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.     

Comparison of molecular and morphological data on St Helena: Elaphoglossum

The endemic elaphoglossoid ferns, Elaphoglossum dimorphum, E. nervosum and Microstaphyla furcata of St Helena, form a closely related group within section Lepidoglossa when analysed phylogenetically using sequences from the chloroplast trnL intron (partial) and trnL-F intergenic spacer. Microstaphyla furcata, traditionally placed in its own genus, is clearly shown to belong to Elaphoglossum confirming the previous transfer of this species to Elaphoglossum as E. bifurcatum. There is hardly any trnL-F sequence divergence between the species, in fact sequences of E. nervosum and E. dimorphum are identical. These results are consistent with the possible origin of E. dimorphum as a hybrid between E. bifurcatum and E. nervosum or with the view that the three species are the result of a recent radiation. The potential conflict between phylogenetic and morphological distinctness in determining species conservation priorities is discussed.     

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。     

浙江省2种蕨类植物新记录

报道浙江蕨类植物2种新记录,即假耳羽短肠蕨Diplazium okudairai Makino、大盖铁角蕨Asplenium bullatum Wall.ex Mett.。凭证标本存于景宁畲族自治县自然资源和规划局(林业局)标本室。     

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.     

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。     

The Rhaetian flora of Rögla,northern Scania,Sweden

Abstract: Rögla is the northernmost locality yielding Mesozoic plant fossils in Scania, southern Sweden, and is one of the northernmost Rhaetian assemblages in Europe. The assemblage consists of over 500 specimens collected 50–60 years ago, of which 139 yielded identifiable plant remains referable to 15 plant species; another 19 specimens are tentatively assigned to four species because of their fragmentary preservation. The flora includes sphenophytes, ferns, cycads, bennettitaleans, seed ferns of uncertain alliance, conifers and some leaf remains that are tentatively assigned to ginkgophytes based on their epidermal anatomy. The species‐level composition of the assemblage is consistent with a Rhaetian age and is similar to well‐known floras from nearby Höganäs and Bjuv, except for the absence of cycads belonging to Nilssonia, which are very common in most other Scanian floras. The fossil assemblage is interpreted to derive from multi‐storey vegetation occupying moist habitats on a coastal plain. Strong affinities are evident with the coeval floras of Jameson Land, Greenland, reinforcing the concept of a distinctive North Atlantic floristic sub‐province at the close of the Triassic.     

云南哀牢山国家级自然保护区附生蕨类植物的多样性与附载植物的关系

为了探讨附生蕨类植物多样性与附载植物的关系,调查了附载植物胸径、树高、枝下高及附载植物种类对附生蕨类植物多样性的影响.按照径级分别对不周径级下的附载植物的附生蕨类植物的多样性进行偏相关分析,并采用方差分析判断附载植物种类对附生蕨类植物多样性的影响.结果表明,附载植物树高和胸径与附生蕨类植物多样性之间存在明显正相关,附载植物的枝下高与附生蕨类植物多样性之间存在着明显的负相关,表明较低的枝下高有利于附生蕨类植物多样性的增加.附载植物种类对附生蕨类植物多样性具有重要的影响.     

The effect of area on local and regional elevational patterns of species richness

Aim To calculate the degree to which differences between local and regional elevational species richness patterns can be accounted for by the effects of regional area. Location Five elevational transects in Costa Rica, Ecuador, La Réunion, Mexico and Tanzania. Methods We sampled ferns in standardized field plots and collated regional species lists based on herbarium and literature data. We then used the Arrhenius function S = cA^z to correct regional species richness (S) for the effect of area (A) using three slightly different approaches, and compared the concordance of local and regional patterns prior to and after accounting for the effect of area on regional richness using linear regression analyses. Results We found a better concordance between local and regional elevational species richness after including the effect of area in the majority of cases. In several cases, local and regional patterns are very similar after accounting for area. In most of the cases, the maximum regional richness shifted to a higher elevation after accounting for area. Different approaches to correct for area resulted in qualitatively similar results. Main conclusions The differences between local and regional elevational richness patterns can at least partly be accounted for by area effects, suggesting that the underlying causes of elevational richness patterns might be the same at both spatial scales. Values used to account for the effect of area differ among the different study locations, showing that there is no generally applicable elevational species–area relationship.