Patrick Kuss

Genetic diversity, phenotypic variation and local adaptation in the alpine landscape: case studies with alpine plant species

Plant survival in alpine landscapes is constantly challenged by the harsh and often unpredictable environmental conditions. Steep environmental gradients and patchy distribution of habitats lead to small size and spatial isolation of populations and restrict gene flow. Agricultural land use has further increased the diversity of habitats below and above the treeline. We studied the consequences of the highly structured alpine landscape for evolutionary processes in four study plants: Epilobium fleischeri, Geum reptans, Campanula thyrsoides and Poa alpina. The main questions were: (1) How is genetic diversity distributed within and among populations and is it affected by altitude, population size or land use? (2) Do reproductive traits such as allocation to sexual or vegetative reproduction vary with altitude or land use? Furthermore, we studied if seed weight increases with altitude. Within-population genetic diversity of the four species was high and mostly not related to altitude and population size. Nevertheless, genetic differentiation among populations was pronounced and strongly increasing with distance. In Poa alpina genetic diversity was affected by land use. Results suggest considerable genetic drift among populations of alpine plants. Reproductive allocation was affected by altitude and land use in Poa alpina and by succession in Geum reptans. Seed weight was usually higher in alpine species than in related lowland species. We conclude that the evolutionary potential to respond to global change is mostly intact in alpine plants, even at high altitude. Phenotypic variability is shaped by adaptive as well as by random evolutionary processes; moreover plastic responses to growth conditions seem to be crucial for survival of plants in the alpine landscape.ZusammenfassungDas Überleben von Pflanzen in der alpinen Landchaft ist durch die harschen Umweltbedingungen eine ständige Herausforderung. Die alpine Landschaft zeichnet sich durch steile Umweltgradienten, fragmentierte Lebensräume und genetisch isolierte, oft kleine Populationen aus. Die landwirtschaftliche Nutzung hat diese kleinräumigen Strukturen noch verstärkt. Wir untersuchten die Auswirkungen der reich strukturierten alpinen Landschaft auf Evolutionsprozesse bei vier Arten: Epilobium fleischeri, Geum reptans, Campanula thyrsoides und Poa alpina. Hauptfragen waren: (1) Ist die genetische Diversität in Populationen abhängig von der Höhe über Meer, von der Populationsgrösse oder der Landnutzung und wie stark sind Populationen genetisch differenziert? (2) Verändern sich reproduktive Eigenschaften wie die Allokation in die sexuelle bzw. die klonale Reproduktion mit der Höhe und der Landnutzung? Weiter untersuchten wir die Auswirkung der Höhe über Meer auf das Samengewicht. Die genetische Diversität war bei allen vier Arten gross und ändert sich in der Regel nicht mit der Höhe oder der Populationsgrösse. Trotzdem fanden wir eine starke Populationsdifferenzierung, die mit der Distanz in allen Fällen deutlich zunahm. Bei Poa alpina wurde die genetische Diversität auch durch die Landnutzung beeinflusst. Genetische Drift war bei allen Arten beträchtlich. Die reproduktive Allokation wurde bei Geum reptans durch das Sukzessionsstadium und bei Poa alpina durch die Höhe und die Landnutzung beeinflusst. Das Samengewicht nimmt bei vielen alpinen Pflanzenarten im Vergleich mit Arten aus dem Tiefland zu. Wir schlussfolgern, dass die Anpassungsfähigkeit an eine sich ändernde Umwelt bei Alpenpflanzen auch in grosser Höhe intakt ist. Phänotypische Variabilität wird nicht nur durch adaptive, sondern auch durch Zufallsprozesse geformt. Phenotypische Plastizität scheint für das Überleben von Alpenpflanzen eine wichtige Rolle zu spielen.

Spatial and temporal patterns of greenness on the Yamal Peninsula, Russia: interactions of ecological and social factors affecting the Arctic normalized difference vegetation index

The causes of a greening trend detected in the Arctic using the normalized difference vegetation index (NDVI) are still poorly understood. Changes in NDVI are a result of multiple ecological and social factors that affect tundra net primary productivity. Here we use a 25 year time series of AVHRR-derived NDVI data (AVHRR: advanced very high resolution radiometer), climate analysis, a global geographic information database and ground-based studies to examine the spatial and temporal patterns of vegetation greenness on the Yamal Peninsula, Russia. We assess the effects of climate change, gas-field development, reindeer grazing and permafrost degradation. In contrast to the case for Arctic North America, there has not been a significant trend in summer temperature or NDVI, and much of the pattern of NDVI in this region is due to disturbances. There has been a 37% change in early-summer coastal sea-ice concentration, a 4% increase in summer land temperatures and a 7% change in the average time-integrated NDVI over the length of the satellite observations. Gas-field infrastructure is not currently extensive enough to affect regional NDVI patterns. The effect of reindeer is difficult to quantitatively assess because of the lack of control areas where reindeer are excluded. Many of the greenest landscapes on the Yamal are associated with landslides and drainage networks that have resulted from ongoing rapid permafrost degradation. A warming climate and enhanced winter snow are likely to exacerbate positive feedbacks between climate and permafrost thawing. We present a diagram that summarizes the social and ecological factors that influence Arctic NDVI. The NDVI should be viewed as a powerful monitoring tool that integrates the cumulative effect of a multitude of factors affecting Arctic land-cover change.

High mountain origin, phylogenetics, evolution, and niche conservatism of arctic lineages in the hemiparasitic genus Pedicularis (Orobanchaceae)

The origin of the arctic flora covering the northernmost treeless areas is still poorly understood. Arctic plants may have evolved in situ or immigrated from the adjacent ecosystems. Frequently arctic species have disjunctive distributions between the Arctic and high mountain systems of the temperate zone. This pattern may result from long distance dispersal or from glacial plant migrations and extinctions of intermediate populations. The hemiparasitic genus Pedicularis is represented in the Arctic by c. 28 taxa and ranks among the six most species-rich vascular plant genera of this region. In this study, we test the hypothesis that these lineages evolved from predecessors occurring in northern temperate mountain ranges, many of which are current centers of diversity for the genus. We generated a nuclear ribosomal and chloroplast DNA phylogeny including almost all of the arctic taxa and nearly half of the genus as a whole. The arctic taxa of Pedicularis evolved 12-14 times independently and are mostly nested in lineages that otherwise occur in the high mountains of Eurasia and North America. It appears that only three arctic lineages arose from the present-day center of diversity of the genus, in the Hengduan Mountains and Himalayas. Two lineages are probably of lowland origin. Arctic taxa of Pedicularis show considerable niche conservatism with respect to soil moisture and grow predominantly in moist to wet soils. The studied characteristics of ecology, morphology, and chromosome numbers of arctic Pedicularis show a heterogeneous pattern of evolution. The directions of morphological changes among the arctic lineages show opposing trends. Arctic taxa are chiefly diploid, the few tetraploid chromosome numbers of the genus were recorded only for arctic taxa. Five arctic Pedicularis are annuals or biennials, life forms otherwise rare in the Arctic. Other genera of the Orobanchaceae consist also of an elevated number of short-lived species, thus hemiparasitism may favor this life form in the Arctic.

Diversity and Frequency of Clonal Traits Along Natural and Land-Use Gradients in Grasslands of the Swiss Alps

The frequency of clonal plants in different vegetation types is known to be influenced by environmental and land-use factors. However, the underlying behavior of individual clonal traits or clonal trait diversity has received little attention. Here, we assess for species- and trait-diverse grasslands of the Swiss Alps the relative importance of temperature, soil moisture, land use and species richness on the diversity and frequency of individual compared with all clonal traits. We further analyzed how cover-weighted data alters the relationships found with commonly used presence-absence data. We combined species compositional, land-use and environmental data from 236 28-m2 grassland plots with clonal trait information for 527 species following the Clonal Growth Organ (CGO) classification. Test results are based on linear models, ANOVAs and ANCOVAs. The grassland sites were 84% dominated by clonal species. Drought-prone grasslands harbored the least clonal species. No increase in clonality was detected with decreasing temperature (= altitude). Mown or pastured grasslands had more clonal species than fallows. Certain sets of traits were correlated. Rhizomatous species especially reacted strongly to climatic and land-use gradients and had highest frequencies in cold, moist and disturbed sites. Clonal diversity was strongly dependent on species richness. Cover-weighted and presence-absence based estimates were largely similar. Overall, our data outlined that common clonal traits react differently to natural and land-use gradients as well as differently to the sum of clonal traits. Also, soil moisture was more decisive than temperature (= altitude) for the presence of clonal species. Lastly, the strong correlation between species-richness and clonal trait diversity needs to be accounted for when interpreting the functional role of clonal traits.

Differentiation in morphology and flowering phenology between two Campanula thyrsoides L. subspecies

Subspecies are usually characterised by sets of morphological discontinuities. By means of common garden experiments, we investigated genetic differentiation in morphological and phenological traits in two geographically disjunct subspecies of Campanula thyrsoides L., i.e. subsp. thyrsoides (=C.* thyrsoides) occurring in the European Alps and Jura Mountains, and subsp. carniolica (=C.* carniolica) occurring in the Southeastern Alps and the Dinaric Arc. Nine out of 16 investigated traits were significantly different between C.* thyrsoides and C.* carniolica. For C.* carniolica inflorescence length was 1.4×, and above-ground biomass 2.7× higher, while flower density was significantly lower. Campanula* carniolica also showed delayed flowering and flower development from bottom to top as compared to C.* thyrsoides which flowered from top to bottom. The inflorescence growth was indeterminate and flowering took several weeks in C.* carniolica, whereas C.* thyrsoides showed determinate flowering, rapidly opening all flowers within a few days. This differentiation in flowering phenology is likely to be adaptive. The submediterranean climate favours indeterminate flowering in C.* carniolica, allowing ongoing growth of the inflorescence throughout the long summer until environmental conditions worsen, whereas determinate and early flowering in C.* thyrsoides is favourable in the short growing season in the high Alps where seed production must be secured before temperature drops. Glacial survival in refugia with different climates (alpine vs. submediterranean) may have caused this regional differentiation.

Cumulative Effects of Rapid Land-Cover and Land-Use Changes on the Yamal Peninsula, Russia

The Yamal Peninsula in northwest Siberia is undergoing some of the most rapid land-cover and land-use changes in the Arctic due to a combination of gas development, reindeer herding, and climate change. Unusual geological condi- tions (nutrient-poor sands, massive ground ice and extensive landslides) exacerbate the impacts. These changes will likely increase markedly as transportation corridors are built to transport the gas to market. Understanding the nature, extent, causes and consequences (i.e., the cumulative effects) of the past and ongoing rapid changes on the Yamal is important for effective, long-term decision-making and planning. The cumulative effects to vegetation are the focus of this chapter because the plants are a critical component of the Yamal landscape that support the indigenous Nenets people and their reindeer and also protect the underlying ice-rich permafrost from melting. We are using a combination of ground-based studies (a transect of five loca- tions across the Yamal), remote-sensing studies, and analyses of Nenets land-use activities to develop vegetation-change models that can be used to help anticipate future states of the tundra and how those changes might affect traditional reindeer herding practices and the thermal state of the permafrost. This chapter provides an overview of the approach, some early results, and recommendations for expanding the concept of cumulative-effects analysis to include examining the simultaneous and interactive effects of multiple drivers of change.

Long-Distance Dispersal after the Last Glacial Maximum (LGM) Led to the Disjunctive Distribution of Pedicularis kansuensis (Orobanchaceae) between the Qinghai-Tibetan Plateau and Tianshan Region

Quaternary climate fluctuations have profoundly affected the current distribution patterns and genetic structures of many plant and animal species in the Qinghai-Tibetan Plateau (QTP) and adjacent mountain ranges, e.g. Tianshan (TSR), Altay, etc. In this greater area disjunct distributions are prominent but have nevertheless received little attention with respect to the historical processes involved. Here, we focus on Pedicularis kansuensis to test whether the current QTP and TSR disjunction is the result of a recent Holocene range expansion involving dispersal across arid land bridge(s) or a Pleistocene range fragmentation involving persistence in refugia. Two chloroplast DNA spacers were sequenced for 319 individuals from 34 populations covering the entire distribution range of this species in China. We found a total of 17 haplotypes of which all occurred in the QTP, and only five in the TSR. Overall genetic diversity was high (HT = 0.882, HS = 0.559) and higher in the QTP than in the TSR. Genetic differentiation among regions and populations was relatively low (GST = 0.366) and little evidence for a phylogeographic pattern emerged. The divergence times for the four main lineages could be dated to the early Pleistocene. Surprisingly, the two ubiquitous haplotypes diverged just before or around the Last Glacial Maximum (LGM) and were found in different phylogenetic lineages. The Species Distribution Model suggested a disappearance of P. kansuensis from the TSR during the LGM in contrast to a relatively constant potential distribution in the QTP. We conclude that P. kansuensis colonized the TSR after the LGM. The improbable long-distance dispersal by wind or water across arid land seed flow may well have had birds or men as vector.

Identity and distribution of weedy Pedicularis kansuensis Maxim. (Orobanchaceae) in Tianshan Mountains of Xinjiang: morphological, anatomical and molecular evidence

Weedy plants affect the biodiversity and ecosystem function as well as the crop and fodder plant production. However, adequate management requires detailed knowledge of the taxonomic identity of these plants. Here, we focused on a hemiparasitic Pedicularis species (Orobanchaceae), which occurs at high densities and results in significant biomass reductions in forage grasses in Bayanbulak Grassland of Xinjiang. The identity of this target species is not clear, with conflicting reports in publications and in herbarium collections. Hence, clear and management-relevant information on demography and reproductive ecology is difficult to be obtained from the literature. Therefore, we analyzed field and archival materials collected from Xinjiang in order to clarify the identity and distribution of the target species. Morphological analyses suggested that the populations at Bayanbulak Grassland should be Pedicularis kansuensis Maxim. rather than P. verticillata L. which has been accepted in the available literature. Phylogenetic analysis with a combination of three barcodes (matK, rbcL and trnH-psbA) uniting a clade of P. kansuensis and individuals from Bayanbulak Grassland populations with 100% bootstrap support, confirmed the target species to be P. kansuensis. Anatomical investigations and field observations showed that the target species is an annual or biennial herb, which also fits with the life cycle as P. kansuensis. Based on archive material and field observations, we verified that the distribution of P. kansuensis is mainly concentrated in the Tianshan Mountains of Xinjiang.

Vegetation on mesic loamy and sandy soils along a 1700-km maritime Eurasia Arctic Transect

Abstract Questions How do plant communities on zonal loamy vs. sandy soils vary across the full maritime Arctic bioclimate gradient? How are plant communities of these areas related to existing vegetation units of the European Vegetation Classification? What are the main environmental factors controlling transitions of vegetation along the bioclimate gradient? Location 1700‐km Eurasia Arctic Transect (EAT), Yamal Peninsula and Franz Josef Land (FJL), Russia. Methods The Braun‐Blanquet approach was used to sample mesic loamy and sandy plots on 14 total study sites at six locations, one in each of the five Arctic bioclimate subzones and the forest–tundra transition. Trends in soil factors, cover of plant growth forms (PGFs) and species diversity were examined along the summer warmth index (SWI) gradient and on loamy and sandy soils. Classification and ordination were used to group the plots and to test relationships between vegetation and environmental factors. Results Clear, mostly non‐linear, trends occurred for soil factors, vegetation structure and species diversity along the climate gradient. Cluster analysis revealed seven groups with clear relationships to subzone and soil texture. Clusters at the ends of the bioclimate gradient (forest–tundra and polar desert) had many highly diagnostic taxa, whereas clusters from the Yamal Peninsula had only a few. Axis 1 of a DCA was strongly correlated with latitude and summer warmth; Axis 2 was strongly correlated with soil moisture, percentage sand and landscape age. Conclusions Summer temperature and soil texture have clear effects on tundra canopy structure and species composition, with consequences for ecosystem properties. Each layer of the plant canopy has a distinct region of peak abundance along the bioclimate gradient. The major vegetation types are weakly aligned with described classes of the European Vegetation Checklist, indicating a continuous floristic gradient rather than distinct subzone regions. The study provides ground‐based vegetation data for satellite‐based interpretations of the western maritime Eurasian Arctic, and the first vegetation data from Hayes Island, Franz Josef Land, which is strongly separated geographically and floristically from the rest of the gradient and most susceptible to on‐going climate change.

Lectotypification of Linnaean names in Pedicularis (Orobanchaceae)

Ten Linnaean names of Pedicularis (Orobanchaceae) are lectotypified: P. comosa, P. flammea, P. hirsuta, P. incarnata, P. lapponica, P. resupinata, P. rostrata, P. tristis, P. tuberosa and P. verticillata. Lectotypes are designated from the Linnaean herbarium at LINN. The type choices made here maintain the present usage of the corresponding Linnaean names for ten Pedicularis species.