茂兰国家级自然保护区石松类和蕨类植物区系特征

本研究通过野外考查、标本采集和分类鉴定，以及文献资料整理，对贵州茂兰国家级自然保护区石松类和蕨类植物区系组成和分布区类型进行统计分析。结果显示：(1)茂兰国家级自然保护区共有石松类和蕨类植物32科76属237种，优势科为鳞毛蕨科(Dryopteridaceae)、蹄盖蕨科(Athyriaceae)、凤尾蕨科(Pteridaceae)、水龙骨科(Polypodiaceae)；优势属为双盖蕨属(Diplazium)、卷柏属(Selaginella)、凤尾蕨属(Pteris)。(2)分布区类型以东亚广布和热带亚洲分布为主，生态类型以阴生植物和土生植物居多。(3)与其他地区的区系比较显示，茂兰与广西花坪物种相似性系数最高，依次是贵州梵净山、四川峨眉山和云南马关县。     

Evolution of the eastern Asian-North American biogeographic disjunctions in ferns and lycophytes

This paper reviews 31 groups in ferns and lycophytes hypothesized to show eastern Asian–North American disjunctions. Fourteen lineages have been supported by recent phylogenetic evidence: Lycopodium nikoense and Lycopodium sitchense; Isoëtes asiatica and the clade of the North American species complex closely allied to I. maritima; Osmundastrum cinnamomeum; Osmunda claytoniana; the Adiantum pedatum complex; the Cryptogramma acrostichoides complex; Diplaziopsidaceae; Cystopteris chinensis and the Cystopteris bulbifera clade; Asplenium rhizophyllum and Asplenium ruprechtii; diploid Phegopteris; Onoclea sensibilis; the Polypodium appalachianum clade; and the Polypodium glycyrrhiza clade. Phylogenetic and/or cytological evidence did not support the biogeographic disjunctions in six cases: (1) Isoëtes asiatica and I. truncata; (2) Botrychium ternatum; (3) Thelypteris beddomei and T. nipponica—Thelypteris noveboracensis and T. nevadensis; (4) Thelypteris glanduligera and Thelypteris. japonica—T. simulata; (5) Woodwardia japonica and W. virginica; and (6) Woodwardia orientalis and Woodwardia fimbriata. Both vicariance and dispersal have been suggested to be the mechanisms for the formation of the disjunct pattern; and the Beringian region has been an active pathway for the migration of ferns and lycophytes between Asia and North America. Disjunctions of ferns and lycophytes reviewed here have been dated in the Tertiary, and are similar to the ages of eastern Asian–North American disjunctions in seed plants, supporting the close biogeographic co‐diversification of ferns and seed plants. Future studies are needed to estimate divergence times and reconstruct biogeographic events in a broad phylogenetic framework, and to test the morphological stasis hypothesis in disjunct ferns and lycophytes.     

Lycophytes from the Middle Triassic (Anisian) locality Kühwiesenkopf (Monte Prà della Vacca) in the Dolomites (northern Italy)

Abstract: Generally, lycophytes are not well represented in Anisian floras. Nonetheless, four different genera, each with one species, have been distinguished in the Anisian flora from Kühwiesenkopf (Monte Prà della Vacca), Italy. They were well preserved and yielded leaf cuticles in all four taxa and in situ spores in two of them. Lycopia dezanchei gen. et sp. nov. is the most common form; it is characterized by a dichotomising creeping prostrate rhizome (representing the primary shoot axis) from which arise aerial axes, apically bifurcated and covered with bundles of long leaves. This taxon cannot be attributed to any of the lycophyte orders, but most resembles Lycopodiales. Two representatives of the Isoetales have been found, of which Isoetites brandneri sp. nov. is abundant. This species is characterized by a short stem, which is unusual for this genus. In situ immature microspores and megaspores were found in some specimens. Lepacyclotes bechstaedtii sp. nov. has a quadrilobe corm with fertile scales inserted in whorls or slightly helicoidally. Short sterile leaves arise distally from the corm in some specimens. Selaginellales are very rare in the flora; Selaginellites leonardii sp. nov. is represented by a strobilus that yielded both micro‐ and megaspores. Some sterile fragments have been found associated, but never in organic connection.     

Why did heterospory evolve?

The primitive land plant life cycle featured the production of spores of unimodal size, a condition called homospory. The evolution of bimodal size distributions with small male spores and large female spores, known as heterospory, was an innovation that occurred repeatedly in the history of land plants. The importance of desiccation‐resistant spores for colonization of the land is well known, but the adaptive value of heterospory has never been well established. It was an addition to a sexual life cycle that already involved male and female gametes. Its role as a precursor to the evolution of seeds has received much attention, but this is an evolutionary consequence of heterospory that cannot explain the transition from homospory to heterospory (and the lack of evolutionary reversal from heterospory to homospory). Enforced outcrossing of gametophytes has often been mentioned in connection to heterospory, but we review the shortcomings of this argument as an explanation of the selective advantage of heterospory. Few alternative arguments concerning the selective forces favouring heterospory have been proposed, a paucity of attention that is surprising given the importance of this innovation in land plant evolution. In this review we highlight two ideas that may lead us to a better understanding of why heterospory evolved. First, models of optimal resource allocation – an approach that has been used for decades in evolutionary ecology to help understand parental investment and other life‐history patterns – suggest that an evolutionary increase in spore size could reach a threshold at which small spores yielding small, sperm‐producing gametophytes would return greater fitness per unit of resource investment than would large spores and bisexual gametophytes. With the advent of such microspores, megaspores would evolve under frequency‐dependent selection. This argument can account for the appearance of heterospory in the Devonian, when increasingly tall and complex vegetative communities presented competitive conditions that made large spore size advantageous. Second, heterospory is analogous in many ways to anisogamy. Indeed, heterospory is a kind of re‐invention of anisogamy within the context of a sporophyte‐dominant land plant life cycle. The evolution of anisogamy has been the subject of important theoretical and empirical investigation. Recent work in this area suggests that mate‐encounter dynamics set up selective forces that can drive the evolution of anisogamy. We suggest that similar dispersal and mating dynamics could have underlain spore size differentiation. The two approaches offer predictions that are consistent with currently available data but could be tested far more thoroughly. We hope to re‐establish attention on this neglected aspect of plant evolutionary biology and suggest some paths for empirical investigation.     

Stomatal innovation and the rise of seed plants

Stomatal valves on the leaves of vascular plants not only prevent desiccation but also dynamically regulate water loss to maintain efficient daytime water use. This latter process involves sophisticated active control of stomatal aperture that may be absent from early-branching plant clades. To test this hypothesis, we compare the stomatal response to light intensity in 13 species of ferns and lycophytes with a diverse sample of seed plants to determine whether the capacity to optimise water use is an ancestral or derived feature of stomatal physiology. We found that in seed plants, the ratio of photosynthesis to water use remained high and constant at different light intensities, but fern and lycophyte stomata were incapable of sustaining homeostatic water use efficiency. We conclude that efficient water use in early seed plants provided them with a competitive advantage that contributed to the decline of fern and lycophyte dominated-ecosystems in the late Paleozoic.     

辽宁省高等植物多样性编目

为给辽宁植物多样性保护提供本底数据, 更准确地评价植物濒危及保护等级, 以及更好地给农学、林学、生态学、系统学等研究提供可靠的基础性数据, 本文首次将几代人在辽宁境内发现的高等植物进行汇总、编目, 包括苔藓植物和维管束植物。苔藓植物根据正式发表的文献和中国科学院沈阳应用生态研究所东北生物标本馆(IFP)馆藏采自辽宁省全域的苔藓植物标本编制。维管束植物主要根据5个方面的信息整理: 作者多年野外调查的一手资料、文献、同行交流、公众提供信息、标本等。编目内容包括辽宁省高等植物总名录、辽宁省栽培植物名录、辽宁省外来植物名录、辽宁省仅有栽培植物的科属目录、辽宁省仅有外来植物的科属目录; 编目中涉及的物种均有科属地位、中文名、拉丁名、生境、产地(县级), 总名录中附有每个物种的信息凭证, 包括标本信息、文献信息或照片依据。本编目共计收录植物254科1,176属3,241种73亚种447变种119变型。其中, 苔藓植物为辽宁地区首次披露名录信息, 计83科205属491种4亚种7变种(含藓类52科156属373种1亚种5变种, 苔类29科46属114种3亚种2变种, 角苔类2科3属4种)。维管束植物虽非首次披露名录信息, 但与以往发布的维管束植物名录相比, 本次收录的种类数最多, 计171科971属2,750种69亚种440变种119变型(含石松类植物2科3属13种, 蕨类植物17科39属83种2亚种7变种, 裸子植物5科19属47种1亚种13变种2变型, 被子植物147科910属2,607种66亚种420变种117变型)。经统计, 辽宁省所有苔藓植物均为本土野生植物。维管束植物中, 野生植物计149科741属2,077种62亚种378变种93变型(含石松类植物2科3属13种, 蕨类植物17科39属83种2亚种7变种, 裸子植物4科8属12种6变种1变型, 被子植物145科133属1,969种60亚种365变种92变型), 栽培植物计104科390属673种7亚种62变种26变型(含裸子植物5科17属35种1亚种7变种1变型, 被子植物99科373属638种6亚种55变种25变型); 本土植物计158科796属2,235种65亚种421变种116变型(含石松类植物2科3属13种, 蕨类植物17科39属83种2亚种7变种, 裸子植物5科14属29种12变种2变型, 被子植物134科740属2,110种63亚种402变种114变型), 外来植物计88科316属515种4亚种19变种3变型(含裸子植物3科10属18种1亚种1变种, 被子植物85科306属497种3亚种18变种3变型)。本编目仅是辽宁省高等植物的阶段性汇总, 尚需要根据野外工作的拓展和研究工作的深入不断完善。     

Molecular evidence for long-distance dispersal in the New Zealand pteridophyte flora

Aim To examine the relative importance of long‐distance dispersal in shaping the New Zealand pteridophyte (ferns and lycophytes) flora and its relationships with other floras, with the null hypothesis that the extant New Zealand pteridophyte flora has been isolated since New Zealand’s separation from Gondwana. Location New Zealand. Methods rbcL DNA sequences were assembled for 31 New Zealand pteridophyte genera, with each genus represented by one New Zealand species and the most closely related non‐New Zealand species for which data were available. Maximum‐likelihood, maximum‐parsimony, and Bayesian analysis phylograms were constructed and used as input for r 8s molecular dating, along with 23 fossil calibrations. Divergence estimates less than conservatively recent ages for New Zealand’s geological isolation, namely H_o > 30 Ma for pairs involving New Caledonian and Norfolk Island species and H_o > 55 Ma for all others, were taken as rejection of the null hypothesis. Results The null hypothesis was rejected for all pairs except, under some parameter conditions, for those involving the New Zealand species Cardiomanes reniforme, Lindsaea trichomanoides, Loxsoma cunninghamii, Lygodium articulatum, Marattia salicina, and Pteris comans. However, the Lindsaea and Pteris results probably reflect the absence in the analyses of closely related non‐New Zealand samples, while the Marattia divergence was highly contingent on which fossil calibrations were used. Main conclusions Rejection of the null hypothesis for the majority of pairs implies that the extant New Zealand lineage has undergone long‐distance dispersal either into or out of New Zealand. The notion of a long isolation since geological separation can, therefore, be dismissed for much of New Zealand’s pteridophyte flora. The analyses do not identify the direction of the long‐distance dispersal, and these New Zealand lineages could have had vicariant origins with subsequent long‐distance emigration. However, the alternative that many extant New Zealand pteridophyte lineages only arrived by long‐distance immigration after geological isolation seems likely.     

石松类和蕨类植物质体基因组结构演化研究进展

近年来, 随着测序技术的发展, 石松类和蕨类植物的核基因组、质体基因组以及线粒体基因组研究发展迅速, 质体基因组研究工作更是呈爆发式增长。截至2019年3月1日, GenBank公布的石松类和蕨类植物的175个质体基因组中, 约3/4为最近两年新增。研究内容从早期对个别质体基因组结构和序列特征的简单报道, 逐渐发展到综合性的比较基因组学和系统发育基因组学研究。目前已发表的质体基因组覆盖了石松类和蕨类植物的所有目和大部分科, 这两大类群的质体基因组结构变异和系统发育的基本框架已逐渐清晰。这些研究为我们理解维管植物的早期演化提供了重要参考。本文对石松类和蕨类植物的质体基因组结构特征进行了系统梳理, 发现其结构变异主要包括大片段倒位、IR区边界变动、基因或内含子丢失等, 其中一些结构变异可作为较高分类阶元的共衍征。RNA编辑和长片段非编码序列插入普遍存在于石松类和蕨类植物的质体基因组中, 但其起源、演化机制和功能等仍不清楚。我们对质体基因组的应用、系统发育研究中质体和核基因组的优劣性, 以及系统发育基因组学的前景进行了评述。     

中国薄叶卷柏复合群的物种划分

卷柏属(Selaginella)是石松类植物中最大的属, 也是分类难度较大的类群之一。该属的物种划分主要基于形态特征, 但许多近缘种在形态上很难区分。近年来, 已有大量分子证据被用于各植物类群的分类学研究, 但目前未发现一套适合卷柏属物种鉴定的分子标记。薄叶卷柏复合群(S. delicatula group)是卷柏属下鉴定较为困难的类群, 包括了薄叶卷柏(S. delicatula)、黑顶卷柏(S. picta)和瓦氏卷柏(S. wallichii) 3个物种, 主要分布于亚洲的热带和亚热带地区。为了探讨薄叶卷柏复合群内物种的亲缘关系和评估不同分子标记在卷柏属分类学研究中的应用价值, 本研究对该复合群物种进行广泛取样, 共收集到73个个体, 并选取3个叶绿体基因(rbcL, psbA和atpI)和2个核基因(26S nrDNA和pgiC)片段进行系统树的构建及叶绿体单倍型分析。研究发现, 基于叶绿体和核基因构建的系统发生关系存在冲突: 叶绿体基因树上薄叶卷柏个体分为两个分支(A和B), 薄叶卷柏B分支与薄叶卷柏A-S. picta分支呈姐妹关系, 并且rbcL单倍型分析结果也表明薄叶卷柏A和B两个分支存在明显分化; 而核基因结果则支持该复合群3个物种各自的单系性, 其中, S. delicatula分支与S. picta分支为姐妹群, S. wallichii与S. delicatula-S. picta分支为姐妹关系。在对复合群分布区大量标本的观察以及野外群体调查的基础上, 评估了植株茎和枝的分枝方式、孢子叶、营养叶(侧叶、中叶和腋叶)和孢子表面纹饰等形态性状的分类学价值。结果表明, 薄叶卷柏A和B分支的样本仅在植株分枝方式和大孢子表面纹饰上存在差异, 但无法依靠小孢子表面纹饰、孢子叶穗和营养叶形态等特征进行区分。基于现有证据, 薄叶卷柏复合群至少可划分为薄叶卷柏、黑顶卷柏和瓦氏卷柏3种, 但彻底澄清该复合群的物种划分还需要获取模式标本产地的材料和细胞学证据。最后, 建议在未来卷柏属的分类学研究集中于该属分类复杂的复合群, 结合使用形态学、细胞学、分子生物学(同时使用核基因和叶绿体分子标记)及地理分布等整合证据来进行物种划分。