The potential for forest canopy litterfall interception by a dense fern understorey, and the consequences for litter decomposition

Most studies on litter decomposition have assumed that all falling plant litter reaches the ground where it then decomposes. In many forests a proportion of this litter may in fact be intercepted by understorey vegetation, but the ecological significance of this remains poorly understood. We performed two experiments in a temperate rainforest in southern New Zealand, in which there was a dense understorey of the crown fern Blechnum discolor. The fronds of this fern originate from a crown, and have a funnel‐like arrangement that can trap falling litter and prevent it from reaching the ground. The first experiment measured the effects of ferns on the spatial distribution of litter accumulation over one year. The ferns intercepted a substantial proportion of the total litterfall, and the fern crowns (from which the fronds originate) retained 10% of the total incoming litterfall (despite occupying only 2% of the ground area). The retained litter had a substantially higher ratio of twig to foliar litter than did the incoming litterfall. Further, much of the litter not retained on the crowns of the ferns accumulated at the base of the fern trunks. The second experiment considered litter decomposition in fern crowns versus the ground under the ferns. The litter that had accumulated in the crowns was characterized by higher microbial basal respiration and active microbial biomass than was the litter that had accumulated on the ground. The use of litterbags revealed that litter decomposition rates were significantly higher on the fern crowns than on the ground at 30 cm and 60 cm from the fern trunks. These results show that litter interception ameliorates the decomposer environment and increases the rate of litter decomposition. In total, this study provides evidence for understorey ferns greatly influencing both the spatial distribution of litterfall and the decomposition of plant litter. Although the ecological role of understorey vegetation in forested ecosystems has received little attention to date, our results point to understorey species as an important driver of forest ecosystem processes.     

附生蕨类植物的克隆性研究进展

石松类及蕨类植物在高等植物中处于比较特殊的进化与系统发育地位, 同时具有孢子植物(孢子)与种子植物(维管束)的双重特征。附生蕨类植物是蕨类植物中占据独特生境的一个大类群, 其生活史策略及进化历史与其附生生长的森林生态系统紧密相关。大部分附生蕨类植物的克隆生长习性及克隆生活史性状在其生态适应中具有重要作用, 但这方面未引起广泛关注。本文主要综述了中国山地森林中附生蕨类植物的根状茎克隆生长、克隆性与生态适应性、不同克隆生长方式与进化等方面, 并展望了蕨类植物克隆性在森林生态系统过程与功能中的作用, 以及今后如何将蕨类植物生态学研究与气候变化、植被恢复、土地利用变化等全球变化的主流方向进行结合。     

蕨类植物的化感作用及其对生物多样性的影响

植物化感作用是植物通过向环境中释放化学物质从而对同种植株和繁殖体或与其他植物之间产生的直接或间接、有益或有害的作用,它影响植物分布、群落形成与演化、作物间作等,与生物多样性保护以及农林和园艺生产实践关系密切,在国际上受到越来越多的关注。但国内这方面的研究起步相对较晚,研究范围有限,对蕨类植物化感作用的报道更少。本文系统介绍了蕨类植物化感作用的研究进展,包括蕨类植物种内的化感作用(即自毒效应)、常见的蕨类植物种间的化感作用(即孢子体对配子体的化感作用和配子体对配子体的化感作用)及蕨类植物对种子植物的化感作用(蕨类植物可通过化感作用与种子植物争夺更多的资源和生长空间)。还介绍了种子植物对蕨类植物的化感作用(主要表现为抑制作用)以及蕨类植物化感作用与动物侵食、微生物侵染的关系,研究发现昆虫侵食能增强或减弱蕨类植物的化感作用,微生物的活动可能增强某些蕨类植物的化感作用。本文从上述不同角度说明蕨类植物化感作用对生物多样性的影响,希望有助于促进我国学者对该领域的深入研究。     

Complete chloroplast genome sequence of a tree fern Alsophila spinulosa : insights into evolutionary changes in fern chloroplast genomes

Background  

Ferns have generally been neglected in studies of chloroplast genomics. Before this study, only one polypod and two basal ferns had their complete chloroplast (cp) genome reported. Tree ferns represent an ancient fern lineage that first occurred in the Late Triassic. In recent phylogenetic analyses, tree ferns were shown to be the sister group of polypods, the most diverse group of living ferns. Availability of cp genome sequence from a tree fern will facilitate interpretation of the evolutionary changes of fern cp genomes. Here we have sequenced the complete cp genome of a scaly tree fern Alsophila spinulosa (Cyatheaceae).     

New guinea salt fern (Asplenium acrobryum complex): Identity,distribution, and chemical composition of its salt

Fronds of a fern of theAsplenium acrobryum complex were traditionally used as a source of salt in the inland areas of Papua New Guinea. All previously published reports of the use of fern salt in the botanical and ethnobotanical literature had erroneously identified the species involved asAsplenium nidus, one of the common bird’s-nest ferns. A chemical analysis of the salts contained in the ash of the salt fern as well as those species related toAsplenium nidus revealed that the bulk of the inorganic component was potassium, calcium and chloride. These results are comparable to those published for the salt-yielding grass,Coix gigantean. There is no obvious chemical reason why onlyAsplenium acrobryum should be used for salt production in preference toAsplenium nidus and related species. In view of the low levels of consumption of such salt it is unlikely that there will be any cases of potassium toxicity attributable to the use of these plants.     

Tree fern trunks facilitate seedling regeneration in a productive lowland temperate rain forest

Seedling regeneration on forest floors is often impaired by competition with established plants. In some lowland temperate rain forests, tree fern trunks provide safe sites on which tree species establish, and grow large enough to take root in the ground and persist. Here we explore the competitive and facilitative effects of two tree fern species, Cyathea smithii and Dicksonia squarrosa, on the epiphytic regeneration of tree species in nutrient-rich alluvial forests in New Zealand. The difficulties that seedlings have in establishing on vertical tree fern trunks were indicated by the following observations. First, seedling abundance was greatest on the oldest sections of tree fern trunks, near the base, suggesting that trunks gradually recruited more and more seedlings over time, but many sections of trunk were devoid of seedlings, indicating the difficulty of establishment on a vertical surface. Second, most seedlings were from small-seeded species, presumably because smaller seeds can easily lodge on tree fern trunks. Deer browsing damage was observed on 73% of epiphytic seedlings growing within 2 m of the ground, whereas few seedlings above that height were browsed. This suggests that tree ferns provide refugia from introduced deer, and may slow the decline in population size of deer-preferred species. We reasoned that tree ferns would compete with epiphytic seedlings for light, because below the tree fern canopy photosynthetically active radiation (PAR) was about 1% of above-canopy PAR. Frond removal almost tripled %PAR on the forest floor, leading to a significant increase in the height growth rate (HGR) of seedlings planted on the forest floor, but having no effects on the HGRs of epiphytic seedlings. Our study shows evidence of direct facilitative interactions by tree ferns during seedling establishment in plant communities associated with nutrient-rich soils.     

Contrasting Predictors of Fern versus Angiosperm Decomposition in a Common Garden

We evaluated differences in the rates and correlates of decomposition among 32 fern and angiosperm litter types collected in Hawai'i. Leptosporangiate ferns were separated into groups based on phylogeny: 'polypod' ferns, a monophyletic clade of ferns that diversified in the Cretaceous, and all other ('non-polypod') ferns that diversified earlier. We measured initial litter chemistry (nutrients and carbon chemistry), and mass loss and nitrogen (N), phosphorus (P), and calcium (Ca) of litter tissue during a 1-yr incubation in a common garden. Nutrient concentrations and carbon (C) chemistry differed significantly among litter types, and litter turnover ( k -values) ranged from 0.29 to 8.31. Decomposition rates were more closely correlated with nutrient concentration than is typically observed. Lignin:N was the best predictor of decomposition across all litter types combined; however, among plant groups different predictors of decomposition were important. Nitrogen and P concentrations best predicted fern decomposition, whereas C chemistry, particularly lignin concentration, was more important for angiosperm (monocot and dicot) decomposition. Among native plants, non-polypod ferns decomposed significantly more slowly than both polypod ferns and angiosperms. Contrary to our hypothesis, fern litter did not decompose more slowly than angiosperm litter overall. Nutrient dynamics in litter were affected by initial litter concentration more than phylogeny; low-nutrient litter immobilized more nutrients than high-nutrient litter. Systematic differences in rates of decomposition, and the importance of nutrients in predicting fern decomposition, imply that changes in species composition within ferns and between ferns and angiosperms could influence the functioning of ecosystems where ferns are important forest components.     

A current perspective on apomixis in ferns

This review provides a synopsis of apogamous reproduction in ferns and highlights important progress made in recent studies of fern apomixis. First, a summary of the apomictic fern life cycle is provided, distinguishing between two pathways to diploid spore production that have been documented in apomictic ferns (premeiotic endomitosis and meiotic first division restitution) and briefly discussing the evolutionary implications of each. Next, recent trends in fern apomixis research are discussed, exposing a shift in focus from the observation and characterization of apomixis in ferns to more integrated studies of the evolutionary and ecological implications of this reproductive mode. Peer-reviewed contributions from the past decade are then summarized, spanning the identification of new apomictic lineages through to the developmental, phylogenetic, and population genetic insights that have been made in studies of fern apomixis during that time. Gaps in our understanding are also discussed, including the extent and implications of recombinant apomixis in ferns, the possible reversibility of reproductive mode (from apomictic to sexual) in ferns, and the genomic causes and consequences of apomixis in seed free vascular plants. To conclude, future directions for fern apomixis research are proposed in the context of modern technological advances and recent insights from studies of apomixis in other groups.     

Invasive feral pigs impact native tree ferns and woody seedlings in Hawaiian forest

Invasive mammals can fundamentally alter native plant communities, especially on isolated islands where plants evolved without them. The globally invasive feral pig (Sus scrofa) can be particularly destructive to native plant communities. Tree ferns are an important understory component in many forests facilitating the establishment of a variety of species. However, the extent and effects of feral pig damage to tree ferns, and associated impacts on plant community regeneration, are largely unknown. We quantified the effect that feral pig damage has on tree fern growth, survival, and epiphytic woody seedling abundance over 1 year on 438 randomly selected tree ferns of three endemic species (Cibotium chamissoi, Cibotium glaucum, and Cibotium menziesii) in a Hawaiian montane wet forest with high tree fern and feral pig densities. Across all tree fern species, feral pigs damaged 13 % of individuals over 1 year. Compared with undamaged tree ferns, moderately- to heavily-damaged individuals had decreases of 4 to 27 % in trunk length increment and lost tenfold more fronds. Tree fern angle (standing, leaning, prone, or semi-prone) and woody seedling abundance co-varied with feral pig damage. Specifically, damaged tree ferns were more often prone or semi-prone and supported more seedlings, but also had annual mortality up to 34 % higher than undamaged tree ferns. Overall, feral pig damage had substantial negative effects on tree ferns by reducing growth and survival. Given the importance of tree ferns as regeneration sites for a variety of native plants, feral pig damage to tree ferns will likely alter future forest composition and structure. Specifically, feral pig damage to tree ferns reduces potential establishment sites for species that either regenerate preferentially as epiphytes or are currently restricted to epiphytic establishment due to ground rooting by feral pigs.     

Four‐level interactions: herbivore use of ferns and subsequent parasitoid–hyperparasitoid performance

Abstract 1. Variables affecting species at the ends of trophic chains may modify the success of members with which they do not directly interact. The majority of such examples involve three trophic levels, but hyperparasitoids provide an excellent opportunity to examine four‐level relationships. 2. The gregarious hyperparasitoid Aprostocetus sp. (Hymenoptera: Eulophidae) commonly attacks the primary parasitoid Alabagrus texanus (Hymenoptera: Braconidae), by far the commonest parasitoid of the moth Herpetogramma theseusalis (Lepidoptera: Crambidae). 3. Larvae of this moth feed on ferns of two families, sensitive fern Onoclea sensibilis (Dryopteridaceae) and marsh fern Thelypteris palustris (Thelypteridaceae), in the study area, an old field in Maine, U.S.A. 4. I test the hypotheses that the ferns indirectly affect the reproductive success of the hyperparasitoids and that the ferns produce similar effects at intermediate links. 5. The moths experienced similar success on the two ferns, and the primary parasitoid performed similarly on moths reared from both ferns. The hyperparasitoid parasitized similar proportions of the primary parasitoid from moths that fed on sensitive fern and marsh fern. 6. However, hyperparasitoid broods on primary parasitoids from moths feeding on marsh fern contained approximately one‐third more offspring, whose individuals were significantly larger than those from sensitive fern, even though their hosts’ sizes did not differ significantly. 7. An indirect effect, related to the primary producers, thus strongly affected Trophic Level 4 in the absence of a significant effect at intermediate levels. To the best of my knowledge, this relationship has not been previously reported in a multi‐year or field‐based study of a natural system.     

The evolution of chloroplast genes and genomes in ferns

Most of the publicly available data on chloroplast (plastid) genes and genomes come from seed plants, with relatively little information from their sister group, the ferns. Here we describe several broad evolutionary patterns and processes in fern plastid genomes (plastomes), and we include some new plastome sequence data. We review what we know about the evolutionary history of plastome structure across the fern phylogeny and we compare plastome organization and patterns of evolution in ferns to those in seed plants. A large clade of ferns is characterized by a plastome that has been reorganized with respect to the ancestral gene order (a similar order that is ancestral in seed plants). We review the sequence of inversions that gave rise to this organization. We also explore global nucleotide substitution patterns in ferns versus those found in seed plants across plastid genes, and we review the high levels of RNA editing observed in fern plastomes.     

Recurrent genome duplication events likely contributed to both the ancient and recent rise of ferns

Ferns, the second largest group of vascular plants, originated ～400 mil ion years ago(Mya). They became dominant in the ancient Earth landscape before the angiosperms and are stil important in current ecosystems.Many ferns have exceptional y high chromosome numbers,possibly resulting from whole-genome duplications(WGDs).However, WGDs have not been investigated molecularly across fern diversity. Here we detected and dated fern WGDs using a phylogenomic approach and by calculating synonymous substitution rates(Ks). We also investigated a possible correlation between proposed WGDs and shifts in species diversification rates. We identified 19 WGDs: three ancient events along the fern phylogenetic backbone that are shared by 66%–97% of extant ferns, with additional lineage-specific WGDs for eight orders, providing strongevidence for recurring genome duplications across fern evolutionary history. We also observed similar Ks peak values for more than half of these WGDs, with multiple WGDs occurring close to the Cretaceous(～145–66 Mya). Despite the repeated WGD events, the biodiversity of ferns declined during the Cretaceous, implying that other factors probably contributed to the floristic turnover from ferns to angiosperms. This study provides molecular evidence for recurring WGDs in ferns and offers important clues to the genomic evolutionary history of ferns.     

蕨类植物花青素和原花青素成分及含量分析

以7目14科共44种蕨类植物为材料,对它们的花青素、原花青素和总黄酮含量进行检测。结果显示,44种蕨类植物均含有花青素,较为进化的水龙骨目植物的总花青素平均含量明显高于其它蕨类植物。矢车菊素、飞燕草素和天竺葵素是蕨类植物主要的花青素类型,其中乌毛蕨科植物富含矢车菊素,鳞毛蕨科植物富含飞燕草素。本研究中大部分蕨类含有原花青素,水龙骨目植物的原花青素平均含量高于其它蕨类。研究结果表明,蕨类植物中花青素和原花青素等黄酮类化合物的分布与植物科属相关,推测花青素与蕨类植物的生长发育和抗逆过程相关。     

蕨类植物配子体育及其性器官分化的研究进展

作者概述了蕨类植物生活周期中不同发育阶段的特点,并着重介绍了不,植物生长调节物质,水和温度等因素对蕨类植物配子体发育的影响以及精子囊素,光,钙离子和其它植物生长调节物质在性器官形成和分化过程中的作用,同时作者对蕨类植物性器官分化的不稳定性进行了探讨,并结合目前蕨类植物性器官分化的遗传学和分子生物学研究的最新进展,说明蕨类植物作为研究系统在植物生物学研究领域中的优势。     

蕨类植物配子体发育及其性器官分化的研究进展

作者概述了蕨类植物生活周期中不同发育阶段的特点,并着重介绍了光、植物生长调节物质、水和温度等因素对蕨类植物配子体发育的影响以及成精子囊素、光、钙离子和其它植物生长调节物质在性器官形成和分化过程中的作用。同时作者对蕨类植物性器官分化的不稳定性进行了探讨、并结合目前蕨类植物性器官分化的遗传学和分子生物学研究的最新进展,说明蕨类植物作为研究系统在植物生物学研究领域中的优势。     

Host-range and cold temperature tolerance of Floracarus perrepae Knihinicki & Boczek (Acari: Eriophyidae), a potential biological-control agent of Lygodium microphyllum (Pteridophyta: Lygodiaceae)

Abstract The leaf galling eriophyid mite Floracarus perrepae is a widespread and damaging herbivore of Old World climbing fern, Lygodium microphyllum , across its native range in tropical and subtropical Asia and Australia; and was therefore selected as a candidate biological-control agent for the invasive fern in Florida, USA. The host testing of F. perrepae focused on Lygodium species from North America and the Neotropics, along with threatened or endangered ferns from Florida. Sporeling ferns were used for the initial no-choice screening and F. perrepae showed normal development on the Florida genotype of L. microphyllum with 33.0 ± 4.6 mites per marginal leaf roll gall, modest development on the North American native Lygodium palmatum with 29.0 ± 9.3 mites per roll, and minor development on six other fern species (4.0–0.3). Leaf rolls were induced only on Lygodium species and full rolls were common only on L. microphyllum . These same six non-target plant species were tested again as more mature plants in both no-choice and choice tests and F. perrepae developed only on L. microphyllum . Lethal minimum temperature and cold stress tests were also conducted on F. perrepae . These tests revealed that it would not likely establish north of USDA hardiness zone 8a (−9.5 to −12.2°C), which means it would not overlap with L. palmatum from temperate North America. Our studies conclude that F. perrepae is specific to L. microphyllum , poses little or no risk to native or cultivated ferns in North and South America, and therefore should be considered for release in Florida.     

Elevational patterns of fern species assemblages and richness in central Japan

We conducted field surveys in 807 quadrats to evaluate the elevational belts, boundary and richness patterns of ferns and lycophytes in the temperate region of central Japan. We analysed fern species assemblages at 100 m elevational steps by cluster analysis and tested the number of upper and lower boundaries for elevational intervals against a null model of random distribution of elevational limits. We compared the pattern of fern species richness along the elevational gradients in central Japan with patterns in several locations to evaluate the fern flora in central Japan in relation to the rest of the world. We recorded 261 ferns species in total, which is one-third of the Japanese ferns. We found clear elevational boundaries of fern assemblages at 900 and 1,800 m and three fern elevational zones, which corresponded well to the elevational limits of forest types in central Japan. The pattern of fern species richness in central Japan was an asymmetric hump-shaped pattern that peaked close to the sea level, with the peak of local richness at lower elevations than that of regional richness. We found that the peak of fern species richness along the elevational gradient in Japan was located at lower elevations than that of fern elevational patterns in several locations around the world.     

Effects of fern thickets on woodland development on landslides in Puerto Rico

Abstract. Thicket-forming ferns are common colonizers of disturbed habitats in the tropics, but little is known about their ecology. The effects of thickets formed by the fern Dicranopteris pectinata on tree seedlings on five landslides in the Luquillo Experimental Forest in northeastern Puerto Rico were both positive and negative. Soil moisture and total soil N were higher under fern thickets than in adjacent open areas and soil bulk density and soil surface temperatures were lower. Germination of seeds of the tree Cecropia schreberiana was higher for seeds sown under fern thickets than for those sown into adjacent open areas. Tree seedlings of Tabebuia hetero-phylla exhibited a threefold reduction in photosynthesis under ferns, probably resulting from a twelvefold reduction of photosynthetic photon flux density. Growth of Tabebuia seedlings was reduced under ferns but the distribution of seedlings of naturally occurring woody plants was not strongly correlated with the presence of fern thickets. Although fern thickets on low-nutrient landslide soils appear to facilitate germination, they inhibit growth of tree seedlings and may, therefore, delay forest development on landslides in Puerto Rico.     

Structure-function constraints of tracheid-based xylem: a comparison of conifers and ferns

The ferns comprise one of the most ancient tracheophytic plant lineages, and occupy habitats ranging from tundra to deserts and the equatorial tropics. Like their nearest relatives the conifers, modern ferns possess tracheid-based xylem but the structure-function relationships of fern xylem are poorly understood. Here, we sampled the fronds (megaphylls) of 16 species across the fern phylogeny, and examined the relationships among hydraulic transport, drought-induced cavitation resistance, the xylem anatomy of the stipe, and the gas-exchange response of the pinnae. For comparison, the results are presented alongside a similar suite of conifer data. Fern xylem is as resistant to cavitation as conifer xylem, but exhibits none of the hydraulic or structural trade-offs associated with resistance to cavitation. On a conduit diameter basis, fern xylem can exhibit greater hydraulic efficiency than conifer and angiosperm xylem. In ferns, wide and long tracheids compensate in part for the lack of secondary xylem and allow ferns to exhibit transport rates on a par with those of conifers. We suspect that it is the arrangement of the primary xylem, in addition to the intrinsic traits of the conduits themselves, that may help explain the broad range of cavitation resistance in ferns.     

21世纪以来蕨类植物研究论文的发表情况: 基于Web of Science的数据统计

蕨类植物是地球上起源最为古老的维管植物, 在陆生植物演化中具有重要意义。本文通过Web of Science核心数据库(Core Collection)对目前所有关于“fern”、“lycophyte”、“pteridophyte”、“pteridophyta”等的科研论文进行了分析。结果表明: (1) 2000年后, 蕨类植物论文数量增加迅速, 特别是2006年以来增加显著, 现在年均已突破600篇; (2) 2010年以来的以蕨类植物为研究材料和对象的学科和研究领域变得越来越多元化, 多学科结合和交叉的研究方向成为新趋势; (3)中国正在成为蕨类植物研究的热点国家并且成果显著; (4)中国蕨类植物研究在快速发展的同时也出现了各研究机构之间的发展不同步。期望本论文能给蕨类植物的相关研究提供新的方向, 为中国蕨类植物工作者提供借鉴。