A New Species Cladophlebis yonganensis sp. nov. from the Lower Cretaceous Bantou Formation of Yong'an, Fujian Province
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摘要: 对福建永安市下白垩统坂头组保存较好的真蕨类营养叶化石枝脉蕨Cladophlebis进行了叶片形态及角质层微细构造特征的分析, 该化石至少二次羽状分裂, 末二次羽轴粗, 小羽片小, 全缘, 排列紧密, 中脉明显, 侧脉密集, 多数一次分叉; 上表皮无气孔器, 细胞轮廓不清晰, 偶见表皮毛; 下表皮具气孔器, 呈纵向带状排列; 表皮细胞长方形或呈条带状; 垂周壁加厚, 较直; 平周壁波状突起; 气孔器近圆形, 具明显拱盖, 保卫细胞弱角质化, 副卫细胞分化不明显.对当前化石小羽片解剖学特征的分析填补了该属植物化石叶片角质层微细构造特征的空白, 进一步将其与现生紫萁Osmunda joponsica以及部分真蕨类化石的表皮特征进行对比, 结合宏观形态和角质层特征的分析及其与相似种类的比较, 将永安的标本命名为一新种, 永安枝脉蕨Cladophlebis yonganensis Dai et B. N. Sun, sp. nov., 并且推断当前化石代表了某一类已经灭绝的真蕨类植物.Abstract: Some fossil remains of morphogenus sterile fern foliage Cladophlebis were collected from the Lower Cretaceous Bantou Formation, Yong'an City, Fujian Province. The gross morphological structure and microstructure of cuticles are studied. These fronds are at least bipinnate, main rachis broad, pinnules small, with entire margin crowdedly attached to the rachis with the whole basis. Their midveins are moderately thick, and the secondary veins at the margin are dense, mostly with a single bifurcation. The upper epidermis is free from stomata, with an indistinct outline of cells, and trichomes can be found sometimes. The stomata on the lower epidermis are longitudinally arranged. The epidermal cells are oblong or elongated, anticlinal cell walls straight or slightly undulant and moderately thick, periclinal walls slightly wrinkled. Stomata are mainly circinal, with well-developed stomata ledges. Guard cells are sunken with weak cutinization, and subsidiary cells are not distinct. The cuticular features described above fill in a gap of the epidermic structure about the genus. Additionally, comparison of epidermic features between present fossils and recent Osmunda joponsica, as well as some filices fossils is carried out. The comparison of gross morphology and cuticular features shows that specimens studied are of a new species and is named as Cladophlebis yonganensis Dai et B. N. Sun, sp. nov., and it is concluded that the fossils studied represent a certain extinct plant of filices.
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Key words:
- Cladophlebis /
- cuticular features /
- epidermic structure /
- stratigraphy /
- Bantou Formation /
- plant fossils
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图 3 Cladophlebis yonganensis sp. nov. 蕨叶形态特征
1.末二次羽片的中间部分，可见末次羽片相互掩复，近对生，标本编号：YAJ3-62-1；2.末二次羽片靠下边的部分，标本编号：YAJ3-62-3；3.末次羽片，标本编号：YAJ3-63-3；4.末二次羽片靠下边的部分，可见末次羽片以较大角度着生于羽轴，标本编号：YAJ3-63-1；5.末二次羽片的中间部分，可见羽轴上的突起及细沟，标本编号：YAJ3-63-2；6.末次羽片，可见其顶端渐尖细，标本编号：YAJ3-62-4；7.末次羽片，可见小羽片近对生，标本编号：YAJ3-63-4；比例尺=1 cm
Fig. 3. Gross morphology of Cladophlebis yonganensis sp. nov.
图 4 Cladophlebis yonganensis sp. nov.小羽片角质层及其形态特征
1.上表皮，无气孔器，细胞轮廓不清晰，比例尺=30 μm；2.上表皮，可见表皮毛，比例尺=30 μm；3.下表皮，示气孔器呈纵向带状排列，比例尺=100 μm；4.下表皮，示垂周壁和平周壁的特征，比例尺=30 μm；5.下表皮，比例尺=50 μm；6.气孔器，可见明显的拱盖，比例尺=15 μm；7~9.立体显微镜下小羽片的形态及脉络特征，7.小羽片基部特征，8.小羽片的侧脉分叉特征，9.小羽片顶端特征，比例尺=1 mm；其中1来自标本YAJ3-62-1，2来自标本YAJ3-62-3，3~6来自标本YAJ3-63-4，7~9来自标本YAJ3-63-2
Fig. 4. Cuticle feature from the pinnules of Cladophlebis yonganensis sp. nov.
图 5 现生Osmunda joponsica表皮特征
1.上表皮，垂周壁弯曲，无气孔器，取自小羽片基部，比例尺=50 μm；2.下表皮，可见气孔器呈带状排列，取自小羽片基部，比例尺=50 μm；3.气孔器，可见肾形的保卫细胞，取自小羽片基部，比例尺=15 μm；4.上表皮，无气孔器，取自小羽片中部，比例尺=30 μm；5下表皮，可见气孔器呈带状排列，取自小羽片中部，比例尺=50 μm；6.气孔器，取自小羽片中部，比例尺=10 μm；1~3为光学显微镜下照片，4~6为扫描电镜下照片
Fig. 5. Cuticle features of modern species Osmunda joponsica
表 1 Cladophlebis yonganensis与其相似种类形态特征的比较
Table 1. Leaf morphological comparisons of Cladophlebis yonganensis with selected similar species of Cladophlebis
特征 C. yonganensis C. argutula C. hsiehiana C. browniana C. ichünensis C. acute C. paltyrachis C. dunkeri 末二次羽轴宽度(mm) 7~10 未知 7 2~3 ≥3 4~5 13 2.5 末次羽轴宽度(mm) 1.3~1.6 未知 未知 0.8~1 较粗 1 未知 较细 末次羽片间距(cm) 互相接触或掩复，少数稍分离 1.5~2.5 相互接触或稍分离 排列紧密但不接触 未知 1.8~2 互相紧靠 排列紧密 末次羽片着生角度 80°~95° 40°~50° 50°~70° 60° 近90° 45°~50° 50°~70° 未知 小羽片形态 长椭圆形或宽线形 披针形，较直 披针形，直 长方形或线形 长方形或长椭圆形 镰刀形 镰刀状，指形 镰刀形或披针形 小羽片顶端特征 钝圆，亚圆或微尖 亚尖 尖 钝圆，或微尖 钝而圆 尖或亚圆 钝圆 方圆或钝圆 小羽片基部特征 耳状，略收缩 较宽，微下延 较宽，微下延 较宽，彼此分离 较宽，微微下延 宽，微连接 下延，不对称 较宽，微下延 小羽片长度(mm) 7~12 8~9 未知 6~12 ≤11 9~10 4 5~8 小羽片宽度(mm) 2.5~4.5 3~4 未知 2~5 ≤4 3.5~4 1.5 2~3 小羽片叶缘特征 全缘 全缘 全缘 全缘 全缘 全缘，少数波状 全缘 全缘 小羽片着生角度 75°~90° 近90° 近90° 近于60° 60°~70° 近90° ≥70° 未知 小羽片间距 彼此紧靠或接触 互相连接或分离 互相连接或分离 紧靠或稍分离 紧挤或接触 紧靠或稍分离 接触或微分离 彼此紧靠 中脉特征 明显，微弯曲，达小羽片顶端 微弱 微弱 明显，达小羽片顶端 弯曲，未达顶端 明显，强直，达顶端 明显，达顶端 适中，达顶端 侧脉分叉次数 多数一次，少数二次 一次或二次 二次 一次或不分叉 一次 一次 一次或不分叉 一次或不分叉 侧脉与中脉夹角 45°~60° 未知 未知 未知 40° 45° 未知 未知 侧脉数量(条/边) 22~28 5~7 紧密 较少 14~20 8~10 6~7 较少 参考文献 本文 斯行健等，1963 斯行健等，1963 曹正尧，1999 斯行健等，1963 张武等，1980; 尹殿奎，2008 曹正尧，1999 Sze，1945 
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