化石及现生真蕨类植物的根系:形态演化及古环境意义

根是植物的重要营养器官,对维持植物生理活动及生态系统物质循环和能量流动极为重要。真蕨类植物的根系以可预测的方式起源于枝,被称为枝生根,通常也被称为不定根。相比于对种子植物根系的广泛研究,对真蕨类植物根系的认识较为薄弱。通过广泛收集各类文献中有关真蕨类根系的信息,以及观察大量化石、现生植物标本,对若干化石及现生真蕨类植物根系的形态学特征进行了综述。在繁盛于中、晚泥盆世的早期真蕨类植物中,密集的枝生根着生于茎轴、或推测的根状茎、或主干膨大的基部上;根系的分枝罕见,若存在,则仅表现为等二歧式分枝;而在真蕨纲冠群植物中,枝生根密集或稀疏,根系呈单轴式分枝,侧根发育,可达2级甚至更多。真蕨类根系统的形态演化可采用"顶枝学说"进行解释:真蕨类植物的根系具有一定的演化保守性,均为枝生根;早期真蕨类的原始根系可视为不分枝的或偶见二歧式分枝的顶枝束;经历顶枝束的越顶生长,侧根系统在真蕨纲冠群中演化出现。真蕨类根系的构型、形态、解剖结构、生物量分配等信息可在化石记录中得以揭示,这些信息可为探讨植物根系的演化及研究地史时期的植物—土壤系统提供重要依据。最后,简述了这方面研究的1个案例,即北京西山侏罗纪地层中的真蕨类根系化石及根系—土壤系统。     

我国晚中生代单缝孢真蕨植物兴衰史及意义

从大化石和孢粉资料两方面总结了我国晚中生代单缝孢类真蕨植物的特点及其兴衰史，认为单缝孢类真蕨植物的在我国北方地区早白垩世植物群中的出现，繁盛和衰退与该地区的潮湿，特别是温暖潮湿型古气候背景关系十分密切，据此阐明了该类植物的潮湿，特别是温暖潮湿型古气候的指示意义，其中多数化石与现生真蕨Ａｔｈｙｒｉｕｍ，Ｄｒｙｏｐｔｅｒｉｓ属等的形态和繁殖器官结构非常相拟，反映真蕨类进入了一个新的演化阶段，并提出Ｄｒ     

黔南地区晚二叠世大隆组植物群生态学研究

本文化石采自贵州南部安顺和惠水地区晚二叠世大隆组地层,共计16属21种,分别属于蕨类植物石松纲、楔叶纲、真蕨和种子蕨纲,裸子植物科达纲、银杏纲、松柏纲,另外有种子化石和分类位置不明类型匙剑叶,对植物群古生态学的研究表明,当前植物群为以松柏类为主的高地植物群落,以裸子植物为主要成分,群落组成为：高层为由以斯氏松为主的松柏类、鳞木类、科达纲、银杏类植物,中层为芦木类等,底层为低矮的真蕨、种子蕨类;对各门类植物群特征及群落分析表明,黔南地区晚二叠世时位于北半球低纬度地区,整体气候炎热,雨量充沛,植物茂盛,但具短暂的干旱无雨季节,为热带-亚热带半落叶季雨林气候     

福建永安下白垩统坂头组枝脉蕨属一新种Cladophlebis yonganensis sp.nov.

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.      

北京西山中侏罗世植物群的古气候意义

在北京市门头沟区九龙山南坡中侏罗统的下窑坡组、上窑坡组和龙门组采集植物化石22属38种。统计资料表明,北京西山中侏罗世植物群以真蕨纲(27.6%)、苏铁纲(25%)和银杏纲(25%)为主,其中真蕨纲的Coniopteris、Cladophlebis分别出现5和17个形态种。本文通过现生近缘种类分析法和植物的生长状态分析研究古气候。研究表明,含量高的植物化石的现生近缘种类紫萁科、蚌壳蕨科和苏铁纲植物主要生长于温热潮湿的热带或亚热带气候区;Cladophlebis较高的分异度,苏铁纲大型的羽状复叶,木贼目宽大的茎干等植物特征,同样指示较为湿热的气候环境。因此,北京西山中侏罗世植物群反映一种温热潮湿的亚热带气候。并且,化石的属种和数量自下窑坡组、上窑坡组到龙门组的降低反映研究区气候由温热潮湿向干热方向的演变。     

中生代主要植物化石的古气候指示意义

中生代植物群以真蕨纲、苏铁纲、银杏纲、松柏纲为主,另有苔藓类、石松纲、楔叶纲及其他种子植物等。从化石的形态和结构、埋藏特点、分布及与现生类型的对比等方面，概要地分析了中生代植物群常见分子的古气候指示意义。其中,反映温暖潮湿气候或生境的主要有苔藓类,石松纲的Lycopodites和Selaginellites属等,楔叶纲,真蕨纲的单缝孢类、早白垩世的蚌壳蕨科、海金砂科的Ruffordia属、紫萁科及可能相关的小羽片大型的Cladophlebis,银杏纲银杏目的主要类型,松柏纲杉科的Athrotaxoites、Cunninghamia、Cephalotaxopsis、Sequoia等,紫杉科的Taxus、Torreya等，苏铁杉科Ferganiella、Podozamites、Swedenborgia等以及形态属Elatocladus；反映温凉气候的主要有松柏纲的松科和银杏纲的茨康目；适应热带、亚热带气候的包括真蕨类的海金沙科、合囊蕨科、马通蕨科、双扇蕨科、桫椤科、里白科以及中国蕨科的Onychiopsis等,苏铁目和本内苏铁目的主要类型；指示偏干旱型气候的主要有石松纲的Pleuromeia和Isoetites, 真蕨纲的Weichiselia, 本内苏铁目的Otozamites、Zamites、Ptilophyllum等,松柏纲掌鳞杉科的Pseudoflenelopsis、Flenelopsis、 Hirmeriella和可能属于掌鳞杉科的Brachyphyllum和Pagiophyllum,柏科的Cupressinocladus以及买麻藤目的Ephedrites等。     

新疆早侏罗世植物群特征

本文根据新疆早侏罗世各组中所产的植物化石,提出了疆新早侏罗世植物群分属于毛羽叶—锥叶蕨植物群与锥叶蕨—拟刺葵植物群的过渡植物群和锥叶蕨—拟刺葵植物群.过渡植物群中苏铁类比较丰富.锥叶蕨—拟刺葵植物群以真蕨类和银杏类为主,真蕨类中锥叶蕨,托第蕨,枝脉蕨最为发育.两个植物群按照不同层位均可分出上下两个组合.本文还探讨了八道湾组时代,认为无疑属早侏罗世,塔里奇克组植物组合也应属早侏罗世(塔里奇克组的地质时代为T_(?)—J_1).早侏罗世早期新疆气侯曾一度炎热,向后转为湿暖潮湿.     

内蒙古乌达地区早二叠世植物群和古生态研究

研究的植物化石采自内蒙古乌达地区早二叠世地层,共计11属20种,分属于石松纲、楔叶纲、瓢叶目、真蕨纲、种子蕨纲和苏铁纲。研究结果表明,植物群含有许多华夏植物群分子和地方性分子,为典型的华夏植物群。根据植物群分子的总体组成,植物群的时代为早二叠世,相当于阿瑟尔期(Asselian)、萨克马尔期(Sakmarian)、亚丁斯克期(Artinskian)和空谷期(Kungurian)。在研究植物群的基础上,对植物群的古生态进行了研究,建立了早二叠世Lepidodendron posthumii Paratingia datongensis植物群落。这一植物群落在二叠纪主要分布在沼泽和低地环境,以大量出现的蕨类和种子蕨分子为特征。植物群落为原地保存,借助于化石资料,植物的生活型得以重建,包括乔木、树蕨、灌木和草本。植物群落指示早二叠世本区为温暖潮湿的气候条件。     

云南临沧晚中新世一种水龙骨科植物化石的修订及其生物学意义

云南现代真蕨类植物资源丰富,蕨类植物多样性的地史起源,必须从化石记录入手。在云南临沧上中新统邦卖组植物化石采集中发现了槲蕨属1块不育叶和2块腐殖叶的新材料,这些标本为修订Drynaria propinqua Wen et al.,2013,以及揭示该种不育叶和腐殖叶的特征提供了新的材料。通过与槲蕨属国内外报道的化石种和现生种的详细比较,将其重新定名为Drynaria cf.propinqua。Drynaria cf.propinqua的发现,表明云南临沧晚中新世的气候与现今中国西南地区温暖湿润的气候类似,这些附生植物的生活习性表明,临沧地区复杂分层的森林生态系统至少在晚中新世已经确立。     

松潘—甘孜造山带容须卡岩浆-穹隆地质特征及构造演化

位于松潘—甘孜造山带雅江穹隆群北东部的容须卡岩浆底辟穹隆区,经历了多期构造演化,内部构造极其复杂。为厘清容须卡地区岩浆与穹隆的演化关系,通过野外地质调查及室内综合研究,探讨了容须卡穹隆的地质特征及构造演化。该穹隆中心发育无根或肠状褶皱,及“S”型、“Z”型褶皱和“A”型平卧褶皱; 穹隆外围发育叠加褶皱,反映造山带早期SN向和EW向收缩挤压。该穹隆主要经历了晚三叠世(印支末期)SN向与EW向的“双向挤压”作用; 成穹期岩浆向上侵位时限为(214.4±1.2) Ma; 成穹后经历了挤压推覆和应力松弛阶段; 早中新世(17~10 Ma),由于青藏高原东南缘快速抬升,松潘—甘孜造山带发育NW-SE向鲜水河左旋走滑断裂,使容须卡地区形成一系列NNW向韧脆性和脆性破碎带。     

鄂尔多斯盆地北岔沟砂岩沉积特征及其对天然气成藏的控制作用*

北岔沟砂岩为华北地台二叠系山西组与太原组划分对比的主要标志层,在鄂尔多斯盆地呈大面积连片分布,是上古生界重要的天然气储集层。依据地面露头调查结果与勘探资料,结合前人有关古构造和岩相古地理研究成果,笔者对鄂尔多斯盆地北岔沟砂岩的沉积特征进行了详细分析。结果表明,受益于物源区大量高石英含量的粗碎屑物质在短时间内的集中供应,北岔沟砂岩在地层中位置稳定,以含砾粗粒石英砂岩为主,石英含量高达90%左右,砂体叠置紧密,单砂体之间很少见泥岩隔夹层。这些砂体分别隶属于冲积扇、砾质辫状河和辫状河三角洲沉积体系,在鄂尔多斯盆地北部高、南部次高、中南部低的古地理背景控制下,由盆地南、北边缘向中部依次推进,沉积中心区以北发育4个规模宏大的三角洲沉积主砂带,以南发育6个规模较小的主砂带。受各主砂带砂体规模、矿物组分含量及其成岩作用差异影响,各砂带的储集物性具有较大变化,其中砂体规模最大、石英含量最高、长石等易溶矿物在酸性地层水作用下溶蚀彻底的杭锦旗—榆林—子洲—清涧—永和—吉县主砂带,储集能力最好,最有利于天然气的大规模聚集成藏,是鄂尔多斯盆地北岔沟砂岩天然气进一步勘探开发的有利场所。     

A review of some fossil plant compressions associated with Mississippian and Pennsylvanian coal deposits in the central Appalachians, Illinois Basin, and elsewhere in the United States

Fossil plant compressions and impressions are abundantly associated with coals of Mississippian and Pennsylvanian age in the central Appalachians, Illinois Basin, and elsewhere in the United States. Remains of the aerial plant parts occur most commonly in shale directly overlying coal, especially in the vicinity of contemporaneous paleochannels. Most roots and a few stumps are found preserved in situ; however, most aerial plant fossils were transported prior to burial. There is good evidence that fossil plants preserved in roof shales are mostly the same types of plants from which the coal formed. Mississippian coal floras are lower in diversity than later ones. Quantitatively important plants include only 2 lycopod genera, 1 arthrophyte genus, 1 fern genus, and 5 seed fern genera. Pennsylvanian floras, on the other hand, include abundant representatives of 5 lycopod genera, 2 arthrophyte genera, 4 fern genera, 5 seed fern genera, and 1 coniferophytic genus. The swamp floras were never in equilibrium, though change appears to have been particulary rapid near the boundary between the Mississippian and the Pennsylvanian.     

Paleoecology of Oligocarpia gothanii Halle from the Middle Permian of Southeastern Shanxi, North China

Fossil ferns are abundant and diverse in the Permian Cathaysian Flora of Shanxi, North China. Although plants assemblages of the Carboniferous and Permian have been extensively investigated, their palaeoecology and especially the paleoecology of ferns has not been studied in detail. Recent reinvestigation of Oligocarpia gothanii has revealed its growth habit and cloning reproduction strategy, but most previous studies lack integrated sedimentological and taphonomic data. In this study, new materials of O. gothanii have been collected from a floodplain setting in the middle Permian Lower Shihhotse Formation of southeastern Shanxi. These plant fossils were found in a lenticular claystone associated with rooting structures. They are characterized by aphlebia at the base of the penultimate rachis. Sedimentological, taphonomic and morphological analyses were conducted to understand the growth habit and ecology of the fern. The result indicates that the Oligocarpia gothanii had a prostrate, ground cover growth habit, and was dominant in pioneer floras that colonized disturbed floodplains.     

Paleoecology of Oligocarpia gothanii Halle from the Middle Permian of Southeastern Shanxi,North China

Fossil ferns are abundant and diverse in the Permian Cathaysian Flora of Shanxi, North China. Although plants assemblages of the Carboniferous and Permian have been extensively investigated, their palaeoecology and especially the paleoecology of ferns has not been studied in detail. Recent reinvestigation of Oligocarpia gothanii has revealed its growth habit and cloning reproduction strategy, but most previous studies lack integrated sedimentological and taphonomic data. In this study, new materials of O. gothanii have been collected from a floodplain setting in the middle Permian Lower Shihhotse Formation of southeastern Shanxi. These plant fossils were found in a lenticular claystone associated with rooting structures. They are characterized by aphlebia at the base of the penultimate rachis. Sedimentological, taphonomic and morphological analyses were conducted to understand the growth habit and ecology of the fern. The result indicates that the Oligocarpia gothanii had a prostrate, ground cover growth habit, and was dominant in pioneer floras that colonized disturbed floodplains.     

The Albian fern flora of Alexander Island,Antarctica

The Albian Alexander Island macrofossil flora from the Antarctic Peninsula preserves a diverse community of liverworts (Marchantiophyta), ferns (Polypodiopsida), Lycopodiales, Equisetales, Cycadales, Ginkgoales, seed-ferns (Bennettitales and Pentoxylales), Coniferales, and the first representatives of angiospermous leaves in Antarctica. Despite the presence of angiosperms in this assemblage, ferns are the most diverse element of the flora and are also ecologically dominant, while angiosperms contribute a smaller component to floristic diversity and have low abundance. Here we describe 11 fern taxa from this assemblage. The fossils are assigned to Cladophlebis, Sphenopteris and two newly created genera. The new genera and species are described under Adiantitophyllum serratum gen. et. sp. nov. and Nunatakia alexanderensis gen. et. sp. nov., and the new species are recognized as Cladophlebis dissecta sp. nov., Cladophlebis drinnanii sp. nov., Cladophlebis macloughlinii sp. nov. and Sphenopteris sinuosa sp. nov. In total, there are 24 fern species known from Alexander Island. In comparison to older floras (Jurassic) there is a greater diversity of ferns, while latest Cretaceous floras preserve significantly fewer fern species and more angiosperms. Possible factors that might account for such high fern diversity are high rainfall or generally humid conditions, regular disturbances by flooding and occasionally fire, and the preservation of a diverse range of fern communities that represent several palaeoenvironments.     

Lower Permian stems as fluvial paleocurrent indicators of the Parnaíba Basin,northern Brazil

A comprehensive biostratinomic study was carried out with abundant stems from the Lower Permian Motuca Formation of the intracratonic Parnaíba Basin, central-north Brazil. The fossils represent a rare tropical to subtropical paleofloristic record in north Gondwana. Tree ferns dominate the assemblages (mainly Tietea, secondarily Psaronius), followed by gymnosperms, sphenophytes, other ferns and rare lycophytes. They are silica-permineralized, commonly reach 4 m length (exceptionally more than 10 m), lie loosely on the ground or are embedded in the original sandstone or siltstone matrix, and attract particular attention because of their frequent parallel attitudes. Many tree fern stems present the original straight cylindrical to slightly conical forms, other are somewhat flattened, and the gymnosperm stems are usually more irregular. Measurements of stem orientations and dimensions were made in three sites approximately aligned in a W–E direction in a distance of 27.3 km at the conservation unit “Tocantins Fossil Trees Natural Monument”. In the eastern site, rose diagrams for 54 stems indicate a relatively narrow azimuthal range to SE. These stems commonly present attached basal bulbous root mantles and thin cylindrical sandstone envelopes, which sometimes hold, almost adjacent to the lateral stem surface, permineralized fern pinnae and other small plant fragments. In the more central site, 82 measured stems are preferentially oriented in the SW–NE direction, the proportion of gymnosperms is higher and cross-stratification sets of sandstones indicate paleocurrents mainly to NE and secondarily to SE. In the western site, most of the 42 measured stems lie in E–W positions. The predominantly sandy succession, where the fossil stems are best represented, evidences a braided fluvial system under semiarid conditions. The low plant diversity, some xeromorphic features and the supposedly almost syndepositional silica impregnation of the plants are coherent with marked dry seasons. Thick mudstones and some coquinites below and above the sandy interval may represent lacustrine facies formed in probably more humid conditions. The taphonomic history of the preserved plants began with exceptional storms that caused fast-flowing high water in channels and far into the floodplains. In the eastern site region, many tree ferns only fell, thus sometimes covering and protecting plant litter and leaves from further fragmentation. Assemblages of the central and western sites suggest that the trees were uprooted and transported in suspension (floating) parallel to the flow. Heavier ends of stems (according to their form or because of attached basal bulbous root mantle or large apical fronds) were oriented to upstream because of inertial forces. During falling water stage, the stems were stranded on riverbanks, usually maintaining the previous transport orientation, and were slightly buried. The perpendicular or oblique positions of some stems may have been caused by interference with other stems or shallow bars. Rare observed stems were apparently waterlogged before the final depositional process and transported as bedload. The differences of interpreted channel orientations between the three sites are expected in a braided fluvial system, considering the very low gradients of the basin and the work scale in the order of tens of kilometers. The mean direction of the drainage probably was to east and the flows apparently became weaker downstream. This study seems to provide reliable data for paleocurrent interpretations, especially considering areas with scarce preserved sedimentary structures.     

早白垩世真蕨植物化石的四个新种

本文据产于内蒙古霍林河盆地早白垩世地层的生殖羽片材料建立了膜蕨科的Hymenophyllites linearifolius sp.nov.和铁线蕨科的Adiantopteris eleganta sp.nov.,并从植物形态、生殖器官和生态特征等方面与现生类别进行了对比。本文还描述了同一产地和层位的Cladophlebis latibasis sp.nov,及产自辽宁铁法盆地早白垩世的Sphenopteris liaoningensis sp.nov.。     

Frullania pinnata spec. nov. (Frullaniaceae,Porellales), a new leafy liverwort in mid-Cretaceous Burmese amber from Myanmar

An inclusion in Cretaceous amber from Myanmar is described as a new fossil species in the extant liverwort genus Frullania. The name Frullania pinnata is proposed for the taxon that is characterized by entire underleaves paired with a conspicuous pinnate branching pattern, two distinctive and stable morphological features. Entire underleaves are known in several extant Frullania species but had not previously been documented in Frullania in amber. The combination of morphological characters in this new fossil species is unknown in any crown group lineage of Frullania, and may very well represent a stem lineage element of the genus. This discovery is important because it expands our understanding of the diversity of Frullaniaceae in the Burmese amber forest as well as important ramifications for the phylogenetic reconstruction of extant Frullania lineages.     

中国白垩纪植物群与生物地层学

早白垩世时中国可划分出北方、南方和藏南3个植物地理区。北方植物地理区可归入瓦赫拉梅耶夫的西伯利亚加拿大植物地理区,发育有热河、阜新和大砬子3个植物群。热河植物群产于辽西义县组和九佛堂组及其他相当地层,时代为早白垩世早期,以苏铁纲和松柏纲占主导地位。阜新植物群赋存于辽西的沙海组和阜新组及相当地层,以真蕨纲、银杏纲和松柏纲共同繁盛,苏铁纲和木贼目较丰富为特点。由早而晚可以进一步划分为Acanthopteris-Ginkgoco riacea组合、Ruffordia goepperti-Dryopterites组合和Ctenis lyrata-Chilinia组合,分别产于辽西的沙海组、阜新组中下部和阜新组上部。大砬子植物群产于吉林延吉盆地的大砬子组和松辽盆地的泉头组,被子植物占优势且掌鳞杉科丰富。南方植物地理区属于瓦赫拉梅耶夫的欧洲中国植物地理区的范畴,苏铁纲、鳞叶或锥叶型松柏和小羽片小而叶膜厚的真蕨类(主要是Cladophlebis)占主导地位,缺少银杏纲、真蕨纲的蚌壳蕨科及单缝孢类型等,为热带、亚热带植物群,可进一步划分为东部、西藏北部和中部3个亚区。其中,东部亚区滨邻古太平洋,以浙江、福建和山东莱阳盆地等的植物为代表,以鳞叶和锥叶型松柏与本内苏铁Ptilophyllum占优势。该亚区植物群可以进一步划分为3或4个植物组合,自早至晚包括Cupressinocladus-Pagiophyllum组合、Cladophlebis-Ptilophyllum组合、Ruffordia-Zamiophyllum组合和Suturovagina-Frenelopsis组合。西藏北部亚区邻近古特提斯洋东北岸,植物群与东部亚区的基本特点一致,但真蕨类更为繁盛,特别是海金沙科Klukia属和里白科的Gleichenites相当丰富,并有海金沙科的Scleropteris属和马通蕨科存在,裸子植物以苏铁纲为主,松柏纲相对较少,可进一步划分为两个组合。中部亚区介于上述两个亚区之间,由于气候干旱,植物群不发育,以甘肃酒泉盆地、民和盆地所产化石为代表,特点是鳞叶、锥叶型松柏类为主,掌鳞杉科较发育,其他类型罕见。藏南植物地理区属于澳大利亚植物地理区的范畴,只发现于喜玛拉雅地区。晚白垩世植物群只发现于东北、华南、西藏等地的少数地点和少数层位,研究程度较低,还不能进一步划分出植物地理区系和组合。以植物化石为主要依据,结合其他生物和非生物证据,建立了中国不同植物地理区白垩纪含植物化石的地层及相关地层的对比关系。