Evaluating plant lineage losses and gains in temperate forest understories: a phylogenetic perspective on climate change and nitrogen deposition.

Global change has accelerated local species extinctions and colonizations, often resulting in losses and gains of evolutionary lineages with unique features. Do these losses and gains occur randomly across the phylogeny? We quantified: temporal changes in plant phylogenetic diversity (PD); and the phylogenetic relatedness (PR) of lost and gained species in 2672 semi-permanent vegetation plots in European temperate forest understories resurveyed over an average period of 40 yr. Controlling for differences in species richness, PD increased slightly over time and across plots. Moreover, lost species within plots exhibited a higher degree of PR than gained species. This implies that gained species originated from a more diverse set of evolutionary lineages than lost species. Certain lineages also lost and gained more species than expected by chance, with Ericaceae, Fabaceae, and Orchidaceae experiencing losses and Amaranthaceae, Cyperaceae, and Rosaceae showing gains. Species losses and gains displayed no significant phylogenetic signal in response to changes in macroclimatic conditions and nitrogen deposition. As anthropogenic global change intensifies, temperate forest understories experience losses and gains in specific phylogenetic branches and ecological strategies, while the overall mean PD remains relatively stable.

Les changements globaux accélèrent les processus de colonisation et d'extinction locales d'espèces, aboutissant à des gains ou à des pertes de lignées évolutives uniques. Ces gains et pertes se produisent-ils de manière aléatoire dans l'arbre phylogénétique ? Nous avons mesuré: les changements de diversité phylogénétique; et la parenté phylogénétique des espèces végétales gagnées ou perdues dans 2672 placettes semi-permanentes disposées dans le sous-bois de forêts tempérées d'Europe sur une période moyenne de 40 ans. Une fois corrigée par la richesse spécifique, la diversité phylogénétique a légèrement augmenté au cours du temps dans les différentes placettes. Les espèces perdues ont une plus grande parenté phylogénétique que les espèces gagnées. Les espèces gagnées sont donc issues d'un plus grand nombre de lignées évolutives que les espèces perdues. Certaines lignées ont gagné ou perdu davantage d'espèces que ce qui est prédit par le hasard : les Ericaceae, les Fabaceae et les Orchidaceae ayant davantage perdu, tandis que les Amaranthaceae, les Cyperaceae, et les Rosaceae ont plus gagné. Il n'y a pas de signal phylogénétique des gains ou pertes d'espèces en réponse aux changements de conditions macroclimatiques ou des dépôts atmosphériques d'azote. Alors que les changements globaux d'origine anthropique s'intensifient, les sous-bois des forêts tempérées connaissent des gains et des pertes de certaines lignées évolutives et de certaines stratégies écologiques, sans que la diversité phylogénétique moyenne ne s'en trouve véritablement affectée.

El cambio global ha acelerado las extinciones y colonizaciones a escala local, lo que a menudo ha supuesto pérdidas y ganancias de linajes evolutivos con características únicas. Ahora bien, ¿estas pérdidas y ganancias ocurren aleatoriamente a lo largo de la filogenia? Cuantificamos: los cambios temporales en la diversidad filogenética de las plantas; y la relación filogenética de las especies perdidas y ganadas en 2.672 parcelas de vegetación semipermanente en sotobosques templados europeos y re-muestreadas durante un período promedio de 40 años. Al controlar por las diferencias en la riqueza de especies, la diversidad filogenética aumentó ligeramente con el tiempo y entre parcelas. Además, las especies perdidas dentro de las parcelas exhibieron un mayor grado de relación filogenética que las especies ganadas. Esto implica que las especies ganadas se originaron en un conjunto de linajes evolutivos más diversos que las especies perdidas. Ciertos linajes también perdieron y ganaron más especies de las esperadas aleatoriamente: Ericaceae, Fabaceae y Orchidaceae experimentaron pérdidas y Amaranthaceae, Cyperaceae y Rosaceae mostraron ganancias. Las pérdidas y ganancias de especies no mostraron ninguna señal filogenética significativa en respuesta a los cambios en las condiciones macro-climáticas y la deposición de nitrógeno. A medida que se intensifica el cambio global antropogénico, los sotobosques temperados experimentan pérdidas y ganancias en ramas filogenéticas y estrategias ecológicas específicas, mientras que la diversidad filogenética media general permanece relativamente estable.

Climate-trait relationships exhibit strong habitat specificity in plant communities across Europe.

Ecological theory predicts close relationships between macroclimate and functional traits. Yet, global climatic gradients correlate only weakly with the trait composition of local plant communities, suggesting that important factors have been ignored. Here, we investigate the consistency of climate-trait relationships for plant communities in European habitats. Assuming that local factors are better accounted for in more narrowly defined habitats, we assigned > 300,000 vegetation plots to hierarchically classified habitats and modelled the effects of climate on the community-weighted means of four key functional traits using generalized additive models. We found that the predictive power of climate increased from broadly to narrowly defined habitats for specific leaf area and root length, but not for plant height and seed mass. Although macroclimate generally predicted the distribution of all traits, its effects varied, with habitat-specificity increasing toward more narrowly defined habitats. We conclude that macroclimate is an important determinant of terrestrial plant communities, but future predictions of climatic effects must consider how habitats are defined.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Disentangling effects of disturbance severity and frequency: Does bioindication really work?

Ecological disturbances are recognized as a crucial factor influencing the attributes of ecological communities. Depending on the specific adaptation or life cycle, plant species show different responses to disturbances of different magnitudes. Herben et al. (Journal of Vegetation Science, 27, 628-636) proposed six disturbance indicator values (DIVs) that describe the niches of Central-European plant species along gradients of disturbance frequency and severity. Here, we ask if the DIVs can be used in community ecology for bioindication of disturbance regime? We used a dataset of riparian forests sampled within mountain catchments (the Sudetes, SW Poland). As the regime of disturbance is driven by changes in floods from the spring toward the mouth, we calculated the position of every plot along longitudinal (upstream-downstream) gradient and used it as a proxy for the disturbance severity and frequency. We then calculated the community-weighted means (CWMs) for each of the six indices for each plot and analyzed whether these indices reflected the position of the plots along the rivers. We expected an increase in the severity indices and a decrease in the frequency indices downstream along the rivers. Moreover, we analyzed relationships between disturbance indices and species optima along longitudinal gradient. Surprisingly, means for all analyzed indices increased along the rivers. Severity indices showed the strongest association with the longitudinal gradient. The disturbance severity index for herbs was the only index that differed significantly among species with different responses along longitudinal gradient. On these results, we identified a strong correlation between the severity and frequency indices as the main problem. We conclude that the DIVs have considerable applicative potential; however, the determination of ecological niches separately for disturbance severity and frequency is difficult because different components interact to shape the realized niche of each species. All analyzed indices encompass different attributes of the disturbance regime including both severity and frequency.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Schall and Heinrichs question our interpretation that the climatic debt in understory plant communities is locally modulated by canopy buffering. However, our results clearly show that the discrepancy between microclimate warming rates and thermophilization rates is highest in forests where canopy cover was reduced, which suggests that the need for communities to respond to warming is highest in those forests.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Bertrand et al question our interpretation about warming effects on the thermophilization in forest plant communities and propose an alternative way to analyze climatic debt. We show that microclimate warming is a better predictor than macroclimate warming for studying forest plant community responses to warming. Their additional analyses do not affect or change our interpretations and conclusions.

Forest microclimate dynamics drive plant responses to warming.

Climate warming is causing a shift in biological communities in favor of warm-affinity species (i.e., thermophilization). Species responses often lag behind climate warming, but the reasons for such lags remain largely unknown. Here, we analyzed multidecadal understory microclimate dynamics in European forests and show that thermophilization and the climatic lag in forest plant communities are primarily controlled by microclimate. Increasing tree canopy cover reduces warming rates inside forests, but loss of canopy cover leads to increased local heat that exacerbates the disequilibrium between community responses and climate change. Reciprocal effects between plants and microclimates are key to understanding the response of forest biodiversity and functioning to climate and land-use changes.

Replacements of small- by large-ranged species scale up to diversity loss in Europe's temperate forest biome.

Biodiversity time series reveal global losses and accelerated redistributions of species, but no net loss in local species richness. To better understand how these patterns are linked, we quantify how individual species trajectories scale up to diversity changes using data from 68 vegetation resurvey studies of seminatural forests in Europe. Herb-layer species with small geographic ranges are being replaced by more widely distributed species, and our results suggest that this is due less to species abundances than to species nitrogen niches. Nitrogen deposition accelerates the extinctions of small-ranged, nitrogen-efficient plants and colonization by broadly distributed, nitrogen-demanding plants (including non-natives). Despite no net change in species richness at the spatial scale of a study site, the losses of small-ranged species reduce biome-scale (gamma) diversity. These results provide one mechanism to explain the directional replacement of small-ranged species within sites and thus explain patterns of biodiversity change across spatial scales.

Response of soil mites (Acari, Mesostigmata) to long-term Norway spruce plantation along a mountain stream.

During the nineteenth and twentieth centuries, coniferous monocultures were introduced, replacing natural broadleaved forests in Central Europe, mainly for economic benefits. In the mountains, Norway spruce [Picea abies (L.) H. Karst] was introduced in large areas previously covered with beech forests and also in natural riverside habitat corridors such as river valleys, despite its negative impact on the soil environment by e.g. organic matter accumulation, decrease of soil pH and changes in C/N ratio. We aimed to check how long-term Norway spruce plantations affect species richness and diversity of soil mites along a mountain river in former mixed and broadleaved forests. The study, based on 342 samples, was carried out in Stolowe Mountains National Park (SW Poland). Understory species biomass, soil pH and soil organic layer thickness significantly affected soil mite communities. Although coniferous forests did not differ from either broadleaved or mixed forests in mite density (number of individuals m-2) and species diversity (H'), they were characterized by low species richness and proportional abundance of Uropodina mites typical for broadleaved forests. In total, 4849 mites classified into 57 species were recorded from all forest types and no unique species were found in the sampled forests. Although the mite communities were dominated by the same common species (Veigaia nemorensis, Paragamasus runcatellus, Leptogamasus obesus and Trachytes aegrota), they still maintain the rare species of broadleaved forests and their high recovery potential may be used in forest conversion.

Natural forest remnants as refugia for bryophyte diversity in a transformed mountain river valley landscape.

Riparian forests are among the most threatened ecosystem types worldwide. Their exploitation and replacement by coniferous plantations affects species pools and contributes to loss of biodiversity. We aimed to investigate bryophyte species pools within different habitat types in a transformed mountain river valley. We especially focused on the contribution of habitat types (relative to their proportional cover) to the species pool of the whole area. The study was conducted along the Czerwona Woda river - a model stream in the Stolowe Mountains National Park (SW Poland, study area: 91.2 ha) - and an example of coniferous plantations replacing natural broadleaved forest vegetation. Our study revealed the presence of 147 bryophyte species. The most valuable habitats in terms of diversity of bryophyte assemblages were remnants of the natural vegetation - broadleaved forests and streams. These habitats, constituting <5% of the study area, hosted ca 40% of the total species pool (61 and 62 species, respectively), while the species pool of Picea abies forests (92 species) was proportional to cover of this habitat type (ca 60%). Remnants of natural vegetation were hotspots of bryophyte diversity within the heavily altered landscape, and may play a future role as sources of recolonization by forest specialists. Our study also confirmed the important role of riparian areas in maintaining bryophyte species diversity at the landscape scale. The river valley studied contributes >20-fold more to the bryophyte species pool of the whole national park than indicated by its size. Thus, river valleys require special treatment - conservation based on natural restoration, and should remain reserved from wood production, as areas providing a wide range of ecosystem services.

Ellenberg's indicator values support prediction of suitable habitat for pre-diapause larvae of endangered butterfly Euphydryas aurinia.

In spite of the great popularity of Ellenberg's Indicator Values (EIVs) in plant ecology, animal ecologists seldom use EIVs to address ecological questions. In this study we used EIVs to test their potential usefulness for the prediction of suitable habitat for pre-diapause larvae of the endangered butterfly species Euphydryas aurinia. Nine transects crossing grasslands in SW Poland with abundant populations of E. aurinia were designed. We sampled 76 vegetation plots along the transects. In addition, the presence of the larval webs of E. aurinia in sampled plots was also recorded. We then calculated the mean community EIVs of light, nitrogen, soil reaction, moisture and temperature for each sample plots. Generalized linear mixed-effects models (GLMMs) were used to assess which factors determine the local occurrence of larval webs of E. aurinia. We found the larval webs only in 12 plots, while the host plant was present in 39 of the examined plots. The presence of the host plant was the most important predictor in both models including all plots or including only plots with host plants. The other significant predictor was the mean EIV of light, and its importance increased in models considering all plots. We attributed the importance of the EIV of light to the site openness and density of the vegetation layer. A positive relationship between this predictor and the presence of larval webs indicates that sites with looser vegetation, a lower contribution of shrubs and tall herbs and better penetration of photosynthetically active radiation to lower vegetation layers are preferred by E. aurinia for oviposition. Moreover, the significance of EIV of light may be linked with management practices. Many light-demanding species decline after cessation of mowing as a result of litter accumulation and the dominance of tall herbs. An absence of light-demanding species decreases the community's mean EIV of light and thus indicates the influence of meadow abandonment.