西双版纳勐腊南贡山季风常绿阔叶林蕨类植物初步研究

勐腊南贡山季风常绿阔叶林(山坡类型)1 hm2面积样地上有蕨类31种,在该森林群落草本层中占有 极其显著的地位。这些蕨类植物的地理成分分析表明它们属于亚洲热带的印度——马来西亚植物区系的一 部分,表现为东南亚热带北缘植物区系的性质,并有热带山地向亚热带山地过渡的特点。在生态表现上,它们 的生活型组成是高位芽蕨类(地生蕨种)占6．45％;地上芽蕨类12．90％;地面芽蕨类32．26％;地下芽蕨类 32．26％;附生蕨类16．13％。由于该地区的季节性干旱气候,高位芽蕨类较少和几种附生蕨类的附生高度也 相对较低,蕨类植物种群的数量与林下空气湿度及土壤表层的湿度一般成正相关。     

Hydrocarbons and fatty acids of ferns

The analysis of hydrocarbons and fatty acids in ten fern species indicate unique differences from plants in a higher phylogenetic order. Significant concentrations of fatty acids above C₂₀ are present. Distributions of hydrocarbons range from C₁₅–C₃₃ with a trend towards two maxima at C₁₇ and C₂₉. Homologous series of n-alkenes are present in all species. Pristane and phytane are large components representing up to 27% of the alkanes. Distinct alkane and fatty acid differences between fern families are observed while species variations within families are slight.     

Diversity variation and tempo-spatial distributions of the Dipteridaceae ferns in the Mesozoic of China

The extant family Dipteridaceae is a remarkable leptosporangiate fern because it includes only one genus with a restricted distribution to tropical regions. The fossil record of this family has been widely reported from the Mesozoic strata in Eurasia, America, Australia, and Greenland. In China, numerous fossils of the Dipteridaceae have been documented, in total, about 74 species of 6 genera. Geographically, they are distributed both in the Southern and Northern Floristic Provinces, and were particularly well developed in the Southern Floristic Province during the Late Triassic and the Early Jurassic intervals. Fossil diversity of Dipteridaceae varies in the different episodes of the Mesozoic in China. It is shown that Dipteridaceae has undergone a diversity development process and a distinct turnover during the Mesozoic. They appear to have diversified in the warm and humid Late Triassic–Early Jurassic, but declined sharply as aridity developed in the Middle Jurassic, and became extinct at the end of the Early Cretaceous. The diversity variation and tempo-spatial distribution pattern is suggested to be linked with paleoclimatic variations during the Mesozoic.     

上海市蕨类植物分布特点及其与环境因子关系分析

在查阅资料和标本基础上,对上海市33个样地蕨类植物进行了野外调查,初步确定了上海市自然分布的蕨类植物23科31属50种,并讨论了归化种与栽培种及曾有记录而可能消亡的种类。采用双向指示种法（TWINSPAN）和典范对应分析（CCA）方法对蕨类植物分布特点及其与环境因子关系进行了分析。结果表明：（1）蕨类分布的样点可分为3组,第1组为市区样点,种数为0～4种;第2组为市郊样点,种数为0～8种;第3组为佘山岛、佘山地区、大金山岛,是上海蕨类最丰富地区,种数达26种以上。（2）影响上海蕨类植物生长的主要环境因子为基质、郁闭度和人为干扰。     

Cainozoic ferns and their distribution

The phytogeographic distribution of Cainozoic ferns is reported based upon a critical re-appraisal of the macrofossil and mesofossil record also taking account of evidence from a few highly diagnostic spores. Well-documented circum-Arctic Cainozoic floras show ferns (Woodwardia, Onoclea, Osmunda, Coniopteris and to a lesser extentAzolla) distributed around the pole to very high paleolatitudes. Some ferns are shared between the mid-paleolatitudes of North America and Europe as would be predicted from the distributions of other biota. Evidence for the composition of Cainozoic fern floras is minimal in some regions (e.g., Antarctica, Central and South America, Africa, India, South East Asia), so the absence of fern fossils from these areas has no biogeographical significance. Matoniaceae were abundant in the preceding Mesozoic. However, the absence of Cainozoic macrofossils, and the fact that no CainozoicMatonisporites spores areMatonia-like, indicates that Matoniaceae had attained their modern relict distribution by, or very early in, the Cainozoic. The important Mesozoic families Marattiaceae and Dipteridaceae are also not represented by Cainozoic macrofossils. They probably also showed Cainozoic restriction but spores are not sufficiently diagnostic to enable testing of this hypothesis. Other ferns, which were also important in the Mesozoic (e.g., Dicksoniaceae, Gleicheniaceae), have patchy, equivocal, or inadequately published Cainozoic records. The dispersed spore record may provide an opportunity to track Cainozoic Gleicheniaceae but this approach is not without problems. Most well-represented Cainozoic fern families, genera and subgenera show widespread Cainozoic ranges, typically with considerable range extensions over their living relatives, both onto other continents and north and south to higher paleolatitudes. These include Schizaeaceae (Lygodium, Anemia, and the extinctRuffordia), Osmundaceae (Osmunda), Pteridaceae (Acrostichum), Thelypteridaceae (Cyclosorus), Lophosoriaceae (Lophosoria), Cyatheaceae (theCnemidaria/Cyathea decurrens clade) and the heterosporous water fernAzolla (Azollaceae). A few well-represented ferns show Cainozoic distributions similar to those of the present day (e.g.,Salvinia [Salviniaceae] andCeratopteris [Pteridaceae] (the latter by the Neogene and based only on spores]) but even these had slightly broader ranges in the Cainozoic. Some Cainozoic ferns have apparently local distributions, e.g.,Blechnum dentatum (Blechnaceae) in Europe; and others are so far represented at only one or few sites, e.g.,Dennstaedtiopsis (Dennstaedtiaceae),Botrychium (Ophioglossales),Grammitis (Grammitidaceae), andMakotopteris andRumohra (Dryopteridaceae). Cainozoic fossils assigned toDryopteris (and some other dryopteroids) require revision along with those of Thelypteridaceae, the latter having high potential to provide useful paleobiogeographic evidence, at least of theCyclosorus group. Cainozoic records of Hymenophyllaceae and Polypodiaceae are here considered unconfirmed.     

Development of primer sets for PCR amplification of the PgiC gene in ferns

Polymerase chain reaction (PCR)-based nuclear DNA markers were developed for fern species. We first determined the partial nucleotide sequence of cDNA of the pgiC gene encoding cytosolic phosphoglucose isomerase from Dryopteris caudipinna, and then PCR primers for exon-primed, intron-crossing (EPIC) amplifications were designed. The EPIC primers are universally applicable to the most derived indusiate fern families such as Dryopteridaceae, Thelypteridaceae, and Woodsiaceae. The PCR products of primers 14F/16R containing two introns are moderate in size (534 bp–ca.1000 bp) and are possibly of value in phylogenetic reconstruction at specific and generic levels. Codominant nuclear DNA markers applicable to the estimation of mating systems and other population genetic studies were also developed by a combination of single-strand conformation polymorphism (SSCP) and EPIC amplification using primers 14F/15R and 15F/16R. In order to provide a case study using these markers, allelic variation of PCR products using 15F/16R was examined in populations of Arachniodes standishii (Dryopteridaceae). Received: July 4, 2001 / Accepted: September 12, 2001     

云南蕨类植物的物种多样性和区系组成

在一定实地调查和分类的基础上,采用统计分析及区系地理分析法,对云南地区蕨类植物进行研究。结果表明:云南地区共有蕨类植物60科193属1 530种(包括变种和变型),科、属、种分别占中国蕨类植物科总数的58.8%,属的83.9%和种的95.2%;在这60个科中含30种以上的有13个,含5个属以上的有11个,分别占该蕨类植物区系属、种数的56.5%和79.2%,特别是鳞毛蕨科、蹄盖蕨科、水龙骨科和金星蕨科,这4个科共拥有70属761种,分别占总属数、总种数的36.3%、49.7%;193属中含30种以上的有11个,分别占总属、总种数的5.7%和41.7%,最具优势的是鳞毛蕨属和耳蕨属、蹄盖蕨属、铁角蕨属、卷柏属,这5个属共有403种,占总种数的26.3%;该区系中无云南特有科,但具有4个亚洲特有科:雨蕨科、稀子蕨科、柄盖蕨科和骨碎补科;该区系以热带、亚热带性质为主,科的区系成分中热带亚热带分布的科占66.6%,热带成分的属有112个,占68.7%(除世界分布类型);该区系是东亚地区蕨类植物区系的重要组成部分,东亚分布31属和中国特有分布6属,分别占总属数(除世界分布属外)的19.0%和3.7%;该地区蕨类植物区系与西藏、台湾具有共同的区系起源和物种分化形成的背景,属的相似性系数约为70%,种的相似性系数约为30%,均起源于热带亚热带地区;属的热带区系成分与温带区系成分所占比例分别为68.7%和27.6%,存在一定的区系过渡性;该区系科的分化强度为3.2,属的分化强度为7.9,在科、属水平上均表现出较强的区系分化特征。     

Epiphytic ferns and bryophytes of Tasmanian tree‐ferns: A comparison of diversity and composition between two host species

Abstract Ferns, bryophytes and lichens are the most diverse groups of plants in wet forests in south‐eastern Australia. However, management of this diversity is limited by a lack of ecological knowledge of these groups and the difficulty in identifying species for non‐experts. These problems may be alleviated by the identification and characterization of suitable proxies for this diversity. Epiphytic substrates are potential proxies. To evaluate the significance of some epiphytic substrates, fern and bryophyte assemblages on a common tree‐fern species, Dicksonia antarctica (soft tree‐fern), were compared with those on a rare species, Cyathea cunninghamii (slender tree‐fern), in eastern Tasmania, Australia. A total of 97 fern and bryophyte species were recorded on D. antarctica from 120 trunks at 10 sites, and 64 species on C. cunninghamii from 39 trunks at four of these sites. The trunks of C. cunninghamii generally supported fewer species than D. antarctica, but two mosses (particularly Hymenodon pilifer) and one liverwort showed significant associations with this host. Several other bryophytes and epiphytic ferns showed an affinity for the trunks of D. antarctica. Species assemblages differed significantly between both sites and hosts, and the differences between hosts varied significantly among sites. The exceptionally high epiphytic diversity associated with D. antarctica suggests that it plays an important ecological role in Tasmanian forests. Evidently C. cunninghamii also supports a diverse suite of epiphytes, including at least one specialist species.     

Tissue-direct PCR, a rapid and extraction-free method for barcoding of ferns

Fern gametophytes and young sporophytes often provide too little material for DNA extraction and are particularly difficult to identify to genus. Here we developed an efficient procedure called 'Tissue-direct PCR', in which a slice of fern tissue is mixed with PCR reagents and primers, allowing certain genomic regions to be amplified directly in the thermal cycler. For these diminutive and featureless stages of ferns, Tissue-direct PCR combined with amplifying plant barcodes promises to make the identification of immature ferns easy and rapid. Tissue-direct PCR would also be very helpful for large-scale ecological studies surveying distribution and population structure.     

蕨类植物性别分化对环境的响应

蕨类植物是维管植物中唯一的孢子体和配子体都能独立生活的类群.同型孢子蕨类配子体的性别分化受到激素和环境因子的影响.生理学研究表明,成精子囊素与赤霉素能诱导雄配子体发育,抑制雌配子体发育;脱落酸阻止成精子囊素诱导的精子器形成;乙烯合成前体ACC促进赤霉素诱导的精子器形成,而乙烯合成抑制因子AOA通过抑制细胞分化来抑制精子器形成.光照对不同种类蕨类配子体分化的影响存在差异.糖类能够促进雄配子体形成,并可加速成熟雌配子体向两性分化.钙离子、钴离子和甲硫氨酸等分别参与了蓝光和赤霉素对配子体性别分化的调控过程.培养密度影响配子体生长及性别表达,高密度下雄性和无性配子体居多,而低密度下两性和雌性配子体居多.近年来的突变体表型分析与分子生物学研究表明,成精子囊素通过影响ANI1、HER、TRA、FEM和MAN等基因的表达调控配子体性别分化.综述了蕨类植物性别分化对环境响应的研究进展.     

DNA amounts for five pteridophyte species fill phylogenetic gaps in C-value data

C-values are important biodiversity characters, yet data for pteridophytes are scarce. A recent survey showed that reliable C-value data were available for only 62 species. This paper reports C-value estimates for a further five pteridophyte species which were selected to fill phylogenetic gaps identified in previous work: Angiopteris lygodiifolia , of the eusporangiate fern family Marattiaceae (1C = 7.10 pg); Lygodium japonicum , of the basal leptosporangiate fern family Schizaeaceae (1C = 11.66 pg); Marsilea quadrifolia , of the heterosporous water fern order Marsileales (1C = 4.01 pg); Isoetes lacustris (1C = 11.97 pg) and Lycopodium clavatum (1C = 2.86 pg), of the lycophyte families Isoetaceae and Lycopodiaceae, respectively. The C-value estimates are discussed in the context of other C-value data. Further, the range of C-values in lycophytes (0.16 pg – 11.97 pg), which comprise Selaginellaceae, Isoetaceae and Lycopodiaceae, is discussed in relation to nucleotypic constraints imposed by sperm flagella number.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 169−173.     

A global comparative analysis of elevational species richness patterns of ferns

Aim Elevational gradients offer an outstanding opportunity to assess factors determining patterns of species richness, but along single transects potential explanatory factors often covary, making it difficult to distinguish between competing hypotheses. Many previous studies on plants have interpreted their results as supporting the mid‐domain effect (MDE) as a major determinant of species richness, even when climatic factors showed similarly high explanatory power. We compared fern species richness along 20 elevational transects to quantify the relative contribution of climate and MDE as drivers of elevational richness patterns. Location Twenty transects world‐wide. Methods Ferns were sampled in 1039 plots of 400–2500 m² each. Mean annual precipitation and temperature, epiphytic bryophyte cover (as a proxy for air humidity) and MDE predictions were included as independent variables. For each transect, we calculated multiple linear models and partitioned the variance to assess the relative contribution of the independent variables, selecting the most parsimonious models based on Akaike weights and multi‐model inference. Results Along most individual gradients, nearly all variance of fern species richness that could be attributed to either space or MDEs was collinear with climatic factors. Yet, the comparison across transects showed that elevational richness patterns are most parsimoniously accounted for by climatic conditions, especially by low water availability at low elevations and in dry regions in general, and by low temperatures at high elevations and in extra‐tropical regions. Main conclusions Fern species richness is most closely related to climatic factors, and while MDE, surface area and metapopulation processes may somewhat modify the patterns, their importance has been overstated in the past. Future research challenges include determining whether the richness–climate relationship reflects: (1) a direct relationship through the physiological tolerance of the plants, (2) an indirect influence of climate on ecosystem productivity, or (3) an evolutionary legacy of longer or faster diversification processes under certain climatic conditions.     

Patterns of ferns community assemblages in some Malaysian and Nigerian tropical forests

Research on fern ecology has gained attention in the last decade, yet there is a paucity of information on the comparison of ferns communities across continents. This study focused on comparing the ferns community assemblages in some tropical forests of Malaysia and Nigeria, thereby assessing the patterns of species richness (SR) and phylogenetic diversity (PD) in relation to the bioclimatic drivers across the continents. The diversity and taxonomic compositions of ferns were assessed using 180 plots of 10 m × 10 m in each country. The species richness and other diversity indices were determined using the combined forests data for each country and for the individual forests. Also, the phylogenetic diversity of the ferns was assessed using the genus‐based molecular sequences downloaded from the GeneBank. The patterns of the ferns SR and PD in the two countries as driven by some bioclimatic factors were evaluated using the regression analysis. The observed and rarefied–extrapolated fern species richness is significantly higher in Malaysian forests than in Nigerian forests. Also, the other diversity indices are significantly higher in Malaysian forests except for the Shannon index which showed no significant difference between the two biogeographic regions. There is a very low similarity (7.41%) in the taxonomic composition of ferns between the two biogeographic areas, although the similarity in composition increased with increasing taxonomic levels (species: 7.41%, genus: 12.77%, family: 70.96%). Terrestrial and epiphytic ferns are more dominant than the other life forms in the two countries. The precipitation variables drive the phylogenetic structure of ferns in Nigeria, whereas both precipitation and temperature variables drive the phylogenetic structure of ferns in Malaysia. This indicates that ferns assemblages in Nigeria and Malaysia are driven by both climatic variables. Besides, we also hypothesize that these observed differences could be due to other historical and evolutionary processes.     

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。     

Stasis and convergence characterize morphological evolution in eupolypod II ferns

Background and Aims

Patterns of morphological evolution at levels above family rank remain underexplored in the ferns. The present study seeks to address this gap through analysis of 79 morphological characters for 81 taxa, including representatives of all ten families of eupolypod II ferns. Recent molecular phylogenetic studies demonstrate that the evolution of the large eupolypod II clade (which includes nearly one-third of extant fern species) features unexpected patterns. The traditional ‘athyrioid’ ferns are scattered across the phylogeny despite their apparent morphological cohesiveness, and mixed among these seemingly conservative taxa are morphologically dissimilar groups that lack any obvious features uniting them with their relatives. Maximum-likelihood and maximum-parsimony character optimizations are used to determine characters that unite the seemingly disparate groups, and to test whether the polyphyly of the traditional athyrioid ferns is due to evolutionary stasis (symplesiomorphy) or convergent evolution. The major events in eupolypod II character evolution are reviewed, and character and character state concepts are reappraised, as a basis for further inquiries into fern morphology.

Methods

Characters were scored from the literature, live plants and herbarium specimens, and optimized using maximum-parsimony and maximum-likelihood, onto a highly supported topology derived from maximum-likelihood and Bayesian analysis of molecular data. Phylogenetic signal of characters were tested for using randomization methods and fitdiscrete.

Key Results

The majority of character state changes within the eupolypod II phylogeny occur at the family level or above. Relative branch lengths for the morphological data resemble those from molecular data and fit an ancient rapid radiation model (long branches subtended by very short backbone internodes), with few characters uniting the morphologically disparate clades. The traditional athyrioid ferns were circumscribed based upon a combination of symplesiomorphic and homoplastic characters. Petiole vasculature consisting of two bundles is ancestral for eupolypods II and a synapomorphy for eupolypods II under deltran optimization. Sori restricted to one side of the vein defines the recently recognized clade comprising Rhachidosoraceae through Aspleniaceae, and sori present on both sides of the vein is a synapomorphy for the Athyriaceae sensu stricto. The results indicate that a chromosome base number of x =41 is synapomorphic for all eupolypods, a clade that includes over two-thirds of extant fern species.

Conclusions

The integrated approach synthesizes morphological studies with current phylogenetic hypotheses and provides explicit statements of character evolution in the eupolypod II fern families. Strong character support is found for previously recognized clades, whereas few characters support previously unrecognized clades. Sorus position appears to be less complicated than previously hypothesized, and linear sori restricted to one side of the vein support the clade comprising Aspleniaceae, Diplaziopsidaceae, Hemidictyaceae and Rachidosoraceae – a lineage only recently identified. Despite x =41 being a frequent number among extant species, to our knowledge it has not previously been demonstrated as the ancestral state. This is the first synapomorphy proposed for the eupolypod clade, a lineage comprising 67 % of extant fern species. This study provides some of the first hypotheses of character evolution at the family level and above in light of recent phylogenetic results, and promotes further study in an area that remains open for original observation.     

Autochory in ferns,not all spores are blown with the wind

Dispersal is a key process in plant population dynamics. In ferns, two successive vectors are needed: the sporangium catapulting mechanism, and wind or gravity. However, some rock ferns have a growth habit that suggests a kind of autochory by placing spores on the rock surface. Moreover, some ferns show modifications of the sporangial dehiscence. To determine the role of growth habit in spore dispersal, we checked the sporangial opening mechanism and explored the spatial distribution of plants on the walls. The presence of spores of Asplenium celtibericum, a rupicolous fern, in the rock surface was checked. In addition, its sporangial dehiscence, plant size and position in the wall were analysed. Spores and indehiscent sporangia were present on walls at each sampling moment. Their highest number was found close to the plants. There was a positive correlation between crack width and plant size. However, most plants occupy the upper half of the cliffs. The growth habit of A. celtibericum is instrumental to deposit the spores over the neighbouring rock surface, thus enhancing the probability of spores to find suitable crevices for germination. Furthermore, dispersal of indehiscent sporangia might promote intergametophytic mating, and the modified sporangial opening mechanism extends the dispersive period.     

金佛山自然保护区蕨类植物资源与区系初步研究

金佛山自然保护区共计有野生蕨类植物46科107属458种(含亚种和变种)。在科的区系成分中单种科14个,少型科(2~5种)16个,多种科(包含6~19种)12个,大科(20~49种)1个,特大科(50种以上)3个,分别占金佛山蕨类植物总科数的30.43%、34.78%、26.09%、2.17%和6.53%;其中热带分布占总科数的54.35%;在属的区系成分中,具12个分布区类型,单种属和少种属数量多;热带成分丰富,包含55属,占总属数的51.40%;特有现象明显,地方特有种十分丰富;属的区系与武陵山和大巴山关系最为密切,与九华山、化龙山、齐云山和云山关系疏远;东亚分布、世界广布、热带亚洲至热带非洲间断分布及北温带分布型对该区系性质的影响最大,热带亚洲分布型和东亚北美间断分布型的影响最小;该区系具有古老、孑遗和原始性等特点,验证了金佛山地区处于我国三大特有现象中心之一的鄂西—川东特有现象中心的观点。