Systematic Revision and Ichnotaxonomy of Zoophycos
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摘要: 三维形态结构复杂、分布广泛的Zoophycos一直是众多遗迹学家研究的热点.通过对1855年至2011年发表的具有详细形态描述并配有图版的动藻迹(Zoophycos)文献进行系统整理, 基于详实的Zoophycos形态结构特征, 系统修订了Zoophycos属, 将Zoophycos主要划分为9个种, 即Zoopycos cauda-galli
Vanuxem, 1842 ；Zoophycos curtainVanuxem, 1842 ；Zoophycos velumVanuxem, 1842 ；Zoophycos brianteusMassalongo, 1855 ；Zoophycos villaeMassalongo, 1855 ；Zoophycos liasinusFischer-Ooster, 1858 ；Zoophycos flabelliformisFischer-Ooster, 1858 ；Zoophycos insignisSquinabol, 1890 和Zoophycos rhodensisBromley and Hanken, 2003 ; 将Spirophyton主要划分为2个种：Spirophyton typumHall, 1863 和Spirophyton eifelienseKayser, 1872 .将Zoophycos与Spirophyton共同归入Zoophycos Group.Abstract: Zoophycos is one of the most widespread but still enigmatic trace fossils, especially with regard to morphology and ethology surrounding its 3-D helically coiled spreite structures. The genus Zoophycos is systematically revised based on the systematic researches of the literature with detailed description and plate from 1855 to 2011. The ichnogenus Zoophycos is revised into 9 ichnospecies, which include Zoophycos cauda-galliVanuxem, 1842 ; Zoophycos curtainVanuxem, 1842 ; Zoophycos velumVanuxem, 1842 ; Zoophycos brianteusMassalongo, 1855 ; Zoophycos villaeMassalongo, 1855 ; Zoophycos liasinusFischer-Ooster, 1858 ; Zoophycos flabelliformisFischer-Ooster, 1858 ; Zoophycos insignisSquinabol, 1890 ; Zoophycos rhodensisBromley and Hanken, 2003 . The ichnogenus Spirophyton is composed of Spirophyton typumHall, 1863 and Spirophyton eifelienseKayser, 1872 . The Zoophycos and Spirophyton are grouped into Zoophycos Group.-
Key words:
- Zoophycos /
- Spirophyton /
- trace fossils /
- revision of genera and species /
- geobiology /
- morphology
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图 1 Zoophycos brianteus、Zoophycos cauda-galli和Fucoides circinatus形态结构特征
a.Zoophycos brianteus Massalongo, 1855原始素描图(转引自Olivero, 2007); b.意大利维罗纳博物馆Zoophycos brianteus Massalongo, 1855补选模式种标本(编号ICN3)(左侧)与原始素描图(右侧)(转引自Olivero, 2007); c.意大利维罗纳博物馆Zoophycos brianteus Massalongo, 1855副模标本(转引自Olivero, 2007); d~e.美国纽约州泥盆纪钙质砂岩水平面中Zoophycos cauda-galli素描图(Vanuxem, 1842), f.Fucoides circinatus(Brongniart, 1828); g.瑞典Husaby地区Bishop城堡废墟石墙上的Fucoides circinatus(Jensen and Bergström, 1995)
Fig. 1. Morphological characteristics of Zoophycos brianteus、Zoophycos cauda-galli and Fucoides circinatus
图 2 修订后世界各地保存较好的Zoophycos brianteus形态
a~c.Zoophycos circinnatus, Pli ka, 1968; d.Zoophycos, Webby, 1969; e.Spirophyton crassum Hall, 1863; f.意大利维罗纳博物馆Zoophycos brianteus Massalongo, 1855副模标本(转引自Olivero, 2007); g.Zoophycos circinnatus, Pli ka, 1969; h.Zoophycos, Olivero, 2003; i.Zoophycos circinnatus, Pli ka, 1969; j~k.Spirophyton Type ‘A’, Plumstead, 1967
Fig. 2. Representative samples of revised Zoophycos brianteus occurred in the world
图 3 Zoophycos curtain和Zoophycos velum素描(Vanuxem, 1842)
Fig. 3. Original drawing of Zoophycos curtain and Zoophycos velum
图 4 Zoophycos villae形态结构特征
a.Zoophycos villae Massalongo, 1855原始素描图(转引自Olivero, 2007); b.意大利维罗纳博物馆Zoophycos villae Massalongo, 1855副模标本(转引自Olivero, 2007); c.Zoophycos villae Massalongo, 1855补选模式种标本(左侧)(编号ICN2)与原始素描图(右侧)(转引自Olivero, 2007); d.Zoophycos villae Massalongo, 1855补选模式种标本(侧面观察)(编号ICN2)与其素描简图(转引自Olivero, 2007)
Fig. 4. Morphological characteristics of Zoophycos villae
图 5 修订后的Zoophycos flabelliformis的形态结构特征
a.Taonurus flabelliformis Fischer-Oster, 1858; b.Zoophycos, Miller and D'Alberto, 2001; c~d.Spirophyton Type ‘B’, Plumstead, 1967; e.Zoophycos, Miller and D'Alberto, 2001
Fig. 5. The illustration showing the representative Zoophycos flabelliformis revised
图 6 修订后的Zoophycos liasinus的形态结构特征
Fig. 6. The illustration showing the representative Zoophycos liasinus revised
a.Taonurus liasinus Fischer-Ooster, 1858; b.Zoophycos ferrum equinum Heer, 1865; c.Zoophycos scoparius Thiol, 1865; d~e.Zoophycos Type D, Patel et al., 2009
图 7 修订后Zoophycos insignis的形态结构特征
a.Zoophycos insignis, Barsanti, 1902; b.Zoophycos insignis, Uchman, 1999
Fig. 7. The illustration showing the representative Zoophycos insignis revised
图 8 修订后Zoophycos rhodensis的主要形态
a.Zoophycos rhodensis Bromley and Hanken, 2003; b.Zoophycos, Lewis, 1970
Fig. 8. The illustration showing the representative Zoophycos rhodensis revised
图 9 修订后Spirophyton typum和Spirophyton eifeliense形态
a.俯视素描图Spirophyton typum Hall, 1863; b.俯视素描图Spirophyton typum Hall, 1863; c.三维形态恢复图Spirophyton typum Hall, 1863; d.俯视图Spirophyton eifeliense Kayser, 1872
Fig. 9. The illustration showing the representative Spirophyton typum and Spirophyton eifeliense revised
图 10 Zoophycos疑问种
a~b.Zoophycos brianteus Fisher-Ooster, 1858; c~d.Cancellophycus marioni Saporta, 1887; e~h.Zoophycos taiyuanensis, 杨式溥, 2004; i~j.Zoophycos xuzhouensis Long, 1987(转杨式溥等，2004); k.Zoophycos circinnatus(转杨式溥等，2004); l.Zoophycos luchengensis(转杨式溥等, 2004); m~n.Zoophycos wudaensis Yang, 1984(转杨式溥等, 2004)
Fig. 10. Questionable Zoophycos species
图 11 Zoophycos与Spirophyton对比
Zoophycos 3D形态据Löwemark and Werner, 2001修改; Spirophyton 3D形态据Hall, 1863修改
Fig. 11. Sketch showing the differences between Zoophycos and Spirophyton
图 12 Rhizocorallium-like Zoophycos与Rhizocorallium对比
Zoophycos 3D形态据Bromley et al., 1999修改; Rhizocorallium 3D形态据Fürsich, 1974修改
Fig. 12. Sketch showing the differences between Rhizocorallium-like Zoophycos and Rhizocorallium
表 1 设想的Zoophycos和Spirophyton特征
Table 1. Assumed morphological codes of Zoophycos and Spirophyton
代码 数字 特征 (1)柱形管 0 无柱形管 1 有柱形管 (2)边缘管 0 无边缘管 1 有边缘管 (3)大蹼纹 0 无大蹼纹 1 有大蹼纹 (4)小蹼纹 0 无小蹼纹 1 有小蹼纹 (5)水平面上形态结构 0 圆形蹼层 1 椭圆形蹼层 2 U形蹼层 3 U形且底端转折或扩散形蹼层 4 漏斗形蹼层 5 螺旋桨形蹼层 6 花瓣形蹼层 7 裙带形蹼层 8 J形蹼层 (6)纵切面柱形管特征 0 纵切面柱形管呈J形 1 纵切面柱形管呈U形 
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表 2 Zoophycos和Spirophyton数字矩阵
Table 2. The digital matrix of Zoophycos and Spirophyton
(1) (2) (3) (4) (5) (6) Z.caudagalli 1 1 1 1 1 0 Z.brianteus 1 0 1 1 0 0 Z.villae 0 1 1 1 8 0 Z.flabelliformis 1 1 1 0 6 1 Z.liasinus 0 1 1 0 2 1 Z.insignis 1 1 1 1 5 1 Z.rhodensis 1 1 1 1 7 1 Z.curtainensis 1 1 1 0 3 1 Z.velum 1 1 1 0 4 0 S.typum 1 1 1 0 0 0 S.eifeliense 1 1 1 0 0 0
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