Reconsidering endemism in the Northeastern Limestone Alps

The restricted distribution of endemic plant species in the Northern Limestone Alps is commonly explained by a severe loss of biotypes and genetic plasticity during Pleistocene glaciation. In the present paper we examine the ecological features of the regional endemic species and plant communities to see if they actually support this interpretation. In particular, we focus on (1) species composition and habitat characteristics of endemic plant communities, (2) the frequency of endemic species in different plant communities and habitat types, (3) the correlation between the frequency of endemic species and the density of the vegetation cover and (4) the altitudinal distribution of endemic species. Concerning endemic plant communities, there is considerable variation in species composition as well as in habitat requirements. Communities of rock, scree and snowbed habitats are characterised by the predominance of endemic or subendemic species, whereas endemic alpine grassland types contain mainly non-endemic plants. Regarding the distribution of endemic plants, no uniform trend could be detected. The endemic species pool contains plants restricted to azonal stands as well as typical species of climax grasslands and widespread generalists. Some endemics distinctly prefer open vegetation types, others predominantly occur in dense grasslands. The bulk of the species considered performs best in the lower alpine zone, some others are especially common in the subalpine zone, whereas only one species showed an occurrence peak in the upper alpine belt. In summary, it may be said that the habitat requirements of endemic plants are rather species-specific and that there are only a few general trends. This fact seems to indicate the complex nature of the phenomenon of regional endemism, which is influenced by a range of factors. The prevailing historical explanation may not be appropriate for every species. Hence, the consideration of current ecological conditions in addition to historical factors may provide a more comprehensive explanation of regional endemism

Pathways to polyploidy : indications of a female triploid bridge in the alpine species Ranunculus kuepferi (Ranunculaceae)

Polyploidy is one of the most important evolutionary processes in plants. In natural populations, polyploids usually emerge from unreduced gametes which either fuse with reduced ones, resulting in triploid offspring (triploid bridge), or with other unreduced gametes, resulting in tetraploid embryos. The frequencies of these two pathways, and male versus female gamete contributions, however, are largely unexplored. Ranunculus kuepferi occurs with diploid, triploid and autotetraploid cytotypes in the Alps, whereby diploids are mostly sexual, while tetraploids are facultative apomicts. To test for the occurrence of polyploidization events by triploid bridge, we investigated 551 plants of natural populations via flow cytometric seed screening. We assessed ploidy shifts in the embryo to reconstruct female versus male gamete contributions to polyploid embryo and/or endosperm formation. Seed formation via unreduced egg cells (B-III hybrids) occurred in all three cytotypes, while only in one case both gametes were unreduced. Polyploids further formed seeds with reduced, unfertilized egg cells (polyhaploids and aneuploids). Pollen was highly variable in diameter, but only pollen > 27 mu m was viable, whereby diploids produced higher proportions of well-developed pollen. Pollen size was not informative for the formation of unreduced pollen. These results suggest that a female triploid bridge via unreduced egg cells is the major pathway toward polyploidization in R. kuepferi, maybe as a consequence of constraints of endosperm development. Triploids resulting from unreduced male gametes were not observed, which explains the lack of obligate sexual tetraploid individuals and populations. Unreduced egg cell formation in diploids represents the first step toward apomixis

Postglacial range expansion of high‐elevation plants is restricted by dispersal ability and habitat specialization

Aim: Species' ecological traits influence their spatial genetic patterns. Bedrock preference strongly shapes the phylogeography of alpine plants, but its interactions with other ecological traits have rarely been disentangled. Here, we explore whether dispersal ability and degree of habitat specialization account for divergent postglacial expansion patterns of high-elevation plants in spite of similar bedrock preference. Location: The Pyrenees, southwestern Europe. Taxon: Cirsium glabrum (Asteraceae), Salix pyrenaica (Salicaceae) and Silene borderei (Caryophyllaceae). Methods: Phylogenetic, genetic structure and demographic modelling analyses based on restriction-site- associated DNA sequencing (RADseq) data from a range-wide populational sampling were conducted. Occurrence data and environmental variables were used to construct species distribution models, which were projected under current and Last Glacial Maximum conditions, and were combined with RADseq data to reconstruct the postglacial history of the study species. The degree of habitat specialization of each species was estimated based on the plant communities within which they occur, and their climatic niche breadth. Results: Salix pyrenaica, which occupies a broad range of habitats, shows a high level of range filling, a blurred genetic structure and an admixture cline between the two main genetic groups, congruent with rapid postglacial expansion. The microsite specialists C. glabrum and S. borderei exhibit a strong genetic structure and low levels of range filling, indicative of slow postglacial expansion. The good disperser C. glabrum shows higher levels of admixture between genetic groups and weaker population differentiation than the poor disperser S. borderei. Main Conclusions: Factors other than bedrock preference have a strong impact on the postglacial range dynamics of high-elevation species. Habitat specialization plays an important role, allowing species occupying a broad range of habitats to more rapidly expand their ranges after environmental change. The effect of dispersal ability is lower than expected for the study species

Climate warming may increase the frequency of cold-adapted haplotypes in alpine plants

The authors incorporate intraspecific variation into a dynamic range model to predict the consequences of twenty-first century warming on six European alpine plants. As well as overall range loss, their model predicts a decrease in the frequency of warm-adapted haplotypes in five out of six species. Modelling of climate-driven range shifts commonly treats species as ecologically homogeneous units. However, many species show intraspecific variation of climatic niches and theory predicts that such variation may lead to counterintuitive eco-evolutionary dynamics. Here, we incorporate assumed intraspecific niche variation into a dynamic range model and explore possible consequences for six high-mountain plant species of the European Alps under scenarios of twenty-first century climate warming. At the species level, the results indicate massive range loss independent of intraspecific variation. At the intraspecific level, the model predicts a decrease in the frequency of warm-adapted haplotypes in five species. The latter effect is probably driven by a combination of leading-edge colonization and priority effects within the species' elevational range and was weakest when leading-edge expansion was constrained by mountain topography The resulting maladaptation may additionally increase the risk that alpine plants face from shrinkage of their ranges in a warming climate.Peer reviewe

Distinct Biogeographic Phenomena Require a Specific Terminology: A Reply to Wilson and Sagoff

In a recent publication, we proposed that the increasing number of range-expanding species that track human induced environmental change warrant specific recognition in science and biodiversity management, and we proposed the term ‘neonative’ for these taxa. Here, we reply to two letters (Wilson 2019, Sagoff 2019) that criticised specific, yet different aspects presented in our publication. While we disagree on several points with both authors, we agree that a broader discourse is needed for developing robust and widely accepted definitions and terms for the ever more important phenomenon of neonative species

What is valued in conservation? A framework to compare ethical perspectives

Perspectives in conservation are based on a variety of value systems. Such differences in how people value nature and its components lead to different evaluations of the morality of conservation goals and approaches, and often underlie disagreements in the formulation and implementation of environmental management policies. Specifically, whether a conservation action (e.g. killing feral cats to reduce predation on bird species threatened with extinction) is viewed as appropriate or not can vary among people with different value systems. Here, we present a conceptual, mathematical framework intended as a tool to systematically explore and clarify core value statements in conservation approaches. Its purpose is to highlight how fundamental differences between these value systems can lead to different prioritizations of available management options and offer a common ground for discourse. The proposed equations decompose the question underlying many controversies around management decisions in conservation: what or who is valued, how, and to what extent? We compare how management decisions would likely be viewed under three idealised value systems: ecocentric conservation, which aims to preserve biodiversity; new conservation, which considers that biodiversity can only be preserved if it benefits humans; and sentientist conservation, which aims at minimising suffering for sentient beings. We illustrate the utility of the framework by applying it to case studies involving invasive alien species, rewilding, and trophy hunting. By making value systems and their consequences in practice explicit, the framework facilitates debates on contested conservation issues, and complements philosophical discursive approaches about moral reasoning. We believe dissecting the core value statements on which conservation decisions are based will provide an additional tool to understand and address conservation conflicts

Capacity of countries to reduce biological invasions

The extent and impacts of biological invasions on biodiversity are largely shaped by an array of socio-economic and environmental factors, which exhibit high variation among countries. Yet, a global analysis of how these factors vary across countries is currently lacking. Here, we investigate how five broad, country-specific socio-economic and environmental indices (Governance, Trade, Environmental Performance, Lifestyle and Education, Innovation) explain country-level (1) established alien species (EAS) richness of eight taxonomic groups, and (2) proactive or reactive capacity to prevent and manage biological invasions and their impacts. These indices underpin many aspects of the invasion process, including the introduction, establishment, spread and management of alien species. They are also general enough to enable a global comparison across countries, and are therefore essential for defining future scenarios for biological invasions. Models including Trade, Governance, Lifestyle and Education, or a combination of these, best explained EAS richness across taxonomic groups and national proactive or reactive capacity. Historical (1996 or averaged over 1996–2015) levels of Governance and Trade better explained both EAS richness and the capacity of countries to manage invasions than more recent (2015) levels, revealing a historical legacy with important implications for the future of biological invasions. Using Governance and Trade to define a two-dimensional socio-economic space in which the position of a country captures its capacity to address issues of biological invasions, we identified four main clusters of countries in 2015. Most countries had an increase in Trade over the past 25 years, but trajectories were more geographically heterogeneous for Governance. Declines in levels of Governance are concerning as they may be responsible for larger levels of invasions in the future. By identifying the factors influencing EAS richness and the regions most susceptible to changes in these factors, our results provide novel insights to integrate biological invasions into scenarios of biodiversity change to better inform decision-making for policy and the management of biological invasions