Palaeoecology and palaeophytogeography of the rhynie chert plants: evidence from integrated analysis of in situ and dispersed spores

The Rhynie cherts yield exceptionally preserved early land plants, and provide a unique insight into the nature of Lower Devonian vegetation. Hitherto they have been poorly age constrained, and the palaeoecology and palaeophytogeography of the flora are poorly understood. Well-preserved dispersed-spore assemblages have been recovered from a number of borehole cores through the stratigraphical sequence of the Rhynie outlier. They are all similar and belong with the polygonalis-emsiensis (PE) spore zone, indicating an Early Devonian age (Early (but not earliest) Pragian to earliest Emsian). Comparisons with PE spore-zone assemblages from elsewhere suggest that the flora of the Rhynie drainage basin was slightly impoverished, with certain plant taxa that occurred at other locations not represented. This probably reflects differences between the flora of an inland intermontaine basin (Rhynie) and that of the lowland flood-plains. In situ spores have been characterized for all seven Rhynie chert plants. Analysis of the distribution of these spore types in the Rhynie sequence, in addition to those of coeval deposits from elsewhere, enables interpretation of the palaeoecology and palaeophytogeography of the Rhynie chert plants. It is concluded that at least some of the plants were not highly specialized or adapted to the peculiar hot-springs environment in which they are preserved. Rather, they were components of a diverse and widespread flora, but were the only elements able to tolerate the inhospitable hot-springs environment (i.e. they were preadapted).     

Wetland megabias: ecological and ecophysiological filtering dominates the fossil record of hot spring floras

Siliceous hot spring deposits form at Earth's surface above terrestrial hydrothermal systems, which create low‐sulphidation epithermal mineral deposits deeper in the crust. Eruption of hot spring waters and precipitation of opal‐A create sinter apron complexes and areas of geothermally influenced wetland. These provide habitat for higher plants that may be preserved in situ, by encrustation of their surfaces and permineralization of tissues, creating T⁰ plant assemblages. In this study, we review the fossil record of hot spring floras from subfossil examples forming in active hot spring areas, via fossil examples from the Cenozoic, Mesozoic and Palaeozoic to the oldest known hot spring flora, the Lower Devonian Rhynie chert. We demonstrate that the well‐known megabias towards wetland plant preservation extends to hot spring floras. We highlight that the record of hot spring floras is dominated by plants preserved in situ by permineralization on geothermally influenced wetlands. Angiosperms (members of the Cyperaceae and Restionaceae) dominate Cenozoic floras. Equisetum and gleicheniaceous ferns colonized Mesozoic (Jurassic) geothermal wetlands and sphenophytes and herbaceous lycophytes late Palaeozoic examples. Evidence of the partitioning of wetland hydrophytic and dryland mesophytic communities, a feature of active geothermal areas, is provided by well‐preserved and well‐exposed fossil sinter apron complexes, which record flooding of dryland environments by thermal waters and decline of local forest ecosystems. Such observations from the fossil record back‐up hypotheses based on active hot springs and vegetation that suggest removal of taphonomic filtering in hot spring environments is accompanied by an increase in ecological and ecophysiological filtering. To this end we also demonstrate that in the hot spring environment, the wetland bias extends beyond broad ecology. We show that ecosystems preserved from the Cenozoic to the Mesozoic provide clear evidence that the dominant plants preserved in situ by hot spring activity are also halophytic, tolerant of high pH and high concentrations of heavy metals. By extension, we hypothesize that this is also the case in Palaeozoic hot spring settings and extended to the early land plant flora of the Rhynie chert.     

Morphologically complex plant macrofossils from the Late Silurian of Arctic Canada

In addition to vegetative remains, fertile remains from ten plants, representing seven distinct taxa whose size and complexity are much greater than most contemporaneous fossils, are reported from late Ludlow (Ludfordian) sediments of Bathurst Island in Nunavut, Canada. Evidence for the age of these beds is gathered from stratigraphic relationships and index fossils including conodonts, graptolites, and brachiopods. Zosterophylls dominate the collection, some of which constitute the earliest record of fertile structures arranged in dense clusters and longitudinal rows along axes. Representatives include a plant that resembles Bathurstia, one species of Zosterophyllum, and two specimens that bear affinity to this genus. Distichophytum is also represented, as is a new zosterophyll named Macivera gracilis. The prevalence of sporangial clustering and reduced sporangial stalks in this flora leads to a discussion of the origins and significance of these morphological features. Following a review of some of the other Silurian floras, particularly the Baragwanathia-bearing Lower Plant Assemblage of Victoria, Australia, which also shows morphological advancement over the rhyniophytoid-dominated floras common to Laurussia, it is concluded that the Bathurst Island flora presents the best evidence to date of substantial morphological diversity, complexity, and stature of vascular land plants in this period.     

The microfossil record of early land plants

Dispersed microfossils (spores and phytodebris) provide the earliest evidence for land plants. They are first reported from the Llanvirn (Mid-Ordovician). More or less identical assemblages occur from the Llanvirn (Mid-Ordovician) to the late Llandovery (Early Silurian), suggesting a period of relative stasis some 40 Myr in duration. Various lines of evidence suggest that these early dispersed microfossils derive from parent plants that were bryophyte-like if not in fact bryophytes. In the late Llandovery (late Early Silurian) there was a major change in the nature of dispersed spore assemblages as the separated products of dyads (hilate monads) and tetrads (trilete spores) became relatively abundant. The inception of trilete spores probably represents the appearance of vascular plants or their immediate progenitors. A little later in time, in the Wenlock (early Late Silurian), the earliest unequivocal land plant megafossils occur. They are represented by rhyniophytoids. It is only from the Late Silurian onwards that the microfossil/ megafossil record can be integrated and utilized in interpretation of the flora. Dispersed microfossils are preserved in vast numbers, in a variety of environments, and have a reasonable spatial and temporal fossil record. The fossil record of plant megafossils by comparison is poor and biased, with only a dozen or so known pre-Devonian assemblages. In this paper, the early land plant microfossil record, and its interpretation, are reviewed. New discoveries, novel techniques and fresh lines of inquiry are outlined and discussed.     

Effects of urban land cover on the local species pool in Britain

Regression models were used to quantify the relationship between the amount of urban land and composition of local species pools. There was no evidence that urban land cover increases the richness of plant species, based on a survey of 785 2-km squares of which 157 had > 10% urban land cover. However, the number of alien plants is significantly higher in urban areas. Complete urbanization approximately doubles the proportion of alien species, and the proportion of aliens is twice as high in southern Britain as in the north. The flora of urban tetrads consists of ubiquitous native species and introduced species characteristic of waste ground, but woodland species are poorly represented. At the tetrad scale. enhanced dispersal by man is not the main factor for maintaining the urban flora; availability of urban habitats and high levels of disturbance are more important. The planned housing expansion to greenfield sites in Britain will increase the proportion of alien species, yet the majority of native species should persist in urban areas if existing woodland is preserved.     

丽水白云山种子植物区系的研究

白云山位于浙江省丽水市境内, 自然植被保存完好, 植物种类丰富, 据统计共有野生种子植物117 科、421 属、831 种。其中10 种以上的科有21 个, 498 种, 233 属, 构成了白云山植物区系的主体。本区系历史起源古老, 区域单型属与寡种属共计405 个, 占属总数的96.2%;我国特有分布贫乏, 珍稀濒危植物丰富;地理成分复杂, 可划分为14 个分布区类型13 个变型, 泛热带成分丰富, 温带成分优势明显, 属亚热带向暖温带过渡的地带。     

Paleofloristic diversity and paleoenvironmental interpretation of new Cisuralian localities in the south-central sector of the Anticlinorium of Huayacocotla,Hidalgo, Mexico

The Anticlinorium of Huayacocotla has several outcrops of geological and paleontological importance. However, reports of localities with paleoflora have been scarce so far. In this study, we report three new localities in the Cisuralian, the diversity and composition of which lead us to propose that the paleoflora may have been derived from vegetation that grew in an arid environment with seasonal wetness. The plant remains from the localities of Papaxtla and Calnali are highly transported and their diversity indicates the presence of several groups of plants that were shared with the La Virgen locality, which yielded the best preserved plant fossils. Changes in paleoenvironmental conditions indicate that the preservation of the flora was affected by the tectonic evolution of the region associated with sea level changes. The data presented here increase our knowledge of both the Cisuralian fossil flora and paleoenvironment in the south-central region of the Huayacocotla Anticlinorium, Hidalgo state.     

Early evolution of life cycles in embryophytes: A focus on the fossil evidence of gametophyte/sporophyte size and morphological complexity

Abstract Embryophytes (land plants) are distinguished from their green algal ancestors by diplobiontic life cycles, that is, alternation of multicellular gametophytic and sporophytic generations. The bryophyte sporophyte is small and matrotrophic on the dominant gametophyte; extant vascular plants have an independent, dominant sporophyte and a reduced gametophyte. The elaboration of the diplobiontic life cycle in embryophytes has been thoroughly discussed within the context of the Antithetic and the Homologous Theories. The Antithetic Theory proposes a green algal ancestor with a gametophyte‐dominant haplobiontic life cycle. The Homologous Theory suggests a green algal ancestor with alternation of isomorphic generations. The shifts that led from haplobiontic to diplobiontic life cycles and from gametophytic to sporophytic dominance are most probably related with terrestrial habitats. Cladistic studies strongly support the Antithetic Theory in repeatedly identifying charophycean green algae as the closest relatives of land plants. In recent years, exceptionally well‐preserved axial gametophytes have been described from the Rhynie chert (Lower Devonian, 410 Ma), and the complete life cycle of several Rhynie chert plants has been reconstructed. All show an alternation of more or less isomorphic generations, which is currently accepted as the plesiomorphic condition among all early polysporangiophytes, including basal tracheophytes. Here we review the existing evidence for early embryophyte gametophytes. We also discuss some recently discovered plants preserved as compression fossils and interpreted as gametophytes. All the fossil evidence supports the Antithetic Theory and indicates that the gametophytic generation/sporophytic generation size and complexity ratios show a gradual decrease along the land plant phylogenetic tree.     

Bennettitales from the Grisethorpe Bed (Middle Jurassic) at Cayton Bay, Yorkshire, UK

Middle Jurassic fossil plants from the Grisethorpe Bed at Cayton Bay and Grisethorpe Bay, Yorkshire, UK, are preserved in a soft claystone, and plant mesofossils recovered by sieving reveal excellent details of external structure. Studies of these mesofossils complement previous work on macrofossils from the Grisethorpe Bed and allow the plant fossils in this classic flora to be studied in a similar way to those preserved in Cretaceous mesofloras. Bennettitales, a key group in discussions of how angiosperms may be related to other seed plants, are especially well represented among mesofossils from the Grisethorpe Bed. Abundant bennettitalean leaves, scale leaves, and fragments of pollen and ovulate organs provide new information on these extinct plants. In particular, a specimen of Williamsoniella coronata (presumed aborted) shows only weak differentiation between interseminal scales and ovules and provides further evidence of homology between these structures.     

亚轮生奇异蕨(Adoketophyton subverticillatum)在贵州都匀泥盆系蟒山群的发现及其意义

Adoketophyton subverticillatum (Li et Cai) Li et Edwards (亚轮生奇异蕨)是云南早泥盆世坡松冲植物群中颇具代表性的、研究较为详细的一种植物, 目前对其整体形态特征和解剖结构都有较深入的认识。本文通过系列针修法, 将采自贵州都匀包阳剖面蟒山群下段的一块植物化石新材料归入A. subverticillatum。基于目前对 Adoketophyton时空分布的认识, 认为该属可作为早泥盆世植物群的代表分子, 并据此推断蟒山群下段的时代应为早泥盆世。Adoketophyton subverticillatum在都匀的发现, 表明贵州都匀蟒山群中的植物与云南坡松冲植物群在属、种一级上存在扩散与交流现象。Adoketophyton以及其他相关植物的研究表明, 在高大气CO₂浓度的早泥盆世, 叶性结构如孢子叶、苞片、枝叶复合体等已广泛分布于华南板块。     

湖南植物区系的特点

在大地构造上,湖南属华南地台东部的杨子准地台和南华准地台两个大地构造单元¹⁾,系渊源于震旦纪杨子陆台和华夏陆台^[1]。此陆台自寒武纪至二迭纪反复受海浸,直至二迭纪印支运动地壳普遍抬升,海水从此全部退出地面,逐渐形成今日地貌雏形,再经侏罗纪-白垩纪燕山运动,第三纪喜马拉亚运动始成为现代湖南地貌。     

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.     

A new genus of Early Devonian plants with novel strobilar structures and vegetative appendages from the Posongchong Formation of Yunnan,China

A new plant, Dibracophyton acrovatum gen. et sp. nov., is described from the Lower Devonian (Pragian) Posongchong Formation of Wenshan District, southeastern Yunnan, China. The plant has creeping axes from which arise vegetative and fertile axes. The vegetative axes helically bear lateral dichotomous appendages with curved or round tips. The fertile axes possess terminal strobili with numerous fertile units arranged in irregular helices. Each fertile unit consists of a stalked long-elliptical sporangium, with dehiscence into two equal valves, and two discrete long-ovate bracts covering sporangium from above–below directions. A new genus and species is thus established based on these characters and is temporarily regarded as incertae sedis of Tracheophyta, although it perhaps closes to the barinophytes in affinity. Detailed comparisons with other plants having a similar architecture, i.e., sporangia closely associated with modified vegetative structures, are made. The discovery of D. acrovatum further enriches the composition of the Posongchong flora and demonstrates great morphological disparity of the Early Devonian vascular plants.     

Cloning and characterization of micro-RNAs from moss

Micro-RNAs (miRNAs) are one class of endogenous tiny RNAs that play important regulatory roles in plant development and responses to external stimuli. To date, miRNAs have been cloned from higher plants such as Arabidopsis, rice and pumpkin, and there is limited information on their identity in lower plants including Bryophytes. Bryophytes are among the oldest groups of land plants among the earth's flora, and are important for our understanding of the transition to life on land. To identify miRNAs that might have played a role early in land plant evolution, we constructed a library of small RNAs from the juvenile gametophyte (protonema) of the moss Physcomitrella patens. Sequence analysis revealed five higher plant miRNA homologues, including three members of the miR319 family, previously shown to be involved in the regulation of leaf morphogenesis, and miR156, which has been suggested to regulate several members of the SQUAMOSA PROMOTER BINDING-LIKE (SPL) family in Arabidopsis. We have cloned PpSBP3, a moss SPL homologue that contains an miR156 complementary site, and demonstrated that its mRNA is cleaved within that site suggesting that it is an miR156 target in moss. Six additional candidate moss miRNAs were identified and shown to be expressed in the gametophyte, some of which were developmentally regulated or upregulated by auxin. Our observations suggest that miRNAs play important regulatory roles in mosses.     

The fossil hornwort described from Dominican amber is an angiosperm flower

Hornworts (Anthocerotophyta) are a main lineage of land plants but they are exceedingly rare as fossils. The only fossil hornwort described from amber has been interpreted as the best preserved fossil of this group. Reinvestigation of this fossil revealed that this Miocene amber inclusion represents a poorly preserved flower that shows some features of the Caesalpinioideae subfamily of the Fabaceae.     

云南武定中泥盆世晚期植物与腹足动物 共生标本的发现及其古生态意义

中泥盆世晚期陆生植物的演化已经达到相当高的水平,我国以云南中泥盆世晚期植物群为代表,称为Lepidodendropsis arborecense-Minarodendron cathaysiense-Eocladoxylou minutum (LME)组合。当前在云南武定中泥盆世晚期地层(西冲组上部)中发现的植物主要有Lepidodendropsis arborescense,Eocladoxylon minutu,Rhipidophyton acanthum等,其中,在R.acanthum的茎干基部密集共生有腹足动物Valvata sp.。根据植物表面无动物咬食痕迹,以及腹足动物生态特性和演化分析,当前陆生植物与水生动物共生的原因是植物茎干飘落水体后,为水生的腹足动物提供了新的栖息场所,并随水流、风浪载运至异地埋藏。这种水生腹足动物寄栖于水中物体的生活方式很普遍,是腹足动物迁徙、扩散的一种重要途径。     

Plants as extreme environments? Ni-resistant bacteria and Ni-hyperaccumulators of serpentine flora

During recent years there has been an increasing interest in the bacterial communities occurring in unusual, often extreme, environments. On serpentine outcrops around the world, a high diversity of plant species showing the peculiar features of metal hyperaccumulation is present. These metal hyperaccumulators have received much attention for their potential biotechnological exploitation in phytoremediation processes, but also as unusual, extreme habitats for the associated bacterial flora, which could reveal novel details concerning bacterial adaptation. This paper will briefly focus on the research topics that have been addressed to date on bacteria associated with serpentine plants and aims to provide a state of the art and to present possible future directions for research which could lead to new insights on microbial adaptation and evolution, and potentially applied in technologies for sustainable use and remediation of contaminated land.     

Ex situ conservation of plant diversity in Romania: A synthesis of threatened and endemic taxa

Romania hosts a relatively high species diversity, including 3,829 vascular and 979 non-vascular spontaneous plant taxa. Multiple national red lists exist, with the number of taxa assessed as threatened varying greatly between them, from 548 to 1,438, and with number of taxa assigned to a given threat category also varying between the different sources. A composite list including all taxa mentioned in at least one of the selected red listings from Romania is required in order to compensate for this lack of consensus and to assess their ex situ conservation status. In this study, we synthesized data from the national red lists and counted 1,220 spontaneous vascular plant species and 201 subspecies, of which 77 are endemic and 76 subendemic for Romania. In addition, 18 non-red-listed endemics and 14 subendemics have been added, bringing the total to 1,453 threatened and (sub)endemic plant taxa, representing almost 38% of the total native vascular flora of Romania. Despite the large network of protected areas in Romania, many taxa are still being threatened with extinction in the region mainly due to anthropogenic pressure. Several ex situ conservation measures have been employed to assure a more substantial buffer against plant extinction in the wild, supported by thorough and adequate conservation strategies and multiple means to reintroduce taxa back to their natural habitats. Consequently, our second aim was to evaluate the ex situ conservation status of these threatened and (sub)endemic plants from Romania, focussing on both conventional methods (cultivation in botanic gardens, seed banking) and biotechnological approaches (in vitro tissue culture, medium-term storage and cryostorage). Of the 1,453 taxa included in our list, 642 (44.2%) are conserved by ex situ approaches. Of these, 524 are harboured in the most important botanic gardens throughout Romania, while 156 are currently held in long-term seed banks locally or in the Millennium Seed Bank of the Royal Botanic Garden, Kew (UK). Conversely, only 64 taxa from the list are preserved at the national level through in vitro cultures, and cryopreservation protocols have been developed for only 8 taxa. Overall, more than half of the threatened and (sub)endemic vascular flora from Romania remains unprotected outside the classical in situ conservation measures. For red-listed bryophytes, only 0.6% are preserved in national ex situ collections. Moreover, some aspects related to population genetic studies and the genetic stability of ex situ conserved plants are also briefly discussed, as essential prerequisites for applied biodiversity conservation programs. Finally, considering the distribution range of targeted taxa, we included a synthesis of biotechnological approaches at both national and international level. Our study presents not only a first assessment of the ex situ conservation status of national red listed flora, but also, to our knowledge, the most comprehensive and updated overview of the rare, threatened and (sub)endemic taxa from Romania. This evaluation will provide a supporting tool for national decision- and policy-making actions for biodiversity conservation, using both in situ and ex situ approaches. We also highlight the need for an updated red list for the Romanian flora that accurately follows the IUCN assessment criteria and protocols.     

Broad phylogenomic sampling and the sister lineage of land plants

The tremendous diversity of land plants all descended from a single charophyte green alga that colonized the land somewhere between 430 and 470 million years ago. Six orders of charophyte green algae, in addition to embryophytes, comprise the Streptophyta s.l. Previous studies have focused on reconstructing the phylogeny of organisms tied to this key colonization event, but wildly conflicting results have sparked a contentious debate over which lineage gave rise to land plants. The dominant view has been that 'stoneworts,' or Charales, are the sister lineage, but an alternative hypothesis supports the Zygnematales (often referred to as "pond scum") as the sister lineage. In this paper, we provide a well-supported, 160-nuclear-gene phylogenomic analysis supporting the Zygnematales as the closest living relative to land plants. Our study makes two key contributions to the field: 1) the use of an unbiased method to collect a large set of orthologs from deeply diverging species and 2) the use of these data in determining the sister lineage to land plants. We anticipate this updated phylogeny not only will hugely impact lesson plans in introductory biology courses, but also will provide a solid phylogenetic tree for future green-lineage research, whether it be related to plants or green algae.