Molecular evidence for long-distance dispersal in the New Zealand pteridophyte flora

Aim To examine the relative importance of long‐distance dispersal in shaping the New Zealand pteridophyte (ferns and lycophytes) flora and its relationships with other floras, with the null hypothesis that the extant New Zealand pteridophyte flora has been isolated since New Zealand’s separation from Gondwana. Location New Zealand. Methods rbcL DNA sequences were assembled for 31 New Zealand pteridophyte genera, with each genus represented by one New Zealand species and the most closely related non‐New Zealand species for which data were available. Maximum‐likelihood, maximum‐parsimony, and Bayesian analysis phylograms were constructed and used as input for r 8s molecular dating, along with 23 fossil calibrations. Divergence estimates less than conservatively recent ages for New Zealand’s geological isolation, namely H_o > 30 Ma for pairs involving New Caledonian and Norfolk Island species and H_o > 55 Ma for all others, were taken as rejection of the null hypothesis. Results The null hypothesis was rejected for all pairs except, under some parameter conditions, for those involving the New Zealand species Cardiomanes reniforme, Lindsaea trichomanoides, Loxsoma cunninghamii, Lygodium articulatum, Marattia salicina, and Pteris comans. However, the Lindsaea and Pteris results probably reflect the absence in the analyses of closely related non‐New Zealand samples, while the Marattia divergence was highly contingent on which fossil calibrations were used. Main conclusions Rejection of the null hypothesis for the majority of pairs implies that the extant New Zealand lineage has undergone long‐distance dispersal either into or out of New Zealand. The notion of a long isolation since geological separation can, therefore, be dismissed for much of New Zealand’s pteridophyte flora. The analyses do not identify the direction of the long‐distance dispersal, and these New Zealand lineages could have had vicariant origins with subsequent long‐distance emigration. However, the alternative that many extant New Zealand pteridophyte lineages only arrived by long‐distance immigration after geological isolation seems likely.     

New Zealand's pteridophyte flora—Plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes arrived in New Zealand relatively recently, by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical regions, 14% with temperate southern Africa and 13% with the circum-Antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area, and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall, and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism than elsewhere and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that at least some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques.     

Multiple intercontinental dispersals shaped the distribution area of Hordeum (Poaceae)

The grass genus Hordeum (Poaceae, Triticeae), comprising 31 species distributed in temperate and dry regions of the world, was analysed to determine the relative contributions of vicariance and long-distance dispersal to the extant distribution pattern of the genus. Sequences from three nuclear regions (DMC1, EF-G and ITS) were combined and analysed phylogenetically for all diploid (20 species) and two tetraploid Hordeum species and the outgroup Psathyrostachys. Ages of clades within Hordeum were estimated using a penalized likelihood analysis of sequence divergence. The sequence data resulted in an almost fully resolved phylogenetic tree that allowed the reconstruction of intrageneric migration routes. Hordeum evolved c. 12 million years ago in South-west Asia and spread into Europe and Central Asia. The colonization of the New World and South Africa involved at least six intercontinental exchanges during the last 4 million years (twice Eurasia-North America, North America-South America, twice South America-North America and Europe-South Africa). Repeated long-distance dispersal between the northern and southern hemisphere were important colonization mechanisms in Hordeum.     

Multi-extent analysis of the relationship between pteridophyte species richness and climate

Aim To determine the relationship between the distribution of climate, climatic heterogeneity and pteridophyte species richness gradients in Australia, using an approach that does not assume potential relationships are spatially invariant and allows for scale effects (extent of analysis) to be explicitly examined. Location Australia, extending from 10° S to 43° S and 112° E to 153° E. Method Species richness within 50 × 50 km grid cells was determined using point distribution data. Climatic surfaces representing the distribution and availability of water and energy at 1 km and 5 km cell resolutions were obtained. Climate at the 50 km resolution of analysis was represented by their mean and standard deviation in that area. Relationships were assessed using geographically weighted linear regression at a range of spatial bandwidths to investigate scale effects. Results The parameters and the predictive strength of all models varied across space at all extents of analysis. Overall, climatic variables representing water availability were more highly correlated to pteridophyte richness gradients in Australia than those representing energy. Their variance in cells further increased the strength of the relationships in topographically heterogeneous regions. Relationships with water were strong across all extents of analysis, particularly in the tropical and subtropical parts of the continent. Water availability explained less of the variation in richness at higher latitudes. Main conclusions This study brings into question the ability of aspatial and single‐extent models, searching for a unified explanation of macro‐scaled patterns in gradients of diversity, to adequately represent reality. It showed that, across Australia, there is a positive relationship between pteridophyte species richness and water availability but the strength and nature of the relationship varies spatially with scale in a highly complex manner. The spatial variance, or actual complexity, in these relationships could not have been demonstrated had a traditional aspatial global regression approach been used. Regional scale variation in relationships may be at least as important as more general relationships for a true understanding of the distribution of broad‐scale diversity.     

Overseas seed dispersal by migratory birds

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.     

Phytogeographic relationships between neotropical and African-Madagascan pteridophytes

The purpose of this study is to determine the floristic affinities of pteridophytes between the neotropics and Africa-Madagascar and examine how these affinities might have arisen. We present an annotated list that contains two kinds of affinities: 1) species in common between both regions (excluding pantropical species) and 2) species pairs (or clusters of species paris) where one of the species (or infrageneric group) occurs in the Neotropics and the other in Africa and/or Madagascar. There are 114 examples on the list, of which 27 are same-species and 87 are species pairs or closely related taxa at some infrageneric level. About 13% of the African pteridoflora and 14% of the Madagascan pteridoflora show affinities with the Neotropics. To determine how these similarities might have originated, we assess three hypotheses: 1) the boreotropics hypothesis, 2) continental drift, and 3) long-distance dispersal. The boreotropics hypothesis is difficult to assess without further phylogenetic information on the groups to which the species belong. Continental drift seems to best explain one example in the geologically old family Schizaeaceae (species inAnemia subgen.Coptophyllum sect.Tomentosae). Nearly all the other examples seem best explained by long-distance dispersal because they belong to families that first appeared during the Paleocene, more than 30 million yearsafter drift had effectively separated South America and Africa. Most of the dispersal events appear to have taken place from the neotropics to Africa-Madagascar, but recent African extinctions may have obscured directionality. Species with green spores or gemmiferous gametophytes were slightly overrepresented on the list compared to pteridophytes as a whole.     

Mediterranean island biogeography: Analysis of fern species distribution in the system of islets around Sicily

Abstract

The aim of this article was to test the way in which geographical factors influence island floras in the Mediterranean basin, using ferns as target organisms, and the islands surrounding Sicily as location. A matrix with presence/absence data concerning fern taxa in the 16 islands studied was compiled. Cluster analysis, principal co-ordinates analysis (PCoA), principal components analysis (PCA) and a Bayesian analysis were performed. For each island, the total number of fern taxa was regressed against three factors: island area, island elevation and isolation. All the analyses pointed to affinities between islands according to their different geological composition, independently from their geographic position. A clear positive island species/area relationship (ISAR) was shown only for the volcanic islands. The island species/(area×elevation) relationship (ISAER), on the contrary, was unsatisfactory. The main features of interest are the following: (1) the clear division of the islands into two groups, volcanic vs. sedimentary; (2) the floristic richness of the volcanic compared to sedimentary islands and (3) the uniqueness of the pteridophyte flora of Pantelleria. This seems to demonstrate that the lower number of taxa in the islands farthest away from the “mainland” (Sicily, Tunisia) is not due to isolation, but due to another factor, probably habitat availability.     

Phylogenetic tests of distribution patterns in South Asia: towards an integrative approach

The last four decades have seen an increasing integration of phylogenetics and biogeography. However, a dearth of phylogenetic studies has precluded such biogeographic analyses in South Asia until recently. Noting the increase in phylogenetic research and interest in phylogenetic biogeography in the region, we outline an integrative framework for studying taxon distribution patterns. While doing so, we pay particular attention to challenges posed by the complex geological and ecological history of the region, and the differences in distribution across taxonomic groups. We outline and compare three widely used phylogenetic biogeographic approaches: topology-based methods (TBMs), pattern-based methods (PBMs) and event-based methods (EBMs). TBMs lack a quantitative framework and utilize only part of the available phylogenetic information. Hence, they are mainly suited for preliminary enquiries. Both PBMs and EBMs have a quantitative framework, but we consider the latter to be particularly suited to the South Asian context since they consider multiple biogeographic processes explicitly, and can accommodate a reticulated history of areas. As an illustration, we present a biogeographic analysis of endemic Sri Lankan agamid lizards. The results provide insights into the relative importance of multiple processes and specific zones in the radiation of two speciose lizard clades.     

Geographic distributions of homosporous ferns: does dispersal obscure evidence of vicariance?

The central problem in biogeography is that interactions between different processes result in the formation of historical patterns, such that it is difficult to discriminate the relative roles of vicariance and dispersal. Ferns are distributed by small wind-dispersed propagules that are produced in very large numbers and capable of dispersing thousands of kilometers. Thus, most taxon distributions in ferns are assumed to be a function of dispersal rather than vicariance. Here, we review some case examples that provide good evidence for vicariance and dispersal in ferns. We then ask whether dispersal is so extensive in ferns that vicariance is no longer detectable in most cases. Although we think that too few studies have been carried out to make generalizations at this stage, we outline the criteria for an effective research programme that can address this issue. Phylogenetic and distributional data are needed, not only because they are lacking in an evolutionarily important group of organisms, but also because data from ferns and other cryptogams are likely to be crucial in making broad biogeographic statements.     

东南亚生物多样性热点地区的形成与演化

东南亚地处热带,生物多样性极为丰富,可分为4个热点地区:印度-缅甸区的中南半岛、巽他区(含马来半岛、婆罗洲、苏门答腊岛)、菲律宾区(菲律宾群岛)、华莱士区(苏拉威西岛、爪哇岛、马鲁古群岛、小巽他群岛等)。中南半岛在泥盆纪便已是欧亚大陆的一部分,在印度板块撞击欧亚大陆之后受挤压而出;巽他区来自于冈瓦纳古陆和澳洲古陆;菲律宾群岛部分来自于劳亚古陆的碎片向南漂移,部分来自于太平洋西南岛弧的向北迁移;华莱士区则是劳亚古陆碎片、太平洋西南岛弧以及澳洲古陆北侧碎片的组合。巽他区地处赤道,常年温湿;菲律宾区、华莱士区、中南半岛则都受到不同程度的季风气候决定的干湿季变动。地质历史和季风气候影响程度的不同,奠定了东南亚4个生物多样性热点地区的雏形。华莱士区保存有大量的早期被子植物原始类群如睡莲目(Nymphaeales)和木兰藤目(Austrobaileyales),是现代被子植物起源地和冰期避难所之一。巽他区(婆罗洲)和中南半岛是亚洲热带植物的现代分布中心和"进化前沿",是整个东南亚地区重要的种源;而华莱士区的爪哇岛和小巽他群岛主要是物种迁入和中转的种库。这样的物种形成历史与迁移格局,塑造了东南亚4个...     

Estimates of the number of mounted pteridophyte specimens in six major herbaria in the United States

A stratified-random sampling method was used to estimate the number of pteridophyte specimens in six of the largest herbaria in the United states (F, GH/A/NEBC, MO, NY, UC, US). The point estimate of the collection size and the 95% confidence intervals for each herbarium were found to be as follows: US 251,000 (±20,000); NY 201,000 (±20,000); MO 160,000 (±15,000); GH/A/NEBC 124,000 (±12,000); UC 119,000 (±9,000); F 97,000 (±10,000). It is hoped that these numbers will help assess the growth of these collections in the future.     

Transoceanic dispersal of terrestrial species by debris rafting

Rare, long-distance dispersal events are a key process in generating and maintaining patterns in biological diversity and species distributions across space and time. The 9.0 magnitude earthquake that struck the eastern coast of Japan in 2011, and the subsequent 38 m high tsunami washed large amounts of shoreline debris into the Pacific Ocean that led to a large-scale biological rafting event carrying nearly 300 marine species to the western shores of North America. Whether oceanic, trans-Pacific dispersal via rafting generates long distance dispersal events for small, flightless, terrestrial species is unknown. By sampling beach debris associated with known hot-spots of tsunami debris along the north and east shores of Graham Island, Haida Gwaii, Canada, I document significantly dissimilar invertebrate communities associated with tide-line beach debris and the occurrence of several putative Japanese species of soil-dwelling mites (Acari: Oribatida). Previous explanations of Haida Gwaii's unique flora and fauna have been attributed to a proximity to the Beringian land bridge and the accumulated evidence of near-offshore glacial refugia during the last glacial period. However, my research also suggests that stochastic, trans-Pacific rafting events contribute to the biodiversity and biogeography of soil communities on the west coast of North America.     

中国石松类和蕨类植物的多样性与地理分布

物种编目及其科属系统排列是了解生物多样性的基础, 本文采用Flora of China使用的分类系统, 结合最新分子分类学研究成果以及近几年发表的新资料, 对中国石松类和蕨类植物多样性和地理分布数据进行了统计和分析。结果表明中国共有石松类和蕨类植物40科178属2,147种5个亚种118个变种, 其中特有种839个, 占总种数的39.08%。种数最多的5个科依次为鳞毛蕨科(505种, 含种下单位, 下同)、蹄盖蕨科(323种)、水龙骨科(280种)、凤尾蕨科(266种)和金星蕨科(209种); 种数最多的5个属依次为耳蕨属(Polystichum, 209种)、鳞毛蕨属(Dryopteris, 176种)、蹄盖蕨属(Athyrium, 137种)、双盖蕨属(Diplazium, 98种)和凤尾蕨属(Pteris, 97种)。在地理分布上, 种数排名前5的省份为云南(1,365种)、四川(875种)、贵州(838种)、广西(785种)和台湾(779种)。含中国特有石松类和蕨类植物的科属中, 排前3位的科分别为鳞毛蕨科(257种)、蹄盖蕨科(169种)和凤尾蕨科(113种); 排前3位的属为耳蕨属(140种)、蹄盖蕨属(82种)和鳞毛蕨属(61种)。     

西藏蕨类植物增补(II)(英文)

报道了 8种蕨类植物为西藏新记录 ,并讨论了西藏蕨类植物的区系特点和分类问题     

Inverted patterns of genetic diversity in continental and island populations of the heather Erica scoparia s.l.

Aim Using the heather Erica scoparia s.l. as a model, this paper aims to test theoretical predictions that island populations are genetically less diverse than continental ones and to determine the extent to which island and continental populations are connected by pollen‐ and seed‐mediated gene flow. Location Macaronesia, Mediterranean, Atlantic fringe of Europe. Methods Patterns of genetic diversity are described based on variation at two chloroplast DNA (cpDNA) loci and one nuclear DNA (nDNA) locus for 109 accessions across the entire distribution range of the species. Global patterns of genetic differentiation were investigated using principal coordinates analysis. Genetic differentiation between island and continental areas, estimations of pollen‐ and seed‐mediated gene flow, and the presence of phylogeographical signal were assessed by means of F_st /N_ST (continental scale) and F_ij/N_ij (local scale). Extant and past distribution ranges of the species were inferred from niche modelling using layers describing present and Last Glacial Maximum (LGM) macroclimatic conditions. Results The Azores exhibited a significantly higher genetic diversity than the continent. The lowest levels of genetic differentiation were observed between the Azores and the western Mediterranean, and the diversity observed in the Azores resulted from at least two colonization waves. Within the Azores, kinship coefficients showed a significant and much steeper decrease with geographical distance in the cpDNA than in the nDNA. The distribution predicted by LGM models was markedly different from the current potential distribution, particularly in western Europe, where no suitable areas were predicted by LGM models, and along the Atlantic coast of the African continent, where LGM models predicted highly suitable climatic conditions. Main conclusions The higher diversity observed in Azorean than in continental populations is inconsistent with MacArthur and Wilson’s equilibrium model and derived theoretical population genetic expectations. This inverted pattern may be the result of extinction on the continent coupled with multiple island colonization events and subsequent allopatric diversification and lineage hybridization in the Azores. The results highlight the role of allopatric diversification in explaining diversification on islands and suggest that this process has played a much more significant role in shaping Azorean biodiversity than previously thought.     

Phylogenetic and biosystematic relationships in four highly disjunct polyploid complexes in the subgenera Ceterach and Phyllitis in Asplenium (Aspleniaceae)

Phylogenetic studies using DNA sequences of two chloroplast regions, rbcL and trnL-F, demonstrate that the proposed genus Ceterach is a small clade within the large genus Asplenium, and sister to the Phyllitis clade. The Ceterach clade is characterised by irregular anastomosing veins and often densely scaled leaf blades. Its taxonomic status as a group nested within Asplenium is confirmed, and it is accepted here as a subgenus with seven species. The Ceterach clade comprises four lineages that correspond to disjunct polyploid complexes: the A. aureum clade forming a polyploid complex (4×, 6×, 8×) in Macaronesia, the A. ceterach clade forming a polyploid complex (2×, 4×, 6×) in the Mediterranean Basin, the A. paucivenosum clade (4×, 6×) in central Asia, and the A. dalhousiae clade (2×) with a disjunct distribution in the Himalaya, Yemen and Eritrea, and southwestern North America. Asplenium paucivenosum is sister to all other members of the Ceterach clade, whereas A. dalhousiae is sister to the A. aureum clade that includes tetraploid A. aureum, hexaploid A. lolegnamense, and octoploid A. parvifolium. Asplenium ceterach and its variations – including the hexaploid A. ceterach subsp. mediterraneum subsp. nov. first described below – form a monophyletic unit, sister to a clade consisting of A. aureum and A. dalhousiae. Asplenium cordatum from Africa and A. haugthonii from the isolated atlantic island of St. Helena are not members of the Ceterach clade, which suggests that leaf blades with dense indumenta have evolved at least twice within asplenioid ferns. The allotetraploid species A. hybridum has the chloroplast DNA from A. ceterach, and therefore the latter species is the maternal ancestor of the former. The other parent of this hybrid species is A. sagittatum that is nested within the sister clade of Ceterach, the Phyllitis clade comprising A. sagittatum and A. scolopendrium. The findings suggest that the current distribution of Ceterach is either the result of long-distance dispersal or represents fragmented relicts of a previously more widely distributed species.     

Costs and benefits of non‐random seed release for long‐distance dispersal in wind‐dispersed plant species

The dispersal ability of plants is a major factor driving ecological responses to global change. In wind‐dispersed plant species, non‐random seed release in relation to wind speeds has been identified as a major determinant of dispersal distances. However, little information is available about the costs and benefits of non‐random abscission and the consequences of timing for dispersal distances. We asked: 1) to what extent is non‐random abscission able to promote long‐distance dispersal and what is the effect of potentially increased pre‐dispersal risk costs? 2) Which meteorological factors and respective timescales are important for maximizing dispersal? These questions were addressed by combining a mechanistic modelling approach and field data collection for herbaceous wind‐dispersed species. Model optimization with a dynamic dispersal approach using measured hourly wind speed showed that plants can increase long‐distance dispersal by developing a hard wind speed threshold below which no seeds are released. At the same time, increased risk costs limit the possibilities for dispersal distance gain and reduce the optimum level of the wind speed threshold, in our case (under representative Dutch meteorological conditions) to a threshold of 5–6 m s^–1. The frequency and predictability (auto‐correlation in time) of pre‐dispersal seed‐loss had a major impact on optimal non‐random abscission functions and resulting dispersal distances. We observed a similar, but more gradual, bias towards higher wind speeds in six out of seven wind‐dispersed species under natural conditions. This confirmed that non‐random abscission exists in many species and that, under local Dutch meteorological conditions, abscission was biased towards winds exceeding 5–6 m s^–1. We conclude that timing of seed release can vastly enhance dispersal distances in wind‐dispersed species, but increased risk costs may greatly limit the benefits of selecting wind conditions for long‐distance dispersal, leading to moderate seed abscission thresholds, depending on local meteorological conditions and disturbances.