EVOLUTIONARY SEQUENCE SYSTEMATICS: CASE ANALYSIS OF PERMIAN WAAGENOPHYLLOID CORALS
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摘要: 进化序列系统学是古生物学中研究物种间和类别间的系统发育关系为目的的学科, 包括研究物种或类别间的亲缘关系, 并确定它们形成的先后序列.表型-分支系统学是该系统学的基础.分支系统学的许多重要概念在该系统学中加入了时间的含义, 例如, 认为最亲近的姐妹种之间的时间间隔不会超过它们母种的生存期.姐妹群和性状镶嵌分析是进化序列研究的主要方法, 而理解性状镶嵌分布是关键.这些分析可由计算机程序实现.以二叠纪瓦岗珊瑚类作为研究进化序列系统学的实例, 该类在特提斯地区广泛分布.通过对瓦岗珊瑚类的特征分析确定了17个性状用于研究, 并且每一性状确定了几个变化状态.根据最广布的性状状态、化石记录和个体发生等原则分辨出近祖性状状态, 并根据性状在系统发育中的重要性确定性状分级.然后, 确定了姐妹群间的关系和物种间的进化序列.在姐妹群与进化序列分析相互检验中可发现性状退化、平行演化和多向分支等古生物学重要现象, 并分辨出物种的异常顺序.在检验生物地层学首次出现事件讨论中推测了一些化石首次出现的可能层位.Abstract: Evolutionary sequence systematics, an academic discipline in paleontology and biology, is oriented toward the research into the systematic growth association between species or taxa, including the relative relations between species or taxa, and also toward the determination of sequence of their formation. The systematics is based on the phenetic-cladistic systematics. Many important theoretical concepts in the cladistics include the time concept in this systematics. For example, the time-span between two closest sister species can not exceed the life span of their mother species. The analysis of sister-group and character mosaic distribution may serve as a major method for the research into the evolutionary sequence. However, the understanding of the character mosaic distribution is the key to this research. These analyses can be executed with the computer programs. The Permian Waagenophylloid coral fauna is taken as an example to illustrate the analysis of evolutionary sequence. These coral fossils are widespread in the Tethyan region. Seventeen characters and several changing states for each character were determined through the analysis of character features of Waagenophylloid corals. The plesiomorph character states for each character were identified in line with the principles of the widest distribution of character states, fossil records and ontogenesis. In addition, these characters are classified as 4 ranks on the basis of their important places in phylogeny. Then the relationship between sister groups and evolutionary sequence between species were determined following the theory and methodology of the evolutionary sequence systematics. In the mutual proof-tests between the sister groups and evolutionary sequence, some important palaeontological phenomena were discovered such as character degeneration, parallel evolution, and poly-directional clades. At the same time, a number of abnormal orders were identified. From the first events that occurred in the biostratigraphical test, possible stratigraphic horizons of some fossils can be deduced.
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图 1 性状镶嵌分布和物种序列
a.性状镶嵌的图解; b.已知物种从假设的祖先种分裂的顺序.以瓦岗珊瑚类的3个化石种(Wku=W. kueichowense Huang, 1932;Wwe=W. wenchengense Huang, 1932;Wvm=W. virgalense mongoliense Grabau, 1931) 之间的性状关系为例.1.衍生性状状态; 2.祖征.箭头表示镶嵌关系.左侧代码为瓦岗珊瑚类的各种性状, 其含义见表 1, 性状的上标代表性状分级.图内的数字为性状状态编号.t1, t2, t3为时间点
Fig. 1. Character mosaic distribution and species sequence
图 2 栖霞期瓦岗珊瑚类系统发育关系
图左下方为亲近度的标尺, 作为确定单系类群的依据.下方字母为类群代号.其他图例见图 1, 化石种的名称省略
Fig. 2. Phylogram of Waagenophylloid corals of Qixia, Permian
图 3 二叠纪瓦岗珊瑚类的系统发育枝形图
1.栖霞期; 2.茅口期; 3.吴家坪期; 4.长兴期. 左下方的标尺示亲近度, 只适用于栖霞期.种名略, 下部字母表示类群划分.虚线表示不确定
Fig. 3. Phylogram of Permian Waagenophylloid corals
图 4 序列异常的姐妹群
a, b.种Llo和Lhu的近祖性状多于其他姐妹种; c, d.种Wlo, 1级性状LD出现衍生状态, 种Lch有更多的衍生性状, 因此成为远离祖征的后序种.图例同图 2
Fig. 4. Sister groups with abnormal sequence
表 1 二叠纪瓦岗珊瑚类性状的代码和变化性状的代号
Table 1. Codes of characters and character states of Permian Waagenophylloid corals
表 2 计算各阶中出现次数最多的性状状态
Table 2. Counting character states with the highest occurrence ratio in each Permian stage
表 3 二叠纪瓦岗珊瑚类71种间的进化序列
Table 3. Evolutionary sequence of 71 species of Permian Waagenophylloid corals
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[1] Hennig W. Phylogenetic systimatics[M]. Urbana: Univ Illinois Press, 1966. 1~263. [2] Simpson G G. Principles of animal taxonomy[M]. New York: Columbia Univ Press, 1961. 1~247. [3] Mayr E. Cladistic analysis or cladistic classification[J]? Zeitschrift fuer Zoologische Systematik und Evolutions forschung, 1974, 12(2): 94~128. [4] Ashlock P D. The use of cladistics[J]. Ann Rev Ecol Syst, 1974, 5: 81~99. doi: 10.1146/annurev.es.05.110174.000501 [5] Xu G R. Phenetic-cladistic systematics and geographic patterns of Triassic rhynchonellids[A]. In: Mackinnon, Lee, Campbell, eds. Brachiopods through Time[C]. Rot terdam: Balkema, 1990. 67~79. [6] 徐桂荣. 三叠纪小嘴贝类(Rhynchonellids)的表型—分支系统学[J]. 现代地质, 1991, 5(2): 239~251. https://www.cnki.com.cn/Article/CJFDTOTAL-XDDZ199103001.htm [7] 徐桂荣, 罗新民, 王永标, 等. 长江中游晚二叠世生物礁的生成模型[M]. 武汉: 中国地质大学出版社, 1997. 1~138. [8] 徐桂荣, Sando W J. 湘西北晚二叠世珊瑚礁中的瓦岗珊瑚类[A]. 见: 徐桂荣, 罗新民, 王永标, 等. 长江中游晚二叠世生物礁的生成模型[C]. 武汉: 中国地质大学出版社, 1997. 108~112. [9] Hayasaka I. On the fauna of the anthracolithic limestone of Om-imura in the western part of Echigo[J]. Sci Rep Tohoku Univ, 2nd Ser (Geol), 1924, Ⅷ(1): 1~83;pls. Ⅰ~Ⅶ. [10] Minato M, Kato M. Waagenophyiildae[J]. Journal of the Faculty of Science, Hokkaido University, Series Ⅳ: Geology and Mineralogy, 1965, Ⅻ(3, 4): 241. [11] Tseng T C. Note on the Liangshanophyllum, a new Subgenus of Waagenophyllum from Permian of China [J]. Bulletin of the Geologica Society of China, 1949, 19 (1): 97~104. [12] Wang H C. A revision of the Zoantharia Rugosa in the light of their minute skeletal structures[J]. Philosoph Trans Roy Soc London, Ser B, 1950, 234(611): 175~246. doi: 10.1098/rstb.1950.0002 [13] Tseng T C. A new Upper Permian tetracoral, Huayunophyllum[J]. Acta Palaeontologica Sinica, 1959, 7(6): 499~502. [14] Hill D. Rugosa and Tabulata[J]. Geol Soc Amer and Univ Kans (Lawrence), 1981, Supplement 1(1~2): 1~762. [15] Nelson G J. Classification as an expression of phylogenetic relationship[J]. Systematic Zoology, 1973, 22: 344~359. doi: 10.2307/2412943 [16] 贵州地质矿产局. 西南古生物图册贵州分册[M]. 北京: 地质出版社, 1978. [17] 赵嘉明. 四川北川、江油及陕西汉中二叠纪珊瑚化石[J]. 中国科学院南京地质古生物研究所所刊, 1981, (15): 233~274;图版1~15. [18] Stevens C H, Miller M M, Nestell M. A new Permian Waagenophylloid coral from the Klamath mountains, California[J]. J Paleontology, 1987, 61(4): 690~699. doi: 10.1017/S002233600002905X -
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