Evolution of Inbreeding and Outcrossing in Ferns and Fern-Allies

Abstract Mating systems of 18 species of homosporous ferns follow a bimodal distribution, similar to that observed for seed plants (Schemske and Lande, 1985). Most species are highly outcrossing, a few are inbreeding, and two species examined to date have mixed mating systems. Equisetum arvense and several species of lycopods are also highly outcrossing. Several mechanisms, including inbreeding depression, antheridiogen, and ontogenetic sequences that result in effectively unisexual gametophytes, promote outcrossing in homosporous ferns and perhaps other homosporous pteridophytes as well. In some species of homosporous ferns, selection has favored the evolution of inbreeding as an adaptation for colonization. High levels of intra- and interpopulational gene flow via spore dispersal, coupled with high levels of intergametophytic crossing, generally lead to genetically homogeneous populations and species of homosporous ferns. However, rock-dwelling ferns and ferns from xeric habitats may exhibit significant population genetic structure due to physically patchy habitats. Reticulate evolution in homosporous ferns may be enhanced by high levels of intergametophytic crossing.     

Trends for Inbreeding in Polyploid Pteridophytes

Abstract Previous studies on the sequence of the gametangium formation in homosporous fern gametophytes (Masuyama, 1975a, b) suggested that diploid and polyploid taxa may favor gametophytic crossing and gametophytic selfing, respectively. In this view, intraspecific polyploids of three fern species ( Phegopteris decursive-pinnata, Lepisorus thunbergianus and Pteris dispar ) have been investigated for the amount of genetic load or the rate of gametophytic selfing. As expected, a marked contrast of mating systems is found between diploids and tetraploids; the former predominantly undergo gametophytic crossing and the latter predominantly undergo gametophytic selfing. It seems likely that diploid homosporous pteridophytes favor gametophytic crossing with some exceptions favoring gametophytic selfing as a derived condition, whereas polyploid homosporous pteridophytes favor gametophytic selfing predominantly.     

Genetic differentiation in populations of the polymorphic fernCeratopteris thalictroides in Japan

Analysis of isozyme variation was carried out for 27 natural populations ofCeratopteris thalictroides in Japan. Of fifteen enzyme loci examined, eight loci were genetically polymorphic. At six loci,Lap, Pgi-2, Pgm-3, Pgm-4, Idh-2, and Skd-2, a marked genetic differentiation was observed between populations to the south of Okinawa Island and those to the north of the island. Okinawa Island contained a mixture of both southern and northern variants. Thus, two genetically distinct types (the south type and the north type) ofC. thalictroides occur allopatrically in Japan. Nei's genetic identity (I) between the two was 0.64, which was within the range of the I values between congeneric pteridophyte species. Regional fixation of a null allele was detected for one duplicated PGI locus in the north type ofC. thalictroides. This finding supports the recent hypothesis of genetic diploidization of polyploids through gene silencing.     

Large-scale diversity patterns of vascular epiphytes in Neotropical montane rain forests

Aim Epiphytes contribute up to 30% to the number of vascular plant species in certain global biodiversity hotspots, e.g. the Ecuadorian Andes. However, their large scale diversity patterns are still discussed on the base of results from a few, local epiphyte inventories. Consequently, explanatory models on epiphyte diversity concentrate on the impact of local climate on small scale epiphyte species richness. Our aim was to analyse large scale elevational patterns of epiphyte diversity integrating data from different geographic scales. Location Tropical America, with special emphasis on the Ecuadorian Andes. Methods Our study is based on two data sources. First, we analysed the elevational patterns of epiphyte diversity based on the Catalogue of the Vascular Plants of Ecuador and the Libro Rojo de las Plantas Endèmicas del Ecuador. Secondly, the floristic turnover between the epiphyte inventories of seven montane and four lowland study sites in the Neotropics was analysed. Results The floristic turnover between Neotropical montane epiphyte floras is higher than the one between lowland epiphyte floras. Montane study sites located only a few kilometres apart from each other show considerable differences in their epiphyte species inventories. Irrespectively of their similar dispersal mode, the floristic turnover is much higher for orchids than for Pteridophyta. The Orchidaceae are the species richest group in all of the examined 11 Neotropical epiphyte floras. At the larger scale of the Ecuadorian Flora, c. 50% of the species in the elevational zone with maximum epiphyte diversity (between 1000 and 1500 m) are orchids. Elevational patterns of epiphyte diversity strongly reflect patterns of Orchidaceae. Main conclusions Our results support the observation of a ‘mid‐elevation bulge’ of epiphyte diversity by Gentry and Dodson. It has been frequently shown that the high humidity in mid‐elevations is suitable to maintan a high epiphyte species richness. Our findings show that in addition, large scale epiphyte diversity in montane rain forest is increased by the high floristic turnover at local and regional scale. Based on the importance of Orchidaceae for epiphyte diversity, we discuss that speciation processes corresponding to the highly diverse environment are a driving force for endemism, floristic heterogeneity and consequently for large scale epiphyte species richness in montane forests.     

Notes on the Scottish Flora

Summary

Pollen analytical investigations are described from a site on the Pass of Drumochter in the central Grampian Highlands. Three pollen assemblage zones were identified in the profile which spans the mid- and late-Postglacial periods. The earliest vegetational record is of birch and hazel woodland, which was in turn succeeded by the establishment of pine forest, possibly around 7000 B.P. Declining arboreal pollen percentages and increased values for open-habitat taxa in the upper reaches of the profile reflect the impact of Neolithic man in the area, with the destruction of the forests, and the spread of dwarf shrub heath and heather moor across formerly wooded hillsides.     

A survey of the mycorrhization of Southeast Asian ferns and lycophytes

The colonisation of land by plants may not have been possible without mycorrhizae, which supply the majority of land plants with nutrients, water and other benefits. In this sense, the mycorrhization of basal groups of land plants such as ferns and lycophytes is of particular interest, yet only about 9% of fern and lycophyte species have been sampled for their mycorrhization status, and no community‐level analyses exist for tropical fern communities. In the present study, we screened 170 specimens of ferns and lycophytes from Malaysia and Sulawesi (Indonesia), representing 126 species, and report the mycorrhization status for 109 species and 19 genera for the first time. Mycorrhizal colonisations were detected in 96 (56.5%) of the specimens, 85 of which corresponded to arbuscular mycorrhizae (AMF), three to dark‐septate endophytes (DSE) and four to mixed colonisations (AMF + DSE). DSE colonisations were lower than in comparable samples of ferns from the Andes, suggesting a geographical or taxonomic pattern in this type of colonisation. Epiphytes had significantly lower levels of colonisation (26.1%) than terrestrial plants (70.7%), probably due to the difficulty of establishment of mycorrhizal fungi in the canopy habitat.     

The occurrence and morphology of Adiantum X variopinnatum (Pteridaceae)

Two populations ofAdiantum Xvariopinnatum were found at the La Selva Biological Field Station in Costa Rica. The plants were morphologically intermediate between their parents, showed additive isozyme banding patterns, and had aborted spores. Herbarium searches for the hybrid turned up specimens from Nicaragua, Costa Rica, Panama, Venezuela, and Colombia. These are the first reports of the hybrid outside of Trinidad.     

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     

中国蕨类植物细胞分类学研究概况

中国蕨类植物类群十分丰富,由63科230属约2600种组成,其中约有10％的种类为中国特有。迄今为止,中国蕨类植物进行过染色体计数的种类约有395种,仅占中国蕨类植物总数的15％,其中48％为多倍体。本文对中国蕨类植物细胞分类学研究的历史和现状进行了综合评述。文中论述了蕨类植物染色体数目和染色体基数在系统学上的应用以及蕨类植物染色体组型分析、多倍化、无融合生殖、多倍体复合体及网状进化的研究概况。中国20多年的蕨类植物细胞分类学研究表明,蕨类植物的染色体数目、基数、组型分析、多倍化及其繁殖方式对蕨类植物的分类、起源和演化的研究都有重要作用,对种及种复合体的鉴定、科属的划分也有重要的参考价值。并对中国蕨类植物细胞分类学研究提出了一些建议。