柏科4个属(Fokienia、Cupressus、Chamaecyparis和Juniperus)植物雌球果的发育(英文)

通过扫描电镜和常规石蜡切片技术,观察了柏科4个属(Fokienia、Cupressus、Chamaecyparis和Juniperus)植物雌球果中胚珠的发育及苞片的结构变化。在所有研究的种类中,可育苞片腋部最先观察到的结构是一扁平的突起,并在其上分化出发育为胚珠的胚珠原基。在雌球果的发育过程中,未观察到独立的珠鳞发育。不同的种中,胚珠的数量和发育顺序有所不同,但苞片的发育是相似的。传粉前,苞片的结构与叶相似。传粉后,由于剧烈的居间生长,苞片发育为盾形,形成球形的球果。另外,在发育后期,苞片内维管系统变得复杂,并且在近轴面有反向的维管束发育。我们还对柏科植物雌球果的形态学特性及其可能的演化趋势进行了讨论。     

北美香柏雌球果的发育

用扫描电镜(SEM)观察了北美香柏 Thuja occidentalis 雌球果的发育过程。在北京,北美香柏的雌球果是在八月初由营养芽转变而来,雌球果一般有4~6对苞片,中间2~3对可育,每一苞片腋部着生两枚胚珠,在可育苞片腋部最先观察到一扁平的隆起,并在其上分化出两个胚珠原基,接着分化出珠被和珠心,最后形成扁平而两侧对称的胚珠。在北美香柏雌球果发育过程中,约一半的雌球果在2~3对可育苞片中位于下面的1~2对的腋部产生3个胚珠原基,中间一个较小,并在以后的发育中逐渐退化。由此推测北美香柏的雌球果可能是由祖先类群中每一苞片具多于2个胚珠的雌球果演化而来。在光镜下对雌球果维管系统的观察发现,传粉前幼小雌球果的苞片内仅有一束维管束,传粉后随着苞片基部的居间生长,有4—8束维管束在苞片内形成,但是新发育的维管束木质部和韧皮部相对位置与正常叶性器官一致,这与在以往报道的柏科植物成熟雌球果的苞片中均有反向维管束的发育不同。北美香柏雌球果早期发育和维管束分析结果支持傅德志和杨亲二提出的解释裸子植物生殖器官形态演化的“苞鳞-种鳞复合体”理论。关键词北美香柏;雌球果的发育;胚珠分化;SEM     

柏科4个属(Fokienia、Cupressus、Chamaeyparis 和Juniperus)植物雌球果的发育

通过扫描电镜和常规石蜡切片技术,观察了柏科4个属(Fokienia、Cupressus、Chamaeyparis 和Juniperus)植物雌球果中胚珠的发育及苞片的结构变化.在所有研究的种类中,可育苞片腋部最先观察到的结构是一扁平的突起,并在其上分化出发育为胚珠的胚珠原基.在雌球果的发育过程中,未观察到独立的珠鳞发育.不同的种中,胚珠怕数量和发育顺序有所不同,但苞片的发育是相似的.传粉前,苞片的结构与叶相似.传粉后,由于剧烈的居间维管束发育.我们还对柏科植物雌球果的形态学特性及其可能的演化趋势进行了讨论.     

松杉类裸子植物的大孢子叶球理论评述

系统评述了迄今为止在裸子植物球果类种鳞形态学本质方面存在过的各种观点及研究进展。将 这些观点归纳为7大类：(1)Delpino和Penzig等主张种鳞为叶性器官的叶性说;(2)Sachs和Eichler认为种鳞相当于叶舌的叶舌说;(3)Bessey认为种鳞不过是胚珠合点端增生的结果的独特结构说;(4)Kubart等提出生胚珠构造的假种皮学说;(5)Arber的半枝学说;(6)Dupler的珠被说认为红豆杉类胚珠的肉质化部分为珠被性质的构造;(7)由Schleiden和Braun提出并由Florin等发展的枝性说则以充分的证据论证了种鳞的枝性本质。关于球果类裸子植物种鳞的性质和来源,本文作者赞同Florin枝性本质的论述;但对该构造现存式样的组成和演化过程,笔者认为该构造是在个体发育中不甚分化的专营生殖的枝性复合构造,由早期裸子植物次级生殖枝及其相关构造经过系统发育变态、融合演化形成。     

侧柏雌球果及其胚珠的发育

观察了侧柏（Ｐｌａｔｙｃｌａｄｕｓ ｏｒｉｅｎｔａｌｉｓ（Ｌ．）Ｆｒａｎｃｅ）胚珠的发育过程及后期球果苞片的结构变化。在北京,雌球果原基７月分化。通常一个球果有４对苞片,中部两对可育,靠球果顶端一对各产生一枚胚珠,其下一对各两枚。胚珠的发育顺序是向顶的,下部可育苞片腑部的两枚胚珠源于同一原基。胚珠原基分化成珠心和珠被,在发育过程中,珠被逐渐包围珠心,最后形成２烧瓶状的胚珠。１１月至次年１月,球果处     

马尾松雌球果的发生和早期发育研究

采用常规石蜡制片技术对马尾松雌球果的发生和早期发育进行了研究。结果表明：雌球果原基发生时间为10月中旬,不同的树龄和着生部位,其发生时间不同。雌球果原基与营养茎端在外部形态及内部细胞组织学分区结构有明显差异。营养茎端外形扁平,内部顶端分生组织结构有顶端原始细胞区、中央母细胞区、形成层状过渡区、周围分生组织区及肋状分生组织区5个明显的分区;而雌球果原基外形呈圆锥状,内部结构只有套层和髓区。12月初,最初的苞片原基在雌球果原基的鳞片的叶腋处产生,之后其由基部向顶部连续发生。翌年1月初,在苞片原基的叶腋处,珠鳞原基发生,发生方向亦为向顶发育。2月底,苞片体积不再发生变化,珠鳞膨大端的基部的近轴面分化出2个倒生胚珠。从雌球果原基发生到胚珠分化历时4个多月。亚热带的冬季气候对马尾松雌球果的生长发育没有明显的抑制作用。     

银杏雌性生殖器官的形态学本质及其系统学意义

关于古老裸子植物银杏Ginkgo biloba L.雌性生殖器官形态学本质的问题争论 很多,因此这一类群的分类位置尚存争议。本文进一步发展了Florin提出的“种鳞复合 体(Seed-scale complex)”概念,提出解释裸子植物生殖器官形态演化的“苞鳞一种鳞复 合体(Bract-scale and seed-scale complex)概念和演化理论,并试图用以解释银杏的生 殖器官形态学本质和裸子植物系统发育的有关问题。由银杏着生胚珠的结构(Ovuliferous structure)生于可育短枝上鳞状叶叶腋的事实,赞同该结构为次级轴性(枝性)性质,提 出该次级轴性结构及连同下部的鳞状叶相当于裸子植物大部分类群所具有的苞鳞—种 鳞复合体(Bract-scale and seed-scale complex),并保留了较多的原始古老性质,可能 代表着苞鳞—种鳞复合体较原始的形态式样。该结构的鳞状叶与可育短枝上正常的营 养叶均为不育苞片的性质,鳞状叶叶腋生出的长柄则是次级轴性结构的次生性伸长,长 柄上部二分叉的着生胚珠结构相当于该次级轴性器官仅存的极度退化的二枚可育叶性器 官,而整个可育短枝相当于一个分化程度很低的复合大孢子叶球。银杏生殖器官中的某 些性状与苏铁十分相似,但由于苞鳞一种鳞复合体的出现,标志着这一类群已远较苏铁特 化,并应与所有具苞鳞—种鳞复合体的类群在同一条演化线上,即苞鳞—种鳞复合体 演化线(Bract-scale and seed-scale complex evolutionary line,BS-line),而不具苞鳞 —种鳞复合体的苏铁类(Cycas)则构成裸子植物中的苏铁演化线(Cycas evolutionary line,C-line)。银杏大孢子叶球的式样,可能代表着裸子植物一种早期的原始大苞子叶 球式样,为古裸子植物不甚分化的可育枝条向高度压缩和变态的可育枝条(苞鳞一种鳞复合体及典型的球果)演化的一种中间过渡式样。     

从云杉属的畸形珠鳞看珠鳞的演化

在云杉属（Ｐｉｃｅａ）畸形现象调查中,发现畸形珠鳞也是常见的一种畸形现象。将畸形珠鳞进行归类、对比、并且做出一个排序：由典型的短枝式珠鳞逐步向正常的鱼鳞状珠鳞过渡。典型的短枝式珠鳞有一个较明显的轴,轴上着生若干枚鳞片,在鳞片的远轴面基部着生孢子囊（胚珠或花粉囊）;过渡形态轴的珠鳞轴逐渐缩短扁化（但并未完全消失）,上部的鳞片逐渐变小、数量减少,下部两枚侧生的鳞片逐渐发达,相背扭转方位约９０°,然后剩下的鳞片之间逐步愈合,最终转变为正常的鱼鳞状。作者认为这类珠鳞的畸变是返祖现象,可能在某些方面反映了珠鳞的历史演变过程。     

封面植物介绍——太白红杉

太白红杉 Larix chinensis Beiss. 亦称太白落叶松 ,为松科落叶松属 Larix Mill. 植物 ,为秦岭特有种之一 .乔木 ,高可达 2 0 m.树皮灰色或灰褐色 ,薄片状剥裂 .小枝下垂 ,一年生小枝常呈淡黄褐色 .叶倒披针状线形 ,扁平柔软 ,长 1 .5～ 2 .5 cm,在长枝上呈螺旋状散生 ,在短枝上簇生 ,叶表面中脉凸起 ,两侧各有 1～ 2条气孔带 .雌雄同株 ,球花单生于短枝顶端 ;雄球花球形至长圆形 ,黄色 ;雌球花近圆形或长圆形 ,每珠鳞具 2枚胚珠 ,苞鳞显著 ,淡紫红色或紫绿色 .球果卵状长圆形 ,长 3～ 4.5 cm,初红色 ,后渐变为紫蓝色或灰褐色 ,成熟后种…     

封面植物介绍——太白红杉

太白红杉 Larix chinensis Beiss. 亦称太白落叶松 ,为松科落叶松属 Larix Mill. 植物 ,为秦岭特有种之一 .乔木 ,高可达 2 0 m.树皮灰色或灰褐色 ,薄片状剥裂 .小枝下垂 ,一年生小枝常呈淡黄褐色 .叶倒披针状线形 ,扁平柔软 ,长 1 .5～ 2 .5 cm,在长枝上呈螺旋状散生 ,在短枝上簇生 ,叶表面中脉凸起 ,两侧各有 1～ 2条气孔带 .雌雄同株 ,球花单生于短枝顶端 ;雄球花球形至长圆形 ,黄色 ;雌球花近圆形或长圆形 ,每珠鳞具 2枚胚珠 ,苞鳞显著 ,淡紫红色或紫绿色 .球果卵状长圆形 ,长 3～ 4.5 cm,初红色 ,后渐变为紫蓝色或灰褐色 ,成熟后种…     

Ontogeny and Metamorphic Patterns of Female Reproductive Organs of Ephedra sinica Stapf (Ephedraceae)

Ontogeny and metamorphic development of female reproductive organs in Ephedra sinica Stapf were surveyed. At the end of April, female cones began to initialize from the vegetative buds. Pollination occurred in mid-May and seeds matured at the beginning of July. The ontogenetic pattern of female reproductive organs of E.sinica is basically similar to that of E.　distachya L. The foliar nature of the outer envelope of the ovule in Ephedra is corroborated. Reduction of ovule number as a tendency of speciation in the genus is substantiated by the occurrence of tri-ovulate cones coupled with comprehensive characters of the genus. The metamorphic patterns as well as the leaf nature of the outer envelope indicate that female cones of Ephedra are compound while the female reproductive units of the ovulate cone have been reduced from secondary reproductive shoots similar to those of cordaites by phylogenetic transformation, fusion and reduction. Each fertile bract together with its axillary female reproductive unit composed the Bract Scale and Seed Scale Complex of Ephedra .     

Ontogeny of triovulate cones of Ephedra intermedia and origin of the outer envelope ofovules of Ephedraceae

The origin of the two envelopes of ovules of extant species of Ephedraceae has been discussed previously in different ways based only on investigation of bimerous female cones. To generalize the characteristics of the two envelopes, ontogenetic and metamorphic patterns of the two envelopes of ovules of trimerous female cones of Ephedra intermedia Schrenk et C. A. Meyer were studied using scanning electron microscopy. The outer envelope of the ovules of triovulate female cones of Ephedra intermedia is initiated as two adaxial lateral protuberances. Thus, the outer envelope of extant Ephedraceae may have been derived from three foliar components of the second proximal whorl of the secondary reproductive shoot of the ancestral compound female cone. Two adaxial lateral foliar components likely gave rise to the outer envelope by fusion, while the third abaxial foliar component was reduced. These new findings suggest that the Gnetales derived their female cones from trimerous multiaxial female reproductive organs of their progenitors. Some evolutionary aspects of female cones of Ephedra are also discussed.     

Morphology and affinities of an Early Cretaceous Ephedra (Ephedraceae) from China

Detailed investigations on Lower Cretaceous Ephedra L. fossils (Gnetopsida) reveal morphological characters similar to those of extant Ephedra rhytidosperma Pachomova, including articulate branches with many fine longitudinal striations, a dichasial branching pattern, uni- or bi-ovulate cones with paired bracts, cones terminal on branchlets, and seeds with a short, straight micropylar tubes, covered by numerous regular and prominent transverse laminar protuberances. Fossils are similar to extant E. rhytidosperma reproductive organs but differ in some vegetative structures and are described and discussed here as Ephedra archaeorhytidosperma Y. Yang et al. Because E. rhytidosperma is currently considered one of the most specialized members in Ephedra L. section Pseudobaccatae Stapf, the occurrence of E. archaeorhytidosperma in the Yixian Formation suggests that Ephedra L. was perhaps a more diverse genus in the Lower Cretaceous. Perhaps the evolution and diversity of Ephedra L. was already in place by the Lower Cretaceous and certainly before the end of the Mesozoic.     

麻黄属种子表面纹饰的多样性及其系统学意义

麻黄属(Ephedra)的分子系统学研究结果明显不同于基于雌球果成熟苞片质地的传统分类,指出麻黄属的形态特征和分子特征存在冲突。对麻黄属53种植物的种子表面微形态特征进行了研究,结果表明麻黄属的种子表面纹饰包括3种主要类型,即横向片层型(T型)、乳突型(P型)和扁平型(S型)。麻黄属的种子表面纹饰特征既与宏观形态特征不相关,也与分子系统学的研究结果不相关,表明该属微形态特征、宏观形态特征和分子特征之间的更多冲突。在欧洲和东亚发现的与买麻藤类有关的几个大化石带有T型种子表面纹饰,暗示具有T型纹饰的麻黄类植物在早白垩纪就十分多样。     

The role of chemical fingerprinting: application to Ephedra

Ephedra sinica, known as Ma Huang, is one of the oldest medicinal herbs in Traditional Chinese Medicine (TCM). Preparations, namely teas, of E. sinica have been used for over 5000 years as a stimulant and as an antiasthmatic. In the West, extracts of E. sinica, E. intermedia or E. equisetina are most commonly used in dietary supplements as a stimulant and to promote weight loss. More than 50 species of Ephedra are native to both hemispheres, but the detection of ephedrine alkaloids has been limited to species in Eurasia. Currently, methods exist to quantitate the ephedrine alkaloids in extracts of plant material or dietary supplements, but the methods are not able to verify the extract is of an Ephedra species. Reverse phase high performance liquid chromatography with photodiode array detection was applied for the chemical fingerprinting of the Ephedra species. Two regions of comparison were determined in the chromatograms at 320 nm. The series of peaks between 52 and 64 min confirms an Ephedra species is being analyzed. The aforementioned peaks also could distinguish between Ephedra species from Eurasia, North America and South America. Peaks at ca. 57 and 59 min were isolated and determined to be two new compounds, 4-(2-eicosyloxycarbonyl-vinyl)-benzoic acid and 4-(2-docosyloxycarbonyl-vinyl)-benzoic acid respectively. Authentication of ground plant material as Ephedra can be achieved by this chemical fingerprinting method.     

Morphology and the evolution of cycadeoidales

OneCycadeoidea stem one cycadeoidalean gynoecium and a bisporangiate cone attached to a slender cycadeoidalean trunkCycadeoidella japonica Ogura from the Cretaceous of Japan shows well-preserved internal structure that provides evidence for a better understanding of the morphological architecture of the cycadeoidalean plant. Structural details of the cone were confirmed. The ovule has an intergument enclosing a free nucellus and a thin outer envelope. Both reproductive and vegetative structures support the medullosan affinity of Cycadeoidales. The cone is interpreted as a compressed fertile shoot. Axillary cones characterizing some Cretaceous genera such asCycadeoidea andMonanthesia consist of a lateral shoot subtended by a frond that is the first leaf of the cone shoot itself. The origin of axillary buds in the Cycadeoidales is discussed. Heterochrony may have mediated the morphological changes that resulted in the establishment of the Cycadeoidales.     

Nageiaceae—A New Gymnosperm Family

A new monotypic gymnosperm family, Nageiaceae D. Z. Fu, is separated from Podocarpaceae. It is characterized by having multinerved leaves without costae, and primitive shoot-like female reproductive organs (female strobili). The new family contains a single genus consisting of 2 sections, 5 species and is distributed along the western coast of the Pacific, from low coastal mountains of eastern and southern Asia to the Phillipines and Papua New Guinea. The first species in the Nageiaceae was described as an angiosperm, Myrica nagi Thunb. (1784), but it was soon recognized to be a gymnosperm belonging to a new genus, and was renamed as Nageia japonica Gaert. (1788). The generic name, Nageia, however, has seldom been used, and the members of Nageia have generally been treated as an isolated section of Podocarpus in the Podocarpaceae. When revising the Podocarpaceae, De Laubenfels (1969) established a new genus Decussocarpus based on Nageia, but several years later (1987) he revived the old generic name, Nageia. Page ( 1988,1990)considered Nageia to be a valid generic name and redefined it as a natural genus. The distinctive,broadly lanceolate, multinerved leaves (without costae) of Nageia are rather unusual in gymnosperms,only being similar to those of Agathis in the Araucariaceae, their leaves are also similar to each other in anatomy. For example, there are many single vascular bundles arranged parallelly, between which occur sclerenchyma cells in the mesophyll. Apparently,leaves in Nageia are rather similar both externally and internally to paleogymnosperm cordaitean leaves, and sclerenchyma cells found in Nageia might be the remains of straps between veins in cordaitean leaves. In addition to leaf characters, the large and nearly round pith of the young shoot in Nageia appears to be a reminiscent of the large pith in cordaitean stem. The female reproductive organs (female strobili ) in Nageia are shoot-like. The female strobilus has a sterile terminal bud, and several opposite or subopposite sterile scaly bracts on its axis; two opposite megasporophylls are found near the axis apex and both have an anatropous ovule which is almost entirely covered by the megasporophyll; a bract is partly adnate to the lower back of the megasporophyll;mature arillate seeds are 1-2 or occasionally 3 in number; the axis becomes woody when the seeds mature, but in some species (N. wallichiana) the upper part of the axis becomes fleshy (in the shape of a receptacle), in which no distinct boundary was found between the fleshy receptacle and the woody part, and both have the same scaly bracts or traces. Many characters in Nageia are distinctly different from those in Podocarpus. Leaves in the Podocarpaceae have distinct midribs; in Podocarpus, the reproductive organ, which was generally thought to be similar to that in Nageia, has no terminal bud, and its bract is entirely free from the lower back of the megasporophyll, the fleshy receptacle is derived from both the axis and the sterile bracts (except the lowest two), and the female strobilus at the seed stage has a secondary stalk. The multinerved leaf in Nageia can rarely be found in most of the living gymnosperms except in some rather isolated groups, such as Araucariaceae, Ephedraceae,Ginkgoaceae and Welwitschiaceae. Paleobotanical evidence shows that multinerved leaves have been found in all of the geological ages from the Paleozoic to the present, and such a shoot-like female reproductive organ as in Nageia was found in some paleogymnosperms. It is very difficult to determine the systematic positions of these fossil plants because of lacks adequate material of reproductive organs or even lack of complete vegetative organs. The vascular system and leaf characters of gymnosperms are considered to be very conservative, and the fact that the common leaf shape and venation exist in both fossil and living gymnosperms could imply that there exists a multinerved-leaved evolutionary line ( M-line ) in gymnosperms, which could be traced back to the paleogymnosperm cordaitean plants or even older ones with multinerved leaves. The different types of the female strobili (female reproductive organs) of living gymnosperms, regardless of having one or only several seeds without a typical cone or many seeds with a cone, might have been derived from shoot-like or spikelike female reproductive organs possessed by their common ancestor.The fossil eviden ce shows that the typical cone similar to those of living gymnosperms first appeared in the Jurassic, much later than the single-seeded fossil plant without cones. The seed fossil appeared in the late Devonian Period. It is very difficult to infer the relationships among living gymnosperms, which are hardly derived from one another. But an analysis of the strobili, including the axis structure and position, number, morphology and degree of adnation of the phyllomes on them, would be helpful to the study of their phylogeny. It is evident, therefore, that the gymnosperms with leaves having a midrib might also have a rather long evolutionary course,but no transition between the midrib and multinerved patterns of leaf venation has so far been found in both living and fossil plants. Finally, it is noteworthy that the Nageiaceae are distributed along the western coast of the Pacific, where many primitive representatives, both in gymnosperms and angiosperms, still survive. This would be advantageous to the consideration of Nageiaceae as a primitive representative, or a descendant of fhe paleogymnos-perm cordaitean plants.     

A New Morphological Interpretation of the Female Reproductive Organs in Ginkgo biloba L., with a Phylogenetic Consideration on Gymnosperms

The ovuliferous structure of Ginkgo biloba L. has been variously interpreted morphologically. As a result the systematic position and the relationship with other gymnosperms of this ancestral gymnosperm have long been under dispute. In the present paper, a brief survey of the main views as to the nature of the ovuliferous structure is givcn. Based on morphological and teratological data previously reported, a new interpretation is proposed. The essential points are summarized as follows: 1. In morphological essence, a fertile dwarf shoot with some ovuliferous structures in Ginkgo biloba L. might as a whole be nothing but a megasporophyll strobilus (female cone), which is shared actually by all the conifers in the gymnosperms. The fertile dwarf shoot has appearance extremely similar to that of the vegetative dwarf shoot, suggesting that in Ginkgo biloba L. the vegetative organs and the reproductive organs have not been yet well differentiated, and thus its megasporophyll strobilus might represent one of the most primitive compound strobilus types. 2. In Ginkgo biloba, the ovuliferous structure borne in the axil of a scale leaf (sometimes a normal leaf) on the dwarf shoot, together with the scale leaf itself, might be the homogenous organ corresponding to the bract-scale and seed-scale complex of the compound female strobilus of the typical conifers. The complex is a relatively isolated reproductive unit on the strobilus. The normal leaves and the scale leaves on the dwarf shoot might be equivalent to the bract-scales in the typical cones, though the normal leaves still retain the vegetative nature as the foliage leaves on the vegetative shoot. The stalk hearing ovules at its top might be equivalent to a seed-scale of the typical cones. 3. The megasporophyll strobilus in Ginkgo biloba, namely a whole fertile dwarf shoot as mentioned above, seems to show much more primitive characteristics than those of typical conifers. In this plant it is very difficult to distinguish the fertile dwarf shoot from the common vegetative dwarf shoot before reproduction time. Moreover, its megasporophyll strobilus often exhibits more atavistic abnormalities than those of other conifers. All the evidence indicates that the primitive ancestor of conifers might have had the fertile organs which might be of basically identical morphology as vegetative shoots, except that in the fertile organs there might exist numerous fertile leaves bearing one or many ovules. 4. The longer stalk of the ovuliferous structure in Ginkgo biloba might have come from mainly a secondary elongation growth of the seed scale, and only a little part of it might be the remains of the original shoot. The fork structure bearing ovules at the top of the stalk might be the rudementary part of the petioles of the only two extremely reduced megasporophylls. The collar around the base of the ovule might be a secondary protective structure. 5. A correct morphological interpretation of the female strobilus in Ginkgo biloba is doubtless of important significance for our better understanding of the evolution of the female reproductive organs in conifers. According to our interpretation mentioned above, together with the concept of the bractscale and seed-scale complex proposed in the present paper, which is mainly based on the concept of the seed-scale complex propose by Florin, here we put forward an evolutionary theory of the bract-scale and seed-scale complex. According to this theory, the female reproductive organs of the ancestral conifers should be very similar, as mentioned above, to the sterile foliage shoot except that the former might have some fertile leaves which could produce ovules at reproduction time. This ancestral female reproductive organ type might have had evolved towards two directions and thus formed two main evolutionary lines. One is represented by the genus Cycas and we may call it the Cycas Evolutionary Line (C-line), in which the megasporophyll strobilus is monopodial, with the fertile leaves and sterile bracts occurring directly on the main axis. The Cycadaceae is the only living gymnosperm member along this evolutionary line. The second line is represented by all the conifers including Ginkgo, which all have the structure of the bract-scale and seed-scale complex, and thus we called it the "Bract-scale and Seed-scale Complex Evolutionary Line" (BS-line). The members along this line have multipodial female strobilus, i.e. compound strobilus. On the main axis occur some sterile vegetative bracts. In the axils of some or most of the bracts occur the seed-scales. The seed-scales are actually the remains of the extremely, or smetimes completely reduced fertile shoots. Each part of the bractt-scale and seed-scale complex and the main axis of the strobilus could have undergone independent or correlated changes, and thus have had formed various types of strobilus which are found in the living conifers. 6. Our theory on the evolution of the bract-scale and seed-scale complex seems to support the division of all the gymnosperms into two major groups as proposed by Chamberlain, and is also in favour of the placement of Ginkgo biloba into the conifers as the most primitive member along BS-line. 7. Based on their similar morphological characters, it can be considered that Ginkgo biloba might have close relationships with the Nageiaceae, Ephedraceae, Welwitschiaceae and Araucariaceae. All these groups have multinerved leaves without costa. These living gymnosperms might have a more direct relationship withthe ancestral cordaites.     

Former diversity of Ephedra (Gnetales): evidence from Early Cretaceous seeds from Portugal and North America

BACKGROUND AND AIMS: The extant species of the seed plant group Gnetales (Ephedra, Gnetum and Welwitschia) have been considered a remnant of a much greater, now extinct, diversity due to the pronounced differences in form and ecology among the genera. Until recently, this hypothesis has not been supported by evidence from the fossil record. This paper adds to the expanding information on Gnetales from the Early Cretaceous and describes coalified seeds from Barremian-Albian localities in Portugal and USA. METHODS: The fossils were extracted from sediment samples by sieving in water. Adhering mineral matrix was removed by chemical treatment. Seeds were investigated using light and scanning electron microscopy. Morphology and anatomy of the seeds were documented and compared with those of extant species. KEY RESULTS: The fossils share characters with extant Ephedra, for example papillae on the inner surface of the seed envelope and in situ polyplicate pollen grains that shed the exine during germination. They differ from extant Ephedra seeds in morphological and anatomical details as well as in their smaller size. Two new species of Ephedra are described together with one species assigned to a new genus of Gnetales. Other Ephedra-like seeds, for which pollen and critical morphological details are currently unknown, are also present in the samples. CONCLUSIONS: These Cretaceous seeds document that key reproductive characters and pollen germination processes have remained unchanged within Ephedra for about 120 million years or more. There is sufficient variety in details of morphology to suggest that a diversity of Ephedra and Ephedra-like species were present in the Early Cretaceous flora. Their presence in Portugal and eastern North America indicates that they were widespread on the Laurasian continent. The fossil seeds are similar to seeds of Erdtmanithecales and this supports the previously suggested relationship between Erdtmanithecales and Gnetales.     

Morphogenesis of the reproductive shoots of Welwitschia mirabilis and Ephedra distachya (Gnetales), and its evolutionary implications

For decades, Gnetales appeared to be closely related to angiosperms, the two groups together forming the anthophyte clade. At present, molecular studies negate such a relationship and give strong support for a systematic position of Gnetales within or near conifers. However, previous interpretations of the male sporangiophores of Gnetales as pinnate with terminal synangia conflict with a close relationship between Gnetales and conifers. Therefore, we investigated the morphogenesis of the male reproductive structures of Welwitschia mirabilis and Ephedra distachya by SEM and light microscopy. The occurrence of reduced apices to both halves of the antherophores of W. mirabilis gives strong support for the assumption that the male ‘flowers’ of W. mirabilis represent reduced compound cones. We assume that each half of the antherophore represents a lateral male cone that has lost its subtending bract. Although both halves of the antherophores of Ephedra distachya lack apical meristems, the histological pattern of the developing antherophores supports interpreting them as reduced lateral male cones as well. Therefore, the male sporangiophores of Gnetales represent simple organs with terminal synangia. Although extant conifers do not exhibit terminal synangia, similar sporangiophores are reported for some Cordaitales, the hypothetical sister group of conifers. Moreover, several Paleozoic conifers exhibit male cones with terminal sporangia or synangia. Therefore, we propose that conifers, Cordaitales and Gnetales originated from a common ancestor that displayed simple sporangiophores with a terminal cluster of sporangia.