Fossil Caddisfly Cases from the Early Cretaceous Aliferous Strata in Liupanshan Basin, China
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摘要: 石蚕(昆虫纲: 毛翅目)巢化石保存在宁夏六盘山盆地早白垩世马东山组和乃家河组含盐地层中, 化石呈群栖、密集、相互平行、叠层状分布, 石蚕巢为直立锥状体, 长约13 mm, 平均直径约2.5 mm.管壁具三层式结构, 包括内、外暗色有机质层和中部胶结颗粒层.暗色有机质层为泥晶结构, 以灰泥质为主；中部胶结颗粒层, 主要由有机质球粒构成, 球粒呈球形、椭球形和卵圆形等, 形态大小一致, 平均长0.3 mm, 直径0.10~0.15 mm.这些有机质球粒在管壁中排列规整, 长轴大多与石蚕巢生长方向垂直.依据管壁中规则排列的有机质球粒将六盘山盆地的石蚕巢化石归为Coprindusia遗迹属.通过对比各种造管生物形态学和生态学方面的特征以及现生造迹昆虫的生活环境、管体形态大小、管壁胶结颗粒组成和排列方式, 综合相同层位上昆虫实体化石的分析, 六盘山盆地的管状化石可能是宁夏石蛾科宁夏石蛾属房氏宁夏石蛾(Ningxiapsyche fangi)的幼虫巢.Abstract: Fossil caddisfly (Insecta: Trichoptera) larval cases have been recently discovered in the Early Cretaceous lacustine deposits (Madongshan Formation and Naijiahe Formation) in Liupanshan basin, Ningxia. The arrays of caddisfly larval cases occur as superimposed layers of vertically oriented cases. The fossils are gregarious, dense, parallel to each other and closely packed. The single larval case is conical and similar in size (generally 13 mm in length and averagely 2.5 mm in diameter). The wall of the caddisfly larval cases is divided into three layers, including internal, external dark organic matter layers which have micrite structures and are composed of lime muds, and the middle layer of cemented aphanitic particles composed of organic pellets. The calcified organic pellets are orbicular, spheroidicity and ovate, with its size ranging from 0.10 to 0.15 mm wide by averagely 0.3 mm long. The pellets are in regular arrangement, and their long axes are mostly perpendicular to the growth direction of the caddisworm cases. The larval cases in Liupanshan basin are attributed to Coprindusia in ichnology according to the pellets along the walls of the cases. By comparing the features of various tube-maker organism in morphology and bionomy, and the habitat, size and shape of cases, the composition of case-building particles, and case construction way, combining the analysis to insect body fossil from the same layers in which the caddisfly cases preserved, it seems like that the caddisfly larval cases in Liupanshan basin are Ningxiapsyche fangi from Ningxiapsychidae.
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Key words:
- Liupanshan basin /
- Early Cretaceous /
- caddisfly cases /
- fossil /
- Trichoptera /
- organic matter /
- sedimentology
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图 1 石蚕巢化石产出地理位置及产出剖面
Fig. 1. Geographical location and section of the caddisfly larval cases
图 2 宁夏同心剖面第4层(a)和第2层(b)的石蚕巢
Fig. 2. Caddisfly larval cases from the forth layer (a) and the second layer (b) in the Tongxin, Ningxia
图 3 石蚕巢的形态及结构特征(化石均采自宁夏同心剖面第4层)
a.单个圆锥状石蚕巢管；b.石蚕巢个体间无联结构造；c, d, e, f, g.管壁的三层式结构, 由内至外(1-3)为: 内暗色有机质层-胶结颗粒层-外暗色有机质层；h.石蚕巢纵切面形态, 胶结颗粒大多圆形, 大小较一致
Fig. 3. Morphological and structural characteristics of caddisfly larval cases
图 4 石蚕巢管壁胶结颗粒排列示意
a.石蚕巢横截面颗粒的排列；b.石蚕巢纵剖面颗粒的排列；c.石蚕巢立体模式图
Fig. 4. Arrangements of cemented particles of the wall of the cases
表 1 石蚕巢管壁粘结颗粒成分对比
Table 1. Composition comparison of cemented particles of the walls of the caddisfly cases
地区和盆地 地质年代 地层 管壁颗粒成分 资料来源 美国Wyoming 始新世 绿河组 介形虫瓣膜 Bradley, 1924 华盛顿东部 中新世 Latah组 叶片 Standley, 1970a Minnesoda中南部 白垩纪 Windrow组 叶片 Standley, 1970b Montana 渐新世 Peterson和Morman Creeks组 叶片 Standley, 1972 英国 早白垩世 Wealden群 叶肢介瓣膜, 鱼骨/鱼鳞, 球粒, 双壳碎片, Jarzembowski, 1995 美国Colorado 晚侏罗世 Morrison组 砂岩颗粒 Hasiotis et al., 1998 美国Wyoming 始新世 绿河组 碳酸盐颗粒 Loewen et al., 1999 美国Wyoming 始新世 绿河组 碳酸盐颗粒, 鲕粒, 砂质石英颗粒 Leggitt and Cushman, 2001; Leggitt and Loewen, 2002 西班牙 现代 其他石蚕巢管, 砂岩颗粒, 植物碎片 Boyero and Barnard, 2004 韩国 早白垩世 Jinju组 细粒砂岩颗粒, 植物碎片, 灰泥碎屑 Paik, 2005 俄罗斯 晚白垩世 Kundur组 植物碎片 Sukatsheva, 2005 英国 早白垩世 Wealden群 鱼骨/鱼鳞 Heads, 2006 俄罗斯Siberia 早白垩世 Baissa 球粒(粪球粒) Ivanov, 2006 中侏罗世 道虎沟动物群 砂质, 植物碎屑 中国 早白垩世 热河动物群 介形虫壳, 双壳类碎片 黄迪颖等, 2009 卢尚坟昆虫群 
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表 2 石蚕巢主要遗迹学分类
Table 2. Ichnological classfication of the caddisfly larval cases
遗迹属 代表种 大小 分布时代 管壁胶结颗粒 文献来源 Conchindusia C. distans 长14~23 mm, 宽4~5 mm K- 叶肢介瓣膜 Jarzembowski, 1995 Piscindusia P. Sukachevae 长17~30 mm, 宽3~4 mm K- 鱼骨/鱼鳞等 Jarzembowski, 1995 Ostracindusia O. vyalovi 长12~14 mm, 宽4 mm K- 介形虫瓣膜形成的球粒 Jarzembowski, 1995 Pelindusia P. percealleni 长35 mm, 宽5 mm K- 双壳和蜗牛壳的碎片 Jarzembowski, 1995 Folindusia F. pinacea 长23 mm, 宽7 mm J- 叶片 Standley, 1970a Scyphindusia S.hydroptiliformis 长40 mm, 宽8 mm J- 分泌腺等物质 Ponomarenko et al., 2009 Terrindusia T. notabilis 长15~19 mm, 宽5 mm K- 少量叶片、木屑和砂岩颗粒 Ponomarenko et al., 2009 Coprindusia C. pallida 长8~14 mm, 宽3.1~4.1 mm K- 水下动物的粪球粒 Ivanov, 2006; Gorman et al., 2008 Indusia I. incredibilis 长12~20 mm, 宽3~4 mm K- 植物和介形虫壳 Ponomarenko et al., 2009 Tektonargus T. kollaspilas 长11~14 mm, 宽0.5~3.0 mm J- 砂岩颗粒 Hasiotis et al., 1998 Secrindusia S. admiranda 长23.0 mm, 宽3.5 mm K- 分泌腺物质和砂岩颗粒 Ponomarenko et al., 2009
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表 3 造管生物对比
Table 3. Comparison of various tube-makers
生物 分类 管状化石形态或大小 管壁特征 分布时代 参考文献 Serpulids 环节动物门多毛纲隐居亚纲 近圆筒状, 直径平均约1.6 mm, 最大长度可达20 mm 管壁呈多层状(约4~7层), 管外表具横纹或纵肋等纹饰 J- 俞昌民和王惠基, 1981；陈哲和孙卫国, 2001；Chen et al., 2008 Vestimentiferans 环节动物门西伯达虫科 圆柱状, 不锥化, 直径0.86~8.75 mm, 平均3.8 mm, 长度最大可达42 mm 隐晶质的碳酸钙组成, 外壁表面光滑, 粘结一些钙球 S- Halanych et al., 1998; Peckmann et al., 2005；陈忠等, 2006, 2007 Dentaliid 软体动物门掘足纲 管体直而微弯, 两端开口 管壁也具有各种形式的装饰 T-Q2 俞昌民和王惠基, 1983 Tentaculitoid 软体动物门Cricoconarida纲 壳体为碳酸钙质, 呈细长直圆锥形, 少数为弯圆锥形, 大多介于1~10 mm, 最长可达70 mm 发育横环、纵肋等 O2-T2 Vinn, 2010 Siphonodendron 腔肠动物门珊瑚纲十字珊瑚目 圆柱状, 平均长约2 cm, 直径约3 mm 管体之间有联结构造 C Aretz et al., 2010 Oocardium 绿藻门双星藻目 管体分叉, 长度不一 ？ Bradley, 1974；俞昌民和王惠基, 1981 Phryganea 节肢动物门昆虫纲毛翅目 长2.5~30.0 mm不等, 两端开口 主要由砂质石英颗粒组成 J2- Leggitt and Loewen, 2002
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