New Material of Galeaspids from the Silurian Llandovery in Wuhan of South China
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摘要: 汉阳鱼作为湖北武汉地区古生代鱼类的代表,其真实形态一直未被确定下来,同时关于汉阳鱼目内部的系统发育学关系也只停留在其下两个属种的讨论上,并未对其开展详细的分析讨论.采用形态学和分支系统学的方法,对湖北武汉兰多维列世清水组上部的盔甲鱼类新材料进行了详细研究,建立了汉阳鱼科一新属新种:意外洪山鱼(Hongshanaspis inexpectatus gen.et sp.nov.),同时修订了汉阳鱼属的部分特征.新属以头甲呈宽大于长的梯形和横长的长条形中背孔等特征而归属于汉阳鱼科,同时又以头甲侧缘从眶孔往后逐渐由光滑变为锯齿状,椭圆形的背位眶孔,发育8对侧横管和细小粒状的突起纹饰等特征区别于汉阳鱼科其他属种.分支系统学显示,汉阳鱼目并未组成一个单系类群,其下的汉阳鱼科的属种也并未组成一个单系类群,而是一个多分支形式,但其下的修水鱼科的属种却组成了一个单系类群.相关的相标志表明,相比于锅顶山汉阳鱼,意外洪山鱼生活在水体更深的潮控三角洲环境中.Abstract: As a representative of the Paleozoic fishes in Wuhan,Hubei Province,Hanyangaspis's true morphology has not been determined,and the phylogenetic relationship within Hanyangaspidida is only discussed in the two genera,without a detailed analysis and discussion. In this paper,morphological and cladistic methods were used to study the new materials of galeaspids. New findings of galeaspids (Agnatha) including Hongshanaspis inexpectatus gen. et sp. nov. and Hanyangaspis guodingshanensis are described from the upper part of the Qingshui Formation (Telychian,Llandovery) in Wuhan of South China. The new genus belongs to Hanyangaspidae based on the trapezoid shape of the head-shield and transverse slit-like median dorsal opening. It differs from other genera and species of Hanyangaspidae by a gradual change from smooth to serrated lateral margin of the head shield,dorsally positioned and oval orbit,8 lateral transverse canals on each side and evenly fine tubercular ornamentation. The cladistics results show that Hanyangaspidida isn't a monophyletic group,but the genera and species of Xiushuiaspidae constitute a monophyletic group. Meanwhile the facies markers of the upper part of the Qingshui Formation imply that Hongshanaspis inexpectatus lived in a deeper tidal delta environment compared with Hanyangaspis guodingshanensis.
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Key words:
- Hanyangaspidida /
- Hongshanaspis /
- Qingshui Formation /
- Silurian Telychian /
- Wuhan /
- stratigraphy
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图 1 盔甲鱼类系统发育位置(a)和内部发育关系(b~g)
a. 甲胄鱼纲系统发育关系,修改自Shan et al.(2020);b~g. 前人对汉阳鱼目系统发育关系综述:图b简化自Janvier(1984);图c简化自Wang(1991);图d简化自Janvier(1996);图e简化自Sansom(2009);图f简化自Zhu and Gai(2007);图g简化自Shan et al.(2020)
Fig. 1. Phylogenetic placement (a) and interrelationships (b-g) of galeaspids
图 3 锅顶山汉阳鱼的化石照片及其形态复原图
a. 一块较为完整的头甲标本(BGEG-G-01);b. 锅顶山汉阳鱼形态复原图,修改自刘玉海等(2015);c、d. 雪花状纹饰特写. 缩写说明:dcm. dorsal commissure,背联络管;ic. inner cornual process,内角;ifc. infraorbital canal,眶下管;ldc. lateral dorsal canal,侧背管;ltca-c. a to c lateral transverse canal issuing from infraorbital canal,从眶下管发出的a~c侧横管;ltc1-5. first to fifth lateral transverse canal issuing from lateral dorsal canal,从侧背管发出的1~5侧横管;md. o. median dorsal opening,中背孔;mtc1. medial transverse canal,中横联络管;orb. orbital opening,眶孔;md.s. middle dorsal spine,中背棘
Fig. 3. Photographs and restoration of Hanyangaspis guodingshanensis
图 4 意外洪山鱼(新属、新种)化石照片及其解释
a. 一块不完整的头甲标本(BGEG-G-02b):a1、a2. 头甲外模照片及其解释图;b. 一件不完整的头甲标本(BGEG-G-02a):b1、b2. 头甲内模照片及其解释图;c. 一块不完整的头甲标本(BGEG-G-02a)中横联络管特写;d. 一块不完整的头甲标本前侧视图(BGEG-G-02a);e1、e2. 头甲锯齿状边缘特写(方框内);f1、f2. 细小粒状纹饰特写. 缩写说明见图 3
Fig. 4. Photographs of Hongshanaspis inexpectatus gen. et sp. nov. and their interpretative drawings
图 6 汉阳鱼目其他属种
a. Latirostraspis chaohuensis Wang,Xia et Chen,1980;b. Kalpinolepis tarimensis Wang,Wang et Zhu,1996;c. Nanjiangaspis kalpingensis Wang,Wang,Zhang,Wang et Zhu,2002;d. Konoceraspis grandoculus Pan,1992;e. Hanyangaspis cf. H. guodingshanensis P’an et Liu,1975;f. Changxingaspis gui Wang,1991;g. Xiushuiaspis jiangxiensis Pan et Wang,1983;h. Xiushuiaspis ganbeiensis Pan et Wang,1983;i. Xiyuaspis zhangi(Lu et al., 2007)Liu,Zhu,Lin,Lu et Gal,2019. 缩写说明:pi. pineal opening,松果孔;soc1. anterior supraorbital canal,前眶上管;soc2. posterior supraorbital canal,后眶上管
Fig. 6. Other genera and species of Hanyangaspidida
图 9 志留纪兰多维列世汉阳鱼类分布
图修改自张翔等(2008)、赵文金等(2009)和Zong et al.(2017);1. 浙江长兴;2. 安徽巢湖;3. 江西修水;4. 湖北武汉;5. 湖北京山;6. 陕西紫阳;7. 湖南湘西;8. 新疆柯坪;9. 新疆巴楚
Fig. 9. Sketch showing the distribution of Hanyangaspididae in the Silurian Landovery
表 1 意外洪山鱼(新属、新种)测量数据(mm)
Table 1. Measurements of Hongshanaspis inexpectatus gen. et sp. nov. (mm)
测量内容 数值 最大长(maximum length of
head-shield)130 最大宽(minimum width of
head-shield)140 眶孔直径(diameter of orbital
opening)5 眶间距(distance between
orbital opening)55 中背孔长(length of median dorsal
opening)8 中背孔宽(width of median dorsal
opening)24 中背孔前端至吻端长(length from anterior end
of median dorsal opening to rostral end)3 中背孔后端至头甲后缘中点长(length from posterior
end of median dorsal opening to midpoint of posterior margin of shield)120 -
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