Morphology informed by phylogeny reveals unexpected patterns of species differentiation in the aquatic moss Rhynchostegium riparioides s.l

Bryophyte floras typically exhibit extremely low levels of endemism. The interpretation, that this might reflect taxonomic shortcomings, is tested here for the Macaronesian flora, using the moss species complex of Rhynchostegium riparioides as a model. The deep polyphyly of R. riparioides across its distribution range reveals active differentiation that better corresponds to geographic than morphological differences. Morphometric analyses are, in fact, blurred by a size gradient that accounts for 80% of the variation observed among gametophytic traits. The lack of endemic diversification observed in R. riparioides in Macaronesia weakens the idea that the low rates of endemism observed in the Macaronesian bryophyte flora might solely be explained by taxonomic shortcomings. To the reverse, the striking polyphyly of North American and European lineages of R. riparioides suggests that the similarity between the floras of these continents has been over-emphasized. Discriminant analyses point to the existence of morphological discontinuities among the lineages resolved by the molecular phylogeny. The global rate of error associated to species identification based on morphology (0.23) indicates, however, that intergradation of shape and size characters among species in the group challenges their identification.     

A numerical analysis of Tasmanian higher plant endemism

KIRKPATRICK, J. B. & BROWN, M. J., 1984. A numerical analysis of Tasmanian higher plant endemism. Tasmanian endemic plant taxa at the species level or below were placed in geographic elements according to the distribution of their genera. These elements are associated with different environments, the endemic and Antarctic elements being most prominent in rainforest and alpine communities; the cosmopolitan element in alpine communities, and the Australian element in the fire-prone lowland communities. A monothetic divisive classification and minimum spanning tree analysis of a matrix of the occurrence of 242 Tasmanian endemic species from 366 areas resulted in groups of species and lists whose distributions were closely related to precipitation, altitude and vegetation type. The proportions of endemic species in local Tasmanian floras were almost totally explained by altitude and precipitation in a stepwise multiple regression analysis. However, it is possible that many of the endemic species have not been able to occupy their potential range in Tasmania as a result of insufficient time having elapsed for them to fully expand from Glacial refugia.     

Composition, geographical affinities and endemism of the Iberian Peninsula orchid flora

The orchid flora of the Iberian Peninsula is relatively well known, but its biogeographical and diversity patterns have until now remained unanalysed. This work compares the richness of this flora with that of 27 other territories in different continents and at different latitudes, with the aim of establishing whether it is richer or poorer than might be expected. Latitude was found to be an excellent predictor of regional orchid species richness. With 122 taxa, the orchid flora of the Iberian Peninsula is more or less as diverse as that of other Mediterranean areas of similar latitude (e.g. France, Greece or Italy), but more diverse than other European or indeed North African orchid floras. In this study, the Iberian orchid species were assigned to eight monophyletic clades and the global distribution of these are mapped to establish continental affinities between the floras. A recent floristic account on the Iberian orchids was also used to assign the orchid taxa to habitats, and the relationship between the number of endemisms and their habitats was analysed. The patterns of endemism differed in different habitats. Very high levels of endemism were found in habitats peculiar to the Mediterranean Basin, indicating the relict status of its orchid flora.     

A comparative study of the distribution of genus size in twenty angiosperm floras

The biodiversity of floras has until recently been measured solely in terms of their species number or species density, with little regard to the breadth of phylogenetic diversity represented by the species. The latter is partly a function of the size of the flora, and partly of the pattern of distribution of the species into higher taxa. To determine whether floras differ in this respect, this study compares the frequency distribution of genus size in 20 island and regional floras. Certain floras (Cape Region, S.W. Australia, New Zealand, Hawaii) are found to have high concentrations of genera containing many species. Others are notably lacking in large genera (Java, Jamaica, Nepal, Niger), though this group tend to be family-rich. In floras with high endemism (Cape, New Zealand, Fiji, Jamaica, Hawaii), the level of endemism is consistently higher in larger genera. Possible reasons for the observed differences between floras are geographic and temporal isolation, level of habitat diversity, climatic history, volcanic, orogenic and tectonic events. Clusters of large genera may indicate recent speciation, possibly following the last glaciation. Genus size may be an important consideration when limited conservation resources have to be targetted to retain the maximum phylogenetic diversity in a threatened flora.     

中国南部石灰岩濒危植物的初步研究

本文所说中国南部石灰岩地区主要指广西、云南、贵州三省的石灰岩地区,这一地区以其植物的多样性和岩溶特有性而闻名。本文对这一地区的植被和植物保护现状、自然屎护区状况、植物濒危原因筹进行了初步考察和研究。     

Threatened Alvar Grasslands in NW Russia and their Relationship to Alvars in Estonia

Alvar grasslands are thin-soil calcareous grasslands on Silurian and Ordovician limestone bedrock. They are considered unique plant communities for northern Europe and America with a high conservation value. Until now, European alvar grasslands have only been described in Sweden and Estonia. We described species-rich alvar grasslands on the Izhora plateau, NW Russia and compared this vegetation with that of Estonian alvar grasslands. Floristic composition and environmental conditions of Russian alvars were similar to Estonian alvar grasslands on thicker dry soil. Russian alvars are isolated from similar communities in Estonia, but extensive transport of domestic animals from Estonia to NW Russia in past centuries has probably enhanced species dispersal. Some characteristic Estonian alvar species were missing in NW Russia because it lies outside their natural ranges. Other differences in vegetation were explained by higher atmospheric nitrogen deposition and younger age of Russian sites. In NW Russia the alvar grasslands were small, but several red-listed plant species were present. In order to preserve European alvar vegetation effectively, Russian alvar grasslands need further study and conservation as an outpost of this threatened vegetation type.     

Modelling the biological aspects of local endemism in South African Fynbos

The biological attributes, dispersal mode, growth form, and regeneration strategy were determined for the endemic and non-endemic flora of the southern Langeberg, Cape Province, South Africa.—Logistic regression analysis was used to assess the simultaneous effects and interactions between these biological attributes on the occurrence of endemism. The model allowed numerical estimation of the probability that a species with a given set of attributes would be endemic.—This approach extends a contingency table analysis of the data, which merely indicated the association between individual biological traits and endemism. Furthermore, the logistic model allows scope for the analysis of the influence of biological traits in determining endemism in other floras, and also tentative prediction of the probability of endemism in species with combinations of biological traits not yet observed in the flora of the southern Langeberg.     

Patterns of species diversity and endemism in comparable temperate brown algal floras

Brown algal species diversity is compared in 100 km sections of the coastlines of four warm temperate regions: southern Australia, California, southwestern Africa, north-central Chile. The highest diversity (over 140 species per section) is found in southern Australia. California has a reasonable diversity (around 70 species per section), and both southern Australia and California have high regional endemism. Sections of north-central Chile and southwestern Africa have similar patterns, with low diversity (< 30 species per 100 km section), low endemism, few or no fucoids, and up to 25% of the brown algal flora are environmentally tolerant species of Scytosiphonales. Species turnover between contiguous sections of coast is generally related to relative change in temperature regime. Thus the high diversity of southern Australia is due to high species diversity within the 100 km sections, with little turnover, except for a rapid reduction in eastern Victoria likely to be related to lack of rocky substatum. It is hypothesized that low diversity and endemism in Chile and southwestern Africa can be explained by the occurrence of major environmental perturbations (upwelling and El Nino effects) in these regions, producing variable inter-annual temperature conditions that select out tolerant species from the local floras.     

The Greater Cape Floristic Region

Aim The Cape Floristic Region (CFR) (Cape Floristic Kingdom) is currently narrowly delimited to include only the relatively mesic Cape fold mountains and adjacent intermontane valleys and coastal plains. We evaluate the floristic support for expanding the delimitation to include the whole winter‐rainfall area (arid and mesic climates) into a Greater CFR. Location Southern Africa, particularly the south‐western tip. Methods The initial divisive hierarchical classification analysis twinspan used the presence/absence of vascular plant genera to obtain major floristic groupings in southern Africa. For the more detailed analyses, we scored the flora as present/absent within a set of centres, among which the floristic relationships were investigated (agglomerative methods, upgma and minimum spanning trees). These analyses were conducted with species, genera and families separately. The centres were grouped into five regions. The species richness and endemism was calculated for the centres, regions and combination of regions. The dominant floristic components of each region were sought by calculating the percentage contribution of each family to the flora. Results The divisive method showed that the winter‐rainfall areas are floristically distinct from the rest of southern Africa. The species‐ and generic‐level analyses revealed five regions: CFR, Karoo Region, Hantam‐Tanqua‐Roggeveld Region, Namaqualand Region and Namib‐Desert Region. The CFR has the highest endemism and richness. However, the combination of the CFR, the Hantam‐Tanqua‐Roggeveld Region and the Namaqualand Region results in a higher total endemism. Combined, these three regions almost match the region delimited by the twinspan analysis, and together constitute the Greater CFR. Main conclusions The CFR constitutes a valid floristic region. This is evident from the endemism and the distinctive composition of the flora. However, the total endemism is higher for the whole winter‐rainfall area, and this supports the recognition of the larger unit. If floristic regions are to be delimited only on endemism, then the Greater CFR is to be preferred. If floristic regions are delimited on the composition of their floras at family level, then the support for such a grouping is weaker.     

THE PENNSYLVANIAN-PERMIAN VEGETATIONAL TRANSITION: A TERRESTRIAL ANALOGUE TO THE ONSHORE-OFFSHORE HYPOTHESIS

An analysis of 68 floras from the Pennsylvanian and Early Permian of Euramerica reveals distinct patterns of environmental distribution. Wetland assemblages are the most commonly encountered floras from the Early and Middle Pennsylvanian. Floras from drier habitats characterize the Permian. Both wetland and dry-site floras occur in the Late Pennsylvanian, but floristic overlap is minimal, which implies strong environmental controls on the distributions of the component species. Drier habitats appear to be the sites of first appearance of orders that become prominent during the Late Permian and Mesozoic. Higher taxa originated in physically heterogeneous, drier habitats, which were geographically marginal throughout most of the Pennsylvanian. They then moved into the lowlands during periods of climatic drying in the Permian, replacing older wetland vegetation. This pattern is analogous to the marine onshore-offshore pattern of origination and migration. The derivation of Mesozoic wetland clades from the Permian dry-lowland vegetation completes the parallel. The similarities of the marine and terrestrial patterns suggest that the combination of evolutionary opportunity, created by physical heterogeneity of the environment, and migrational opportunity, created by changing extrinsic conditions, may be underlying factors that transcend the specifics of organism and environment.     

State and tendencies in development of floras of man-transformed ecosystems

Basing on published data, authors estimate the current state and tendencies in the development of man-transformed floras. Special attention is given to the flora of cities and such man-transformed habitats, as open-cast mines, dumps, hydrotechnical constructions and railway embankments. Floras of man-made forests and plant communities of cultivated lands were also analyzed. Structure of urbanized landscapes are considered as the aggregates of man-transformed landscapes with fragments of natural vegetation. Floras of urbanized areas are characterized by high species diversity, inconsistency of species composition and diminishing influence of natural vegetation characteristic for particulate zone. Species variety and structure of floras of man-transformed habitats depend on specificity of substrate, geographical latitude, neighboring natural communities and intensity of technical plants exploitation. In the floras of railway embankments species of 10 main families are more abundant then those in natural ecosystems of the same latitude. Floras of man-made forests are characterized by very high species diversity, this type of habitats plays the main compensating role. Analysis of segetal floras reveals the replacement of oligotrophic weeds by eutrophic ones and the increase in diversity of Poaceae species.     

The plant cover of Russia in the area of the Braun-Blanquet syntaxonomy: the development of an approach and the results

Theoretical basis of ecologo-floristic classification according to Braun-Blanquet approach is discussed with the special attention to the contribution of Russian scientists. Principal results concern presentation of vegetation continuum in discrete classification units: polymodal conception of plant communities, principle of plural syntaxonomic decisions, pragmatic approach to classification, method of homological rows of plant communities, etc. Current ecologo-floristic classification of vegetation in Russia consists of 77 classes. This system is a good basement for estimation of rarity of plant communities and biodiversity protection. The first variant of "Red Book" of plant communities of Russia and other eastern European countries is discussed.     

The latitudinal gradient of species-area relationships for vascular plants of North America

The species-area relationship (SAR), describing the increase in species richness with increasing area, and the latitudinal diversity gradient (LDG), describing the decrease in species richness with increasing latitude, are the oldest and most robust patterns in biogeography, yet connections between them remain poorly understood. Here, using 1,742 floras covering the entirety of North America north of Mexico (NAM) and including all of NAM's native species of vascular plants, we show that the slope of the SAR consistently decreases with increasing latitude. This trend is general and holds for subsets of the floras in eastern and western NAM. The southernmost latitudinal quarter of NAM exhibits SARs more than twice as steep as those of the northernmost quarter for both eastern and western regions. This decrease in SAR slope with increasing latitude is consistent with the environmental texture hypothesis and Rapoport's rule, and it suggests that more detailed studies of species endemism in relation to environmental and historical factors will yield significant insights into the underlying causes of SAR and LDG patterns.     

A biogeographic delineation of the European Alpine System based on a cluster analysis of Carex curvula-dominated grasslands

Biogeographic delineations within the European temperate mountains remain poorly understood, as there has been little effort to assemble and analyze vegetation relevés covering Pyrenees, Alps, Carpathians and Balkans altogether. Our study tackles this issue by focusing on the widely distributed alpine acidic grasslands dominated by Carex curvula. Cluster analysis of more than 800 vegetation relevés revealed the European-scale spatial patterns of vascular plant diversity in these alpine grasslands. The geographical distribution of floristic clusters was partly congruent with the physiography of European mountains. Southern European ranges (Southern Balkans and Pyrenees) exhibit a high level of endemism and corresponding floristic clusters are well separated from the others. Marked floristic similarities between the Easternmost Alps, the Carpathians, and the Northern Balkans (Stara Planina) supported a major floristic boundary that runs through the Austrian Alps and that is likely the legacy of a shared Quaternary history. Within the Alps, floristic clustering was mainly driven by ecological drivers and not geography. This paper presents the first detailed study of spatial patterns of species distribution within the European Alpine System, based on a comprehensive analysis of within- and between-community species diversity. It shows that the quantitative analysis of large and consistent data sets may question the traditional delineations of biogeographic regions within European mountains.     

中国常绿阔叶林分类试行方案

植被分类是植被生态学研究中最复杂、充满着争论的问题之一,直到现在并没有一个能为植被学家共同接受的、统一的分类系统。常绿阔叶林的复杂性仅次于热带雨林,加之在人类长期干扰下,变化极大,过渡性群落极多,更增加了分类的困难。中国常绿阔叶林分类经历了漫长的道路,20世纪50年代学习苏俄学派,60年代初引进威斯康星学派的“重要值"概念,在研究云南滇青冈(Cyclobalanopsis glaucoides)林时也曾试用过法瑞学派的理论和方法。80年代《中国植被》(1980)一书的作者们提出,对于“南方某些类型"在“群系(Formation)"和“群丛(Association)"划分时应采用“标志种(Diagnostic species)"的原则,当时由于资料所限,这一原则并未得到贯彻。本文在总结以往分类方案的基础上,根据高级单位以生态外貌、中级单位以优势度类型、低级单位以特征种组的分类原则,构建了从“群丛"、“群系"、“群系组"到“植被亚型"和“植被型"的中国常绿阔叶林分类系统。将中国常绿阔叶林划分出3个植被型、8个植被亚型、14个群系组和53个群系。这些群系大都占据一定的地理区域,并与一定的生境相联系。     

Distance decay of similarity among European urban floras: the impact of anthropogenic activities on β diversity

Aim We examine how two categories of non‐native species (archaeophyte and neophyte, introduced before and after ad 1500, respectively) have had different impacts on β diversity across European urban floras. Our goal is to use the unique biological perspective provided by urban areas, and the contrasting historical and geographical perspectives provided by archaeophytes and neophytes, to infer how non‐native species will impact upon β diversity in the future. Location Twenty‐two urban areas located in seven European countries. Methods We used the β‐sim dissimilarity index to estimate the level of β diversity for 231 unique pair‐wise combinations of 22 urban floras. We examined bivariate plots of dissimilarity by geographical separation of city centres to evaluate distance decay of similarity for native species, archaeophytes and neophytes. Results Based on average percentages, 52.8% (SD = 8.2%) of species in the urban floras were identified as non‐native with 28.3% (SD = 6.9%) classified as neophytes and 24.5% (SD = 4.9%) as archaeophytes. Relative to native species, across urban floras, archaeophytes were associated with higher compositional similarity and weaker distance decay patterns, whereas neophytes were associated with lower compositional similarity and stronger distance decay patterns. Main conclusions Across European urban floras, archaeophytes and neophytes occurred in similar numbers but archaeophytes were consistently associated with lower β diversity and neophytes with higher β diversity. Thus, the impact of non‐native species on β diversity can be determined, at least in part, through their historical and geographical associations with anthropogenic activities. If archaeophytes represent the long‐term biogeographical outcome for human commensal species, neophytes could develop similar patterns. The consequences, however, are likely to be more substantial ecologically and geographically due to the increasing numbers of neophytes and their global anthropogenic associations. Nevertheless, at present, our findings suggest that, based on occurrence information, neophytes have not achieved this state with European urban floras retaining regionally distinct assemblages of neophytes.     

Evolutionary history of the flora of Mexico: Dry forests cradles and museums of endemism

Mexico is considered an exceptional biogeographic area with a varied endemic flora, however spatial phylogenetic measures of biodiversity have not yet been estimated to understand how its flora assembled to form the current vegetation. Patterns of species richness, endemism, phylogenetic diversity, phylogenetic endemism and centers of neo‐ and paleo‐endemism were determined to examine differences and congruence among these measures, and their implications for conservation. Of 24 360 vascular plant species 10 235 (42%) are endemic. Areas of endemism and phylogenetic endemism were associated with dry forests in zones of topographic complexity in mountain systems, in deserts, and in isolated xeric vegetation. Every single locality where seasonally tropical dry forests have been reported in Mexico was identified as an area of endemism. Significant phylogenetic diversity was the most restricted and occurred in the Trans‐Mexican Volcanic Belt and in the Sierra de Chiapas. Notably, the highest degree of phylogenetic clustering comprising neo‐, paleo‐, and super‐endemism was identified in southernmost Mexico. Most vascular plant lineages diverged in the Miocene (5–20 mya) when arid environments expanded across the world. The location of Mexico between two very large landmasses and the fact that more than fifty percent of its surface is arid favored the establishment of tropical lineages adapted to extreme seasonality and aridity. These lineages were able to migrate from both North and South America across Central America presumably during the Miocene and to diversify, illustrating the signature of the flora of Mexico of areas of endemism with a mixture of neo‐ and paleo‐endemism.     

Using legumes as indicators in the seasonally dry vegetation types in South America

The South American corridor of seasonally dry vegetation (SACSV) includes different types of physiognomies forming a continuous corridor with high biodiversity and endemism; however, little attention has been paid to the conservation of the SACSV. As this is an area with great diversity, cataloguing all the species is challenging. Thus, we suggest the use of Leguminosae species (trees and shrubs) as bioindicators of the different types of vegetation present in the area and to identify priority areas for conservation of the SACSV, since the family is highly represented in this vegetation. The study area was divided into 358 grid cells with recorded specimens. For each grid cell, species richness, taxonomic diversity, number of species restricted to one type of vegetation, and threatened and indicator species of phytogeographic domain were calculated. To determine the phytogeographic domains and indicator species, analysis of similarity, cluster and indicator species (ISA) were performed. The results show that 43% of the grid cells (154) have high biological importance for conservation (high taxonomic diversity, species richness and number of restricted species), all of which lie outside of protected areas. We identified 72 indicator species for seven floristic units, which, in general, include areas of the same phytogeographic domain, supporting the existing classification systems. We suggest that for effective conservation of biodiversity present in the SACSV, it is necessary to establish protected areas throughout the SACSV.     

A comparison of fynbos and non-fynbos coenoclines in the lower Gamtoos River Valley,southeastern Cape,South Africa

Patterns in the relative importance of structural attributes and growth forms along fynbos and non-fynbos coenoclines were studied to test the hypothesis that there would be less structural variation in the former because the overriding influence of low levels of soil nutrients would be manifest in a great deal of structural convergence in fynbos. The coenoclines were ranged along identical environmental gradients of increasing altitude, rainfall and soil moisture and decreasing climatic variability. Results showed that along the entire fynbos coenocline vegetation was structurally a small-leaved sclerophyllous shrubland with a graminoid understorey and, usually, a large-leaved (proteoid) shrub overstorey. Fynbos structure was interpreted largely as a response to low levels of soil nutrients. Non-fynbos vegetation ranged from mixed succulent-sclerophyllous and spiny large-leaved thicket at lower altitudes to tall mesic forest at the upper end of the gradient. Non-fynbos structure was explained in terms of variations in soil moisture and climate. An analysis of the biogeographical affinities of sample floras at sites along the coenoclines showed that fynbos vegetation was dominated by taxa endemic to the Cape phytochorion, although phytochorological mixing was pronounced at the lower altitude sites. The level of local endemism in the fynbos coenocline was relatively high; nearly all endemics were Cape fynbos taxa and their incidence increased with increasing altitude. These data indicate that fynbos vegetation has had a lengthy history in the southeastern Cape and that high altitude sites would have comprised a refuge for Cape taxa during unfavourable climatic periods. Non-fynbos vegetation ranged from dry subtropical Tongaland-Pondoland thicket with a strong Karoo-Namib component to temperate Afromontane forest. Levels of endemism were lower than the fynbos coenocline and decreased with increasing altitude. The high number of karroid endemics found in both coenoclines at low altitudes suggests that karroid vegetation would have been more widespread in the past, probably during the last glacial which was considerably drier than the present Holocene interglacial.     

Phylogenetic beta diversity of native and alien species in European urban floras

Aim Human activities have weakened biogeographical barriers to dispersal, increasing the rate of introduction of alien plants. However, their impact on beta diversity and floristic homogenization is poorly understood. Our goal is to compare the phylogenetic beta diversity of native species with that of two groups of alien species, archaeophytes and neophytes (introduced before and after ad 1500, respectively), across European urban floras to explore how biological invasions affect phylogenetic turnover at a continental scale. Location Twenty European cities located in six countries between 49 and 53° N latitude in continental Europe and the British Isles. Methods To compare the phylogenetic beta diversity of native and alien species we use the average phylogenetic dissimilarity of individual floras from their group centroid in multivariate space. Differences in phylogenetic beta diversity among different species groups are then assessed using a randomization test for homogeneity of multivariate dispersions. Results Across European urban floras, and when contrasted with natives, archaeophytes are usually associated with lower levels of phylogenetic beta diversity while neophytes tend to increase phylogenetic differentiation. Main conclusions While archaeophytes tend to promote limited homogenization in phylogenetic beta diversity, because of their diverse geographical origin together with short residence times in the invaded regions, neophytes are not promoting biotic homogenization of urban floras across Europe. Therefore, in spite of the increasing rate of alien invasion, an intense phylogenetic homogenization of urban cities is not to be expected soon.