海洋生物分类与系统演化实验室知识库

原足目物种是一类营底栖生活的小型甲壳动物，隶属于节肢动物门（Arthropoda）、甲壳动物亚门（Crustacea）、软甲纲（Malacostraca）、真软甲亚纲（Eumalacostraca）、囊虾总目（Peracarida）的原足目（Tanaidacea），现生物种分为长尾亚目（Apseudomorpha）与原足亚目（Tanaidomorpha）2亚目。它们体长通常不超过15毫米，海生，栖息于世界各纬度、各深度、各类型的海洋栖地中，极少数为淡水性物种。原足目物种在特定的栖息地，如河口红树林湿地等环境具有很高的丰度，为当地食物链重要的一环。我国对于原足目的分类研究十分稀少且局限，已有研究集中于香港周围海域及台湾东岸海域，其他地区海域的研究处于空白状态，亟需更广域且系统性的分类学研究。

本研究以中国海域，特别是胶州湾、舟山群岛、庙湾岛、海南岛、西沙群岛、东沙群岛、南沙群岛、以及台湾东岸等地采集的样品作为材料，经过分选、解剖、观察、鉴定、拍摄，共鉴定原足目10科19属21种，其中包括2个新记录科（背肢虫科、伪原足虫科），7个新记录属（锐丽尾虫属、异细螯虫属、秀细螯虫属、拟细螯虫属、拟岛原足虫属、多棘原足虫属、六节虫属），2个新记录种（鲜明粗螯虫、佐助六节虫），以及8个新种（齿指长尾虫、山东锐丽尾虫、平额太古尾虫、镰形异细螯虫、多节秀细螯虫、毛尾拟细螯虫、刻痕拟岛原足虫、怪足多棘原足虫）。各新种中，齿指长尾虫（长尾虫科）发现于南海，特征为头胸甲左右前侧缘各有1个很大的棘状突起、第3到第6胸节左右侧缘各有1个尖而弯曲的棘状突起、并且螯足不可动指及指节切缘皆有1个齿状突起；山东锐丽尾虫（丽尾虫科）发现于胶州湾及黄海，与同属物种最大辨别特征为第1触角柄第1节腹面及内缘共有7个刺状刚毛、尾肢基节内侧末端有1个明显的刺状刚毛；平额太古尾虫（拟尾虫科）仅发现于胶州湾，与同属物种最大差异为额角平坦无突起、第4胸肢指节具端刺且腹侧末端有很长的突起；镰形异细螯虫（细螯虫科）发现于东沙群岛及海南岛，其最大特征为雄性第4到第6胸肢指节与端刺愈合成细长镰刀状；多节秀细螯虫（细螯虫科）仅发现于西沙群岛，与同属物种的最大差异为雄性第1触角鞭有10到11节；毛尾拟细螯虫（细螯虫科）发现于台湾东岸，与同属相似物种的最大差异为颚足内叶有2个偶联刺、且雌雄尾肢外肢都为2节、内肢都为5节；刻痕拟岛原足虫（背肢虫科）仅发现于东沙群岛，与同属物种最大差异为雌性螯足掌节内侧有1根很强壮的刚毛、以及雄性螯足腕节有1条很长的凹痕由外侧近端延伸至腹侧末端；怪足多棘原足虫（伪原足虫科）发现于台湾东岸，其独有特征为第2和第3胸肢指节呈S形、端刺末端呈扁平鸭嘴形。本研究总结了此10科19属21种的分类特征，对新种及台湾东岸物种进行了详细的分类特征描述并绘制解剖图；并汇整过往发表之分类文献及物种名录，修订为中国浅海原足目物种名录总表共13科29属37种，包含物种修订前后之学名与中文名、物种于世界之分布、以及参考文献。

本研究以传统形态学方法，对我国海域原足目首度进行了系统的分类学研究，分析评述了我国海域已有的原足目种类记录，初步澄清了我国海域的原足目种类组成情况，总结了我国海域原足目种类的分布和区系特性。此研究成果期望开启我国海域原足目分类学与生态学研究的大门，对我国小型底栖无脊椎动物的多样性与生态有更多的了解。

Tanaidaceans are small benthic crustaceans, belonging to Phylum Arthropoda, Subphylum Crustacea, Class Malacostraca, Subclass Eumalacostraca, Superorder Peracarida, and Order Tanaidacea. The order contains two main extant groups, i.e., Suborder Apseudomorpha and Suborder Tanaidomorpha. Species of Tanaidacea barely exceed 15 mm in body length, occuring in the full range of latitude and depth, inhabiting various marine habitats. In some particular habitats, such as estuarine mangrove wetlands, tanaidacean species could reach extraordinarily high densities and is believed to be ecologically important. However, studies on taxonomy of Tanaidacea from the China seas are quite scarce, limited to littoral regions of Hong Kong and eastern Taiwan. Knowledge of species diversity and composition from the extensive coastline of China remains unknown and more investigation is required.

In the present study, materials from China seas mainly Jiaozhou Bay, Zhoushan Archipelago, Miaowan Island, Hainan Island, the Xisha Islands, the Dongsha Islands, the Nansha Islands, and eastern Taiwan were identified, including 10 families (Nototanaidae and Pseudotanaidae were first recorded from China), 19 genera (Phoxokalliapseudes, Alloleptochelia, Kalloleptochelia, Paraleptochelia, Paranesotanais, Akanthinotanais and Hexapleomera were first recorded from China), and 21 species (Apseudes spinidigitus, Phoxokalliapseudes shandongensis, Swireapseudes planafrontis, Alloleptochelia falciformis, Kalloleptochelia multiarticulata, Paraleptochelia setosa, Paranesotanais incisus and Akanthinotanais pedecerritulus were new species; Chondrochelia distincta and Hexapleomera sasuke were first recorded from China). The new species Apseudes spinidigitus (family Apseudidae), from the South China Sea, was characterized by one large spine-like anterior apophysis on lateral margin of carapace, one curved spine-like anterolateral apophysis on pereonites 3–6, and cheliped fixed finger and dactylus both having one conspicuous apophysis on incisive margin. Phoxokalliapseudes shandongensis (family Kalliapseudidae), from Jiaozhou Bay and the Yellow Sea, could be differentiated from its congeners by having seven spiniform setae on article 1 of antennule peduncle and one inner distal spiniform seta on uropod basal article. Swireapseudes planafrontis (family Parapseudidae), only known from Jiaozhou Bay, could be clearly separated from its congeners by the wide and flat rostrum without apophysis, and pereopod 4 dactylus with unguis and one particularly long ventrodistal prolongation. Alloleptochelia falciformis (family Leptocheliidae), from the Dongsha Islands and Hainan Island, was characterized by the long falciform dactylus fused with unguis on male pereopods 4–6. Kalloleptochelia multiarticulata (family Leptocheliidae), only known from the Xisha Islands, could be distinguished from its congeners by having 10 to 11-articled flagellum on male antennule. Paraleptochelia setosa (family Leptocheliidae), from eastern Taiwan, could be differentiated from its most similar congener by having two coupling hooks on maxilliped endite, 2-articled uropod exopod, and 5-articled uropod endopod. Paranesotanais incisus (family Nototanaidae), only known from the Dongsha Islands, could be separated from its only congener by female having one robust inner seta on cheliped propodus and male having one long and conspicuous notch from outer surface to ventrodistal corner. Akanthinotanais pedecerritulus (family Pseudotanaidae), from eastern Taiwan, could be characterized by the unusual sigmoid dactylus and unguis with blunt and duckbill-like tip in pereopods 2 and 3. The diagnosis of each family, genus, and species, as well as the descriptions and figures of each new species or species from eastern Taiwan were provided. A list of all the 37 tanaidacean species (of 13 families and 29 genera) known from shallow seas of China, including their original names, emended names, and distributions, was also provided as table.

Through morphological method, this thesis deals with comprehensive research of taxonomy and provides literature for further study on taxonomy and ecology of tanaidaceans from Chinese waters.

目  录

第1章 总论 ……………………………………………………………………….1

1.1 原足目的分类地位 ……………………………………….……………….......1

1.2 原足目的形态特征 ………………………………………………………........3

1.3 原足目的生物学特性与生态重要性 ………………………………………....8

1.4 原足目的国内外研究概况 …………………………………………………..11

1.4.1 国外研究概况 …………………………………………………………...11

1.4.2 国内研究概况 …………………………………………………………...14

1.5 材料与方法 …………………………………………………………………..14

1.5.1 研究材料 …………………………………………………………….......14

1.5.2 研究方法 …………………………………………………………….......15

第2章 中国浅海原足目的分类学研究 ……………………………………17

2.1 长尾亚目 Apseudomorpha Sieg, 1980 ……………………………………….17

2.1.1 长尾虫科 Apseudidae Leach, 1814 ………………………………….......22

2.1.2 丽尾虫科 Kalliapseudidae Lang, 1956 ...…………………………….......46

2.1.3 后尾虫科 Metapseudidae Lang, 1970 ...……………………………........63

2.1.4 蛣尾虫科 Pagurapseudidae Lang, 1970 ...…………………………….....78

2.1.5 拟尾虫科 Parapseudidae Guţu, 1981 ...…………………………………..89

2.2 原足亚目 Tanaidomorpha Sieg, 1980 ……………………………………....107

2.2.1 细螯虫科 Leptocheliidae Lang, 1973 ………………………………......113

2.2.2 背肢虫科 Nototanaidae Sieg, 1976 ……………………………….........172

2.2.3 拟原足虫科 Paratanaidae Lang, 1949 ……………………………….....183

2.2.4 伪原足虫科 Pseudotanaidae Sieg, 1976 …………………………..........194

2.2.5 原足虫科 Tanaididae Nobili, 1906 …………………………………......203

第3章 总结 …………………………………………………………………….233

3.1  中国浅海原足目物种名录…………………………………………………233

3.2  中国海域原足目的物种多样性……………………………………………240

3.3  中国海域原足目研究的未来展望…………………………………………240

参考文献 ………………………………………………………………………...243

致谢 ………………………………………………………………………………271

作者简历及攻读学位期间发表的学术论文与研究成果 ……………..273

图  目  录

图1.1 原足目整体侧面观及构造名称……………………………………………….5

图1.2 原足目触角形态及构造名称………………………………………………….6

图1.3 原足目口器构造名称………………………………………………………….6

图1.4 原足目口器各构造形态及构造名称………………………………………….7

图1.5 原足目螯足和胸肢形态及构造名称………………………………………….8

图1.6 原足目常见刚毛或感觉毛形态……………………………………………….8

图2.1 中国海长尾亚目物种分布…………………………………………………...21

图2.2–2.5 齿指长尾虫…………………………………………………………..31–34

图2.6–2.8 邦查奇异尾虫………………………………………………………..42–44

图2.9–2.14 山东锐丽尾虫………………………………………………………55–60

图2.15–2.18 毛触须伪尾形虫…………………………………………………..70–73

图2.19 汉氏合尾虫………………………………………………………………….77

图2.20–2.23 多瘤印度尾虫……………………………………………………..84–86

图2.24 麦克碟尾虫………………………………………………………………….91

图2.25–2.30 平额太古尾虫…………………………………………………..100–105

图2.31 中国海原足亚目物种分布………………………………………………...122

图2.32–2.36 镰形异细螯虫…………………………………………………..123–127

图2.37 鲜明粗螯虫………………………………………………………………...133

图2.38 萨氏粗螯虫………………………………………………………………...136

图2.39–2.43 台东粗螯虫……………………………………………………..143–147

图2.44–2.46 多节秀细螯虫…………………………………………………..154–156

图2.47–2.51 毛尾拟细螯虫…………………………………………………..166–170

图2.52–2.56 刻痕拟岛原足虫………………………………………………..173–177

图2.57–2.59 光齿非拟原足虫………………………………………………..189–191

图2.60–2.61 怪足多棘原足虫………………………………………………..200–201

图2.62 佐助六节虫………………………………………………………………...206

图2.63 深水湾无结虫……………………………………………………………...208

图2.64–2.66 东界原足虫……………………………………………………..215–217

图2.67–2.73 石梯坪联虫……………………………………………………..226–232

表  目  录

表2.1 中国及南海产长尾虫属物种形态比较……………………………………...36

表2.2 奇异尾虫属物种雌性的形态比较……………………………………...........45

表2.3 锐利尾虫属物种区分性别的形态比较………………………………….61–62

表2.4 伪尾形属物种形态比较……………………………………………………...74

表2.5 印度尾虫属物种形态比较…………………………………………………...88

表2.6 太古尾虫属物种形态比较………………………………………………….106

表2.7 异细螯虫属物种区分性别的形态比较……………………………….128–129

表2.8 中国海域及邻近区域粗螯虫属相似物种形态比较……………………….148

表2.9 秀细螯虫属物种雄性的形态比较………………………………………….157

表2.10 拟细螯虫属物种的形态比较……………………………………………...171

表2.10 非拟原足虫属物种形态比较………………………………………...192–193

表2.12 多棘原足虫属物种的形态比较…………………………………………...202

表2.13 原足虫属物种形态比较…………………………………………………...218

表3.1 中国浅海原足目物种名录………………………………………….....234–239