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作者简介:

夏梦丽,女,1978年生。高级工程师,从事博物馆管理与研究。E-mail:menglixia666@163.com。

通讯作者:

朱旭峰,男,1995年生。博士,主要从事恐龙蛋分类及显微结构研究。E-mail:zhuxufeng@bmnh.org.cn。

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目录contents

    摘要

    汝阳盆地是豫西地区的一个中—新生代断陷盆地,自2006年以来发现了一个以巨型蜥脚类恐龙为代表的恐龙动物群。本文描述了近期发现于汝阳盆地下白垩统郝岭组中的若干恐龙蛋蛋壳碎片。基于以下对蛋壳宏观形态和显微特征的分析,这些蛋壳碎片可以归入长形蛋科(Elongatoolithidae)的长形蛋属(Elongatoolithus):蛋壳外表面具分散瘤点状或线性脊状的纹饰;蛋壳自内而外分为锥体层和柱状层,二层之间的界线不清晰;锥体层中锥体排列紧密;柱状层中具波浪状的平行生长纹。这一发现代表了长形蛋属在早白垩世地层中的首次记录,进一步拓展了该蛋属的地史和古地理分布。由以往已知的化石关联推测,这些蛋壳可能与该层位中的窃蛋龙类如刘店洛阳龙(Luoyanggia liudianensis)等具有相关性,表明汝阳盆地在早白垩世晚期具有适宜该类恐龙生存与繁殖的环境。

    Abstract

    The Ruyang basin is a Meso-Cenozoic graben basin in western Henan Province. A dinosaur fauna, represented by giant sauropod dinosaurs was discovered in 2006. This paper describes several dinosaur eggshells recently discovered from the Lower Cretaceous Haoling Formation in the Ruyang basin. Based on analysis of the following macro and micro characteristics, these eggshells can be assigned to Elongatoolithus (Elongatoolithidae): dispersituberculate or linearituberculate ornamentation; two-layer eggshell structure that consists of a cone layer and a columnar layer from the bottom to top; cone layer composed of closely arranged cones; undulating parallel accretion line in the columnar layer. This discovery marks the first record of Elongatoolithus in the stratum of Early Cretaceous, which expands the geohistorical and geographical distribution of this oogenus. According to the previously known fossil connections, these eggshells may be related to the oviraptorrosaurs in the stratum such as Luoyanggia liudianensis, indicating a suitable living and breed environment for these dinosaurs at the late stage of Early Cretaceous in Ruyang basin.

  • 汝阳盆地自2006年以来发现了一个以巨型蜥脚类恐龙为代表的、属种丰富的恐龙动物群(Lü Junchang et al.,2009)。在该动物群中,已命名有蜥脚类4种,包括世界上迄今为止发现的体腔最大的恐龙——汝阳黄河巨龙(Huanghetitanruyangensis)(Lü Junchang et al.,2007),世界上最大的恐龙之一巨型汝阳龙(Ruyangosaurus giganteus)(吕君昌等,2009),大型蜥脚类恐龙——汝阳云梦龙(Yunmenglongruyangensis)(Lü Junchang et al.,2013),以及一种中等体型的蜥脚类恐龙——史家沟岘山龙(Xianshanosaurusshijiagouensis)(Lü Junchang et al.,2009); 命名有中国第一个有确凿证据的结节龙类——洛阳中原龙(Zhongyuansaurusluoyangensis)(徐莉等,2007); 命名一种窃蛋龙类——刘店洛阳龙(Luoyanggialiudianensis)(Lü Junchang et al.,2009)。此外还发现有未命名的似鸟龙类(Lü Junchang et al.,2009)、棘龙类(Lü Junchang et al.,2009)、鲨齿龙类(Lü Junchang et al.,2016)、禽龙类(张逸阳等,2013)以及可能的驰龙类化石(Lü Junchang et al.,2009)。从恐龙动物群的组合面貌来看,其时代应为早白垩世晚期至晚白垩世早期,结合微体古生物化石证据,其时代进一步被缩小至早白垩世晚期(徐莉等,2012; 王德有,2013)。

  • 汝阳盆地曾发现有恐龙蛋(徐莉等,2017),但尚未有正式的分类描述。2021年,本文作者在洛阳市汝阳县刘店镇曹家村郝岭组中采集到若干蛋壳碎片,经切片鉴定后归入长形蛋科(Elongatoolithidae)长形蛋属(Elongatoolithus),代表了该蛋属在早白垩世地层中的首次发现,本文予以记述。

  • 1 地质背景

  • 汝阳盆地是豫西地区的一个中—新生代断陷盆地,以往地质学界把汝阳盆地九店组以上的一套碎屑岩系划分为古近系陈宅沟组、蟒川组、石台街组。2006年以来,河南自然博物馆在汝阳三屯—刘店一带的原“蟒川组”内发现了密集分布的恐龙化石。徐莉等(2012)在该区开展了大比例尺地质填图和剖面测制,综合微体、无脊椎动物和脊椎动物化石鉴定及同位素测年、稳定同位素、地球化学、岩石簿片等样品分析,认为分布于汝阳盆地中部(上店—刘店地区)的这套碎屑岩原“陈宅沟组、蟒川组”应该划为白垩系,新建白垩系下河东组、郝岭组、上东沟组来代表这一套含恐龙化石的碎屑岩系(图1)。

  • 下河东组相当于原“陈宅沟组”,平行不整合于九店组之上,下部以紫红色复成分砾岩为主,中部以淡紫红色砾岩为主,夹棕红色含砾泥质粉砂岩、灰白色岩屑砂岩、砂砾岩,上部为淡紫红色、灰绿色砾岩与紫红色泥质粉砂岩、紫红色粉砂质泥岩互层。郝岭组相当于原“蟒川组”的中下部,整合于下河东组之上,下部为淡紫红色砾岩与棕红色泥质粉砂岩互层,夹灰白色岩屑砂岩,中部为绿灰色砾岩、砂砾岩与棕红色泥质粉砂岩、粉砂质泥岩互层,夹棕黄色岩屑砂岩、灰色、灰绿色泥岩,上部为紫红、绿灰色砾岩、黄绿色岩屑砂岩与紫红色泥质粉(细)砂岩互层,夹灰白色砂砾岩。上东沟组相当于原“蟒川组”的上部,整合于郝岭组之上,以棕红色泥质粉砂岩、灰白色砂砾岩为主,夹含砾泥质粉砂岩、砾岩,与上覆石台街组呈平行不整合接触。(徐莉等,2017)。

  • 徐莉等(2012)在郝岭组下部灰绿色粉砂质泥岩和靠近恐龙化石层位附近的紫红色粉砂质泥岩中采集了大量的微体化石,获得一些介形类、孢粉类和轮藻类。介形类主要属种有:ZiziphocypriscostataZiziphocypris sp.,CyprideaunicostataCyprideaconcinaCypridea sp.,CandonashangshuiensisCandonaauritaCandona sp.,Candoniella sp.,DarwinulaleguminellaDarwinulaContractaEucyprisinfantilisEucypris debilis,多数分子为早白垩世常见分子的组合特征化石; 蕨类植物孢子以HsuisporitesCicatricosisporitesDensoisporites为优势分子,裸子植物花粉以Classopollis占优势,孢粉组合的地质时代更倾向于早白垩世中晚期; 轮藻化石Clypeator-Flabellochara-Mesochara-Aclistochara组合反映了早白垩世中晚期的特征(朱红卫等,2020)。结合恐龙动物群的组合特征,郝岭组的地质时代大致为早白垩世晚期。

  • 2 材料与方法

  • 汝阳盆地恐龙蛋发现于洛阳市汝阳县刘店镇曹家村东岩郝岭组(K1h)(图1b),野外仅发现恐龙蛋壳,未发现完整的恐龙蛋。蛋壳外表面纹饰清晰,棱角分明(图2),磨蚀不显著,表明为近原地埋藏,可能为恐龙出壳后遗留的破碎蛋壳。

  • 首先对这些蛋壳样品进行超声波清洗,取保存较好的5片蛋壳进行硬组织学切片。蛋壳被包埋在EXAKT Technovit 7200单组分树脂中,使用EXAKT 300CP切割机进行切割,并使用EXAKT 400CS变速磨片机配合水磨砂纸磨至适合镜下观察的厚度后抛光。再使用偏光显微镜对制作完成的薄片进行观察和照相,使用ImageJ软件在显微照片中进行测量。硬组织学切片的制作和观察均在中国科学院脊椎动物演化与人类起源重点实验室完成。

  • 图1 化石产地及地层分布图

  • Fig.1 Fossil site and stratum distribution

  • (a)—汝阳盆地在河南省的地理位置;(b)—汝阳盆地中部地质略图,红色三角形表示蛋壳产出位置

  • (a) —geological position of the Ruyang basin in Henan Province; (b) —simplified geological map of the middle part of the Ruyang basin, the red triangle indicates the eggshell site

  • 图2 汝阳盆地下白垩统蛋壳外表面二次电子图像

  • Fig.2 SEM image of outer surface of the eggshells from the Lower Cretaceous of Ruyang basin

  • (a)~(d)—室内样品编号N2032105-1;(e)~(i)—室内样品编号N2032105-2

  • (a) ~ (d) —specimen No. N2032105-1; (e) ~ (i) —specimen No. N2032105-2

  • 3 系统古生物学

  • 长形蛋科  Elongatoolithidae,Zhao,1975

  • 长形蛋属  Elongatoolithus,Zhao,1975

  • 长形蛋未定种  Elongatoolithus sp.

  • 标本编号 室内编号为N2032105-1(野外编号为RY210517-1)的4片蛋壳(其中2片对应的切片编号为210607-4、5); 室内编号为N2032105-2(野外蛋壳编号为RY210518-1)的5片蛋壳(其中3片对应的切片编号为210607-1、6、7)(图3)。

  • 产地及层位 河南省洛阳市汝阳县刘店镇曹家村,下白垩统郝岭组。

  • 描述 蛋壳呈灰黑色。外表面具分散瘤点状(dispersituberculate; 图2b、c、f、g、i)或线性脊状纹饰(linearituberculate; 图2a、d、e、h)。蛋壳厚度较薄,不含纹饰的厚度平均为697 μm,范围为595~782 μm,含纹饰的厚度平均为902 μm,范围为834~965 μm。蛋壳由内而外分为锥体层和柱状层,界线不截然。锥体层由一层紧密排列的锥体组成,厚度平均为124 μm,范围为113~136 μm。柱状层呈马赛克状消光,具发达的平行生长纹,起伏与外表面一致。

  • 4 讨论

  • 根据分散瘤点状纹饰和线性脊状纹饰、锥体层和柱状层的两层结构、平行起伏的生长纹等特征,这些蛋壳可以归入长形蛋类(包括长形蛋科Elongatoolithidae和巨型长形蛋科Macroelongatoolithidae)(赵资奎,1975)。在长形蛋类中,现存10个蛋属,本次发现的这些蛋壳的锥体层与柱状层界线不截然,不同于巨形蛋属(Macroolithus)(赵资奎,1975; Mikhailov,1994)、巨型长形蛋属(Macroelongatoolithus)(王强等,2010b)中锥体层与柱状层之间的截然界线; 蛋壳纹饰高度一般,与TrachoolithusMikhailov,1994)、波纹蛋属(Undulatoolithus)(Wang Qiang et al.,2013)中接近甚至大于蛋壳厚度(不含纹饰)的纹饰高度不符,也与比较光滑的南雄蛋属(Nanhsiungoolithus)(赵资奎,1975)显著不同; 柱状层生长纹分布较为均匀,与黑山蛋属(Heishanoolithus)(赵宏等,1999)集中于蛋壳近外表面的生长纹不同,后者的蛋壳外表仅见分散瘤点状纹饰,且瘤点一般较其他长形蛋类较细; EllipsoolithusMohabey,1998)的纹饰也为分散瘤点状或线性脊状,但分布明显较其他蛋属密集,难以与本文标本比较; 蛋壳壳单元结构不明显,与保留明显壳单元边界的网纹副长形蛋(Paraelongatoolithus reticulatus)不同,后者的外表纹饰呈网状的浅脊,锥体较为宽圆,锥体层与柱状层界线较为清晰(王强等,2010a),也可以此与之区分; 巨型纺锤蛋属(Megafusoolithus)是巨型长形蛋科的另一蛋属,虽然具有与本文标本相似的蛋壳结构(同样与长形蛋属相似),但其蛋壳最小厚度也达到了1.45 mm(王强等,2010b),较本文标本超出过多。根据以上对比,这些蛋壳仅与长形蛋属(Elongatoolithus)的特征较为吻合(赵资奎,1975),在该属内,其蛋壳厚度范围与长形长形蛋(E. elongatus)和大长形蛋(E. magnus)的变异范围基本符合(表1),但由于这两个蛋种的差异主要体现在蛋体大小上,仅凭蛋壳特征是无法区分的(赵资奎等,2015),在缺少完整蛋体的情况下,暂不便将本文标本归入任一蛋种。

  • 表1 汝阳盆地下白垩统产恐龙蛋蛋壳与现有长形蛋类蛋壳的厚度对比

  • Table1 Thickness comparison of current elongatoolithid eggshells and dinosaur eggshells from the Lower Cretaceous of Ruyang basin

  • 汝阳发现的这批蛋壳标本的保存状况整体较好,蛋壳近外表面结构较为完整,但在锥体层完整程度上略有差异,其中编号为210607-6、7的径切面切片(图3c、d)中观察到的锥体结构较为完整。纹饰的定向性在不同蛋壳标本上具有些微差异,反映了蛋壳在蛋体上位置的不同,这一现象在长形蛋类中较为常见,分散瘤点状纹饰常出现在蛋体两端,向蛋体中部逐渐过渡为线性脊状纹饰(Mikhailov,1997)。纹饰相对高度的不同可能因蛋壳位置和切片位置不同而受到影响,如210607-4的径切面切片(图3e)中观察到的蛋壳厚度无纹饰厚度相对较高,而纹饰高度较低,可能代表其为靠近蛋的端部位置的蛋壳,而其余几片蛋壳结合其外表纹饰的特征,也可大体判断为来自于蛋体上的不同位置(图4)。

  • 长形蛋类的分布较广,是亚洲和北美洲的白垩纪陆相地层中较为常见的蛋化石类型(Wang Qiang et al.,2013; Simon et al.,2019),主要发现于晚白垩世地层中,此前仅有中国辽宁省的常氏黑山蛋(Heishanoolithuschangii)(赵宏和赵资奎,1999),蒙古南戈壁地区的MacroolithusmutabilisMikhailov,1994)以及美国犹他州的Macroelontaoolithus carlyleiZelenitsky et al.,2000)曾报道产于早白垩世地层中。长形蛋属(Elongatoolithus)于早白垩世地层中尚未发现。此次在河南省汝阳地区下白垩统中发现长形蛋属(Elongatoolithus)的蛋壳碎片尚属首次,拓展了该蛋属的地史和古地理分布范围。

  • 图3 汝阳盆地下白垩统长形蛋未定种(Elongatoolithus sp.)径切面显微照片

  • Fig.3 Micrographs of radial thin section of Elongatoolithus sp. from the Lower Cretaceous of Ruyang basin

  • (a)、(c)、(d)—室内样品编号N2032105-2;(a)—切片编号210607-1,见图2f;(c)—切片编号210607-6,见图2h;(d)—切片编号210607-7,见图2g;(b)、(e)—室内样品编号N2032105-1;(b)—切片编号210607-5,见图2a;(e)—切片编号210607-4,见图2b

  • (a) , (c) , (d) —specimen No. N2032105-2; (a) —thin section No.210607-1, see Fig.2f; (c) —thin section No.210607-6, see Fig.2h; (d) —thin section No.210607-7, see Fig.2g; (b) , (e) —specimen No.N2032105-1; (b) —thin section No.210607-5, see Fig.2a; (e) —thin section No.210607-4, see Fig.2b

  • 图4 汝阳盆地下白垩统蛋壳在长形蛋类蛋体上的部位复原

  • Fig.4 Position reconstruction of the eggshells from the Lower Cretaceous of Ruyang basin on a complete elongatoolithid egg

  • 根据已有的化石发现,长形蛋类被认为是属于窃蛋龙类或与其亲缘关系相近的恐龙所产(Varricchio et al.,2004; 王强等,2010; Pu Hanyong et al.,2017)。汝阳盆地下白垩统郝岭组中发现有属窃蛋龙类的刘店洛阳龙(Luoyanggialiudianensis)(Lü Junchang et al.,2009),是这些蛋可能的产蛋者。这也表明早白垩世晚期的汝阳盆地的环境对此类恐龙的生存与繁殖是较为理想的。

  • 5 结论

  • (1)本次在汝阳盆地中发现的若干蛋壳可归入长形蛋属未定种(Elongatoolithus sp.)。

  • (2)该批材料代表了长形蛋属(Elongatoolithus)在下白垩统中的首次记录。

  • (3)丰富的恐龙骨骼与新发现的蛋壳表明,早白垩世晚期汝阳盆地的环境不仅适宜窃蛋龙类等恐龙的生存,对其繁殖也十分有利。

  • 致谢:中国科学院古脊椎动物与古人类研究所尤海鲁与刘永红在野外采集了标本,刘永红完成了蛋壳的显微结构切片与显微照相,本文作者在此一并表示感谢。

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