Limestone Alps Research Articles

Using barcoding to reveal ecological patterns of nivicolous myxomycetes in the German Alps: How do they deal with varying snow conditions?

A transect in the German limestone Alps was monitored over ten years for nivicolous myxomycetes to test if species display stable altitudinal belts for fruiting. The data set comprised 1368 barcoded specimens assigned to 112 ribotypes forming 51 ribogroups. Ribogroups were largely consistent with 35 identified morphospecies, although in eleven cases a morphospecies included several ribogroups. Fructification abundance correlated with duration of the snow cover inferred from data loggers placed at ground height. Morphospecies, ribogroups, and ribotypes showed a peak of fructification abundance at different elevations in different years. Species composition, not abundances, showed a high overlap with soil metabarcoding data. Thirteen ribogroups detected in the metabarcoding data set were never found as fructifications. This survey demonstrates that nivicolous myxomycetes are opportunists, which are likely to persist as trophic or resting stages independent from snow cover, but fruit only in altitudes and years with snow cover stable over several months.

Arrested Succession on Fire-Affected Slopes in the Krummholz Zone and Subalpine Forest of the Northern Limestone Alps

Fire in the Northern Alps is comparatively rare. Yet, previous human-ignited fire events in subalpine forests up to the treeline have triggered severe fire damage to vegetation and soil. Here, we investigate post-fire vegetation dynamics in the Northern Limestone Alps about 80 years after disturbance. We observed higher species richness in burned compared to unburned vegetation and clearly distinct floristic communities emerging after fire-driven forest removal, with several alpine specialist species uniquely found in the burned subalpine sites. The functional composition of vegetation was also distinct, with higher relative forb cover in burned plots. This difference was likely driven by disturbance-related environmental changes, such as increased light availability, offering safe sites for subalpine and alpine species. Due to a general lack of tree encroachment, we consider this a case of arrested succession after fire. We conclude that the recovery of fire-affected subalpine forests is modulated by complex interactions of climatic and biotic filters producing extreme site conditions, controlling the recolonization of the disturbed areas by forest species while providing safe sites for the establishment of a rich subalpine and alpine low-statured flora. The coupling of disturbance and abiotic filters makes high-elevation treeline ecotones very vulnerable to climate change.

Comparative Understory Development in Fenced and Unfenced Beech Forest Stands in the Southern Alps

This study examines the impact of ungulate browsing on the understory dynamics of the Cansiglio Forest, the only European beech high forest in the Southern Limestone Alps. Utilizing seven pairs of fenced versus unfenced forest stands established over two decades, we assessed how fencing affects understory structure and tree regeneration. Significant differences were found in cover, height and vascular species composition in both tree regeneration and shrub layers. Fenced areas showed a significantly higher number and greater average height of young trees, supporting the hypothesis that ungulate browsing negatively impacts tree regeneration. Despite previous studies attributing the lack of silver fir (Abies alba Mill.) regeneration primarily to red deer (Cervus elaphus Linnaeus, 1758) browsing, our findings did not show a significant association of this species with fenced conditions. While Rubus abundance was higher inside fenced areas, overall vascular species richness did not differ. Our results highlight the evident effects of ungulate browsing and suggest the need to consider other factors influencing forest regeneration and understory development. Future research should establish additional study replicates, balanced across diverse site and stand structure conditions, to further explore these dynamics.

Repeated colonization of the Northern Limestone Alps from the Southern Limestone Alps by the rock‐dwelling snail species Cochlostoma henricae

AbstractHausdorf, B., Xu, J., BAMBERGER, S. (2024) Repeated colonization of the Northern Limestone Alps from the Southern Limestone Alps by the rock‐dwelling snail species Cochlostoma henricae—Zoologica Scripta, 00, 000–000. The evolution of biogeographic disjunctions in the Southern and Northern Limestone Alps has been studied only in a few taxa. Cochlostoma henricae (Strobel, 1851) is a land snail species restricted to limestone rocks showing such disjunctions. We studied the phylogeny and population structure of this species using genomic ddRAD data to better understand the differentiation of the species in the Southern Alps and the origin of the isolated occurrences in the Northern Limestone Alps. Phylogenetic and population‐genetic analyses of the ddRAD data revealed a subdivision of the C. henricae populations into a western cluster and an eastern cluster. These clusters were probably separated by glaciers that extended from the Piave Valley to the margin of the Alps. Such a longitudinal subdivision is a common pattern of several species in the Southern Limestone Alps. However, the boundaries between western and eastern population groups differ between taxa. Cochlostoma henricae colonized the Northern Limestone Alps at least twice. The population from Breitenwang in Tyrol originated from the western group, whereas the occurrences in the Salzkammergut are from the eastern group. These and similar disjunctions of several other land snails show that even species with limited dispersal abilities could cross the Alps repeatedly by passive long‐distance dispersal events. A subdivision of C. henricae into three subspecies, proposed based on shell characters, does not correspond to its population‐genetic structure. Instead, the eastern and western population groups represent distinct evolutionary units.

Microbial perspective of inhibited carbon turnover in Tangel humus of the Northern Limestone Alps.

Tangel humus primarily occurs in montane and subalpine zones of the calcareous Alps that exhibit low temperatures and high precipitation sums. This humus form is characterized by inhibited carbon turnover and accumulated organic matter, leading to the typical thick organic layers. However, the reason for this accumulation of organic matter is still unclear, and knowledge about the microbial community within Tangel humus is lacking. Therefore, we investigated the prokaryotic and fungal communities along with the physical and chemical properties within a depth gradient (0-10, 10-20, 20-30, 30-40, 40-50 cm) of a Tangel humus located in the Northern Limestone Alps. We hypothesized that humus properties and microbial activity, biomass, and diversity differ along the depth gradient and that microbial key players refer to certain humus depths. Our results give the first comprehensive information about microbiota within the Tangel humus and establish a microbial zonation of the humus. Microbial activity, biomass, as well as microbial alpha diversity significantly decreased with increasing depths. We identified microbial biomarkers for both, the top and the deepest depth, indicating different, microbial habitats. The microbial characterization together with the established nutrient deficiencies in the deeper depths might explain reduced C-turnover and Tangel humus formation.

The Potential of Deep Geothermal Energy in Tyrol—Based on a Pre-feasibility Study

The economic use of deep geothermal energy is fundamentally controlled by the factors of rock permeability, temperature gradient, and depth. The carbonates of the Northern Limestone Alps are considered possible hydrothermal deep aquifers in Tyrol. This is the so-called main dolomite and Wetterstein limestone. For an initial assessment of the geothermal potential, information from the deep Kramsach Th1 borehole was used. With a temperature gradient of approx. 1.8 °C/100 m, which could be derived from the Kramsach Th1 borehole, temperatures of 65 °C at depths of approx. 3000 m and 100 °C at a depth of 5000 m occur in the Inn Valley expect. In addition, it is noted that further in the northwest of the Limestone Alps, at the deep boreholes Vorderriß 1 and Hindelang 1, higher temperature gradients of 2.2 °C/100 m and 2.6 °C/100 m were observed, respectively. Successful thermal water development at these depths requires that hydraulically well-permeable rocks are present. To clarify this question, extensive investigations of the reservoir rocks through exploration drilling are still required. Deep geothermal energy can lead to associated seismicity. In order to quickly detect associated seismicity and to be able to react in a timely manner, seismic monitoring is required during drilling activities and during operation of the systems, whereby the accompanying seismic monitoring must be able to distinguish between natural and induced seismicity.

Preliminary observation of Marmolada glacier collapse of July 2022 with space-based cameras

ABSTRACT On 3 July 2022, a section of the Marmolada glacier (Marmolada Group, Fassa Dolomites, Southern Limestone Alps) collapsed, causing an ice avalanche resulting in the death of 11 alpinists. Before the event, there were no clear signs of any potential collapse, and no warning notices were broadcasted; due to the velocity of the avalanche, no escape was possible for the involved persons. In this work, a preliminary analysis of the Marmolada glacier collapse is performed with four optical channels space imaging provided by Planet Labs, with resolution of 3 m per pixel. Pre- and post-event satellite observations are employed to assess the areas involved in the event: multispectral images can be compared to identify the collapse area and the debris conoid. A first estimation suggests a collapse area of about 4800 m2 and a collapse volume of about 96,000 m3. In addition, historical data from the Planet Labs optical pictures database can be employed to compare the status of the detached glacier and the evolution of the crevasses on its surface.

Phytogeographic Characteristics of Montane Coniferous Forests of the Central Balkan Peninsula (SE Europe)

We investigated taxonomic and endemic richness, patterns of spatial distribution, cenotic and spatial diversification, and chorological and life form spectra of montane coniferous forests in the central part of the Balkan Peninsula. We collected information on 1435 taxa (1351 at the level of species and 84 subspecies) with 65,289 species-occurrence data, published in 1930 original plots with a total area of about 215 ha in the analysis. All statistical analyses (univariate and multivariate) were performed on binary matrices prepared for different levels of analysis. Our main results showed that the montane coniferous forests of the central Balkan Peninsula represent very species-rich vegetation. At the same time, the high proportion of endemics indicated that the montane coniferous forests of the central Balkan Peninsula differ significantly from Central European and boreal forests of a similar type. Furthermore, we found that there were regional differences in the species composition of the coniferous forests of the Balkan Peninsula, and that the primary centers of floristic richness are located in the area of the central and continental Dinarides. This latter finding suggested that the true centers of the richness of European coniferous forests are located south of the Limestone Alps-Western Dinarides-Carpathian Foothills line in Romania, which used to be considered the center of the richness of the coniferous forests in Europe.

Molecular phylogeny of Alpine groundwater Niphargidae (Crustacea, Amphipoda) reveals the effects of Quaternary climate fluctuations

The groundwater amphipod genus Niphargus comprises over 425 described species mainly distributed in the Western Palearctic. Representatives of the genus are quite rare in areas formerly occupied by Quaternary glaciers. Although a couple of species were previously discovered in high-elevation caves and karstic springs, it is only recently that the genus was found to be widely distributed along the Alpine chain (Fišer et al. 2018; Stoch et al. 2020). Using a molecular phylogeny approach, we tested the hypothesis of multiple origins of Alpine Niphargus species followed by dispersal versus allopatric speciation in glacial refugia. We sampled all type localities of species present in the Alps and collected at several new sites, both in formerly glaciated areas and in areas very close to the last Quaternary glacial border. All sequences of Alpine species stored in GenBank were downloaded and included in the analysis as well. Our results show that several phyletic lineages independently colonized the Alpine chain at different periods. We found examples of preglacial colonization, with species surviving as glacial relicts with a very limited range expansion, as well as of recent post-glacial dispersal allowing colonization of very large areas on the northern slope of the Alps. In the Southern Limestone Alps as well as in the Western Alps, most species have a distribution restricted to small karstic massifs south of the area occupied by glaciers during the Last Glacial Maximum (Stoch et al. 2022), except for a single putative relict. Three clades show a relict distribution with highly isolated species found also in caves more than 2,000 m above sea level. The reconstructed time-calibrated phylogeny strongly supports the hypothesis that the evolutionary history of most clades is pre-Pleistocenic on the southern slope of the Alps, while on the northern slope both phylogenetic relicts and recent dispersers co-exist.

Long-term soil warming alters fine root dynamics and morphology, and their ectomycorrhizal fungal community in a temperate forest soil.

Climate warming is predicted to affect temperate forests severely, but the response of fine roots, key to plant nutrition, water uptake, soil carbon, and nutrient cycling is unclear. Understanding how fine roots will respond to increasing temperature is a prerequisite for predicting the functioning of forests in a warmer climate. We studied the response of fine roots and their ectomycorrhizal (EcM) fungal and root-associated bacterial communities to soil warming by 4°C in a mixed spruce-beech forest in the Austrian Limestone Alps after 8 and 14years of soil warming, respectively. Fine root biomass (FRB) and fine root production were 17% and 128% higher in the warmed plots, respectively, after 14years. The increase in FRB (13%) was not significant after 8years of treatment, whereas specific root length, specific root area, and root tip density were significantly higher in warmed plots at both sampling occasions. Soil warming did not affect EcM exploration types and diversity, but changed their community composition, with an increase in the relative abundance of Cenoccocum at 0-10cm soil depth, a drought-stress-tolerant genus, and an increase in short- and long-distance exploration types like Sebacina and Boletus at 10-20cm soil depth. Warming increased the root-associated bacterial diversity but did not affect their community composition. Soil warming did not affect nutrient concentrations of fine roots, though we found indications of limited soil phosphorus (P) and potassium (K) availability. Our findings suggest that, in the studied ecosystem, global warming could persistently increase soil carbon inputs due to accelerated fine root growth and turnover, and could simultaneously alter fine root morphology and EcM fungal community composition toward improved nutrient foraging.

Log Transport in the Limestone Alps, Austria: Where Did the Timber Go?

Abstract Wood is one of the most important sustainable natural resources. Throughout centuries, large cities as well as regions of very high wood demand (for example, areas of salt production and iron processing) had to be supplied with wood for building and burning. One of the largest forests in Austria can be found in the National Park covering the range of mountains called “Limestone Alps.” Within this region, a wide network of dams was set up to transport single logs (“Holztrift”). Due to the dendrochronological dating of the surviving dams, regional chronologies spanning a length of almost 600 years for Norway spruce, Silver fir and European larch were set up. These chronologies serve as the background for dendro-provenancing timber. Dendro-provenancing checks the similarities between samples and different regional chronologies. The highest statistical figures give hints of the origins of the wood. Archival analyses show that many logs were transported to Vienna as rafts. For example, in 1865, more than 1504 rafts arrived in Vienna, all from different origins. The logs were used as building material as well as for burning. Further archival analyses showed that logs from this region were also used for building the roof of the Hofburg Castle in Vienna. Several hundred samples from the roof construction were dendrochronologically dated and used for dendro-provenancing. The origin of the logs was confirmed to be the wider region of the Northern alpine foothills. This meant that rafts from the rivers Alm and Traun as well as from the rivers Steyr and Enns had floated down to Vienna.

Algal Community Change in Mountain Lakes of the Alps Reveals Effects of Climate Warming and Shifting Treelines1.

The biological communities of mountain lakes are suspected to be highly sensitive to global warming and associated catchment changes. To identify the parameters determining algal community responses, subfossil pigments from 21 different mountain lakes in the Bavarian-Tyrolean Limestone Alps were investigated. Sediment cores were radio-isotopically dated, and their pigment preservation evaluated. General additive models (GAM) of pigment compositions were calculated with temperature as the explanatory variable and generalized linear models with several lake parameters explaining log-transformed GAM P-values. Lake depth and trophic state were identified as major control variables of the algal community and productivity changes. Shifts in a deep oligotrophic alpine lake (lg(P)=-1.04) were half as strong as in a shallow mesotrophic alpine lake (lg(P)=-1.86) with faster warming and higher productivity forcing the development of biomass. Phytoplankton and macrophyte pigments increased clearly with warming, at lower altitudes, and decreased at the treeline, so that periphytic pigments dominated alpine sediments. This pattern is probably the result of interactions of UV radiation and allochthonous inputs of DOM. Our findings suggest that (sub)alpine shallow lakes with higher nutrient levels are most vulnerable to climate change-driven changes whereas deep, nutrient-poor lakes appear more resilient.

Evidence for Glacial Refugia of the Forest Understorey Species Helleborus niger (Ranunculaceae) in the Southern as Well as in the Northern Limestone Alps.

Glacial refugia of alpine and subnival biota have been intensively studied in the European Alps but the fate of forests and their understory species in that area remains largely unclear. In order to fill this gap, we aimed at disentangling the spatiotemporal diversification of disjunctly distributed black hellebore Helleborus niger (Ranunculaceae). We applied a set of phylogeographic analyses based on restriction-site associated DNA sequencing (RADseq) data and plastid DNA sequences to a range-wide sampling of populations. These analyses were supplemented with species distribution models generated for the present and the Last Glacial Maximum (LGM). We used exploratory analyses to delimit genomically coherent groups and then employed demographic modeling to reconstruct the history of these groups. We uncovered a deep split between two major genetic groups with western and eastern distribution within the Southern Limestone Alps, likely reflecting divergent evolution since the mid-Pleistocene in two glacial refugia situated along the unglaciated southern margin of the Alps. Long-term presence in the Southern Limestone Alps is also supported by high numbers of private alleles, elevated levels of nucleotide diversity and the species’ modeled distribution at the LGM. The deep genetic divergence, however, is not reflected in leaf shape variation, suggesting that the morphological discrimination of genetically divergent entities within H. niger is questionable. At a shallower level, populations from the Northern Limestone Alps are differentiated from those in the Southern Limestone Alps in both RADseq and plastid DNA data sets, reflecting the North-South disjunction within the Eastern Alps. The underlying split was dated to ca. 0.1 mya, which is well before the LGM. In the same line, explicit tests of demographic models consistently rejected the hypothesis that the partial distribution area in the Northern Limestone Alps is the result of postglacial colonization. Taken together, our results strongly support that forest understory species such as H. niger have survived the LGM in refugia situated along the southern, but also along the northern or northeastern periphery of the Alps. Being a slow migrator, the species has likely survived repeated glacial-interglacial circles in distributional stasis while the composition of the tree canopy changed in the meanwhile.

Geodetic-Gravimetric Monitoring of Mountain Uplift and Hydrological Variations at Zugspitze and Wank Mountains (Bavarian Alps, Germany)

In 2004, first absolute gravity (AG) measurements were performed on the top of Mt. Zugspitze (2 sites) and at the foot (1 site) and top (1 site) of Mt. Wank. Mt. Wank (summit height 1780 m) and Mt. Zugspitze (2960 m) are about 15 km apart from each other and belong geologically to different parts of the Northern Limestone Alps. Bridging a time span of 15 years, the deduced gravity variations for Zugspitze are in the order of −0.30 μm/s2 with a standard uncertainty of 0.04 μm/s2. The Wank stations (foot and top) show no significant gravity variation. The vertical stability of Wank summit is also confirmed by results of continuous GNSS recordings. Because an Alpine mountain uplift of 1 or 2 mm/yr cannot explain the obtained gravity decline at Zugspitze, the dominating geophysical contributions are assumed to be due to the diminishing glaciers in the vicinity. The modelled gravity trend caused by glacier retreat between epochs 1999 and 2018 amounts to −0.012 μm/s2/yr at both Zugspitze AG sites. This explains more than half of the observed gravity decrease. Long-term variations on inter-annual and climate-relevant decadal scale will be investigated in the future using as supplement superconducting gravimetry (installed in 2019) and GNSS equipment (since 2018).

The late pleistocene cave bear fauna of the Torrener Bärenhöhle in the northern alps (Salzburg, Austria)

ABSTRACT The Torrener Bärenhöhle (Cave of Torren) is an alpine bear cave in the limestone Alps near Salzburg (Austria). The entrance of this cave is located in the riverbed of a periodically flowing stream that floods the caverns during snowmelt or after heavy rainfall. Due to these flooding events, the fossiliferous layers were repeatedly destroyed and the fossil remains – mainly cave bear bones – were distributed over the cave’s entrance hall. The fossil collections have taken place since 1924. During the last decades, numerous bones scattered over the cave floor have been collected again and again. In this study, we conduct metric and morphological analyses of this fossil material as well as DNA analyses in an attempt to clarify the taxonomic position of the cave bear remains. The chronological status of the bear remains has not yet been clarified, because the few samples that have been analysed so far were beyond the range of 14C dating, indicating they are older than 49,000 cal yr BP. The following taxa are represented in small numbers: Brown bear (Ursus arctos L.), wolf (Canis lupus L.), cave lion (Panthera spelaeus Goldfuss, 1810), European bison (Bison bonasus L.) and beaver (Castor fiber L.).

Dialektik der Erschließung: The German–Austrian Alps between Exploration and Exploitation

Focusing on the so-called Nördliche Kalkalpen or Northern Limestone Alps of Germany and Austria, I will discuss how human interaction with these mountains during the age of the Anthropocene shifts from scientific and athletic exploration to commercial and industrial exploitation. More specifically, I will examine travel narratives by the nineteenth-century mountaineers Friedrich Simony and Hermann von Barth, juxtaposing their respective experiences in diverse Alpine subranges with the environmental history of those regions. This juxtaposition harbors a deeper paradox, one that can be formulated as follows: Whereas Simony and Barth both rank as historically important Erschließer of the German and Austrian Alps, having explored their crags and glaciers in search of somatic adventure and geoscientific knowledge, these very sites of rock and ice were about to become so erschlossen by modernized tourism that one wonders where the precise boundaries between individual-based discovery and technology-driven development lie. In other words, during the nineteenth century a kind of Dialektik der Erschließung (a variation on Horkheimer and Adorno’s Dialektik der Aufklärung) manifests itself in the increasing anthropogenic alteration of the Alps.

Biotic and abiotic controls on carbon storage in aggregates in calcareous alpine and prealpine grassland soils

Alpine and prealpine grasslands provide various ecosystem services and are hotspots for the storage of soil organic C (SOC) in Central Europe. Yet, information about aggregate-related SOC storage and its controlling factors in alpine and prealpine grassland soils is limited. In this study, the SOC distribution according to the aggregate size classes large macroaggregates (> 2000 μm), small macroaggregates (250–2000 μm), microaggregates (63–250 μm), and silt-/clay-sized particles (< 63 μm) was studied in grassland soils along an elevation gradient in the Northern Limestone Alps of Germany. This was accompanied by an analysis of earthworm abundance and biomass according to different ecological niches. The SOC and N stocks increased with elevation and were associated with relatively high proportions of water-stable macroaggregates due to high contents of exchangeable Ca2+ and Mg2+. At lower elevations, earthworms appeared to act as catalyzers for a higher microaggregate formation. Thus, SOC stabilization by aggregate formation in the studied soils is a result of a joined interaction of organic matter and Ca2+ as binding agents for soil aggregates (higher elevations), and the earthworms that act as promoters of aggregate formation through the secretion of biogenic carbonates (low elevation). Our study highlights the importance of aggregate-related factors as potential indices to evaluate the SOC storage potential in other mountainous grassland soils.Graphical abstract

The Fate of Bryophyte Sporophytes-Phenology and Vectors of Buxbaumia viridis in the Kalkalpen National Park, Austria.

Knowledge about the epixylic moss Buxbaumia viridis has increased significantly due to the monitoring obligation under the Habitats Directive. However, there are still open questions about its dispersal, as the wind plays a limited role in forest ecosystems, and vectors have been suspected but not yet studied systematically for this species. Here, we present data on potential vectors of Buxbaumia viridis collected for the first time with the help of cameras, completed by monitoring the fate of sporophytes during their life cycle in the Limestone Alps National Park in Austria over a period of two years. Young, green sporophytes appeared mainly in autumn, with the highest number in October. Most of them survived winter and spring but did not exceed the age of 14 months. The sharpest decline in capsules occurred in summer when mature, and the lowest number of sporophytes appeared at the end of August. Most likely, mice seem to be responsible for this loss, as the photos from the wildlife cameras suggest, and should be considered both as predators and vectors. Birds should be considered as vectors, too. In summary, most of the reproductive biomass is sacrificed in favor of more effective dispersal, including over longer distances.

The Eastern Austrian Alps – Their Exceptional Demographic Status in the Alpine Region

Abstract In most Alpine regions with current negative migratory balance, new in-migration from urban areas has meanwhile become a relevant factor. However, this phenomenon is insignificant in Austria’s Eastern Alps. The aim of this thesis-led work is to empirically demonstrate the main reasons for this demographically exceptional position. The Styrian-Lower Austrian Limestone Alps, representing the peripheral core area of Austria's Eastern Alps, were chosen as the study area, as in this region the population decline and above all, migratory losses are particularly high. Through various qualitative and quantitative data acquisition techniques it was shown that new inmigration is still insignificant. The main reasons for this are the perseverance of large estates and the dominance of forestry.

Temporal and spatial trade-offs between resistance and performance traits in herbaceous plant species

Frost resistance (FR) is a highly adaptive trait and important for plant performance, survival and distribution. While overall seasonal changes in the frost resistance of herbaceous species are well documented, knowledge of the variability during the growth period is scarce. Responses could be expected due to differences in temperature yet investment in frost resistance might be at the expense of plant performance. To analyse temporal and spatial (i.e. same date but differing temperatures) variability, FR of leaves of six herbaceous species on five sampling dates were assessed along an elevational gradient in the northern limestone Alps. We used chlorophyll fluorescence techniques to calculate the lethal temperature of 90% of the population (LT90) thereof. To test the association with plant performance, we measured eco-morphological leaf traits (specific leaf area (SLA), leaf dry matter content, leaf phosphorous and magnesium content as well as stomatal pore area index (SPI)) in parallel. We found that FR as well as leaf traits exhibit a strong temporal variation whereas spatial variability was low. When analysing the relationship of FR to leaf traits we found that SLA as a proxy of growth rate was negatively associated with FR indicating a trade-off between growth and resistance, whereas SPI showed a positive relationship to FR. This finding gives further insight into the variability of traits and will help to improve predictions concerning plant performance and distribution under changing climate regimes.