gobiiforms from Ningming Basin; and cypriniforms from Baise Basin. Among all these fossil fishes, cypriniforms, a group of primary freshwater fishes, were the dominant fishes in terms of specimen numbers and taxonomic diversity, which spanned from the Paleocene to the Oligocene and occurred in every basin of this region. Described cypriniforms from this region include(1) Cyprinidae: Eoprocypris maomingensis from late Eocene Youganwo Formation of Maoming Basin; Huashancyprinus robustispina and Ecoparpia ningmingensis from Oligocene Ningming Formation of Ningming Basin, and Nanningocyprinus wui from Oligocene Yongning Formation of Naning Basin;(2) Jianghanichthyidae(the only fossil family of the order Cypriniformes and only occurred in South China): Jianghanichthys sanshuiensis from Paleocene Buxin Formation of Sanshui Basin; and(3) Cobitidae: Cobitis nanningensis from Oligocene Yongning Formation of Nanning Basin. These fishes are among the oldest cypriniform fossils, which provide critical information on the origin and evolution of the largest extant freshwater fish order Cypriniformes. Moreover, Oligocene ellimmichthyiforms(Clupeomorpha) and gobiiforms(Percomorpha) are important discoveries from this region. The former represents the first discovery of ellimmichthyiform from the Oligocene worldwide which is younger than its known fossil records. The latter is the first occurrence of fossil gobiiforms in China. Furthermore, the most abundant Paleogene fishes of this region are cypriniforms, which are strict freshwater fish. Meanwhile, shark materials are also discovered along with possibly marine ellimmichthyiform and siluriforms. The combination of above fishes indicates that the freshwater bodies of this region were invaded occasionally by marine water during the Paleogene. To conclude, the Paleogene ichthyofauna from onshore basins of Beibu Gulf consists of oldest fossils of major clades of Cenozoic fishes and a relict of typical Mesozoic fish taxa, which suggest that this region is an evolutionary hotspot of Cenozoic freshwater fishes, especially of cypriniforms. The vertebrate paleontology, systematics, taxonomy, and paleoecology of the fish assemblages from this region are pending further study. To understand the origin and divergence of the major clades of the Paleogene fish, as well as hypothesize on the development of the recent fish fauna of East and South Asia, we suggest conducting more comprehensive fieldwork to collect scientifically valuable specimens in the onshore basins around Beibu Gulf, and performing in-depth laboratory research using traditional paleontological methods, 2-D and 3 D computational tools, and molecular data of related extant fishes.
引文
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