李罡. (2023). 燕辽生物群叶肢介动物群组成和演化* [J]. 古地理学报, 25(2): 368-381.
LI Gang. (2023). Clam shrimp fauna of the Jurassic Yanliao Biota: composition and evolution[J]. Journal Of Palaeogeography, 25(2): 368-381.   
Doi: 10.7605/gdlxb.2023.02.044
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燕辽生物群叶肢介动物群组成和演化*
李罡1,2
1 现代古生物学和地层学国家重点实验室,中国科学院南京地质古生物研究所,江苏南京 210008
2 中国科学院生物演化与环境卓越创新中心,江苏南京 210008

第一作者简介 李罡,男,1967年生,中国科学院南京地质古生物研究所研究员。E-mail: gangli@nigpas.ac.cn。

收稿日期:2023-02-27 修回日期:2023-03-01
基金资助:* 中国科学院战略性科技先导专项(B类)(编号: XDB26000000)和国家国家自然科学基金项目(编号: 41972007,41572006,42288201)资助
摘要

中国北方著名中、晚侏罗世的燕辽生物群含有丰富的叶肢介动物群。早期燕辽生物群也称为道虎沟生物群,主要产出层位为海房沟组,含有三饰叶肢介动物群,主要分子包括: 海房沟三饰叶肢介( Triglypta haifanggouensis)、滦平三饰叶肢介( T.luanpingensis)和平泉三饰叶肢介( T. pingquanensis)。晚期燕辽生物群又可称为玲珑塔生物群,主要产出层位为髫髻山组,含有丰富的叶肢介化石,主要分子包括: 建昌三饰叶肢介( Triglypta jiancangensis)、玲珑塔辽西叶肢介( Liaoxiestheria linglongtaensis)、大西山玲珑塔叶肢介( Linglongtaestheria daxishanensis)和青龙玲珑塔叶肢介( Linglongtaestheria qinglongensis)。文中总结了北方雕饰叶肢介( Aquilonoglypta)、柴达木叶肢介( Qaidamestheria)和针孔叶肢介( Punctatestheria)的最新研究成果,支持将柴达木叶肢介、辽西叶肢介和玲珑塔叶肢介划归北方雕饰叶肢介科。同时,三饰叶肢介是由针孔叶肢介演化而来,与柴达木叶肢介并无直接演化关系。

关键词: 侏罗纪; 叶肢介动物群; 道虎沟生物群; 玲珑塔生物群; 燕辽生物群
中图分类号:Q915 文献标志码:A 文章编号:1671-1505(2023)02-0368-14
Clam shrimp fauna of the Jurassic Yanliao Biota: composition and evolution
LI Gang1,2
1 State Key Laboratory of Palaeobiology and Stratigraphy,Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences,Nanjing 210008, China
2 Center for Excellence in Life and Palaeoenvironment,Chinese Academy of Sciences,Nanjing 210008,China

About the first author LI Gang,born in 1967,is a researcher in Nanjing Institute of Geology and Palaeontology,Chinese Academy of Sciences. E-mail: gangli@nigpas.ac.cn.

Fund:Strategic Priority Research Program of Chinese Academy of Sciences(No.XDB26000000),by the National Natural Science Foundation of China(Nos.41972007,41572006,42288201). This is a contribution to UNESCO-IUGS IGCP Project 679
Abstract

The well-known Middle-Late Jurassic Yanliao Biota contains abundant and excellently preserved terrestrial fossils. It can be subdivided into the early Yanliao Biota, i.e. the Daohugou Biota,and the late Yanliao Biota, i.e. the Linglongta Biota,respectively. The Daohugou Biota contains a triglyptids clam shrimp fauna,including Triglypta haifanggouensis, T. luanpingensis and T.pingquanensis. The Linglongta Biota contains a triglyptids-aquilonoglyptids clam shrimp fauna,comprising Triglypta jianchangensis, Liaoxiestheria linglongtaensis, Linglongtaestheria daxishanensis and Linglongtaestheria qinglongensis. According to the latest SEM research results, Qaidamestheria, Liaoxiestheria and Linglongtaestheria should be assigned to the family Aquilonoglyptidae. Punctatestheria,ornamented with evenly distributed punctae,first occurred in the Lower Jurassic Badaowan Formation in Junggar Basin,and gave rise to Triglypta in the late Middle Jurassic. Qaidamestheria does not have direct evolutionary relationship with Triglypta.

Key words: Jurassic; Clam shrimp fauna; Daohugou Biota; Linglongta Biota; Yanliao Biota
1 概述

中国北方燕辽地区中、晚侏罗世之交的燕辽生物群化石宝库发育了精美的水生和陆生化石, 包含植物、双壳类、介形类、叶肢介、昆虫、蛛形纲、两栖类、有鳞类、长羽毛恐龙、翼龙和哺乳类(Zhou et al., 2010)。燕辽生物群的名字源自洪友崇(1983)提出的中侏罗世燕辽昆虫群, 包括了产自冀北辽西地区海房沟组(或九龙山组)、髫髻山组(或蓝旗组)和土城子组(或后城组)的昆虫化石。后来随着鱼类、爬行类和哺乳类等脊椎动物的发现, 燕辽昆虫群发展成燕辽动物群(任东等, 1995)。近年来, 内蒙古宁城道虎沟、辽西建昌玲珑塔和河北青龙满族自治县等地区持续不断地有珍惜昆虫、脊椎动物和植物化石的发现(图1)(Gao and Shubin, 2003; Ji et al., 2006; Meng et al., 2006, 2017; Hu et al., 2009; Ren et al., 2009; Lü et al., 2011a; Luo et al., 2011; Huang et al., 2012, 2013; 郑少林等, 2012; Yuan et al., 2013; Sullivan et al., 2014; Xu et al., 2015; Han et al., 2016; Liu and Wang, 2016, 2017; Zhou et al., 2019)。同时, 在海房沟组、髫髻山组和土城子组陆续获得了新的精确测年数据(Swisher et al., 2002; 陈文等, 2004; He et al., 2004; Yang and Li, 2008; Chang et al., 2009; Zhang et al., 2009; Liu et al., 2012; 王亮亮等, 2013; 王思恩等, 2013; Yu et al., 2021)。目前大家接受的燕辽生物群仅包括中、晚侏罗世之交的陆相生物群。它被进一步划分为早期的道虎沟生物群(产出层位为海房沟组)和晚期的玲珑塔生物群(产出层位为髫髻山组)(黄迪颖, 2015; Xu et al., 2016; Li, 2020b)。

图1 燕辽生物群的重要化石产地(修改于 Xu et al., 2016)Fig.1 Fossil localities of the Yanliao Biota(modified from Xu et al., 2016)

叶肢介是燕辽生物群常见的化石门类。它们的化石在野外很容易被找到。如在道虎沟地区可以见到丰富的叶肢介沿层面密集分布, 化石密度可达3000~8000个/m2(沈炎彬等, 2003)。叶肢介作为燕辽生物群的重要组成部分, 对它们的研究一直受到地层古生物工作者的关注。对它们的组成和时代认识也是循序渐进的。早年的研究者将燕辽地区海房沟组(或九龙山组)的叶肢介归为巴柔阶(Bajocian)或巴柔阶— 巴通阶(Bajocian-Bathonian)自流井真叶肢介动物群(Euestheria ziliujingensis fauna)(张文堂等, 1976, 1987; 陈丕基和沈炎彬, 1979; 沈炎彬等, 2003; Li and Matsuoka, 2012), 或者中侏罗统真叶肢介— 三饰叶肢介动物群(Euestheria-Triglypta fauna)(王思恩, 1984)。近年来, 在辽西建昌玲珑塔髫髻山组发现的新叶肢介化石首先被鉴定为真叶肢介(Euestheria)和可疑的点列叶肢介(Polygrapta?)(段冶等, 2009)。后来该层位的叶肢介连同河北青龙霸王沟髫髻山组发现的新叶肢介化石被鉴定为柴达木叶肢介(Qaidamestheria)(廖焕宇和黄迪颖, 2014)。最近, 辽西玲珑塔和河北青龙又有新叶肢介化石的报道, 包括辽西叶肢介(LiaoxiestheriaLiao et al., 2017)和玲珑塔叶肢介(LinglongtaestheriaLi, 2020b)。但是对它们科的归属出现分歧。文中将详细讨论燕辽生物群叶肢介动物群的组成、演化和归属。

2 地质背景和燕辽生物群的组成

燕辽地区位于华北克拉通东北缘, 具有丰富的煤炭资源, 发育了陆相侏罗系和白垩系, 产出著名的燕辽生物群和热河生物群(Zhou et al., 2010)。燕辽地区是著名的燕山运动的命名地(Wong, 1927), 也是华北克拉通破坏与陆相生物演化研究的关键地区(Zhu et al., 2012; Li and Wang, 2018)。中、晚侏罗世之交在冀北、辽西、内蒙东南部地区沉积了海房沟组(或九龙山组)和髫髻山组(或蓝旗组), 其中蕴含了著名的燕辽生物群。海房沟组最初叫做海房沟砾岩, 在辽西不整合覆盖在煤系地层北票组之上(图2)。层型剖面位于北票市海丰沟村, 是一套灰白色、黄灰色复成分砾岩夹黄灰色长石石英砂岩、粉砂岩及页岩, 局部夹碳质页岩、流纹质凝灰熔岩和凝灰岩。以往研究者将海房沟组与京西和冀北的九龙山组对比。近年来, 在海房沟组中部凝灰岩获得了166.7 Ma同位素年龄(Chang et al., 2014), 在京西九龙山组底部凝灰岩获得2期锆石同位素年龄, 分别是154 Ma 和161 Ma(李海龙等, 2014)。从而印证了京西九龙山组不会老于161 Ma, 和髫髻山组底部年龄相当, 暗示了它与髫髻山组的可能相变关系(黄汲清, 1956, 1960)。因而作者采用新的划分意见, 将冀北、内蒙等地含化石的九龙山组对比海房沟组。

图2 冀北辽西地区地层层序和燕辽生物群的产出层位Fig.2 Stratigrphic sequences in northern Hebei Province and western Liaoning Province, and the Yanliao Biota bearing strata

2.1 道虎沟生物群

海房沟组蕴含早期燕辽生物群, 即道虎沟生物群(图2), 重要的化石点包括宁城五化镇五化西沟、陈台子、姜杖子、道虎沟, 凌源热水汤无白丁, 建平棺材山, 北票海丰沟等(图1)。含有锥叶蕨— 拟刺葵(Coniopteris-Phoenicopsis)植物群的中期组合, 苏铁类含量居首位, 包括异羽叶属(Anomozamites)、侧羽叶属(Pterophyllum)、基尔米亚叶属(Tyrmia)、蕉羽叶属(Nilssonia)和篦羽叶属(Ctenis)等; 其次是真蕨类(包括蚌壳蕨科和双扇蕨科)和银杏类, 松柏类较少(邓胜徽等, 2003)。这一植物组合与俄罗斯中亚地区同期植物群面貌相似, 而与贫苏铁的西伯利亚植物群形成了鲜明对照(张武和郑少林, 1987)。

道虎沟生物群含有丰富的动物化石(表1), 包括双壳类费尔干蚌(Ferganoconcha)(姜宝玉, 2006)、三饰叶肢介(Triglypta)、多样性非常丰富的燕辽划蝽(Yanliaocorixa)昆虫群和珍稀的脊椎动物群。鱼类以软骨硬鳞鱼北票寻科的洪氏辽鲟(Liaosteus hongi)为代表(任东等, 1995)。道虎沟化石点的有尾两栖类出现了现代小鲵科祖先类型道虎沟辽西螈(Liaoxitriton daohugouensis)(王原, 2004), 隐鳃鲵科的基干成员天义初螈(Chunerpeton tianyiensis)(Gao and Shubin, 2003), 以及较原始的基干型非冠群的奇异热河螈(Jeholotriton paradoxus)(王原, 2000)和产自辽西凌源无白丁的身体肥胖的中华胖螈(Pangerpeton sinensis)(Wang and Evans, 2006)。

表1 燕辽动物群属种及产地 Table1 Taxa and fossil localities of the Yanliao Biota

建平棺材山的建平北燕螈(Beiyanerpeton jianpingensis)代表了蝾螈亚目的基部成员(Gao and Shubin, 2012), 但是有人认为它可能是初螈的同义名(Sullivan et al., 2014)。棺材山还出土了谢氏红山蜥(Hongshanxi xiei)(Dong et al., 2019)。

道虎沟生物群的翼龙包括喙嘴龙亚目的宁城热河翼龙(Jeholopterus ningchengensis)(汪筱林等, 2002)和沃氏翼手喙龙(Pterorhynchus wellnhoferi)(Czerkas and Ji, 2002)。兽脚类恐龙包括擅攀龙科宁城树息龙(Epidendrosaurus ningchengensis)(Zhang et al., 2002)和胡氏耀龙(Epidexipteryx hui)(Zhang et al., 2008)。

道虎沟生物群的哺乳动物群充分展现了早期哺乳类运动和生活方式的多样性。半水生游泳的柱齿兽类獭形狸尾兽(Castorocauda lutrasimilis)拥有便于游泳的蹼状足和像河狸一样的尾巴(Ji et al., 2006)。树栖生活的攀援灵巧柱齿兽(Agilodocodon scansorius)具有特化的铲形门齿, 指示了它是最早的以植物汁液为食的动物(Meng et al., 2015)。树栖的灵巧微小柱齿兽(Microdocodon gracilis)在咽喉部位保存了鞍状舌骨(Zhou et al., 2019)。空中滑翔的远古翔兽(Volaticotherium antiquus)有毛茸茸的皮肤翼膜结构(Meng et al., 2006)。蜀兽类的粗壮假碾磨齿兽(Pseudotribos robustus)具有高度发育的假三磨楔齿和相当原始的下颌, 其肩带与单孔类相似, 四肢展现了掘穴特征(Luo et al., 2007)。贼兽目陆地栖息的哺乳形巨齿兽(Megaconus mammaliaformis)身披毛发, 后脚有毒刺, 具有原始颌骨和踝骨(Zhou et al., 2013)。

2.2 玲珑塔生物群

髫髻山组来源于叶良辅1920命名于北京西山的“ 髫髻山层” , 广泛分布于冀北、辽西地区, 以大规模火山岩为特征, 夹页岩和砾岩, 局部地区底部还夹有煤线。本组的含义与米家榕等(1980)提出的辽西蓝旗组相当。在冀北辽西地区, 髫髻山组沉积岩夹层发育了晚期燕辽生物群, 即玲珑塔生物群(图2)。重要化石产地除了辽西北票地区以外(Jiang et al., 2016), 近年来新增了辽西建昌玲珑塔、要路沟、河北青龙满族自治县木头凳霸王沟和干沟乡南石门等化石产地。玲珑塔生物群含有锥叶蕨— 拟刺葵(Coniopteris-Phoenicopsis)植物群的晚期组合。其中苏铁类植物继续发展, 几乎占到植物组合种数的一半。同时出现了以往仅见于西欧、中亚、中国南方和日本外带的似查米亚属(Zamites), 查米羽叶属(Zamiophyllum), 毛羽叶属(Ptilophyllum), 苏铁鳞片属(Cycadolepis), 威廉逊花属(Williamsonia)等。虽然真蕨类植物仍然位居第二, 但双扇蕨科大大衰退。银杏类也表现出拟刺葵属(Phoenicopsis)大量减少, 拜拉属(Baiera)和茨康叶属(Czekanowskia)已不见踪迹(张武和郑少林, 1987)。同时玲珑塔生物群还含有丰富的木化石, 种类包括苏铁、银杏和松柏类等(Jiang et al., 2019)。

玲珑塔生物群含有丰富的动物化石, 包括双壳类、介形类、叶肢介、昆虫、鱼类、两栖类、翼龙、长羽毛兽脚类恐龙、哺乳动物等。玲珑塔化石层的介形类化石可以归为萨雷提缅达尔文介— 季米里亚介(Darwinulla sarytirmenensis-Timiriasevia)介形类组合(段冶等, 2009)。该介形类组合常见于中国北方侏罗系, 如九龙山组、王家山组、头屯河组、七克台组、窑街组和下花园组等(邓胜徽等, 2003)。玲珑塔化石层的双壳类首先被鉴定为北方晚三叠世常见分子陕西蚌(Shaanxiconcha), 后来被改定为额尔古纳蚌(Arguniella)(黄迪颖, 2015), 以区别于早期燕辽生物群的费尔干蚌(Ferganoconcha)。玲珑塔化石层含有丰富的昆虫化石, 以划蝽最常见, 与中华燕辽划蝽不同。双翅目和鞘翅目含量较高, 尚未发现在道虎沟化石层常见的革翅目(黄迪颖, 2015)。玲珑塔化石层的鱼类在大西山和青龙木头凳霸王沟异常丰富, 被初步鉴定为褶鳞鱼科(Ptycholepidae)(段冶等, 2009; 郭相奇等, 2012), 也有人认为是古鳕(Palaeonisciforms)(黄迪颖, 2015)。

玲珑塔生物群含有丰富的翼龙化石群, 包括原始的喙嘴龙亚目和进步的翼手龙亚目过渡类型的李氏悟空翼龙(Wukongopterus lii)(Wang et al., 2009), 潘氏长城翼龙(Changchengopterus pani)(Lü , 2009), 模块达尔文翼龙(Darwinopterus modularis)(Lü et al., 2010b), 玲珑塔达尔文翼龙(Darwinopterus linglongtaensis)(Wang et al., 2010), 粗齿达尔文翼龙(Darwinopterus robustodens)(Lü et al., 2011b), 中华鲲鹏翼龙(Kunpengopterus sinensis)(Wang et al., 2010), 对握拇指鲲鹏翼龙(Kunpengopterus antipollicatus)(Zhou et al., 2021), 以及船颌翼龙类的李氏凤凰翼龙(Fenghuangopterus lii)(Lü et al., 2010a), 赵氏建昌翼龙(Jianchangopterus zhaoianus)(Lü and Bo, 2011)和粗壮建昌颌翼龙(Jianchangnathus robustus)(Cheng et al., 2012), 和最早的帆翼龙类的玲珑塔始帆翼龙(Archaeoistiodactylus linglongtaensis)(Lü and Fucha, 2010)。青龙木头凳产出奇特的蛙嘴翼龙类的木头凳树翼龙(Dendrorhynchoides mutoudengensis)(Lü and Hone, 2012), 它不但有长尖弯齿, 还出现了短粗直齿, 更特别的是具有相对较长的尾巴。

玲珑塔化石层含有赫氏近鸟龙(Anchiornis huxleyi)(Xu et al., 2009; Hu et al., 2009)和郑氏晓廷龙(Xiaotingia zhengi)(Xu et al., 2011)。这2类兽脚类恐龙的发现对研究鸟类起源和飞行起源具有重要意义。建昌要路沟出土的徐氏始鸟龙(Aurornis xui)代表了鸟类的最早祖先(Godefroit et al., 2013a); 要路沟的短羽始中华羽龙(Eosinopteryx brevipenna)代表了最原始的伤齿龙科分子(Godefroit et al., 2013b)。

玲珑塔生物群含有多样化的哺乳动物, 如辽西建昌玲珑塔镇大西山发现了最早的有胎盘真兽类中华侏罗兽(Juramaia sinensis)(Luo et al., 2011), 它活动敏捷, 适合攀爬。大西山的欧亚皱纹齿兽(Rugosodon eurasiaticus)(Yuan et al., 2013)代表了最早的多瘤齿兽, 牙齿和脚部特征显示它是杂食陆栖动物。掘穴生活的短指挖掘柱齿兽(Docofossor brachydactylus)产于河北青龙干沟乡, 具有适合挖掘的脚掌和爪子(Luo et al., 2015)。贼兽类种群分异度较高, 包括攀援树栖和滑翔类型。树栖类型已经报道了5个种, 包括出土于河北青龙县木头凳镇的金氏树贼兽(Arboroharamiya jenkinsi), 它是目前发现的最大贼兽, 具有毛发和单块齿骨构成的下颌(Zheng et al., 2013), 产自玲珑塔大西山的陆氏神兽(Shenshou lui)、玲珑仙兽(Xianshou linglong)、宋氏仙兽(Xianshou songae)(Bi et al., 2014)以及济赞堂奇兽(Qishou kizantang)(Mao and Meng, 2019)都显示了树栖特征。目前报道了3种滑翔的贼兽类, 它们都发育了皮翼膜, 包括产自辽西建昌玲珑塔大西山的似叉骨祖翼兽(Maiopatagium furculiferum)(Meng et al., 2017)、河北青龙干沟乡南石门出土的双钵翔齿兽(Vilevolodon diplomylos)(Luo et al., 2017)和阿霍氏树贼兽(Arboroharamiya allinhopsoni)(Han et al., 2017)。双钵翔齿兽中耳听小骨发育了上下紧密叠覆的砧骨— 锤骨关节(Wang et al., 2021)。阿霍氏树贼兽听小骨发育了独特的上隅骨。这些新发现为解开中耳演化之谜提供了新素材。

3 叶肢介动物群
3.1 道虎沟生物群叶肢介动物群

海房沟组叶肢介最早被鉴定为海房沟真叶肢介(Euestheria haifanggouensisChen in Zhang et al., 1976)和自流井真叶肢介(Euestheria ziliujingensisChen in Zhang et al., 1976)。它们壳体外形相似, 近方圆形, 原始壳瓣凸度较大, 压平保存成化石后壳面常产生褶皱。海房沟种生长线较密, 总数在30条以上。自流井种生长线较稀疏, 在20条左右。当时通过光学显微镜观察, 它们的壳面装饰被描述成小网或外模上表现为密集的点粒装饰。自流井真叶肢介也出现在重庆下沙溪庙组。该叶肢介动物群以南北方共有种命名, 所以叫做自流井真叶肢介动物群(Euestheria ziliujingensis fauna)(张文堂等, 1976)。它广泛分布在中国南方和北方的中侏罗统地层中, 除了上述层位, 它还出现在云南的张河组与和平乡组、福建的漳平组、甘肃天祝窑街组、靖远的王家山组、潮水盆地的宁远堡组、内蒙石拐子煤田长汗沟组、满洲里中蒙边界地区的万宝组、陕北的延安组、山西云岗组、河南马凹组、新疆准噶尔盆地西山窑组、头屯河组以及吐鲁番盆地七克台组等(张文堂等, 1976; 乌统昱, 1980; 曹宝森, 1986; 沈炎彬等, 2002; 邓胜徽等, 2003; 宝音乌力吉等, 2011)。

以往报道的自流井真叶肢介动物群的主要分子除了自流井种和海房沟种以外, 还含有扁形真叶肢介(E.complanata)、弱小真叶肢介(E.exilis)、曼庄真叶肢介(E.manzhuangensis)、椭圆真叶肢介(E.fabiformis)、靖远真叶肢介(E.jingyuanensis)和严家湾真叶肢介(E.yanjiawanensis)。通过新化石地点的发现, 这个叶肢介动物群后来又增加了新成员, 包括冀北滦平周营子九龙山组的滦平真叶肢介(Euestheria luanpingensisShen and Niu in Zhang et al., 1987), 平泉松树台营子九龙山组的平泉三饰叶肢介(Triglypta pingquanensisWang, 1984), 青海柴达木盆地大柴旦大煤沟组的大煤沟柴达木叶肢介(Qaidamestheria dameigouensisWang, 1983)。虽然王思恩等(1984)在《华北地区古生物图册(二)中生代分册》上描述了营子三饰叶肢介, 但是它和平泉种产于同层。原始描述说平泉种外形为卵圆形至椭圆形, 而营子种的外形为卵圆形, 由此看来二者可能为同一物种。随后三饰叶肢介和柴达木叶肢介在中国中侏罗统中被发现, 如新疆吐鲁番盆地七克台组、准噶尔盆地头屯河组及塔里木盆地北缘克孜勒努尔组。因此, 中国北方自流井真叶肢动物群又被叫做三饰叶肢介— 柴达木叶肢介(Triglypta-Qaidamestheria)组合, 并与英国苏格兰大河口群巴通阶的 Qaidamestheria-Neopolygrapta 叶肢介组合对比(Chen and Hudson, 1991; 王思恩, 1998)。

内蒙宁城道虎沟化石层含有极丰富叶肢介化石, 它们的壳瓣密集成群出现, 常常保存珍贵的软体化石。这一叶肢介动物群最早由沈炎彬等(2003)描述。根据所含4种真叶肢介, 即自流井真叶肢介(Euestheria ziliujingensis)、海房沟真叶肢介(E.haifanggouensis)、 靖远真叶肢介(E. jingyuanensis)和滦平真叶肢介(E. luanpingensis), 他们将该叶肢介动物群归为自流井真叶肢介动物群, 并将道虎沟化石层归到九龙山组, 即辽西的海房沟组, 从而肯定了道虎沟生物群中侏罗世的时代属性。

近年来, 道虎沟化石层的叶肢介开展了扫描电镜研究。首先, 扫描电镜研究发现靖远真叶肢介的模式标本壳面具有不规则的网、线或瘤脊装饰, 这种装饰特征的种类在道虎沟化石层并没有出现(廖焕宇等, 2014; Liao, 2022; Teng, 2022)。同时在海房沟真叶肢介的副模标本(NIGP30111)上发现了针孔和带有针孔的小网装饰, 在外模上表现为均匀分布的细小点粒和成簇状的小点粒(Liao et al., 2017, fig.9; Liao, 2022, fig. 5.1-5.3)。同样的针孔和带针孔的小网装饰也出现在道虎沟化石层海房沟真叶肢介标本中(Liao et al., 2017, fig. 5; Liao, 2022, fig. 5.4-5.6)。因此, 海房沟种被改归到三饰叶肢介属。滦平真叶肢介也因具有针孔和带针孔的网饰被修订并改归到三饰叶肢介(Liao et al., 2017, 2022)。连同河北平泉松树台营子村发现的平泉三饰叶肢介, 早期燕辽生物群的叶肢介动物群由3种三饰叶肢介组成: 即海房沟三饰叶肢介(Triglypta haifanggouensis)、滦平三饰叶肢介(T.luanpingensis)和平泉三饰叶肢介(T.pingquanensis)(Liao, 2022)。

3.2 玲珑塔生物群叶肢介动物群

晚期燕辽生物群的叶肢介产于髫髻山组。主要化石点包括辽西建昌玲珑塔大西山、河北青龙满族自治县木头凳霸王沟和干沟南石门等地。玲珑塔大西山化石层的叶肢介最初被鉴定为真叶肢介未定种(Euestheria sp.)和可疑的点列叶肢介(Polygrapta?sp.)(段冶等, 2009)。随后又有新种被描述: 玲珑塔辽西叶肢介(Liaoxiestheria linglongtaensisLiao et al., 2017)(图3-A— 3-D), 建昌天祝叶肢介(Tianzhuestheria jianchangensisWang, 2014), 大西山玲珑塔叶肢介(Linglongtaestheria daxishanensisLi, 2020b)(图3-E— 3-H)。河北青龙霸王沟报道了柴达木叶肢介未定种(Qaidamestheria sp.)(廖焕宇和黄迪颖, 2014)和玲珑塔辽西叶肢介(Liao et al., 2017)。青龙干沟乡南石门化石点最新报道了青龙玲珑塔叶肢介(Linglongtaestheria qinglongensisTeng et al., 2022)。

图3 玲珑塔辽西叶肢介和大西山玲珑塔叶肢介壳面装饰特征
A-D Liaoxiestheria linglongtaensisLiao et al., 2017: A— 右壳, NIGPCAS163671, 采自建昌玲珑塔大西山上侏罗统髫髻山组; B— 近胎壳生长带上的多边形小网装饰, 网孔内无针孔; C— 壳体中部和腹部生长带上均匀分布的针孔装饰; D— 近腹边缘生长带上的针孔装饰。E-H Linglongtaestheria daxishanensisLi, 2020b: E— 叠复的壳体, 正模, NIGPCAS 163667, 产地层位同上; F— 胎壳附近生长带上多边形小网装饰, 网孔内无针孔; G— 壳瓣后腹部生长带上的2种装饰: 生长带的上部均匀分布的针孔装饰和生长带下部线脊夹针孔装饰, 可见横耙; H— 壳瓣 前腹部生长带上的多边形网状装饰, 网孔内具有针孔装饰。A和E为Zeiss V20光学显微镜照片, 其他为LEO1530 VP扫描电镜照片Fig.3 Carapace ornamentation in Liaoxiestheria linglongtaensis and Linglongtaestheria daxishanensis

段冶等(2009)的研究成果只给出了化石名单, 但是没有相关图片发表。根据作者最新研究, 点列叶肢介近胎壳部位生长带具有多边形小网装饰, 壳瓣中部和腹部生长带具有规则分布的稀疏放射线脊和横耙装饰(图4)(Li, 2020a)。目前根据已经报道的种类和作者采集到的玲珑塔化石层的叶肢介标本, 在髫髻山组还没有发现点列叶肢介。

图4 点列叶肢介壳面装饰特征
A-D Polygrapta chatangensisNovojilov, 1946; Tescan Vaga II 扫描电镜照片: A— 右壳, PIN No.401/118(俄罗斯科学院古生物研究所), 采自俄罗斯东北哈坦加湾上二叠统; B— 近胎壳生长带上具有多边形小网装饰(网孔15~30 μ m); C— 壳瓣后腹部生长带上的放射线脊及横耙组成放射状排列的网状装饰; D— 壳瓣中部生长带上的放射线脊装饰, 线脊间发育微弱的横耙Fig.4 Carapace ornamentation in Polygrapta

青龙木头凳霸王沟报道的柴达木叶肢介(Qaidamestheria sp.)很可能是辽西叶肢介, 只是当时没有发现壳瓣背部的小网装饰。王思恩(2014)依据大西山化石点的标本描述了建昌天祝叶肢介(Tianzhuestheria jianchangensis)。根据发表的特征描述和图片信息, 该种壳瓣背部光滑无饰, 中部发育均匀分布的针孔装饰, 到腹部生长带发育了小网和放射线脊装饰, 小网内有4~6个针孔, 线脊间的凹沟有2列放射状排列的针孔。根据天祝叶肢介(Tianzhuestheria)(沈炎彬等, 2002)的原始描述, 它的每条生长带的背部是均匀分布的针孔装饰, 而每条生长带的腹部则发育了线脊夹针孔装饰。这在建昌天祝叶肢介生长带上并没有看到, 所以作者建议将其修订为三饰叶肢介。另外, 王思恩(2014)提到髫髻山组出现了营子三饰叶肢介, 也许说明平泉三饰叶肢介可以上延至髫髻山组, 这是今后工作需要证实的。玲珑塔生物群的叶肢介目前共描述了4种叶肢介: 即三饰叶肢介科的建昌三饰叶肢介, 和北方雕饰叶肢介科的玲珑塔辽西叶肢介, 大西山玲珑塔叶肢介, 青龙玲珑塔叶肢介。这个动物群面貌有别于道虎沟叶肢介动物群, 后者只有三饰叶肢介科的分子。

4 讨论

由于光学显微镜观察手段放大倍数低, 使得叶肢介壳表面的微细装饰特征无法辨认清晰。燕辽生物群的多数叶肢介是通过光学显微镜观察被鉴定为真叶肢介(EuestheriaDeperet and Mazeran, 1912)。扫描电镜的应用提高了壳面微小装饰的辨识度。以往在光学显微镜下看不清的针孔装饰在电镜下面清晰可见, 因此描述了柴达木叶肢介(Qaidamestheria)和三饰叶肢介(Triglypta)。前者因具有针孔(或麻坑)和线脊装饰被归到北方雕饰叶肢介科(Aquilonoglyptidae)(王思恩, 1983), 后者具有针孔、小网和线脊装饰被归到点列叶肢介科(Polygraptidae)(王思恩, 1984)。随后的研究发现这2个属的叶肢介广泛分布于新疆准噶尔盆地的头屯河组和吐鲁番盆地的七克台组(王思恩, 1985)。王思恩(2014)将三饰叶肢介属从点列叶肢介科移出, 并以它为模式属建立了三饰叶肢介科。最近点列叶肢介属模式种扫描电镜研究也证实点列叶肢介并不具有针孔装饰(图4), 所以将三饰叶肢介从点列叶肢介科移出是正确选择(Li, 2020a)。

叶肢介是蜕皮节肢动物。它的侧扁似虾的软体外面披着一个几丁质的双瓣壳。每次蜕皮时软体脱皮, 但是它的壳瓣并不脱落, 而是在壳瓣远端增加1条新的生长带。叶肢介的这种生长方式保留了壳瓣发育的全部信息。从背部到腹部生长带装饰的演变过程再现了它的演化历史, 为研究叶肢介的系统演化和分类提供了有力证据(王思恩, 2014)。三饰叶肢介壳体背部生长带上发育了均匀分布的针孔装饰, 向壳体中部和腹部出现了带针孔的小网装饰和夹有针孔的放射线脊装饰。以往文献报道的发育针孔装饰的叶肢介种类包括陕西下三叠统和尚沟组的戏楼沟北方雕饰叶肢介(Aquilonoglypta xilougouensis Wang in Wang and Liu, 1980)、斜方形北方雕饰叶肢介(A. clinoquadrata Wang in Wang and Liu, 1980)、青海大柴旦中侏罗统大煤沟组大煤沟柴达木叶肢介(Qaidamestheria dameigouensisWang, 1983)和苏格兰大河口群中侏罗世的Euestheria trotternishensisChen and Hudson, 1991。王思恩(1998)认为苏格兰大河口群的真叶肢介具有单一针孔装饰应该归入柴达木叶肢介。所以他提出了由北方雕饰叶肢介(AquilonoglyptaNovojilov, 1958)经柴达木叶肢介(Qaidamestheria)演化到三饰叶肢介的假说(王思恩, 2014)。即由戏楼沟北方雕饰叶肢介经Qaidamestheria trotternishensis 再分别演化出大煤沟柴达木叶肢介和三饰叶肢介。但是, 最新研究进展并不支持这种演化路径。

首先, 最新扫描电镜研究结果显示北方雕饰叶肢介(图5)和柴达木叶肢介壳体背部生长带上发育的不是针孔装饰, 而是多边形小网装饰, 他们壳体中部的生长带上才开始出现针孔装饰。到达腹部后, 北方雕饰叶肢介的生长带上发育了向远端逐渐增大的小网状装饰(图5-E)(Li, 2022)。而柴达木叶肢介的腹部生长带上发育了线脊夹单列针孔装饰(Teng and Li, 2021)。这种分布于背部生长带上的小网装饰特征支持了王思恩(1983)将柴达木叶肢介归到北方雕饰叶肢介科的观点。同时北方雕饰叶肢介从早三叠世开始出现, 而柴达木叶肢介在中侏罗世开始出现, 可以推测后者由前者演化而来。前文提到产于苏格兰大河口群中侏罗统的Euestheria trotternishensis 壳体表面只发育了针孔装饰。这说明该种并非北方雕饰叶肢介的直接后裔种群。不能把它归到真叶肢介或柴达木叶肢介属内。相反, 应该将它划归针孔叶肢介属(PunctatestheriaZhang et al., 2017)。针孔叶肢介壳面只发育了针孔装饰, 它最早出现在新疆准噶尔盆地下侏罗统八道湾组, 也见于吐鲁番盆地上侏罗统七克台组。根据针孔叶肢介壳体只发育了针孔装饰, 可以推测它是早期燕辽生物群三饰叶肢介的直接祖先类型。而晚期燕辽生物群出现了辽西叶肢介和玲珑塔叶肢介。这2个属的壳体背部近胎壳生长带上都发育了多边形小网装饰, 根据这一重要特征这2个属应该划归北方雕饰叶肢介科。

图5 北方雕饰叶肢介壳面装饰特征
A-E Aquilonoglypta meta (Novojilov, 1946)emend. Li, 2022; 采自俄罗斯拉普帖夫海岸下三叠统; Tescan Vaga II 扫描电镜照片; A-B 全型标本(PIN No.401/31, 俄罗斯科学院古生物研究所): A— 近胎壳生长带上的多边形小网装饰; B— 壳瓣背部生长带上的由小网向针孔装饰演变; C-E 另一标本(PIN No.401/30, 俄罗斯科学院古生物研究所); C— 壳瓣中部生长带上针孔装饰; D— 壳瓣后腹部生长带由针孔向网状装饰演变过程; E— 壳瓣腹部生长带上由针孔向逐渐增大的网状装饰演变Fig.5 Carapace ornamentation in Aquilonoglypta

文中研究材料只限于燕辽地区。目前研究结果显示早期和晚期燕辽生物群发育了不同的叶肢介动物群, 两者没有发现相同的物种。前文提到燕辽叶肢介动物群在中国南方和北方广泛分布, 但以往研究将多数物种归到真叶肢介属。今后需要进一步开展扫描电镜研究以期确定它们的确切分类学归属, 从而为深入探讨燕辽叶肢介动物群演替、生活环境和生物古地理打下基础。

5 结论

中国北方燕辽地区发育了著名的燕辽生物群, 可以划分为早期的道虎沟生物群和晚期的玲珑塔生物群。早期燕辽生物群含有三饰叶肢介动物群, 包括海房沟三饰叶肢介、滦平三饰叶肢介和平泉三饰叶肢介, 重要产出层位为海房沟组。晚期燕辽生物群含建昌三饰叶肢介、玲珑塔辽西叶肢介、大西山玲珑塔叶肢介和青龙玲珑塔叶肢介。同时根据最新研究成果三饰叶肢介并非由柴达木叶肢介演化而来, 而是由针孔叶肢介演化而来。

致谢 感谢审稿专家和编辑提出的宝贵修改意见和建议。

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